MP Van de Graaff Human Anatomy

Van De Graaff: Human
Anatomy, Sixth Edition
Front Matter
Preface
The McGrawHill
Companies, 2001
Preface
H
uman Anatomy was written to serve as a foundation and

resource for students pursuing health-related careers in
fields such as medicine, dentistry, nursing, physician assistant,
podiatry, optometry, chiropractic, medical technology, physical
therapy, athletic training, massage therapy, and other healthrelated professions. Created to accompany the one-semester
human anatomy course, this text presents a basic introduction to
human anatomy for students enrolled in medical, allied-health,
and physical education programs, or for those majoring in biological science. The focus of Human Anatomy is to provide applicable knowledge of the structure of the human body and
foundation information for understanding physiology, cell biology, developmental biology, histology, and genetics. Practical information is presented in this text that will enable students to
apply pertinent facts to the real-world situations they might encounter in their chosen profession.
Many changes have been made in the sixth edition of
Human Anatomy to provide students with a high-quality text for
their course of study. Because human anatomy is such a visual science, many refinements and additions have been made in a continuing effort to provide an effective art program. Many new
illustrations, radiographs, and photographs (including images of
cadaver dissections) make this text even more useful. Strengthening clinical aspects of the text has been another major focus in the
sixth edition. Additional Clinical Practicums have been added at
the ends of the chapters throughout the text. These case studies
and their accompanying images test student knowledge and
demonstrate the application of anatomical information in a clinical setting. A final task in creating the sixth edition of Human
Anatomy has been to revise the content for currentness and accuracy. In keeping with the pace of research, updated information is
presented on the history of current human genome research, the
structure of DNA and RNA, protein synthesis, utilization of stem
cells from red bone marrow and fetal tissue, and the classification
of hair. The comprehensive nature of the sixth edition of this text
and its current clinical information enable it to be used as a valuable reference resource regarding the structure, function, development, senescence, and possible dysfunctions of the human body.
OBJECTIVES
In preparing and updating a text and its ancillaries (website, laboratory manual, instructors manual, test bank, and so forth), it is essential to consider both the needs of the student and the needs of
the instructor. A well-written and inviting text is at the heart of an
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effective educational package. With this in mind, the following objectives were formulated for the sixth edition of Human Anatomy:
To provide a text that is inviting and attractivea text that
is readable and informative with accurate, up-to-date information of practical concern. Human Anatomy aims to entice
readers to study the material and thereby enhance their appreciation of life through a better understanding of the
structure, function, and magnificence of their own bodies.
To provide a conceptual framework of learning through
the use of concise concept statements, learning objectives,
and chapter review questions.
To express the beauty of the body through spectacular art
that is anatomically accurate. Anatomy is a visual science
where exactness is essential. The numerous high-quality illustrations prepared expressly for this edition augment the
acclaimed art program of the previous editions.
To stimulate student interest in anatomy and related subjects through a series of thematic commentaries, highlighted by topic icons.
To provide a systematic, balanced presentation of anatomical concepts at the developmental, cellular, histological,
clinical, and gross anatomy levels.
To build students technical vocabularies to the point
where they feel comfortable with basic medical terminology, enabling them to become conversant with health-care
providers and understand current medical literature.
To encourage proper care of the body in order to enjoy a
healthier, more productive life, and to provide a foundation of knowledge students can share to help enrich the
lives of others.
To acquaint students with the history of anatomy, from its
primitive beginnings to recent advances in the field.
Only with the realization of how long it took to build up
knowledge that is now taken for grantedand with what
difficultycan students appreciate the science of anatomy
in its proper proportion.
TEXT ORGANIZATION
The 22 chapters in this text are grouped into seven units that are
identified by colored tabs on the outside page margins.
Unit 1: Historical Perspective In this unit, the stage is set
for studying human anatomy by providing a historical perspective

Front Matter
Preface
on how this science has developed over the centuries. Anatomy is

an exciting and dynamic science that remains vital as it continues
to broaden its scope. It is hoped that this unit will make the
reader feel a part of the heritage of human anatomy.
Unit 2: Terminology, Organization, and the Human Organism In this unit, the anatomical characteristics that define humans as a distinct species are described. The various levels of
organization of the human body are also described, and the basic
terminology necessary for understanding the structure and functioning of the body is introduced.
Unit 3: Microscopic Structure of the Body The microscopic
aspect of body organization is considered at the cellular and histological levels in this unit. Cellular chemistry is emphasized as an
integral aspect of learning about how the body functions.
Unit 4: Support and Movement Support, protection, and
movement of the human body are the themes of this unit. The
integumentary system provides the body with external support
and protection, and the skeletal system provides internal support
and protection for certain organs of the body. Movement is possible at the joints of the skeleton as the associated skeletal muscles are contracted. Surface anatomy and regional anatomy are
given detailed coverage in chapter 10 of this unit. Atlas-quality
photographs of dissections of human cadavers are included in
this chapter.
Unit 5: Integration and Coordination This unit includes
chapters on the nervous system, endocrine system, and sensory
organs. The concepts identified and discussed in these chapters
are concerned with the integration and coordination of body
functions and the perception of environmental stimuli.
Unit 6: Maintenance of the Body In this unit, the structure and function of the circulatory, respiratory, digestive, and
urinary systems are discussed as they contribute in their individual ways to the overall functioning and general welfare of the organism. All of these systems work together in maintaining a
stable internal environment in which the cells of the body can
thrive on a day-to-day basis.
Unit 7: Reproduction and Development The male and female reproductive systems are described in this unit, and the
continuance of the human species through sexual reproduction is
discussed. Unit 7 provides an overview of the entire sequence of
human life, including prenatal development and postnatal
growth, development, and aging. Basic concepts of genetics and
inheritance are also explained.
The McGrawHill
Companies, 2001
LEARNING AIDS
Each of the 22 chapters of this text incorporates numerous pedagogical devices that organize and underscore the practicality of
the material, clarify important concepts, help assess student
learning, and stimulate students natural curiosity about the
human body. In short, these aids make the study of human
anatomy more effective and enjoyable.
Chapter Introductions
The beginning page of each chapter contains an outline of the
chapter contents and a Clinical Case Study pertaining to the
subject matter of the chapter. Each case study is elucidated with
a related photograph. These hypothetical situations underscore
the clinical relevance of anatomical knowledge and entice students to watch for information contained within the chapter
that may be needed to answer the case study questions. The solution to the case study is presented at the end of the chapter, following the last major section.
Understanding Anatomical Terminology

Each technical term is set off in boldface or italic type, and is
often followed by a phonetic pronunciation in parentheses, at
the point where it first appears and is defined in the narrative.
The roots of each term can be identified by referring to the glossary of prefixes and suffixes found on the inside of the front
cover. In addition, the derivations of many terms are provided in
footnotes at the bottom of the page on which the term is introduced. If students know how a term was derived, and if they can
pronounce the term correctly, the term becomes more meaningful and is easier to remember.
Chapter Sections
Each chapter is divided into several major sections, each of
which is prefaced by a concept statement and a list of learning
objectives. A concept statement is a succinct expression of the
main idea, or organizing principle, of the information contained
in a chapter section. The learning objectives indicate the level of
competency needed to understand the concept thoroughly and
be able to apply it in practical situations. The narrative that follows discusses the concept in detail, with reference to the objectives. Knowledge Check questions at the end of each chapter
section test student understanding of the concept and mastery of
the learning objectives.
xi

Front Matter
Preface
Commentaries and Clinical Information

Set off from the text narrative are short paragraphs highlighted
by accompanying topic icons. This interesting information is relevant to the discussion that precedes it, but more important, it
demonstrates how basic scientific knowledge is applied. The five
icons represent the following topic categories:
Clinical information is indicated by a stethoscope. The information
contained in these commentaries provides examples of the applied
medical nature of the information featured in the topic discussion.
Aging information is indicated by an hourglass. The information
contained in these commentaries is relevant to normal aging and
indicates how senescence (aging) of body organs impacts body
function.
Developmental information of practical importance is indicated by a
human embryo. Knowledge of pertinent developmental anatomy
contributes to understanding how congenital problems develop and
impact body structure and function.
Homeostasis information is indicated by a gear mechanism. The
information called out by this icon is relevant to the body processes that
maintain a state of dynamic equilibrium. These commentaries point out
that a disruption of homeostasis frequently accompanies most diseases.
Academic interest commentaries discuss topics that are relevant to
human anatomy that are quite simply of factual interest.
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Companies, 2001
challenged to evaluate the clinical findings, explain the origin of

symptoms, diagnose the patient, recommend treatment, etc.
Each body system chapter contains one or two Clinical
Practicums, placed before the chapter summary. Detailed answers
to the Clinical Practicum questions are provided in Appendix B.
Chapter Summaries
A summary, in outline form, at the end of each chapter reinforces
the learning experience. These comprehensive summaries serve as
a valuable tool in helping students prepare for examinations.
Review Activities
Following each chapter summary, sets of objective, essay, and
critical thinking questions give students the opportunity to measure the depth of their understanding and learning. The critical
thinking questions have been updated and expanded in the sixth
edition to further challenge students to use the chapter information in novel ways toward the solution of practical problems.
The correct responses to the objective questions are provided in
Appendix A. Each answer is explained, so students can effectively use the review activities to broaden their understanding of
the subject matter.
Illustrations and Tables

In addition to the in-text commentaries, selected developmental disorders, aging, clinical procedures, and diseases or dysfunctions of specific organ systems are described in Clinical
Considerations sections that appear at the end of most chapters.
Photographs of pathological conditions accompany many of
these discussions.
Developmental Expositions
In each body system chapter, a discussion of prenatal development follows the presentation on gross anatomy. Each of these
discussions includes exhibits and explanations of the morphogenic events involved in the development of a body system.
Placement near the related text discussion ensures that the
anatomical terminology needed to understand the embryonic
structures has been introduced.
Clinical Practicums
These focused clinical scenarios present a patient history and
supporting diagnostic imagesuch as a radiograph, ultrasound,
or photographfollowed by a series of questions. Students are
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Because anatomy is a descriptive science, great care has been

taken to continuously enhance the photographs and illustrations
in Human Anatomy. A hallmark feature of the previous editions
of this text has been the quality art program. In keeping with the
objective of forever improving and refining the art program, over
150 full-color illustrations were substantially revised or rendered
entirely new for the sixth edition. Each illustration has been
checked and rechecked for conceptual clarity and precision of the
artwork, labels, and captions. Color-coding is used in certain art
sequences as a technique to aid learning. For example, the bones
of the skull in chapter 6 are color-coded so that each bone can be
readily identified in the many renderings included in the chapter.
These illustrations represent a collaborative effort between author
and illustrator, often involving dissection of cadavers to ensure
accuracy. Illustrations are combined with photographs whenever
possible to enhance visualization of anatomical structures. Light
and scanning electron micrographs are used throughout the text
to present a true picture of anatomy from the cellular and histological levels. Surface anatomy and cadaver dissection images
help students understand the juxtaposition of anatomical structures and help convey the intangible anatomical characteristics
that can be fully appreciated only when seen in a human speci-

Front Matter
Preface
men. Many of the cadaver dissection photographs have been

modified or replaced with new, high-quality images shot expressly
for the sixth edition. All of the figures are integrated with the text
narrative to maximize student learning.
Numerous tables throughout the text summarize information
and clarify complex data. Many tables have been enhanced with
the addition of illustrations to communicate information in the
most effective manner. Like the figures, all of the tables are referenced in the text narrative and placed as close to the reference as
possible to spare students the trouble of flipping through pages.
Appendixes, Glossary, and Index

Appendixes A and B provide answers and explanations for the
objective questions at the end of each chapter and for the questions that accompany the Clinical Practicum boxes. The glossary
provides definitions for the important technical terms used in
the text. Phonetic pronunciations are included for most of the
terms, and an easy-to-use pronunciation guide appears at the beginning of the glossary. Synonyms, including eponymous terms,
are indicated, and for some terms antonyms are given as well.
TEACHING AND LEARNING

SUPPLEMENTS
There is much more to Human Anatomy than this book. Numerous study and teaching aids round out the complete package.
Students can order supplemental study materials by contacting
the McGraw-Hill Customer Service Department at 800-3383987. Instructors can obtain teaching aids by calling the Customer Service Department or by contacting your McGraw-Hill
sales representative.
Online Learning Center

The Online Learning Center (OLC) at www.mhhe.com/vdg offers an extensive array of learning and teaching tools. This website includes chapter-specific quizzes and web links, clinical
applications, interactive activities, art labeling exercises, case
studies, and more. Teaching resources at the instructor site include image and animations libraries, PowerPoint lecture presentations, technology resources, and the online Instructors Manual
for Human Anatomy. In addition, the OLC provides online access to the following premium interactive products:
Essential Study Partner for Anatomy and Physiology is a
complete, interactive student study tool packed with hundreds of animations and more than 800 learning activities. Interactive diagrams and quizzes make learning core concepts
stimulating and fun.
The McGrawHill
Companies, 2001
adam Online Anatomy is a comprehensive database of detailed anatomical images that allows users to point, click, and
identify more than 20,000 anatomical structures within fully
dissectible male and female bodies in anterior, lateral, medial,
and posterior views. Exhaustively reviewed by panels of leading anatomists, adam Online Anatomy is recognized as the
standard anatomical database in computer-based medical education worldwide.
BioCourse.com delivers rich, interactive content to fortify
the learning and teaching experience in the life sciences. In
addition to over 10,000 animations, images, case studies, and
video presentations, discussion boards and laboratory exercises
foster collaboration and infinite learning and teaching opportunities. Biocourse.com contains these specific areas:
The Faculty Club gives new and experienced instructors access to a variety of resources to help increase their effectiveness in lecture, discover groups of instructors with similar
interests, and find information on teaching techniques and
pedagogy. A comprehensive search feature allows instructors
to search for information using a variety of criteria.
The Student Center allows students the opportunity to
search BioCourse for information specific to the course area
they are studying, or by using specific topics or keywords.
Information is also available for many aspects of student
life, including tips for studying and test taking, surviving
the first year of college, and job and internship searches.
BioLabs helps laboratory instructors, who often face a special set of challenges. BioLabs addresses those challenges
by providing laboratory instructors and coordinators with a
source for basic information on suppliers, best practices,
professional organizations, and lab exchanges.
Briefing Room is where to go for current news in the life
sciences. News feeds from The New York Times, links to
prominent journals, commentaries from popular McGrawHill authors, and XanEdu journal search service are just a
few of the resources you will find here.
The Quad utilizes a powerful indexing and searching tool
to provide the user with a guided review of specific course
content. Information is available from a variety of McGraw-Hill sources, including textbook material, Essential
Study Partner modules, Online Learning Centers, and images from Visual Resource Libraries.
R&D Center is the opportunity to see what new textbooks, animations, and simulations McGraw-Hill is working on and to send McGraw-Hill your feedback. You can
also learn about other opportunities to review as well as
submit ideas for new projects.
xiii

Front Matter
Preface
Laboratory Manual to accompany

Human Anatomy, Sixth Edition
Kent Van De Graaff has authored a comprehensive laboratory
manual specifically designed to accompany Human Anatomy, sixth
edition. This laboratory manual emphasizes learning anatomical
structures through visual observation, palpation, and knowledge of
the functional relationship of one body system to another. It focuses primarily on the human organism, but also contains cat dissections and selected organ dissections. Closely integrated with the
Human Anatomy text, the companion lab manual utilizes a wellrounded pedagogical system that helps students organize the background information and materials needed to complete each lab
exercise. Coloring and labeling activities placed throughout the
chapters reinforce recognition of anatomical structures, and laboratory reports at the end of each chapter encourage students to
synthesize concepts covered in both lab and lecture.
Instructors Manual
for the Laboratory Manual
This online manual is housed within the instructor Online
Learning Center. It provides answers to the lab report questions,
as well as overviews on how to present each laboratory exercise,
materials lists, and additional topics for discussion.
The McGrawHill
Companies, 2001
strategies, discussion and demonstration ideas for lectures, and suggestions for laboratory exercises. This manual also includes a listing of transparencies and multimedia resources that correlate with
each text chapter and provides answers to the Knowledge Check,
Essay, and Critical Thinking questions that appear in the text.
Test Item File

The Test Item File contains fill-in-the-blank, multiple choice,
and true/false questions specifically designed to complement
each chapter of the text. Instructors using WebCT, Blackboard,
or PageOut can access the Test Item File online.
MicroTest
MicroTest is a computerized test generator that is free upon request to qualified adopters. The test generator contains the complete Test Item File on CD-ROM. MicroTest requires no
programming experience and is designed to work on both Windows and Macintosh platforms.
PageOut is McGraw-Hills exclusive tool for creating your own

website for your anatomy course. It requires no knowledge of
coding. Simply type your course information into the templates
provided. PageOut is hosted by McGraw-Hill.
Transparencies
This set of transparency acetates includes 200 full-color illustrations from the text that have been chosen for their value in reinforcing lecture presentations.
Visual Resource Library

Accessed through the instructor site at the Online Learning
Center and also available on CD-ROM, the Visual Resource Library contains labeled and unlabeled versions of the key illustrations and photos from the book, as well as all tables. You can
quickly preview images and incorporate them into PowerPoint or
other presentation programs to create your own multimedia presentations. You can also remove and replace labels to suit your
own preferences in terminology or level of detail.
Instructors Manual
Accessed via the Online Learning Center, the instructors manual
by Jeffrey S. Prince, M.D. and Karianne N. Prince provides instructional support in the use of the textbook. It includes teaching
xiv
In addition to the materials specifically designed to accompany Human Anatomy, McGraw-Hill offers the following supplemental resources to enrich the study and instruction of anatomy
and physiology.
Regional Human Anatomy: A Laboratory Workbook For
Use With Models and Prosections by Frederick E. Grine,
State University of New YorkStony Brook. Organized with a
regional approach to human anatomy, this workbook utilizes
coloring and labeling activities to simplify the learning of
anatomy. Brief text descriptions of key anatomical structures
are grouped with detailed illustrations that can be colored and
labeled to reinforce the material presented. Critical thinking
questions encourage students to think about how anatomical
structures work together, and boxed clinical insights highlight
facts of interest to students pursuing health-related professions.
Anatomy and Physiology Laboratory Manual-Fetal Pig by
Terry R. Martin, Kishwaukee College. Provides excellent fullcolor photos of the dissected fetal pig with corresponding labeled
art. It includes World Wide Web activities for many chapters.

Front Matter
Preface
Web-Based Cat Dissection Review for Human Anatomy

and Physiology by John Waters, Pennsylvania State University. This online multimedia program contains vivid, highquality labeled cat dissection photographs. The program helps
students easily identify and review the corresponding structures and functions between the cat and the human body.
Dynamic Human, Version 2.0. A set of two interactive CDROMs that cover each body system and demonstrate clinical
concepts, histology, and physiology with animated threedimensional and other images.
Interactive Histology CD-ROM by Bruce Wingerd and Paul
Paolini, San Diego State University. This CD contains 135
full-color, high-resolution light micrograph images and 35 scanning electron micrograph images of selected tissue sections typically studied in anatomy and physiology. Each image has labels
that can be clicked on or off, has full explanatory legends, offers
views at two magnifications, and has links to study questions.
The CD also has a glossary with pronunciation guides.
Life Science Animation VRL 2.0 contains over 200 animations of major biological concepts and processes, such as the
sliding filament mechanism, active transport, genetic transcription and translation, and other topics that may be difficult for students to visualize.
Life Science Animations 3D Videotape contains 42 key biological processes that are narrated and animated in vibrant
full color with dynamic three-dimensional graphics.
Life Science Animations (LSA) videotape series contains 53
animations on five VHS videocassettes: Chemistry, the Cell,
and Energetics; Cell Division, Heredity, Genetics, Reproduction, and Development; Animal Biology No. 1; Animal Biology No. 2; and Plant Biology, Evolution, and Ecology.
Another available videotape is Physiological Concepts of Life
Science.
Atlas to Human Anatomy by Dennis Strete, McLennan
Community College, and Christopher H. Creek. This atlas
takes a systems approach with references to regional anatomy,
thereby making it a great complement to your regular course
structure, as well as to your laboratory.
Atlas of the Skeletal Muscles, third edition, by Robert and
Judith Stone, Suffolk County Community College. This atlas
is a guide to the structure and function of human skeletal
muscles. The illustrations help students locate muscles and
understand their actions.
Laboratory Atlas of Anatomy and Physiology, third edition,
by Eder et al. This full-color atlas contains histology, human
skeletal anatomy, human muscular anatomy, dissections, and
reference tables.
The McGrawHill
Companies, 2001
Coloring Guide to Anatomy and Physiology by Robert and

Judith Stone, Suffolk County Community College. This guide
emphasizes learning through the process of color association.
The Coloring Guide provides a thorough review of anatomical
and physiological concepts.
ACKNOWLEDGMENTS
Preparing a new edition of a text is a formidable task that involves a number of colleagues, students, and publishing professionals. And in the case of this text, even family members were
involved. My sincere gratitude is extended to faculty and students who have used previous editions of this text and have
taken the time to suggest ways to improve it. They are indeed
thinking of others who will be using the text in the future, and at
the same time, ensuring a future for the text.
I am especially appreciative of Samuel I. Zeveloff and
Ronald Galli, colleagues at Weber State University, who were
especially supportive of my efforts in preparing this edition. A
number of professors who taught from the previous edition
shared suggestions that have been incorporated into this one.
Furthermore, some students who used the text offered suggestions for improvement. Melissa J. Bentley, Eric F. Stakebake,
and Amber Bennett were particularly helpful in the preparation
of this edition. Feedback from conscientious students is especially useful and appreciated.
Several physicians contributed clinical input to this edition. I especially appreciate the assistance of Dr. Jeffrey S. Prince
and Karianne N. Prince for their contributions of additional
Clinical Practicums and the accompanying radiographic images.
Their involvement is especially rewarding to me, in that they are
former students. A fathers request to three of his sons resulted in
additional clinical input. A heartfelt thanks is extended to Drs.
Kyle M. Van De Graaff, Eric J. Van De Graaff, and Ryan L. Van
De Graaff for their generous suggestions and genuine interest in
what their dad does. My good friend and collaborator John L.
Crawley has continued to be supportive of my writing endeavors.
The visual appeal and accuracy provided by quality photographs and illustrations are essential in an anatomy text. I have
enjoyed my years of professional interaction with Christopher
Creek, the talented artist who rendered many of the illustrations
in the previous editions and a number of new ones for this edition. His anatomical art is engaging and realistic. Dr. Gary M.
Watts, Department of Radiology at the Utah Valley Regional
Medical Center, provided many of the radiographic images used
in the previous editions of this text and some new ones for this
edition. Thanks are also extended to Don Kincaid and Rebecca
Gray of Ohio State University, who dissected and photographed
the new cadaver images for this edition.
xv

Front Matter
Preface
Sincere gratitude is extended to the editors at McGrawHill for their talent, dedication, and encouragement of my efforts. Sponsoring Editors Marty Lange and Kristine Tibbetts and
Developmental Editor Kristine Queck were superb to work with.
I enjoyed my association with Jane Matthews, Project Manager,
and John Leland, Photo Research Coordinator. Both of these
people spent countless hours attending the myriad details that a
technical text such as this involves.
Marion Alexander
University of Manitoba
Frank Baker
Golden West College
Leann Blem
Virginia Commonwealth University
Carolyn W. Burroughs
Bossier Parish Community College
Russ Cagle
Willamette University
Paul V. Cupp, Jr.
Eastern Kentucky University
Brian Curry
Grand Valley State University
Shirley Dillaman
Penn StateShenango
Cathryn R. Dooly
Ball State University
Ruth E. Ebeling
Biola University
Charles A. Ferguson
University of Colorado at Denver
xvi
The McGrawHill
Companies, 2001
McGraw-Hill dutifully assembled a panel of competent

anatomists to review the previous text and the new manuscript
as it was being developed for the sixth edition. These professionals aided my work immeasurably, and I am especially grateful for
their frank criticism, comments, and reassurance.
David K. Ferris
University of South CarolinaSpartanburg
Allan Forsman
East Tennessee State University
Carl D. Frailey
Johnson County Community College
Glenn A. Gorelick
Citrus College
Douglas J. Gould
University of Kentucky Chandler Medical
Center
Melanie Gouzoules
University of North CarolinaGreensboro
Phyllis C. Hirsch
East Los Angeles College
Bert H. Jacobson
Oklahoma State University
Glenn E. Kietzmann
Wayne State College
Dennis Landin
Louisiana State University
Bryan G. Miller
Eastern Illinois University
Virginia L. Naples
Northern Illinois University
Daniel R. Olson
Northern Illinois University
Scott Pedersen
South Dakota State University
Russell L. Peterson
Indiana University of Pennsylvania
Larry A. Reichard
Maple Woods Community College
Alexander Sandra
University of Iowa
David J. Saxon
Morehead State University
Stephen P. Schiffer
Georgetown University Medical Center
Leeann Sticker
Northwestern State University of Louisiana
R. Brent Thomas
University of South CarolinaSpartanburg
Judy A. Williams
Southeastern Oklahoma State University

Front Matter
A Visual Guide
The McGrawHill
Companies, 2001
Visual Guide
Chapter Outline
A page-referenced preview of major topics is included on the

opening page of each chapter, allowing you to see at a glance what
the upcoming chapter covers.
Body Organization and

Anatomical Nomenclature
Classification and Characteristics of

Humans 23
Body Organization 28
Anatomical Nomenclature 30
Planes of Reference and Descriptive
Terminology 33
Body Regions 35
Body Cavities and Membranes 41
Clinical Case Study Answer 45
Chapter Summary 46
Review Activities 46
Clinical Case Study
FIGURE: Radiographic anatomy is

important in assessing trauma to bones and
visceral organs.
A young woman was hit by a car while crossing a street. Upon arrival at the scene, paramedics
found the patient to be a bit dazed but reasonably lucid, complaining of pain in her abdomen
and the left side of her chest. Otherwise, her vital signs were within normal limits. Initial evaluation in the emergency room revealed a very tender abdomen and left chest. The chest radiograph demonstrated a collapsed left lung resulting from air in the pleural space
(pneumothorax). The emergency room physician inserted a drainage tube into the left chest
(into the pleural space) to treat the pneumothorax. Attention was then turned to the abdomen. Because of the finding of tenderness, a peritoneal lavage was performed. This procedure
involves penetrating the abdominal wall and inserting a tube into the peritoneal cavity. Clear
fluid such as sterile water or normal saline is then instilled into the abdomen and siphoned out
again. The fluid used in this procedure is called lavage fluid. A return of lavage fluid containing
blood, fecal matter, or bile indicates injury to an abdominal organ that requires surgery. The return of lavage fluid from this patient was clear. However, the nurse stated that lavage fluid was
draining out of the chest tube.
From what you know about how the various body cavities are organized, do you suppose
this phenomenon could be explained based on normal anatomy? What might have caused it to
occur in our patient? Does the absence of bile, blood, etc., in the peritoneal lavage fluid guarantee that no organ has been ruptured? If it does not, explain why in terms of the relationship of
the various organs to the membranes within the abdomen.
Clinical Case Study

A hypothetical medical situation sets the stage for the chapter by
underscoring the clinical relevance of the chapter content. As you
read the chapter, watch for the background information needed to
solve the case study, then check your answer against the solution
given at the end of the chapter.
DEFINITION AND
CLASSIFICATION OF TISSUES
Shaft of a
hair within
a hair follicle
Histology is the specialty of anatomy that involves study of the microscopic structure of tissues. Tissues are assigned to four basic
categories on the basis of their cellular composition and histological appearance.
Objective 1
Define tissue and discuss the importance of
histology.
Concept Statement
Describe the functional relationship between

cells and tissues.
Objective 3
List the four principal tissue types and briefly

describe the functions of each type.
A carefully worded expression of the main idea, or organizing

principle, of the information contained in a chapter section gives
you a quick overview of the material that will follow.
Learning Objectives
Each chapter section begins with a set of learning objectives that
indicate the level of competency you should attain in order to
thoroughly understand the concept and apply it in practical
situations.
Vocabulary Aids
New terms appear in boldface print as they are introduced and

immediately defined in context. Definitions and phonetic
pronunciations for boldfaced terms are gathered in the glossary at
the end of the book.
The Greek or Latin derivations of many terms are provided in
footnotes at the bottom of the page on which the term first appears.
Objective 2
Although cells are the structural and functional units of the

body, the cells of a complex multicellular organism are so specialized that they do not function independently. Tissues are aggregations of similar cells and cell products that perform specific
functions. The various types of tissues are established during
early embryonic development. As the embryo grows, organs form
from specific arrangements of tissues. Many adult organs, including the heart, brain and muscles, contain the original cells and
tissues that were formed prenatally, although some functional
changes occur in the tissues as they are acted upon by hormones
or as their effectiveness diminishes with age.
The study of tissues is referred to as histology. It provides a
foundation for understanding the microscopic structure and
functions of the organs discussed in the chapters that follow.
Many diseases profoundly alter the tissues within an affected
organ; therefore, by knowing the normal tissue structure, a physician can recognize the abnormal. In medical schools a course in
histology is usually followed by a course in pathology, the study of
abnormal tissues in diseased organs.
Although histologists employ many different techniques
for preparing, staining, and sectioning tissues, only two basic
kinds of microscopes are used to view the prepared tissues. The
light microscope is used to observe overall tissue structure
(fig. 4.1), and the electron microscope to observe the fine details of
tissue and cellular structure. Most of the histological photomicrographs in this text are at the light microscopic level. However, where fine structural detail is needed to understand a
particular function, electron micrographs are used.
Many tissue cells are surrounded and bound together by a
nonliving intercellular matrix (matriks) that the cells secrete.
Matrix varies in composition from one tissue to another and may
take the form of a liquid, semisolid, or solid. Blood, for example,
histology: Gk. histos, web (tissue); logos, study

pathology: Gk. pathos, suffering, disease; logos, study
matrix: L. matris, mother
(a)
Shaft of hair
emerging from
the exposed
surface of
the skin
(b)
FIGURE 4.1 The appearance of skin (a) magnified 25 times, as

seen through a compound light microscope, and (b) magnified 280
times, as seen through a scanning electron microscope (SEM).
has a liquid matrix, permitting this tissue to flow through vessels.

By contrast, bone cells are separated by a solid matrix, permitting
this tissue to support the body.
The tissues of the body are assigned to four principal types
on the basis of structure and function: (1) epithelial (ep-thele-al)
tissue covers body surfaces, lines body cavities and ducts, and
forms glands; (2) connective tissue binds, supports, and protects
body parts; (3) muscle tissue contracts to produce movement; and
(4) nervous tissue initiates and transmits nerve impulses from one
body part to another.
Knowledge Check
1. Define tissue and explain why histology is important to the
study of anatomy, physiology, and medicine.
2. Cells are the functional units of the body. Explain how the
matrix permits specific kinds of cells to be even more effective and functional as tissues.
3. What are the four principal kinds of body tissues? What are
the basic functions of each type?
xvii
Front Matter
A Visual Guide
The McGrawHill
Companies, 2001

Beautifully Rendered Full-Color Art

Carefully prepared, accurate illustrations are a hallmark of this text.
Human anatomy is a visual science, and realistic art is essential.
Vibrant four-color illustrations are often paired with photographs,
reinforcing the detail conveyed in the drawings with direct
comparisons of actual structures.
Secretion
Lumen
Mucus
Liver
Stomach
Cell membrane
Gallbladder
Golgi complex
Large intestine
Small intestine
Nucleus of goblet cell

Rough endoplasmic
reticulum
Creek
Right lung
Diaphragm muscle
Left lung
Heart
Inferior vena cava
Right kidney
Left renal artery
Celiac trunk
Left kidney
Common hepatic
artery
Abdominal aorta
Right common
iliac artery
Right external
iliac artery
Right external
iliac vein
xviii
Inferior mesenteric
artery
Superior mesenteric
artery
Atlas-Quality Cadaver Images

Precisely labeled photographs of dissected human cadavers provide
detailed views of human anatomy that allow students concrete
visualization of anatomical structures and their position relative to
other parts of the body.

Front Matter
Illustrated Tables
A Visual Guide
Selected tables combine artwork with

summarized content to provide
comprehensive topic coverage in an easy-tofollow format.
The McGrawHill
Companies, 2001
TABLE 11.6 Septa of the Cranial Dura Mater

Septa
Location
Falx cerebri
Extends downward into the longitudinal fissure to partition the right and left cerebral hemispheres; anchored
anteriorly to the crista galli of the ethmoid bone and posteriorly to the tentorium
Tentorium cerebelli
Separates the occipital and temporal lobes of the cerebrum from the cerebellum; anchored to the tentorium, petrous
parts of the temporal bones, and occipital bone
Falx cerebelli
Partitions the right and left cerebellar hemispheres; anchored to the occipital crest
Diaphragma sellae
Forms the roof of the sella turcica
Superior sagittal sinus

Dura mater
Inferior sagittal sinus
Cerebral veins
Falx cerebri
Tentorium cerebelli
Cerebral arterial
circle
Cranium
Pituitary gland
Sella turcica
Transverse sinus
Falx cerebelli
Diaphragma sellae
Monocular field
Binocular field
Macular field
pathetic neurons by fibers from the superior colliculi. Postganglionic

neurons in the ciliary ganglia behind the eyes, in turn, stimulate
constrictor fibers in the iris. Contraction of the ciliary body during
accommodation also involves stimulation of the superior colliculi.
Processing of Visual Information
Lens
Retina
Optic nerve
Optic chiasma
Optic tract
Superior
colliculus
Optic radiation
Lateral geniculate
nucleus of
thalamus
Visual cortex of
occipital lobes
For visual information to have meaning, it must be associated

with past experience and integrated with information from other
senses. Some of this higher processing occurs in the inferior temporal lobes of the cerebral cortex. Experimental removal of these
areas from monkeys impairs their ability to remember visual tasks
that they previously learned and hinders their ability to associate
visual images with the significance of the objects viewed. Monkeys with their inferior temporal lobes removed, for example,
will fearlessly handle a snake. The symptoms produced by loss of
the inferior temporal lobes are known as KlverBucy syndrome.
In an attempt to reduce the symptoms of severe epilepsy,
surgeons at one time would cut the corpus callosum in some patients. This fiber tract, as previously described, transmits impulses
between the right and left cerebral hemispheres. The right cerebral hemisphere of patients with such split brains would therefore,
receive sensory information only from the left half of the external
world. The left hemisphere, similarly cut off from communication
with the right hemisphere, would receive sensory information
only from the right half of the external world. In some situations,
these patients would behave as if they had two separate minds.
Experiments with split-brain patients have revealed that the two
hemispheres have separate abilities. This is true even though
each hemisphere normally receives input from both halves of the external world through the corpus callosum. If the sensory image of an object, such as a key, is delivered only to the left hemisphere (by
showing it only to the right visual field), the object can be named. If the
object is presented to the right cerebral cortex, the person knows what
the object is but cannot name it. Experiments such as this suggest that
(in right-handed people) the left hemisphere is needed for language
and the right hemisphere is responsible for pattern recognition.
Topic Icons
Eyeball
Knowledge Check
Knowledge Check
vision. An overlapping of the visual field of each eye provides binocular visionthe ability to perceive depth.
superior colliculi stimulate the extrinsic ocular muscles (see

table 15.3), which are the skeletal muscles that move the eyes.
Two types of eye movements are coordinated by the superior colliculi. Smooth pursuit movements track moving objects and
keep the image focused on the fovea centralis. Saccadic (sakad'ik) eye movements are quick (lasting 2050 msec), jerky
movements that occur while the eyes appear to be still. These
saccadic movements are believed to be important in maintaining
visual acuity.
The tectal system is also involved in the control of the intrinsic ocular musclesthe smooth muscles of the iris and of the ciliary
body. Shining a light into one eye stimulates the pupillary reflex in
which both pupils constrict. This is caused by activation of parasym-
15. List the accessory structures of the eye that either cause the
eye to move or protect it within the orbit.
16. Diagram the structure of the eye and label the following:
sclera, cornea, choroid, retina, fovea centralis, iris, pupil,
lens, and ciliary body. What are the principal cells or tissues in each of the three layers of the eye?
17. Trace the path of light through the two cavities of the eye
and explain the mechanism of light refraction. Describe
how the eye is focused for viewing distant and near objects.
18. List the different layers of the retina and describe the path
of light and of nerve activity through these layers. Continue tracing the path of a visual impulse to the cerebral
cortex, and list in order the structures traversed.
Creek
FIGURE 15.27 Visual fields of the eyes and neural pathways for
Topic icons highlight information of practical

application and special interest. These commentaries
reinforce the importance of learning the preceding
facts. The five icon images and the topics they represent
are: clinical information (stethoscope), aging
(hourglass), developmental information (embryo),
homeostasis (gear mechanism), and academic interest
information (mortarboard).
Placed at the end of each major section, Knowledge

Check questions help you test your understanding of
the material and encourage concept application.
KlverBucy syndrome: from Heinrich Klver, German neurologist, 18971979

and Paul C. Bucy, American neurologist, b. 1904
xix
Front Matter
Developmental Exposition
The Axial Skeleton
A Visual Guide
Each systems chapter includes a discussion of the morphogenic

events involved in the prenatal development of the profiled body
system.
certain bones of the cranium are formed this way. Sesamoid bones
are specialized intramembranous bones that develop in tendons.
The patella is an example of a sesamoid bone.
EXPLANATION
The McGrawHill
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Developmental Expositions

DEVELOPMENT OF THE SKULL
Development of Bone
Bone formation, or ossification, begins at about the fourth week of
embryonic development, but ossification centers cannot be readily observed until about the tenth week (exhibit I). Bone tissue
derives from specialized migratory cells of mesoderm (see
fig. 4.13) known as mesenchyme. Some of the embryonic mesenchymal cells will transform into chondroblasts (kon'dro-blasts)
and develop a cartilage matrix that is later replaced by bone in a
process known as endochondral (en''do-kon'dral) ossification.
Most of the skeleton is formed in this fashionfirst it goes
through a hyaline cartilage stage and then it is ossified as bone.
A smaller number of mesenchymal cells develop into bone
directly, without first going through a cartilage stage. This type
of bone-formation process is referred to as intramembranous
(in''tra-mem'bra-nus) ossification. The clavicles, facial bones, and
The formation of the skull is a complex process that begins during the fourth week of embryonic development and continues
well beyond the birth of the baby. Three aspects of the embryonic skull are involved in this process: the chondrocranium, the
neurocranium, and the viscerocranium (exhibit II). The chondrocranium is the portion of the skull that undergoes endochondral ossification to form the bones supporting the brain. The
neurocranium is the portion of the skull that develops through
membranous ossification to form the bones covering the brain
and facial region. The viscerocranium (splanchnocranium) is the
portion that develops from the embryonic visceral arches to form
the mandible, auditory ossicles, the hyoid bone, and specific
processes of the skull.
chondrocranium: Gk. chondros, cartilage; kranion, skull

chondroblast: Gk. chondros, cartilage; blastos, offspring or germ
viscerocranium: L. viscera, soft parts; Gk. kranion, skull
Parietal
bones
Occipital
bone
Frontal
bones
Temporal
bone
Humerus
Zygomatic
bone
Maxilla
Nasal bone
Mandible
Metacarpal
bones
Phalanges
Carpal bones
Ribs
Radius
Ulna
Chondrocranium
Vertebrae
Clavicle
Scapula
Femur
Tibia
Fibula
Ilium
Sacrum
Coccyx
Phalanges
Metatarsal bones
Tarsal bones
Creek
(a)
(b)
EXHIBIT I Ossification centers of the skeleton of a 10-week-old fetus. (a) The diagram depicts endochondrial ossification in red and intramembranous ossification in a stippled pattern. The cartilaginous portions of the skeleton are shown in gray. (b) The photograph shows
the ossification centers stained with a red indicator dye.
These special sections appearing at the end of most chapters

describe selected developmental disorders, diseases, or dysfunctions
of specific organ systems, as well as relevant clinical procedures. The
effects of aging in regard to specific body systems are also profiled.
Clinical Considerations
CLINICAL CONSIDERATIONS
The clinical aspects of the central nervous system are extensive
and usually complex. Numerous diseases and developmental
problems directly involve the nervous system, and the nervous
system is indirectly involved with most of the diseases that afflict
the body because of the location and activity of sensory pain receptors. Pain receptors are free nerve endings that are present
throughout living tissue. The pain sensations elicited by disease
or trauma are important in localizing and diagnosing specific diseases or dysfunctions.
Only a few of the many clinical considerations of the central nervous system will be discussed here. These include neurological assessment and drugs, developmental problems, injuries,
infections and diseases, and degenerative disorders.
Creek
(a)
Third lumbar vertebra
Neurological Assessment and Drugs

Neurological assessment has become exceedingly sophisticated and
accurate in the past few years. In a basic physical examination, only
the reflexes and sensory functions are assessed. But if the physician
suspects abnormalities involving the nervous system, further neurological tests may be done, employing the following techniques.
A lumbar puncture is performed by inserting a fine needle
between the third and fourth lumbar vertebrae and withdrawing
a sample of CSF from the subarachnoid space (fig. 11.45). A cisternal puncture is similar to a lumbar puncture except that the
CSF is withdrawn from a cisterna at the base of the skull, near
the foramen magnum. The pressure of the CSF, which is normally about 10 mmHg, is measured with a manometer. Samples of
CSF may also be examined for abnormal constituents. In addition, excessive fluid, accumulated as a result of disease or trauma,
may be drained.
The condition of the arteries of the brain can be determined through a cerebral angiogram (an'je-o-gram). In this technique, a radiopaque substance is injected into the common
carotid arteries and allowed to disperse through the cerebral vessels. Aneurysms and vascular constrictions or displacements by
tumors may then be revealed on radiographs.
The development of the CT scanner, or computerized
axial tomographic scanner, has revolutionized the diagnosis of
brain disorders. The CT scanner projects a sharply focused, detailed tomogram, or cross section, of a patients brain onto a television screen. The versatile CT scanner allows quick and
accurate diagnoses of tumors, aneurysms, blood clots, and hemorrhage. The CT scanner may also be used to detect certain types
of birth defects, brain damage, scar tissue, and evidence of old or
recent strokes.
A machine with even greater potential than the CT scanner is the DSR, or dynamic spatial reconstructor. Like the
CT scanner, the DSR is computerized to transform radiographs
into composite video images. However, with the DSR, a threedimensional view is obtained, and the image is produced much
faster than with the CT scanner. The DSR can produce 75,000
cross-sectional images in 5 seconds, whereas the CT scanner can
xx
Coccyx
Spinal cord
(b)
Subarachnoid
space
Dura mater
Inserted
needle
Sacrum
FIGURE 11.45 (a) A lumbar puncture is performed by inserting a

needle between the third and fourth lumbar vertebrae (L3L4) and
(b) withdrawing cerebrospinal fluid from the subarachnoid space.
produce only one. With that speed, body functions as well as

structures may be studied. Blood flow through vessels of the brain
can be observed. These types of data are important in detecting
early symptoms of a stroke or other disorders.
Certain disorders of the brain may be diagnosed more simply by examining brain-wave patterns using an electroencephalogram (see Table 11.5). Sensitive electrodes placed on the
scalp record particular EEG patterns being emitted from evoked
cerebral activity. EEG recordings are used to monitor epileptic
patients to predict seizures and to determine proper drug therapy,
and also to monitor comatose patients.
The fact that the nervous system is extremely sensitive to
various drugs is fortunate; at the same time, this sensitivity has
potential for disaster. Drug abuse is a major clinical concern because of the addictive and devastating effect that certain drugs
have on the nervous system. Much has been written on drug
abuse, and it is beyond the scope of this text to elaborate on the
effects of drugs. A positive aspect of drugs is their administration
in medicine to temporarily interrupt the passage or perception of
sensory impulses. Injecting an anesthetic drug near a nerve, as in
dentistry, desensitizes a specific area and causes a nerve
block. Nerve blocks of a limited extent occur if an appendage is
cooled or if a nerve is compressed for a period of time. Before the

Front Matter
A Visual Guide
The McGrawHill
Companies, 2001
Clinical Practicums
These focused clinical scenarios challenge you to put your
knowledge of anatomy to work in a clinical setting. Given a brief
patient history and accompanying diagnostic images, you must apply
the chapter material to diagnose a condition, explain the origin of
symptoms, or even recommend a course of treatment. Detailed
answers to the Clinical Practicum questions are provided in
Appendix B.
CLINICAL PRACTICUM 16.1

A 75-year-old male with a long history of
hypertension presents to the emergency
room with complaints of stabbing chest pain
that goes through to his back. On physical
exam, the patients lungs are clear, and heart
sounds are also normal with regular rate and
rhythm. An electrocardiogram is also normal. Because of his symptoms, you suspect
an aortic dissection and order a CT scan.
(MRA = main pulmonary artery, AA = ascending aorta, DA = descending aorta.)
QUESTIONS
1. What is the dark line noted within the
contrast-filled aorta?
2. Which portions of the aorta are
involved?
3. You also note that the patient has a
difference in blood pressure between
the left and right arm, with the left arm
having a significantly lower blood
pressure. What could be the cause?
AA MPA
DA
At the end of each chapter, a summary in outline form reinforces

your mastery of the chapter content.
Objective, essay, and critical thinking

questions at the end of each chapter allow
you to test the depth of your understanding
and learning. Answers and explanations
to the objective questions are given
in Appendix A. The essay and critical
thinking exercises are answered
in the Instructors Manual.
Review Activities
Review Activities
Objective Questions
1. Viscera are the only body organs that are
(a) concerned with digestion.
(b) located in the abdominal cavity.
(c) covered with peritoneal membranes.
(d) located within the thoracic and
abdominal cavities.
2. Which of the following types of teeth are
found in the permanent but not in the
deciduous dentition?
(a) incisors
(c) premolars
(b) canines
(d) molars
3. The double layer of peritoneum that
supports the GI tract is called
(a) the visceral peritoneum.
(b) the mesentery.
(c) the greater omentum.
(d) the lesser omentum.
4. Which of the following tissue layers in
the small intestine contains the lacteals?
(a) the submucosa
(b) the muscularis mucosae
(c) the lamina propria
(d) the tunica muscularis
5. Which of the following organs is not
considered a part of the digestive system?
(a) the pancreas (c) the tongue
(b) the spleen
(d) the gallbladder
6. The numerous small elevations on the
surface of the tongue that support taste
buds and aid in handling food are called
(a) cilia.
(c) intestinal villi.
(b) rugae.
(d) papillae.
7. Most digestion occurs in
(a) the mouth.
(b) the stomach.
(c) the small intestine.
(d) the large intestine.
8. Stenosis (constriction) of the sphincter of
ampulla (of Oddi) would interfere with
(a) transport of bile and pancreatic juice.
(b) secretion of mucus.
(c) passage of chyme into the small
intestine.
(d) peristalsis.
9. The first organ to receive the blood-borne
products of digestion is
(a) the liver.
(c) the heart.
(b) the pancreas. (d) the brain.
Chapter Summary
Chapter Summary
Introduction to the Digestive System
(pp. 635636)
2. The incisors and canines have one root

each; the bicuspids and molars have two
or three roots.
(a) Humans are diphyodont; they have
deciduous and permanent sets of teeth.
(b) The roots of teeth fit into sockets
called dental alveoli that are lined
with a periodontal membrane. Fibers
in the periodontal membrane insert
into the cementum covering the
roots, firmly anchoring the teeth in
the sockets.
(c)
Enamel
forms the outer layer of the
Serous Membranes and Tunics of the
tooth
crown; beneath the enamel is
Gastrointestinal Tract (pp. 636640)
dentin.
1. Peritoneal membranes line the abdominal
(d) The interior of a tooth contains a
wall and cover the visceral organs. The
pulp cavity, which is continuous
GI tract is supported by a double layer of
through the apical foramen of the
peritoneum called the mesentery.
root with the connective tissue
(a) The lesser omentum and greater
around the tooth.
omentum are folds of peritoneum that
3. The major salivary glands are the parotid
extend from the stomach.
glands, the submandibular glands, and the
(b) Retroperitoneal organs are positioned
sublingual glands.
behind the parietal peritoneum.
4. The muscular pharynx provides a
2. The layers (tunics) of the abdominal GI
passageway connecting the oral and nasal
tract are, from the inside outward, the
cavities to the esophagus and larynx.
mucosa, submucosa,
tunica muscularis,
10. Which of the following statements
about
13. Describe the location and gross structure
and serosa.
andofStomach
(pp. 648652)
hepatic portal blood is true?
of the liver. Draw aEsophagus
labeled diagram
a
simple
(a) It contains absorbed(a)
fat. The mucosa consists of aliver
lobule.
1.
Swallowing
(deglutition) occurs in three
columnar epithelium, a thin layer of
(b) It contains ingested proteins.
14. Describe how the gallbladder
filledinvolves
with structures of the oral
phasesis and
connective
the lamina
(c) It is mixed with bile in the
liver. tissue calledand
emptied of bile fluid.cavity,
What pharynx,
is the and esophagus.
propria,
of smooth
(d) It is mixed with blood from
the and thin layers function
of bile?
2. Peristaltic waves of contraction push food
muscle called the muscularis
hepatic artery in the liver.
15. List the functions of the through
large intestine.
the lower esophageal sphincter
mucosae.
What are the biomechanical
movements
into the
stomach.
(b) The submucosa is composed of
Essay Questions
of the large intestine 3.
thatThe
make
these consists of a cardia, fundus,
stomach
connective tissue; the tunica
functions possible?
body, and pylorus. It displays greater and
1. Define digestion. Differentiatemuscularis
between consists of layers
of
16. Define cirrhosis and explain
why
this
lesser
curvatures,
and contains a pyloric
the mechanical and chemicalsmooth
aspectsmuscle;
of
and the serosa is
condition is so devastating
to the liver.
sphincter
at its junction with the
digestion.
composed of connective tissue
What are some of the causes
of cirrhosis?
duodenum.
2. Distinguish between the gastrointestinal
covered with the visceral peritoneum.
(a) The mucosa of the stomach is thrown
tract, viscera, accessory digestive
(c) The organs,
submucosa contains the
Critical-Thinking Questions into distensible gastric folds; gastric
and gut.
submucosal plexus, and the tunica
gastric glands are present in
3. List the specific portions or structures
1. Technically,
notand
in the
muscularisofcontains the
myenteric ingested food ispits
the mucosa.
the digestive system formed by
eachofofautonomic
the
body. Neither are feces excreted
from
plexus
nerves.
parietal cells of the gastric glands
three embryonic germ layers.
within the body (except (b)
bile The
residue).
secrete
HCl,
4. Define serous membrane.
HowPharynx,
are the and Associated
Explain these statements. Why
would
thisand the principal cells
Mouth,
serous membranes ofStructures
the abdominal
a drug pepsinogen.
(pp. 640648) information be important to secrete
cavity classified and what are their
company interested in preparing a new
1. The oral cavity is formed by the cheeks,
functions?
oral medication? Small Intestine (pp. 652656)
lips,
and
hard
palate
and
soft
palate.
The
5. Describe the structures of the four tunics
2. The deciduous (milk)1.teeth
dontofmatter
Regions
the small intestine include the
tongue and teeth are contained in the
in the wall of the GI tract.
because they fall out anyway.
Do youjejunum, and ileum; the
duodenum,
oral cavity.
6. Why are there two autonomic
agree or disagree with this
statement?
common
bile duct and pancreatic duct
(a) Lingual
with
innervations to the GI tract?
Identifytonsils
the and papillae
Explain.
empty into the duodenum.
taste budsin
are located on the tongue.
specific sites of autonomic stimulation
3. Which surgery do you2.think
would
have
Fingerlike extensions of mucosa, called
(b) Structures of the palate include
the tunic layers.
the most profound effectintestinal
on digestion:
villi, project into the lumen,
palatal folds, a cone-shaped
7. Define the terms dental formula,
(a) removal of the stomach
and(gastrectomy),
at the bases of the intestinal villi the
projection called the palatine uvula,
diphyodont, deciduous teeth, permanent
(b) removal of the pancreas
mucosa forms intestinal glands.
and palatine tonsils.
teeth, and wisdom teeth.
(pancreatectomy), or (c)(a)
removal
of the
New epithelial
cells are formed in the
8. Outline the stages of deglutition. What
gallbladder (cholecystectomy)?
Explaincrypts.
intestinal
biomechanical roles do the tongue, hard
your reasoning.
palate and soft palate, pharynx, and hyoid
4. Describe the adaptations of the GI tract
bone perform in deglutition?
that make it more efficient by either
9. How does the stomach protect itself from
increasing the surface area for absorption
the damaging effects of HCl?
or increasing the time of contact between
10. Describe the kinds of movements in the
food particles and digestive enzymes.
small intestine and explain what they
5. During surgery to determine the cause of
accomplish.
an intestinal obstruction, why might the
11. Diagram an intestinal villus and explain
surgeon elect to remove a healthy
why intestinal villi are considered the
appendix?
functional units of the digestive system.
6. Explain why a ruptured appendix may
12. What are the regions of the large
result in peritonitis, while an inflamed
intestine? In what portion of the
kidney (nephritis) generally does not
abdominal cavity and pelvic cavity is each
result in peritonitis.
region located?
1. The digestive system mechanically and
chemically breaks down food to forms
that can be absorbed through the
intestinal wall and transported by the
blood and lymph for use at the cellular
level.
2. The digestive system consists of a
gastrointestinal (GI) tract and accessory
digestive organs.
(b) The membrane of intestinal

epithelial cells is folded to form
microvilli; this brush border of the
mucosa increases the absorptive
surface area.
3. Movements of the small intestine include
rhythmic segmentation, pendular
movement, and peristalsis.
Large Intestine (pp. 656660)

1. The large intestine absorbs water and
electrolytes from the chyme and passes
fecal material out of the body through the
rectum and anal canal.
2. The large intestine is divided into the
cecum, colon, rectum, and anal canal.
(a) The appendix is attached to the
inferior medial margin of the cecum.
(b) The colon consists of ascending,
transverse, descending, and sigmoid
portions.
(c) Haustra are bulges in the walls of the
large intestine.
3. Movements of the large intestine include
peristalsis, haustral churning, and mass
movement.
Liver, Gallbladder, and Pancreas

(pp. 660669)
1. The liver is divided into right, left,
quadrate, and caudate lobes. Each lobe
contains liver lobules, the functional units
of the liver.
(a) Liver lobules consist of plates of
hepatic cells separated by modified
capillaries called sinusoids.
(b) Blood flows from the periphery of
each lobule, where branches of the
hepatic artery and hepatic portal vein
empty, through the sinusoids and out
the central vein.
(c) Bile flows within the hepatic plates,
in bile canaliculi, to the biliary
ductules at the periphery of each
lobule.
2. The gallbladder stores and concentrates
the bile; it releases the bile through the
cystic duct and common bile duct into
the duodenum.
3. The pancreas is both an exocrine and an
endocrine gland.
(a) The endocrine portion, consisting of
the pancreatic islets, secretes the
hormones insulin and glucagon.
(b) The exocrine acini of the pancreas
produce pancreatic juice, which
contains various digestive enzymes.
Visit our Online Learning Center at http://www.mhhe.com/vdg

for chapter-by-chapter quizzing, additional study resources, and related web links.
xxi

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Resources
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The Online Learning Center (OLC) that accompanies this text is
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extensive array of learning tools that are tailored to coincide with
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Learning Activities
Among the activities awaiting you at the OLC are chapter quizzes, crossword puzzles, art labeling exercises, vocabulary flashcards, and
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CHAPTER 1

Unit 1
I. Historical Perspective
1. History of Anatomy
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Historical Perspective
Like the people of Mesopotamia and Greece, the ancient

Egyptians were concerned with a controlling spirit of the
body. In fact, they even had a name for this life forcethe Ba spirit
and they believed that it was associated with the bowels and the
heart. Food was placed in the tomb of a mummy to feed the Ba spirit
during the journey to Osiris, Egyptian god of the underworld.
China and Japan

China
In ancient China, interest in the human body was primarily
philosophical. Ideas about anatomy were based on reasoning
rather than dissection or direct observation. The Chinese
revered the body and abhorred its mutilation. An apparent exception was the practice of binding the feet of young girls and
women in an attempt to enhance their beauty. Knowledge of the
internal organs came only from wounds and injuries. Only in recent times have dissections of cadavers been permitted in Chinese medical schools.
The ancient Chinese had an abiding belief that everything
in the universe depended on the balance of the two opposing
cosmic principles of yin and yang. As for the circulatory system,
the blood was the conveyor of the yang, and the heart and vessels represented the yin. Other structures of the body were composed of lesser forces termed zo and fu.
The Chinese were great herbalists. Writings more than
5,000 years old describe various herbal concoctions and potions
to alleviate a wide variety of ailments, including diarrhea, constipation, and menstrual discomfort. Opium was described as an excellent painkiller.
Until recently, the Chinese have been possessive of their
beliefs, and for this reason Western cultures were not influenced
by Chinese thoughts or writings to an appreciable extent. Perhaps the best known but least understood of the Chinese contributions to human anatomy and medicine is acupuncture.
Acupuncture is an ancient practice that was established to
maintain a balance between the yin and the yang. Three hundred
sixty-five precise meridian sites, or vital points, corresponding to
the number of days in a year, were identified on the body (fig. 1.7).
Needles inserted into the various sites were believed to release
harmful secretions and rid the tissues of obstructions. Acupuncture
is still practiced in China and has gained acceptance with some
medical specialists in the United States and other countries as a
technique of anesthesia and as a treatment for certain ailments.
The painkilling effect of acupuncture has been documented and is
more than psychological. Acupuncture sites have been identified
on domestic animals and have been used to a limited degree in
veterinary medicine. Why acupuncture is effective remains a mystery, although it has recently been correlated with endorphin production within the brain (see chapter 11).
Japan
The advancement of anatomy in Japan was strongly influenced
by the Chinese and Dutch. The earliest records of anatomical
interest in Japan date back to the sixth century. Buddhist
monks from Japan were trained in China where they were exposed to Chinese philosophy, and so Chinese beliefs concerning the body became prevalent in Japan as well. By the
eighteenth century, Western influences, especially the Dutch,
were such that the Japanese sought to determine for themselves
which version of anatomy was correct. In 1774, a five-volume
work called Kaitai Shinsho (A New Book of Anatomy), published
by a Japanese physician, Genpaku Sugita, totally adopted the
Dutch conceptualization of the body. This book marked the beginning of a modern era in anatomy and medicine for the
Japanese people.
For several hundred years, Western nations were welcome in
Japan. In 1603, however, the Japanese government banned
all contact with the Western world because they feared the influences
of Christianity on their society. Although this ban was strictly enforced
and Japan became isolated, Japanese scholars continued to circulate Western books on anatomy and medicine. These books eventually prompted Japanese physicians to reassess what they had been
taught concerning the structure of the body.
Grecian Period
It was in ancient Greece that anatomy first gained wide acceptance as a science. The writings of several Greek philosophers
had a tremendous impact on future scientific thinking. During
this period, the Greeks were obsessed with the physical beauty of
the human body, as reflected by their exquisite sculptures.
The young people of Greece were urged to be athletic and
develop their physical abilities, but at about age 18 they were directed more to intellectual pursuits of science, rhetoric, and philosophy. An educated individual was expected to be acquainted
with all fields of knowledge, and it was only natural that great
strides were made in the sciences.
Perhaps the first written reference to the anatomy of
wounds sustained in battle is contained in the Iliad, written by
Homer in about 800 B.C. Homers detailed descriptions of the
anatomy of wounds were exceedingly accurate. However, he described clean woundsnot the type of traumatic wounds that
would likely be suffered on a battlefield. This has led to speculation that human dissections were conducted during this period
and that anatomical structure was well understood. Victims of
human sacrifice may have served as subjects for anatomical study
and demonstration.
Hippocrates
Hippocrates (460377 B.C.), the most famous of the Greek physicians of his time, is regarded as the father of medicine because of
the sound principles of medical practice that his school established (fig. 1.8). His name is memorialized in the Hippocratic
oath, which many graduating medical students repeat as a
promise of professional stewardship and duty to humankind.
Hippocrates probably had only limited exposure to human
dissections, but he was well disciplined in the popular humoral
theory of body organization. Four body humors were recognized,
humor: L. humor, fluid

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(a)
History of Anatomy
(b)
FIGURE 1.7 Acupuncture has long been practiced for diagnostic or therapeutic purposes. Acupressure is application of finger-point pressure at
specific meridian sites to manage pain. (a) An acupuncture chart from the Ming dynasty of ancient China and (b) a patient receiving acupuncture.
and each was associated with a particular body organ: blood with
the liver; choler, or yellow bile, with the gallbladder; phlegm
with the lungs; and melancholy, or black bile, with the spleen. A
healthy person was thought to have a balance of the four humors. The concept of humors has long since been discarded, but
it dominated medical thought for over 2,000 years.
Perhaps the greatest contribution of Hippocrates was that
he attributed diseases to natural causes rather than to the displeasure of the gods. His application of logic and reason to medicine was the beginning of observational medicine.
The four humors are a part of our language and medical
practice even today. Melancholy is a term used to describe
depression or despondency in a person, whereas melanous refers to
a black or sallow complexion. The prefix melano- means black.
Cholera is an infectious intestinal disease that causes diarrhea and

vomiting. Phlegm (pronounced flem) within the upper respiratory system is symptomatic of several pulmonary disorders. Sanguine, a term
that originally referred to blood, is used to describe a passionate
temperament. This term, however, has evolved to refer simply to the
cheerfulness and optimism that accompanied a sanguine personality, and no longer refers directly to the humoral theory.
Aristotle
Aristotle (384322 B.C.), a pupil of Plato, was an accomplished
writer, philosopher, and zoologist (fig. 1.9). He was also a renowned
teacher and was hired by King Philip of Macedonia to tutor his son,
Alexander, who later became known as Alexander the Great.
cholera: Gk. chole, bile

melancholy: Gk. melan, black; chole, bile
phlegm: Gk. phlegm, inflammation

sanguine: L. sanguis, bloody
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The Hippocratic Oath

I swear by Apollo Physician and Aesculapius and Hygeia and Panacea
and all the gods and goddesses making them my witnesses, that I will
fulfill according to my ability and judgment this oath and this covenant:
To hold him who has taught me this art as equal to my parents and to
live my life in partnership with him, and if he is in need of money to give
him a share of mine, and to regard his offspring as equal to my brothers
in male lineage and to teach them this artif they desire to learn it
without fee and covenant; to give a share of precepts and oral instruction
and all the other learning to my sons and to the sons of him who has
instructed me and to pupils who have signed the covenant and have
taken an oath according to the medical law, but to no one else.
I will apply dietetic measures for the benefit of the sick according to
my ability and judgment; I will keep them from harm and injustice.
I will neither give a deadly drug to anybody if asked for it, nor will I make
a suggestion to this effect. Similarly I will not give to a woman an abortive
remedy. In purity and holiness I will guard my life and my art.
I will not use the knife, not even on sufferers from stone, but will
withdraw in favor of such men as are engaged in this work.
Whatever houses I may visit, I will come for the benefit of the sick,
remaining free of all intentional injustice, of all mischief, and in particular
of sexual relations with both female and male persons, be they free or
slaves.
What I may see or hear in the course of the treatment or even outside of
the treatment in regard to the life of men, which on no account one must
spread abroad, I will keep to myself, holding such things shameful to be
spoken about.
If I fulfill this oath and do not violate it, may it be granted to me to
enjoy life and art, being honored with fame among all men for all time
to come; if I transgress it and swear falsely, may the opposite of all this
be my lot.
FIGURE 1.8 A fourteenth-century painting of the famous Greek physician Hippocrates. Hippocrates is referred to as the father of medicine;
his creed is immortalized as the Hippocratic oath (left).
Aristotle made careful investigations of all kinds of animals, which included references to humans, and he pursued a
limited type of scientific method in obtaining data. He wrote the
first known account of embryology, in which he described the
development of the heart in a chick embryo. He named the aorta
and contrasted the arteries and veins. Aristotles best known zoological works are History of Animals, On the Parts of Animals, and
On the Generation of Animals (see table 1.2). These books had a
profound influence on the establishment of specialties within
anatomy, and they earned Aristotle recognition as founder of
comparative anatomy.
In spite of his tremendous accomplishments, Aristotle perpetuated some erroneous theories regarding anatomy. For example, the doctrine of the humors formed the boundaries of his
thought. Plato had described the brain as the seat of feeling and
thought, but Aristotle disagreed. He placed the seat of intelligence in the heart and argued that the function of the brain,
which was bathed in fluid, was to cool the blood that was
pumped from the heart, thereby maintaining body temperature.
Aesculapius: Gk. (mythology) son of Apollo and god of medicine

Hygeia: Gk. (mythology) daughter of Aesculapius; personification of health;
hygies, healthy
Panacea: Gk. (mythology) also a daughter of Aesculapius; assisted in temple rites
and tended sacred serpents; pan, all, every; akos, remedy
Alexandrian Era
Alexander the Great founded Alexandria in 332 B.C. and established it as the capital of Egypt and a center of learning. In addition to a great library, there was also a school of medicine in
Alexandria. The study of anatomy flourished because of the acceptance of dissections of human cadavers and human vivisections (viv-sekshunz) (dissections of living things). This brutal
procedure was commonly performed on condemned criminals.
People reasoned that to best understand the functions of the
body, it should be studied while the subject was alive, and that a
condemned man could best repay society through the use of his
body for a scientific vivisection.
Unfortunately, the scholarly contributions and scientific
momentum of Alexandria did not endure. Most of the written
works were destroyed when the great library was burned by the
Romans as they conquered the city in 30 B.C. What is known
about Alexandria was obtained from the writings of later scientists, philosophers, and historians, including Pliny, Celsus,
Galen, and Tertullian. Two men of Alexandria, Herophilus and
Erasistratus, made lasting contributions to the study of anatomy.
vivisection: L. vivus, alive; sectio, a cutting

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History of Anatomy
11
rate, he also had notions that were primitive and mystical. He

thought, for example, that the cranial nerves carried animal spirits and that muscles contracted because of distention by spirits.
He also believed that the left ventricle of the heart was filled
with a vital air spirit (pneuma) that came in from the lungs, and
that the arteries transported this pneuma rather than blood.
Both Herophilus and Erasistratus were greatly criticized later
in history for the vivisections they performed. Celsus (about
30 B.C.) and Tertullian (about A.D. 200) were particularly critical of the
practice of vivisection. Herophilus was described as a butcher of
men who had dissected as many as 600 living human beings, sometimes in public demonstrations.
Roman Era
FIGURE 1.9 This Roman copy of a Greek sculpture is believed to

be of Aristotle, the famous Greek philosopher.
Herophilus
Herophilus (about 325 B . C .) was trained in the Hippocratic
school but became a great teacher of anatomy in Alexandria.
Through vivisections and dissections of human cadavers, he provided excellent descriptions of the skull, eye, various visceral organs and organ relationships, and the functional relationship of
the spinal cord to the brain. Two monumental works by
Herophilus are entitled On Anatomy and Of the Eyes. He regarded the brain as the seat of intelligence and described many of
its structures, such as the meninges, cerebrum, cerebellum, and
fourth ventricle. He was also the first to distinguish nerves as either sensory or motor.
Erasistratus
In many respects, the Roman Empire stifled scientific advancements and set the stage for the Dark Ages. The approach to science shifted from theoretical to practical during this time. Few
dissections of cadavers were performed other than in attempts to
determine the cause of death in criminal cases. Medicine was not
preventive but was limited, almost without exception, to the
treatment of soldiers injured in battle. Later in Roman history,
laws were established that attested to the influence of the
Church on medical practice. According to Roman law, for example, no deceased pregnant woman could be buried without prior
removal of the fetus from the womb so that it could be baptized.
The scientific documents that have been preserved from
the Roman Empire are mostly compilations of information obtained from the Greek and Egyptian scholars. New anatomical
information was scant, and for the most part was derived from
dissections of animals other than human. Two important
anatomists from the Roman era were Celsus and Galen.
Celsus
Most of what is currently known about the Alexandrian school
of medicine is based on the writings of the Roman encyclopedist
Cornelius Celsus (30 B.C.A.D. 30). He compiled this information into an eight-volume work called De re medicina. Celsus had
only limited influence in his own time, however, probably because of his use of Latin rather than Greek. It was not until the
Renaissance that the enormous value of his contribution was
recognized.
Galen
Erasistratus (about 300 B.C.) was more interested in body functions than structure and is frequently referred to as the father of
physiology. In a book on the causes of diseases, he included observations on the heart, vessels, brain, and cranial nerves. Erasistratus noted the toxic effects of snake venom on various visceral
organs and described changes in the liver resulting from various
diseases. Although some of his writings were scientifically accu-
Claudius Galen (A.D. 130201) was perhaps the best physician

since Hippocrates. A Greek living under Roman domination, he
was certainly the most influential writer of all times on medical
subjects. For nearly 1,500 years, the writings of Galen represented the ultimate authority on anatomy and medical treatment. Galen probably dissected no more than two or three
human cadavers during his career, of necessity limiting his
anatomical descriptions to nonhuman animal dissections. He
visceral: L. viscus, internal organ
pneuma: Gk. pneuma, air
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Middle Ages
The Middle Ages, frequently referred to as the Dark Ages,
began with the fall of Rome to the Goths in A.D. 476 and lasted
nearly 1,000 years, until Constantinople was conquered by the
Turks in 1453. The totalitarian Christian Church suppressed
science and medical activity stagnated. Human existence continued to be miserably precarious, and people no longer felt capable of learning from personal observation. Rather, they
accepted life on faith.
Dissections of cadavers were totally prohibited during this
period, and molesting a corpse was a criminal act frequently punishable by burning at the stake. If mysterious deaths occurred, examinations by inspection and palpation were acceptable. During
the plague epidemic in the sixth century, however, a few necropsies (nekrop-sez) and dissections were performed in the hope of
determining the cause of this dreaded disease (see fig. 1.4).
FIGURE 1.10 These surgical and gynecological instruments were

found in the House of the Surgeon (about A.D. 6779) in Pompeii.
They are representative of the medical equipment used throughout
the Roman Empire during this time.
During the Crusades soldiers cooked the bones of their dead

comrades so that they could be returned home for proper
burial. Even this act, however, was eventually considered sacrilegious and strongly condemned by the Church. One of the ironies of
the Middle Ages was that the peasants received far less respect and
had fewer rights when they were alive than when they were dead.
Contributions of Islam
compiled nearly 500 medical papers (of which 83 have been preserved) from earlier works of others, as well as from his personal
studies. He perpetuated the concept of the humors of the body
and gave authoritative explanations for nearly all body functions.
Galens works contain many errors, primarily because of
his desire to draw definitive conclusions regarding human body
functions on the basis of data obtained largely from animals
such as monkeys, pigs, and dogs. He did, however, provide some
astute and accurate anatomical details in what are still regarded
as classic studies. He proved to be an experimentalist, demonstrating that the heart of a pig would continue to beat when
spinal nerves were transected so that nerve impulses could not
reach the heart. He showed that the squealing of a pig stopped
when the recurrent laryngeal nerves that innervated its vocal
cords were cut. Galen also tied off the ureter in a sheep to prove
that urine was produced in the kidney, not in the urinary bladder as had been falsely assumed. In addition, he proved that the
arteries contained blood rather than pneuma.
Galen compiled a list of many medicinal plants and used
medications extensively to treat illnesses. Although he frequently used bloodletting in an effort to balance the four humors, he cautioned against removing too much blood. He
accumulated a wide variety of medical instruments and suggested
their use as forceps, retractors, scissors, and splints (fig. 1.10). He
was also a strong advocate of helping nature heal through good
hygiene, a proper diet, rest, and exercise.
The Arabic-speaking people made a profound contribution to

the history of anatomy in a most unusual way. It was the Islamic
world that saved much of Western scholarship from the ruins of
the Roman Empire, the oppression of the Christian Church, and
the onset of the Middle Ages. With the expansion of Islam
through the Middle East and North Africa during the eighth
century, the surviving manuscripts from Alexandria were taken
back to the Arab countries, where they were translated from
Greek to Arabic.
As the Dark Ages enshrouded Europe, the Christian
Church attempted to stifle any scholarship or worldly knowledge
that was not acceptable within Christian dogma. The study of
the human body was considered heretical, and the Church
banned all writings on anatomical subjects. Without the Islamic
repository of the writings of Aristotle, Hippocrates, Galen, and
others, the progress of centuries in anatomy and medicine would
have been lost. It wasnt until the thirteenth century that the
Arabic translations were returned to Europe and, in turn, translated to Latin. During the translation process, any Arabic terminology that had been introduced was systematically removed, so
that today we find few anatomical terms of Arabic origin.
epidemic: Gk. epi, upon; demos, people

necropsy: Gk. nekros, corpse; opsy, view

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History of Anatomy
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Renaissance
The period known as the Renaissance was characterized by a rebirth of science. It lasted roughly from the fourteenth through
the sixteenth centuries and was a transitional period from the
Middle Ages to the modern age of science.
The Renaissance was ushered in by the great European
universities established in Bologna, Salerno, Padua, Montpellier, and Paris. The first recorded human dissections at these
newly established centers of learning were the work of the surgeon William of Saliceto (121580) from the University of
Bologna. The study of anatomy quickly spread to other universities, and by the year 1300 human dissections had become an integral part of the medical curriculum. However, the Galenic
dogma that normal human anatomy was sufficiently understood
persisted, so interest at this time centered on methods and techniques of dissection rather than on furthering knowledge of the
human body.
The development of movable type in about 1450 revolutionized the production of books. Celsus, whose De re medicina
was rediscovered during the Renaissance, was the first medical
author to be published in this manner. Among the first anatomy
books to be printed in movable type was that of Jacopo Berengario of Carpi, a professor of surgery at Bologna. He described
many anatomical structures, including the appendix, thymus,
and larynx. The most influential text of the period was written
by Mondino de Luzzi, also of the University of Bologna, in 1316.
First published in 1487, it was more a dissection guide than a
study of gross anatomy, and in spite of its numerous Galenic errors, 40 editions were published, until the time of Vesalius.
Because of the rapid putrefaction of an unembalmed corpse,
the anatomy textbooks of the early Renaissance were organized so that the more perishable portions of the body were considered first. Dissections began with the abdominal cavity, then the
chest, followed by the head, and finally the appendages. A dissection was a marathon event, frequently continuing for perhaps 4 days.
With the increased interest in anatomy during the Renaissance, obtaining cadavers for dissection became a serious problem. Medical students regularly practiced grave robbing until
finally an official decree was issued that permitted the bodies of
executed criminals to be used as specimens.
Corpses were embalmed to prevent deterioration, but this
was not especially effective, and the stench from cadavers
was apparently a persistent problem. Anatomy professors lectured
from a thronelike chair at some distance from the immediate area
(fig. 1.11). The phrase, I wouldnt touch that with a 10-foot pole
probably originated during this time in reference to the smell of a decomposing cadaver.
The major advancements in anatomy that occurred during

the Renaissance were in large part due to the artistic and scientific abilities of Leonardo da Vinci and Andreas Vesalius. Working in the fifteenth and sixteenth centuries, each produced
monumental studies of the human form.
FIGURE 1.11 The scene of a cadaver dissection, 1500, from Fasciculus Medicinae by Johannes de Ketham. The anatomy professor
removed himself from the immediate area to a thronelike chair overlooking the proceedings. The dissections were performed by hired
assistants. One of them, the ostensor, pointed to the internal structures with a wand as the professor lectured.
Leonardo
The great Renaissance Italian Leonardo da Vinci (14521519) is
best known for his artistic works (e.g., Mona Lisa) and his scientific contributions. He displayed genius as a painter, sculptor, architect, musician, and anatomistalthough his anatomical
drawings were not published until the end of the nineteenth century. As a young man, Leonardo regularly participated in cadaver
dissections and intended to publish a textbook on anatomy with
the Pavian professor Marcantonio della Torre. The untimely
death of della Torre at the age of 31 halted their plans. When
Leonardo died, his notes and sketches were lost and were not discovered for more than 200 years. The advancement of anatomy
would have been accelerated by many years if Leonardos notebooks had been available to the world at the time of his death.
Leonardos illustrations helped to create a new climate of
visual attentiveness to the structure of the human body. He was
intent on accuracy, and his sketches are incredibly detailed
(fig. 1.12). He experimentally determined the structure of complex body organs such as the brain and the heart. He made wax
casts of the ventricles of the brain to study its structure. He constructed models of the heart valves to demonstrate their action.
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FIGURE 1.13 A painting of the great anatomist Andreas Vesalius,

as he dissects a cadaver. From his masterpiece De Humani Corporis
Fabrica.
FIGURE 1.12 Only a master artist could achieve the detail and
accuracy of the anatomical sketches of Leonardo da Vinci. The
painter who has acquired a knowledge of the nature of the sinews,
muscles and tendons, Leonardo wrote, will know exactly in the
movement of any limb how many and which of the sinews are the
cause of it, and which muscle by its swelling is the cause of the
sinews contracting.
Vesalius
The contribution of Andreas Vesalius (151464) to the science
of human anatomy and to modern medicine is immeasurable.
Vesalius was born in Brussels into a family of physicians. He received his early medical training at the University of Paris and
completed his studies at the University of Padua in Italy, where
he began teaching surgery and anatomy immediately after graduation. At Padua, Vesalius participated in human dissections and
initiated the use of live models to determine surface landmarks
for internal structures (fig. 1.13).
Vesalius apparently had enormous energy and ambition. By
the time he was 28 years old, he had already completed the masterpiece of his life, De Humani Corporis Fabrica, in which the various body systems and individual organs are beautifully illustrated
and described (fig. 1.14). His book was especially important in
that it boldly challenged hundreds of Galens teachings. Vesalius
wrote of his surprise upon finding numerous anatomical errors in
the works of Galen that were taught as fact, and he refused to accept Galens explanations on faith. Because he was so outspo-
FIGURE 1.14 A plate from De Humani Corporis Fabrica, which

Vesalius completed at the age of 28. This book, published in 1543,
revolutionized the science of anatomy.

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FIGURE 1.15 The Anatomy Lesson of Dr. Tulp, a famous Rembrandt painting completed in 1632, depicts one of the public anatomies that
were popular during this period. Dr. Nicholas Tulp was a famous Dutch anatomist who described the congenital defect in the spinal column
known as spinal bifida aperta.
kenly opposed to Galen, he incurred the wrath of many of the traditional anatomists, including his former teacher Sylvius (Jacques
Dubois) of Paris. Sylvius even went so far as to give him the nickname Vesanus (madman). Vesalius became so unnerved by the
relentless attacks that he destroyed much of his unpublished work
and ceased his dissections.
Fortunately there were also serious, scientific-minded

anatomists who made significant contributions during this period. Two of the most important contributions were the explanation of blood flow and the development of the microscope.
Although Vesalius was the greatest anatomist of his epoch,

others made significant contributions and to an extent paved
the way for Vesalius. Michelangelo pursued anatomy in 1495, being
supplied with corpses by the friar of a local monastery. Mondino de
Luzzi and the surgeon Jacopo Berengario of Carpi also corrected
many of Galens errors. Fallopius (152362) and Eustachius
(152474) completed detailed dissections of specific body regions.
In 1628, the English physician William Harvey (15781657)

published his pioneering work On the Motion of the Heart and
Blood in Animals. Not only did this brilliant research establish
proof of the continuous circulation of blood within contained
vessels, it also provided a classic example of the scientific
method of investigation (fig. 1.16). Like Vesalius, Harvey was
severely criticized for his departure from Galenic philosophy.
The controversy over circulation of the blood raged for 20 years,
until other anatomists finally repeated Harveys experiments and
concurred with his findings.
Seventeenth and Eighteenth Centuries

During the seventeenth and eighteenth centuries, the science of
anatomy attained an unparalleled acceptance. In some of its aspects, it also took on a somewhat theatrical quality. Elaborate
amphitheaters were established in various parts of Europe for
public demonstrations of human dissections (fig. 1.15). Exorbitantly priced tickets of admission were sold to the wealthy, and
the dissections were performed by elegantly robed anatomists
who were also splendid orators. The subjects were usually executed prisoners, and the performances were scheduled during
cold weather because of the perishable nature of the cadavers.
Harvey
Leeuwenhoek
Antoni van Leeuwenhoek (laven-hook) (16321723) was a
Dutch optician and lens grinder who improved the microscope
to the extent that he achieved a magnification of 270 times. His
many contributions included developing techniques for examining tissues and describing blood cells, skeletal muscle, and
the lens of the eye. Although he was the first to accurately describe sperm cells, Leeuwenhoek did not understand their role in
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salivary glands and lymph nodes within the neck and facial regions. In 1664, Thomas Willis published a summary of what was
then known about the nervous system.
A number of anatomical structures throughout the body are
named in honor of the early anatomists. Thus we have
graafian follicles, Stensens and Whartons ducts, fallopian tubes,
Bartholins glands, the circle of Willis, and many others. Because
these terms have no descriptive basis, they are not particularly useful
to a student of anatomy.
Nineteenth Century
FIGURE 1.16 In the early seventeenth century, the English physician William Harvey demonstrated that blood circulates and does not
flow back and forth through the same vessels.
fertilization. Rather, he thought that a spermatozoan contained a

miniature human being called a homunculus.
The development of the microscope added an entirely new
dimension to anatomy and eventually led to explanations of
basic body functions. In addition, the improved microscope was
invaluable for understanding the etiologies of many diseases, and
thus for discovering cures for many of them. Although Leeuwenhoek improved the microscope, credit for its invention is usually
given to the Dutch spectacle maker, Zacharius Janssen. The first
scientific investigation using a microscope was performed by
Francisco Stelluti in 1625 on the structure of a bee.
Malpighi and Others

Marcello Malpighi (mal-pege) (162894), an Italian anatomist,
is sometimes referred to as the father of histology. He discovered
the capillary blood vessels that Harvey had postulated and described the pulmonary alveoli of lungs and the histological structure of the spleen and kidneys.
Many other individuals made significant contributions to
anatomy during this 200-year period. In 1672, the Dutch
anatomist Regnier de Graaf described the ovaries of the female
reproductive system, and in 1775 Lazzaro Spallanzani showed
that both ovum and sperm cell were necessary for conception.
Francis Glisson (15971677) described the liver, stomach, and
intestines, and suggested that nerve impulses cause the emptying
of the gallbladder. Thomas Wharton (161473) and Niels
Stensen (163886) separately contributed to knowledge of the
homunculus: L. diminutive form of homo, man
The major scientific contribution of the nineteenth century was

the formulation of the cell theory. It could be argued that this
theory was the most important breakthrough in the history of biology and medicine because all of the bodys functions were
eventually interpreted as the effects of cellular function.
The term cell was coined in 1665 by an English physician,
Robert Hooke, as he examined the structure of cork under his microscope in an attempt to explain its buoyancy. What Hooke actually observed were the rigid walls that surrounded the empty cavities
of the dead cells. The significance of cellular structure did not become apparent until approximately 150 years after Hookes work.
With improved microscopes, finer details were observed. In
1809, a French zoologist, Jean Lamarck, observed the jellylike
substance within a living cell and speculated that this material
was far more important than the outside structure of the cell. Fifteen years later, Ren H. Dutrochet described the differences between plant and animal cells.
Two German scientists, Matthias Schleiden and Theodor
Schwann, are credited with the biological principle referred to as
the cell theory. Schleiden, a botanist, suggested in 1838 that each
plant cell leads a double lifethat is, in some respects it behaves
as an independent organism, but at the same time it cooperates
with the other cells that form the whole plant. A year later,
Schwann, a zoologist, concluded that all organisms are composed
of cells that are essentially alike. Nineteen years later, the addition of another biological principle seemed to complete the explanation of cells. In 1858, the German pathologist Rudolf
Virchow wrote a book entitled Cell Pathology in which he proposed that cells can arise only from preexisting cells. The mechanism of cellular replication, however, was not understood for
several more decades.
Johannes Mller (180158), a comparative anatomist, is
noted for applying the sciences of physics, chemistry, and psychology to the study of the human body. When he began his
teaching career, science was sufficiently undeveloped to allow
him to handle numerous disciplines at once. By the time of his
death, however, knowledge had grown so dramatically that several professors were needed to fill the positions he had held alone.
Twentieth Century
Contributions to the science of anatomy during the twentieth
century have not been as astounding as they were when little was
known about the structure of the body. The study of anatomy
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Like the people of Mesopotamia and Greece, the ancient

Egyptians were concerned with a controlling spirit of the
body. In fact, they even had a name for this life forcethe Ba spirit
and they believed that it was associated with the bowels and the
heart. Food was placed in the tomb of a mummy to feed the Ba spirit
during the journey to Osiris, Egyptian god of the underworld.
China and Japan

China
In ancient China, interest in the human body was primarily
philosophical. Ideas about anatomy were based on reasoning
rather than dissection or direct observation. The Chinese
revered the body and abhorred its mutilation. An apparent exception was the practice of binding the feet of young girls and
women in an attempt to enhance their beauty. Knowledge of the
internal organs came only from wounds and injuries. Only in recent times have dissections of cadavers been permitted in Chinese medical schools.
The ancient Chinese had an abiding belief that everything
in the universe depended on the balance of the two opposing
cosmic principles of yin and yang. As for the circulatory system,
the blood was the conveyor of the yang, and the heart and vessels represented the yin. Other structures of the body were composed of lesser forces termed zo and fu.
The Chinese were great herbalists. Writings more than
5,000 years old describe various herbal concoctions and potions
to alleviate a wide variety of ailments, including diarrhea, constipation, and menstrual discomfort. Opium was described as an excellent painkiller.
Until recently, the Chinese have been possessive of their
beliefs, and for this reason Western cultures were not influenced
by Chinese thoughts or writings to an appreciable extent. Perhaps the best known but least understood of the Chinese contributions to human anatomy and medicine is acupuncture.
Acupuncture is an ancient practice that was established to
maintain a balance between the yin and the yang. Three hundred
sixty-five precise meridian sites, or vital points, corresponding to
the number of days in a year, were identified on the body (fig. 1.7).
Needles inserted into the various sites were believed to release
harmful secretions and rid the tissues of obstructions. Acupuncture
is still practiced in China and has gained acceptance with some
medical specialists in the United States and other countries as a
technique of anesthesia and as a treatment for certain ailments.
The painkilling effect of acupuncture has been documented and is
more than psychological. Acupuncture sites have been identified
on domestic animals and have been used to a limited degree in
veterinary medicine. Why acupuncture is effective remains a mystery, although it has recently been correlated with endorphin production within the brain (see chapter 11).
Japan
The advancement of anatomy in Japan was strongly influenced
by the Chinese and Dutch. The earliest records of anatomical
interest in Japan date back to the sixth century. Buddhist
monks from Japan were trained in China where they were exposed to Chinese philosophy, and so Chinese beliefs concerning the body became prevalent in Japan as well. By the
eighteenth century, Western influences, especially the Dutch,
were such that the Japanese sought to determine for themselves
which version of anatomy was correct. In 1774, a five-volume
work called Kaitai Shinsho (A New Book of Anatomy), published
by a Japanese physician, Genpaku Sugita, totally adopted the
Dutch conceptualization of the body. This book marked the beginning of a modern era in anatomy and medicine for the
Japanese people.
For several hundred years, Western nations were welcome in
Japan. In 1603, however, the Japanese government banned
all contact with the Western world because they feared the influences
of Christianity on their society. Although this ban was strictly enforced
and Japan became isolated, Japanese scholars continued to circulate Western books on anatomy and medicine. These books eventually prompted Japanese physicians to reassess what they had been
taught concerning the structure of the body.
Grecian Period
It was in ancient Greece that anatomy first gained wide acceptance as a science. The writings of several Greek philosophers
had a tremendous impact on future scientific thinking. During
this period, the Greeks were obsessed with the physical beauty of
the human body, as reflected by their exquisite sculptures.
The young people of Greece were urged to be athletic and
develop their physical abilities, but at about age 18 they were directed more to intellectual pursuits of science, rhetoric, and philosophy. An educated individual was expected to be acquainted
with all fields of knowledge, and it was only natural that great
strides were made in the sciences.
Perhaps the first written reference to the anatomy of
wounds sustained in battle is contained in the Iliad, written by
Homer in about 800 B.C. Homers detailed descriptions of the
anatomy of wounds were exceedingly accurate. However, he described clean woundsnot the type of traumatic wounds that
would likely be suffered on a battlefield. This has led to speculation that human dissections were conducted during this period
and that anatomical structure was well understood. Victims of
human sacrifice may have served as subjects for anatomical study
and demonstration.
Hippocrates
Hippocrates (460377 B.C.), the most famous of the Greek physicians of his time, is regarded as the father of medicine because of
the sound principles of medical practice that his school established (fig. 1.8). His name is memorialized in the Hippocratic
oath, which many graduating medical students repeat as a
promise of professional stewardship and duty to humankind.
Hippocrates probably had only limited exposure to human
dissections, but he was well disciplined in the popular humoral
theory of body organization. Four body humors were recognized,
humor: L. humor, fluid

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(a)
History of Anatomy
(b)
FIGURE 1.7 Acupuncture has long been practiced for diagnostic or therapeutic purposes. Acupressure is application of finger-point pressure at
specific meridian sites to manage pain. (a) An acupuncture chart from the Ming dynasty of ancient China and (b) a patient receiving acupuncture.
and each was associated with a particular body organ: blood with
the liver; choler, or yellow bile, with the gallbladder; phlegm
with the lungs; and melancholy, or black bile, with the spleen. A
healthy person was thought to have a balance of the four humors. The concept of humors has long since been discarded, but
it dominated medical thought for over 2,000 years.
Perhaps the greatest contribution of Hippocrates was that
he attributed diseases to natural causes rather than to the displeasure of the gods. His application of logic and reason to medicine was the beginning of observational medicine.
The four humors are a part of our language and medical
practice even today. Melancholy is a term used to describe
depression or despondency in a person, whereas melanous refers to
a black or sallow complexion. The prefix melano- means black.
Cholera is an infectious intestinal disease that causes diarrhea and

vomiting. Phlegm (pronounced flem) within the upper respiratory system is symptomatic of several pulmonary disorders. Sanguine, a term
that originally referred to blood, is used to describe a passionate
temperament. This term, however, has evolved to refer simply to the
cheerfulness and optimism that accompanied a sanguine personality, and no longer refers directly to the humoral theory.
Aristotle
Aristotle (384322 B.C.), a pupil of Plato, was an accomplished
writer, philosopher, and zoologist (fig. 1.9). He was also a renowned
teacher and was hired by King Philip of Macedonia to tutor his son,
Alexander, who later became known as Alexander the Great.
cholera: Gk. chole, bile

melancholy: Gk. melan, black; chole, bile
phlegm: Gk. phlegm, inflammation

sanguine: L. sanguis, bloody
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The Hippocratic Oath

I swear by Apollo Physician and Aesculapius and Hygeia and Panacea
and all the gods and goddesses making them my witnesses, that I will
fulfill according to my ability and judgment this oath and this covenant:
To hold him who has taught me this art as equal to my parents and to
live my life in partnership with him, and if he is in need of money to give
him a share of mine, and to regard his offspring as equal to my brothers
in male lineage and to teach them this artif they desire to learn it
without fee and covenant; to give a share of precepts and oral instruction
and all the other learning to my sons and to the sons of him who has
instructed me and to pupils who have signed the covenant and have
taken an oath according to the medical law, but to no one else.
I will apply dietetic measures for the benefit of the sick according to
my ability and judgment; I will keep them from harm and injustice.
I will neither give a deadly drug to anybody if asked for it, nor will I make
a suggestion to this effect. Similarly I will not give to a woman an abortive
remedy. In purity and holiness I will guard my life and my art.
I will not use the knife, not even on sufferers from stone, but will
withdraw in favor of such men as are engaged in this work.
Whatever houses I may visit, I will come for the benefit of the sick,
remaining free of all intentional injustice, of all mischief, and in particular
of sexual relations with both female and male persons, be they free or
slaves.
What I may see or hear in the course of the treatment or even outside of
the treatment in regard to the life of men, which on no account one must
spread abroad, I will keep to myself, holding such things shameful to be
spoken about.
If I fulfill this oath and do not violate it, may it be granted to me to
enjoy life and art, being honored with fame among all men for all time
to come; if I transgress it and swear falsely, may the opposite of all this
be my lot.
FIGURE 1.8 A fourteenth-century painting of the famous Greek physician Hippocrates. Hippocrates is referred to as the father of medicine;
his creed is immortalized as the Hippocratic oath (left).
Aristotle made careful investigations of all kinds of animals, which included references to humans, and he pursued a
limited type of scientific method in obtaining data. He wrote the
first known account of embryology, in which he described the
development of the heart in a chick embryo. He named the aorta
and contrasted the arteries and veins. Aristotles best known zoological works are History of Animals, On the Parts of Animals, and
On the Generation of Animals (see table 1.2). These books had a
profound influence on the establishment of specialties within
anatomy, and they earned Aristotle recognition as founder of
comparative anatomy.
In spite of his tremendous accomplishments, Aristotle perpetuated some erroneous theories regarding anatomy. For example, the doctrine of the humors formed the boundaries of his
thought. Plato had described the brain as the seat of feeling and
thought, but Aristotle disagreed. He placed the seat of intelligence in the heart and argued that the function of the brain,
which was bathed in fluid, was to cool the blood that was
pumped from the heart, thereby maintaining body temperature.
Aesculapius: Gk. (mythology) son of Apollo and god of medicine

Hygeia: Gk. (mythology) daughter of Aesculapius; personification of health;
hygies, healthy
Panacea: Gk. (mythology) also a daughter of Aesculapius; assisted in temple rites
and tended sacred serpents; pan, all, every; akos, remedy
Alexandrian Era
Alexander the Great founded Alexandria in 332 B.C. and established it as the capital of Egypt and a center of learning. In addition to a great library, there was also a school of medicine in
Alexandria. The study of anatomy flourished because of the acceptance of dissections of human cadavers and human vivisections (viv-sekshunz) (dissections of living things). This brutal
procedure was commonly performed on condemned criminals.
People reasoned that to best understand the functions of the
body, it should be studied while the subject was alive, and that a
condemned man could best repay society through the use of his
body for a scientific vivisection.
Unfortunately, the scholarly contributions and scientific
momentum of Alexandria did not endure. Most of the written
works were destroyed when the great library was burned by the
Romans as they conquered the city in 30 B.C. What is known
about Alexandria was obtained from the writings of later scientists, philosophers, and historians, including Pliny, Celsus,
Galen, and Tertullian. Two men of Alexandria, Herophilus and
Erasistratus, made lasting contributions to the study of anatomy.
vivisection: L. vivus, alive; sectio, a cutting

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rate, he also had notions that were primitive and mystical. He

thought, for example, that the cranial nerves carried animal spirits and that muscles contracted because of distention by spirits.
He also believed that the left ventricle of the heart was filled
with a vital air spirit (pneuma) that came in from the lungs, and
that the arteries transported this pneuma rather than blood.
Both Herophilus and Erasistratus were greatly criticized later
in history for the vivisections they performed. Celsus (about
30 B.C.) and Tertullian (about A.D. 200) were particularly critical of the
practice of vivisection. Herophilus was described as a butcher of
men who had dissected as many as 600 living human beings, sometimes in public demonstrations.
Roman Era
FIGURE 1.9 This Roman copy of a Greek sculpture is believed to

be of Aristotle, the famous Greek philosopher.
Herophilus
Herophilus (about 325 B . C .) was trained in the Hippocratic
school but became a great teacher of anatomy in Alexandria.
Through vivisections and dissections of human cadavers, he provided excellent descriptions of the skull, eye, various visceral organs and organ relationships, and the functional relationship of
the spinal cord to the brain. Two monumental works by
Herophilus are entitled On Anatomy and Of the Eyes. He regarded the brain as the seat of intelligence and described many of
its structures, such as the meninges, cerebrum, cerebellum, and
fourth ventricle. He was also the first to distinguish nerves as either sensory or motor.
Erasistratus
In many respects, the Roman Empire stifled scientific advancements and set the stage for the Dark Ages. The approach to science shifted from theoretical to practical during this time. Few
dissections of cadavers were performed other than in attempts to
determine the cause of death in criminal cases. Medicine was not
preventive but was limited, almost without exception, to the
treatment of soldiers injured in battle. Later in Roman history,
laws were established that attested to the influence of the
Church on medical practice. According to Roman law, for example, no deceased pregnant woman could be buried without prior
removal of the fetus from the womb so that it could be baptized.
The scientific documents that have been preserved from
the Roman Empire are mostly compilations of information obtained from the Greek and Egyptian scholars. New anatomical
information was scant, and for the most part was derived from
dissections of animals other than human. Two important
anatomists from the Roman era were Celsus and Galen.
Celsus
Most of what is currently known about the Alexandrian school
of medicine is based on the writings of the Roman encyclopedist
Cornelius Celsus (30 B.C.A.D. 30). He compiled this information into an eight-volume work called De re medicina. Celsus had
only limited influence in his own time, however, probably because of his use of Latin rather than Greek. It was not until the
Renaissance that the enormous value of his contribution was
recognized.
Galen
Erasistratus (about 300 B.C.) was more interested in body functions than structure and is frequently referred to as the father of
physiology. In a book on the causes of diseases, he included observations on the heart, vessels, brain, and cranial nerves. Erasistratus noted the toxic effects of snake venom on various visceral
organs and described changes in the liver resulting from various
diseases. Although some of his writings were scientifically accu-
Claudius Galen (A.D. 130201) was perhaps the best physician

since Hippocrates. A Greek living under Roman domination, he
was certainly the most influential writer of all times on medical
subjects. For nearly 1,500 years, the writings of Galen represented the ultimate authority on anatomy and medical treatment. Galen probably dissected no more than two or three
human cadavers during his career, of necessity limiting his
anatomical descriptions to nonhuman animal dissections. He
visceral: L. viscus, internal organ
pneuma: Gk. pneuma, air
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Middle Ages
The Middle Ages, frequently referred to as the Dark Ages,
began with the fall of Rome to the Goths in A.D. 476 and lasted
nearly 1,000 years, until Constantinople was conquered by the
Turks in 1453. The totalitarian Christian Church suppressed
science and medical activity stagnated. Human existence continued to be miserably precarious, and people no longer felt capable of learning from personal observation. Rather, they
accepted life on faith.
Dissections of cadavers were totally prohibited during this
period, and molesting a corpse was a criminal act frequently punishable by burning at the stake. If mysterious deaths occurred, examinations by inspection and palpation were acceptable. During
the plague epidemic in the sixth century, however, a few necropsies (nekrop-sez) and dissections were performed in the hope of
determining the cause of this dreaded disease (see fig. 1.4).
FIGURE 1.10 These surgical and gynecological instruments were

found in the House of the Surgeon (about A.D. 6779) in Pompeii.
They are representative of the medical equipment used throughout
the Roman Empire during this time.
During the Crusades soldiers cooked the bones of their dead

comrades so that they could be returned home for proper
burial. Even this act, however, was eventually considered sacrilegious and strongly condemned by the Church. One of the ironies of
the Middle Ages was that the peasants received far less respect and
had fewer rights when they were alive than when they were dead.
Contributions of Islam
compiled nearly 500 medical papers (of which 83 have been preserved) from earlier works of others, as well as from his personal
studies. He perpetuated the concept of the humors of the body
and gave authoritative explanations for nearly all body functions.
Galens works contain many errors, primarily because of
his desire to draw definitive conclusions regarding human body
functions on the basis of data obtained largely from animals
such as monkeys, pigs, and dogs. He did, however, provide some
astute and accurate anatomical details in what are still regarded
as classic studies. He proved to be an experimentalist, demonstrating that the heart of a pig would continue to beat when
spinal nerves were transected so that nerve impulses could not
reach the heart. He showed that the squealing of a pig stopped
when the recurrent laryngeal nerves that innervated its vocal
cords were cut. Galen also tied off the ureter in a sheep to prove
that urine was produced in the kidney, not in the urinary bladder as had been falsely assumed. In addition, he proved that the
arteries contained blood rather than pneuma.
Galen compiled a list of many medicinal plants and used
medications extensively to treat illnesses. Although he frequently used bloodletting in an effort to balance the four humors, he cautioned against removing too much blood. He
accumulated a wide variety of medical instruments and suggested
their use as forceps, retractors, scissors, and splints (fig. 1.10). He
was also a strong advocate of helping nature heal through good
hygiene, a proper diet, rest, and exercise.
The Arabic-speaking people made a profound contribution to

the history of anatomy in a most unusual way. It was the Islamic
world that saved much of Western scholarship from the ruins of
the Roman Empire, the oppression of the Christian Church, and
the onset of the Middle Ages. With the expansion of Islam
through the Middle East and North Africa during the eighth
century, the surviving manuscripts from Alexandria were taken
back to the Arab countries, where they were translated from
Greek to Arabic.
As the Dark Ages enshrouded Europe, the Christian
Church attempted to stifle any scholarship or worldly knowledge
that was not acceptable within Christian dogma. The study of
the human body was considered heretical, and the Church
banned all writings on anatomical subjects. Without the Islamic
repository of the writings of Aristotle, Hippocrates, Galen, and
others, the progress of centuries in anatomy and medicine would
have been lost. It wasnt until the thirteenth century that the
Arabic translations were returned to Europe and, in turn, translated to Latin. During the translation process, any Arabic terminology that had been introduced was systematically removed, so
that today we find few anatomical terms of Arabic origin.
epidemic: Gk. epi, upon; demos, people

necropsy: Gk. nekros, corpse; opsy, view

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Renaissance
The period known as the Renaissance was characterized by a rebirth of science. It lasted roughly from the fourteenth through
the sixteenth centuries and was a transitional period from the
Middle Ages to the modern age of science.
The Renaissance was ushered in by the great European
universities established in Bologna, Salerno, Padua, Montpellier, and Paris. The first recorded human dissections at these
newly established centers of learning were the work of the surgeon William of Saliceto (121580) from the University of
Bologna. The study of anatomy quickly spread to other universities, and by the year 1300 human dissections had become an integral part of the medical curriculum. However, the Galenic
dogma that normal human anatomy was sufficiently understood
persisted, so interest at this time centered on methods and techniques of dissection rather than on furthering knowledge of the
human body.
The development of movable type in about 1450 revolutionized the production of books. Celsus, whose De re medicina
was rediscovered during the Renaissance, was the first medical
author to be published in this manner. Among the first anatomy
books to be printed in movable type was that of Jacopo Berengario of Carpi, a professor of surgery at Bologna. He described
many anatomical structures, including the appendix, thymus,
and larynx. The most influential text of the period was written
by Mondino de Luzzi, also of the University of Bologna, in 1316.
First published in 1487, it was more a dissection guide than a
study of gross anatomy, and in spite of its numerous Galenic errors, 40 editions were published, until the time of Vesalius.
Because of the rapid putrefaction of an unembalmed corpse,
the anatomy textbooks of the early Renaissance were organized so that the more perishable portions of the body were considered first. Dissections began with the abdominal cavity, then the
chest, followed by the head, and finally the appendages. A dissection was a marathon event, frequently continuing for perhaps 4 days.
With the increased interest in anatomy during the Renaissance, obtaining cadavers for dissection became a serious problem. Medical students regularly practiced grave robbing until
finally an official decree was issued that permitted the bodies of
executed criminals to be used as specimens.
Corpses were embalmed to prevent deterioration, but this
was not especially effective, and the stench from cadavers
was apparently a persistent problem. Anatomy professors lectured
from a thronelike chair at some distance from the immediate area
(fig. 1.11). The phrase, I wouldnt touch that with a 10-foot pole
probably originated during this time in reference to the smell of a decomposing cadaver.
The major advancements in anatomy that occurred during

the Renaissance were in large part due to the artistic and scientific abilities of Leonardo da Vinci and Andreas Vesalius. Working in the fifteenth and sixteenth centuries, each produced
monumental studies of the human form.
FIGURE 1.11 The scene of a cadaver dissection, 1500, from Fasciculus Medicinae by Johannes de Ketham. The anatomy professor
removed himself from the immediate area to a thronelike chair overlooking the proceedings. The dissections were performed by hired
assistants. One of them, the ostensor, pointed to the internal structures with a wand as the professor lectured.
Leonardo
The great Renaissance Italian Leonardo da Vinci (14521519) is
best known for his artistic works (e.g., Mona Lisa) and his scientific contributions. He displayed genius as a painter, sculptor, architect, musician, and anatomistalthough his anatomical
drawings were not published until the end of the nineteenth century. As a young man, Leonardo regularly participated in cadaver
dissections and intended to publish a textbook on anatomy with
the Pavian professor Marcantonio della Torre. The untimely
death of della Torre at the age of 31 halted their plans. When
Leonardo died, his notes and sketches were lost and were not discovered for more than 200 years. The advancement of anatomy
would have been accelerated by many years if Leonardos notebooks had been available to the world at the time of his death.
Leonardos illustrations helped to create a new climate of
visual attentiveness to the structure of the human body. He was
intent on accuracy, and his sketches are incredibly detailed
(fig. 1.12). He experimentally determined the structure of complex body organs such as the brain and the heart. He made wax
casts of the ventricles of the brain to study its structure. He constructed models of the heart valves to demonstrate their action.
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FIGURE 1.13 A painting of the great anatomist Andreas Vesalius,

as he dissects a cadaver. From his masterpiece De Humani Corporis
Fabrica.
FIGURE 1.12 Only a master artist could achieve the detail and
accuracy of the anatomical sketches of Leonardo da Vinci. The
painter who has acquired a knowledge of the nature of the sinews,
muscles and tendons, Leonardo wrote, will know exactly in the
movement of any limb how many and which of the sinews are the
cause of it, and which muscle by its swelling is the cause of the
sinews contracting.
Vesalius
The contribution of Andreas Vesalius (151464) to the science
of human anatomy and to modern medicine is immeasurable.
Vesalius was born in Brussels into a family of physicians. He received his early medical training at the University of Paris and
completed his studies at the University of Padua in Italy, where
he began teaching surgery and anatomy immediately after graduation. At Padua, Vesalius participated in human dissections and
initiated the use of live models to determine surface landmarks
for internal structures (fig. 1.13).
Vesalius apparently had enormous energy and ambition. By
the time he was 28 years old, he had already completed the masterpiece of his life, De Humani Corporis Fabrica, in which the various body systems and individual organs are beautifully illustrated
and described (fig. 1.14). His book was especially important in
that it boldly challenged hundreds of Galens teachings. Vesalius
wrote of his surprise upon finding numerous anatomical errors in
the works of Galen that were taught as fact, and he refused to accept Galens explanations on faith. Because he was so outspo-
FIGURE 1.14 A plate from De Humani Corporis Fabrica, which

Vesalius completed at the age of 28. This book, published in 1543,
revolutionized the science of anatomy.

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FIGURE 1.15 The Anatomy Lesson of Dr. Tulp, a famous Rembrandt painting completed in 1632, depicts one of the public anatomies that
were popular during this period. Dr. Nicholas Tulp was a famous Dutch anatomist who described the congenital defect in the spinal column
known as spinal bifida aperta.
kenly opposed to Galen, he incurred the wrath of many of the traditional anatomists, including his former teacher Sylvius (Jacques
Dubois) of Paris. Sylvius even went so far as to give him the nickname Vesanus (madman). Vesalius became so unnerved by the
relentless attacks that he destroyed much of his unpublished work
and ceased his dissections.
Fortunately there were also serious, scientific-minded

anatomists who made significant contributions during this period. Two of the most important contributions were the explanation of blood flow and the development of the microscope.
Although Vesalius was the greatest anatomist of his epoch,

others made significant contributions and to an extent paved
the way for Vesalius. Michelangelo pursued anatomy in 1495, being
supplied with corpses by the friar of a local monastery. Mondino de
Luzzi and the surgeon Jacopo Berengario of Carpi also corrected
many of Galens errors. Fallopius (152362) and Eustachius
(152474) completed detailed dissections of specific body regions.
In 1628, the English physician William Harvey (15781657)

published his pioneering work On the Motion of the Heart and
Blood in Animals. Not only did this brilliant research establish
proof of the continuous circulation of blood within contained
vessels, it also provided a classic example of the scientific
method of investigation (fig. 1.16). Like Vesalius, Harvey was
severely criticized for his departure from Galenic philosophy.
The controversy over circulation of the blood raged for 20 years,
until other anatomists finally repeated Harveys experiments and
concurred with his findings.
Seventeenth and Eighteenth Centuries

During the seventeenth and eighteenth centuries, the science of
anatomy attained an unparalleled acceptance. In some of its aspects, it also took on a somewhat theatrical quality. Elaborate
amphitheaters were established in various parts of Europe for
public demonstrations of human dissections (fig. 1.15). Exorbitantly priced tickets of admission were sold to the wealthy, and
the dissections were performed by elegantly robed anatomists
who were also splendid orators. The subjects were usually executed prisoners, and the performances were scheduled during
cold weather because of the perishable nature of the cadavers.
Harvey
Leeuwenhoek
Antoni van Leeuwenhoek (laven-hook) (16321723) was a
Dutch optician and lens grinder who improved the microscope
to the extent that he achieved a magnification of 270 times. His
many contributions included developing techniques for examining tissues and describing blood cells, skeletal muscle, and
the lens of the eye. Although he was the first to accurately describe sperm cells, Leeuwenhoek did not understand their role in
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salivary glands and lymph nodes within the neck and facial regions. In 1664, Thomas Willis published a summary of what was
then known about the nervous system.
A number of anatomical structures throughout the body are
named in honor of the early anatomists. Thus we have
graafian follicles, Stensens and Whartons ducts, fallopian tubes,
Bartholins glands, the circle of Willis, and many others. Because
these terms have no descriptive basis, they are not particularly useful
to a student of anatomy.
Nineteenth Century
FIGURE 1.16 In the early seventeenth century, the English physician William Harvey demonstrated that blood circulates and does not
flow back and forth through the same vessels.
fertilization. Rather, he thought that a spermatozoan contained a

miniature human being called a homunculus.
The development of the microscope added an entirely new
dimension to anatomy and eventually led to explanations of
basic body functions. In addition, the improved microscope was
invaluable for understanding the etiologies of many diseases, and
thus for discovering cures for many of them. Although Leeuwenhoek improved the microscope, credit for its invention is usually
given to the Dutch spectacle maker, Zacharius Janssen. The first
scientific investigation using a microscope was performed by
Francisco Stelluti in 1625 on the structure of a bee.
Malpighi and Others

Marcello Malpighi (mal-pege) (162894), an Italian anatomist,
is sometimes referred to as the father of histology. He discovered
the capillary blood vessels that Harvey had postulated and described the pulmonary alveoli of lungs and the histological structure of the spleen and kidneys.
Many other individuals made significant contributions to
anatomy during this 200-year period. In 1672, the Dutch
anatomist Regnier de Graaf described the ovaries of the female
reproductive system, and in 1775 Lazzaro Spallanzani showed
that both ovum and sperm cell were necessary for conception.
Francis Glisson (15971677) described the liver, stomach, and
intestines, and suggested that nerve impulses cause the emptying
of the gallbladder. Thomas Wharton (161473) and Niels
Stensen (163886) separately contributed to knowledge of the
homunculus: L. diminutive form of homo, man
The major scientific contribution of the nineteenth century was

the formulation of the cell theory. It could be argued that this
theory was the most important breakthrough in the history of biology and medicine because all of the bodys functions were
eventually interpreted as the effects of cellular function.
The term cell was coined in 1665 by an English physician,
Robert Hooke, as he examined the structure of cork under his microscope in an attempt to explain its buoyancy. What Hooke actually observed were the rigid walls that surrounded the empty cavities
of the dead cells. The significance of cellular structure did not become apparent until approximately 150 years after Hookes work.
With improved microscopes, finer details were observed. In
1809, a French zoologist, Jean Lamarck, observed the jellylike
substance within a living cell and speculated that this material
was far more important than the outside structure of the cell. Fifteen years later, Ren H. Dutrochet described the differences between plant and animal cells.
Two German scientists, Matthias Schleiden and Theodor
Schwann, are credited with the biological principle referred to as
the cell theory. Schleiden, a botanist, suggested in 1838 that each
plant cell leads a double lifethat is, in some respects it behaves
as an independent organism, but at the same time it cooperates
with the other cells that form the whole plant. A year later,
Schwann, a zoologist, concluded that all organisms are composed
of cells that are essentially alike. Nineteen years later, the addition of another biological principle seemed to complete the explanation of cells. In 1858, the German pathologist Rudolf
Virchow wrote a book entitled Cell Pathology in which he proposed that cells can arise only from preexisting cells. The mechanism of cellular replication, however, was not understood for
several more decades.
Johannes Mller (180158), a comparative anatomist, is
noted for applying the sciences of physics, chemistry, and psychology to the study of the human body. When he began his
teaching career, science was sufficiently undeveloped to allow
him to handle numerous disciplines at once. By the time of his
death, however, knowledge had grown so dramatically that several professors were needed to fill the positions he had held alone.
Twentieth Century
Contributions to the science of anatomy during the twentieth
century have not been as astounding as they were when little was
known about the structure of the body. The study of anatomy

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(c)
FIGURE 1.17 Different techniques for viewing microscopic anatomy have greatly enhanced our understanding of the structure and function of
the human body. (a) The appearance of hair under a simple magnifying glass, (b) an observation of a stained section of a hair through a light microscope, and (c) a hair emerging from skin as viewed through an electron microscope (340at 35-mm size).
grew increasingly specialized, and research became more detailed

and complex.
One innovation that gained momentum early in the twentieth century was the simplification and standardization of
nomenclature. Because of the proliferation of scientific literature
toward the end of the nineteenth century, over 30,000 terms for
structures in the human body were on record, many of which
were redundant. In 1895, in an attempt to reduce the confusion,
the German Anatomical Society compiled a list of approximately 500 terms called the Basle Nomina Anatomica (BNA).
The terms on this list were universally approved for use in the
classroom and in publications.
Other conferences on nomenclature have been held
throughout the century, under the banner of the International
Congress of Anatomists. At the Seventh International Congress
held in New York City in 1960, a resolution was passed to eliminate all eponyms (tombstone names) from anatomical terminology and instead use descriptive names. Structures like Stensens
duct and Whartons duct, for example, are now properly referred
to as the parotid duct and submandibular duct, respectively. Because eponyms are so entrenched, however, it will be extremely
difficult to eliminate all of them from anatomical terminology.
But at least there is a trend toward descriptive simplification.
In this text, the preferred descriptive terms are used in
both the text narrative and the accompanying illustrations.
Where the term first appears in the narrative, however, the preferred form is followed by a parenthetical reference to the traditional name honoring an individualfor example, uterine
(fallopian) tube. The terminology used in this text is in accordance with the official anatomical nomenclature presented in
the reference publication, Nomina Anatomica, Sixth Edition.
In response to the increased technology and depth of understanding in the twentieth century, new disciplines and specialties have appeared in the science of human anatomy in an
attempt to categorize and use the new knowledge. The techniques of such cognate disciplines as chemistry, physics, electronics, mathematics, and computer science have been incorporated
into research efforts.
There are several well-established divisions of human
anatomy. The oldest, of course, is gross anatomy, which is the
study of body structures that can be observed with the unaided eye. Stringent courses in gross anatomy in professional
schools provide the foundation for a students entire medical
or paramedical training. Gross anatomy also forms the basis
for the other specialties within anatomy. Surface anatomy
(see chapter 10) deals with surface features of the body that
can be observed beneath the skin or palpated (examined by
touch).
Microscopic Anatomy
Structures smaller than 0.1 mm (100 m) can be seen only
with the aid of a microscope. The sciences of cytology (the
study of cells), or cellular biology, developmental anatomy
(the study of prenatal development) and histology (the study of
tissues) are specialties of anatomy that have provided additional insight into structure and function of the human body.
One can observe greater detail with the electron microscope than
with the light microscope (fig. 1.17). New techniques in staining
and histochemistry have aided electron microscopy by revealing the fine details of cells and tissues that are said to compose
their ultrastructure.
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CHAPTER 1

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(b)
(c)
FIGURE 1.18 The versatility of radiology makes this technique one of the most important tools in diagnostic medicine and provides a unique
way of observing specific anatomical structures within the body. (a) A radiograph of a healing fracture, (b) a radiograph of gallstones within a
gallbladder, and (c) a radiograph of a stomach filled with a radiopaque contrast medium.
Radiographic Anatomy
Radiographic anatomy, or radiology, provides a way of observing
structures within the living body. Radiology is based on the principle that substances of different densities absorb different
amounts of X rays, resulting in a differential exposure on film.
Radiopaque substances such as barium can be ingested (swallowed) or injected into the body to produce even greater contrasts (fig. 1.18). Angiography involves making a radiograph
after injecting a dye into the bloodstream. In angiocardiography,
the heart and its associated vessels are x-rayed. Cineradiography
permits the study of certain body systems through the use of motion picture radiographs. Traditional radiographs have had limitations as diagnostic tools for understanding human anatomy
because of the two-dimensional plane that is photographed. Because radiographs compress the body image with an overlap of
organs and tissues, diagnosis is often difficult.
X rays were discovered in 1895 by Wilhelm Konrad Roentgen
(rentgen). The radiograph image that is produced on film is
frequently referred to as a roentgenograph. The recent development
of the computerized axial tomography technique has been hailed as
the greatest advancement in diagnostic medicine since the discovery of X rays themselves.
The computerized axial tomography technique (CT, or

CAT, scan) has greatly enhanced the versatility of X rays. It uses
a computer to display a cross-sectional image similar to that
which could only be obtained in an actual section through the
body (fig. 1.19a).
Another technique of radiographic anatomy is the dynamic spatial reconstructor (DSR) scan (fig. 1.19b). The DSR
functions as an electronic knife that pictorially slices an organ,
such as the heart, to provide three-dimensional images. The
DSR can be used to observe movements of organs, detect defects,

assess the extent of a disease such as cancer, or determine the extent of trauma to tissues after a stroke or heart attack.
Magnetic resonance imaging (MRI), also called nuclear
magnetic resonance (NMR), provides a new technique for diagnosing diseases and following the response of a disease to chemical treatment (fig. 1.19c). An MRI image is created rapidly as
hydrogen atoms in tissues, subjected to a strong magnetic field,
respond to a pulse of radio waves. MRI has the advantage of
being noninvasivethat is, no chemicals are introduced into the
body. It is better than a CT scan for distinguishing between soft
tissues, such as the gray and white matter of the nervous system.
A positron emission tomography (PET) scan is a radiological technique used to observe the metabolic activity in organs
(fig. 1.19d) following the injection of a radioactive substance,
such as treated glucose, into the bloodstream. PET scans are very
useful in revealing the extent of damaged heart tissue and in identifying areas where blood flow to the brain is blocked.
Human anatomy will always be a relevant science. Not
only does it enhance our personal understanding of body functioning, it is also essential in the clinical diagnosis and treatment
of disease. Human anatomy is no longer confined to the observation and description of structures in isolation, but has expanded
to include the complexities of how the body functions as an integrated whole. The science of anatomy is dynamic and has remained vital because the two aspects of the bodystructure and
functionare inseparable.
One of the important aspects of human anatomy and medicine
is the autopsya thorough postmortem examination of all of
the organs and tissues of a body. Autopsies were routinely performed in the early part of the twentieth century, but their frequency
has declined significantly in the last three decades. Currently, only

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History of Anatomy
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(b)
(d)
FIGURE 1.19 Different techniques in radiographic anatomy provide unique views of the human body. (a) A CT scan through the head,
(b) a DSR scan through the trunk (torso), (c) an MRI image through the head, and (d ) a PET scan through the head.
15% of corpses are autopsied in the United States, which is down

from over 50% 30 years ago. Autopsies are of value because (1) they
often determine the cause of death, which may confirm or disconfirm
preliminary death statements; (2) they frequently reveal diseases or
structural defects that may have gone undetected in life; (3) they
check the effectiveness of a particular drug therapy for a patient or
the success of a particular surgery; and (4) they serve as a means of
training medical students.
An interesting piece of information regarding the value of an
autopsy in confirming the cause of death was revealed in a study of
2,557 autopsies conducted over a 30-year period to determine
the accuracy of physicians diagnoses of deaths (see Autopsy,
S. A. Geller, Scientific American, March 1983). In this study, the
causes of death had been improperly or inaccurately recorded in
42% of the cases. This means that because of a decline in the num-
ber of autopsies, there may be over a million death certificates filed

in the United States each year that are in error. This has important
implications for criminal justice and medical genetics, as well as for
the insurance industry.
An objective of this text is to enable students to become

educated and conversant in anatomy. An excellent way to keep
up with anatomy during and after completing the formal course
is to subscribe to and read magazines such as Science, Scientific
American, Discover, and Science Digest. These publications and
others include articles on recent scientific findings, many of
which pertain to anatomy. If you are to be an educated contributor to society, it is essential to stay informed.
CHAPTER 1
Chapter 1
CHAPTER 1

20
Unit 1
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Knowledge Check
Clinical Case Study Answer
7. Briefly discuss the impact of each of the following on the

science of human anatomy: the humoral theory of body organization, vivisections, the Middle Ages, human dissections, movable type, the invention of the microscope, the
cell theory, and the development of X-ray techniques.
8. Explain why knowledge of anatomy is personally relevant.
9. Why is it important to stay abreast of new developments in
human anatomy? How might this be accomplished?
In this case, the doctor made the right diagnosis and provided a therapy
that helped the patient. However, he did so with a flawed understanding
of human anatomy and physiology and disease. The application of sound
scientific methods has corrected classic misconceptions of disease, such
as the humoral theory.
Chapter Summary
Definition of the Science (p. 2)
1. Human anatomy is the science concerned
with the structure of the human body.
2. The terms of anatomy are descriptive and
are generally of Greek or Latin derivation.
3. The history of human anatomy parallels
that of medicine and has also been greatly
influenced by various religions.
Prescientific Period (pp. 24)

1. Prehistoric interest in anatomy was
undoubtedly limited to practical
information necessary for survival.
2. Trepanation was a surgical technique that
was practiced by several cultures.
3. Paleopathology is the science concerned
with diseases of prehistoric people.
Scientific Period (pp. 419)

1. A few anatomical descriptions were
inscribed in clay tablets in cuneiform
writing by people who lived in
Mesopotamia in about 4000 B.C.
2. Egyptians of about 3400 B.C. developed a
technique of embalming. It was not
recorded, however, and therefore was not
of value in furthering the study of
anatomy.
3. The belief in a balance between yin and
yang was a compelling influence in
Chinese philosophy and provided the
rationale for the practice of acupuncture.
4. The advancement of anatomy in Japan
was largely due to the influence of the
Chinese and Dutch.
5. Anatomy first found wide acceptance as a

science in ancient Greece.
(a) Hippocrates is regarded as the father
of medicine because of the sound
principles of medical practice he
established.
(b) The Greek philosophy of body
humors dominated medical thought for
over 2,000 years.
(c) Aristotle pursued a limited type of
scientific method in obtaining data; his
writings contain some basic anatomy.
6. Alexandria was a center of scientific
learning from 300 to 30 B.C.
(a) Human dissections and vivisections
were performed in Alexandria.
(b) Erasistratus is referred to as the father
of physiology because of his
interpretations of various body functions.
7. Theoretical data was deemphasized during
the Roman era.
(a) Celsuss eight-volume work was a
compilation of medical data from the
Alexandrian school.
(b) Galen was an influential medical
writer who made some important advances
in anatomy; at the same time he
introduced serious errors into the literature
that went unchallenged for centuries.
(c) Science was suppressed for nearly
1,000 years during the Middle Ages, and
dissections of human cadavers were
prohibited.
(d) Anatomical writings were taken from
Alexandria by Arab armies, and thus
8.
9.
10.
11.
saved from destruction during the Dark

Ages in Europe.
During the Renaissance, many great
European universities were established.
(a) Andreas Vesalius and Leonardo da
Vinci were renowned Renaissance men
who produced monumental studies of the
human form.
(b) De Humani Corporis Fabrica, written by
Vesalius, had a tremendous impact on the
advancement of human anatomy. Vesalius
is regarded as the father of human anatomy.
Two major scientific contributions of the
seventeenth and eighteenth centuries
were the explanation of blood flow and
the development of the microscope.
(a) In 1628, William Harvey correctly
described the circulation of blood.
(b) Shortly after the microscope had
been perfected by Antoni van
Leeuwenhoek, many investigators added
new discoveries to the rapidly changing
specialty of microscopic anatomy.
The cell theory was formulated during the
nineteenth century by Matthias
Schleiden and Theodor Schwann, and
cellular biology became established as a
science separate from anatomy.
A trend toward simplification and
standardization of anatomical
nomenclature began in the twentieth
century. In addition, many specialties
within anatomy developed, including
cytology, histology, embryology, electron
microscopy, and radiology.

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History of Anatomy
21
Review Activities
Objective Questions
1. Anatomy is derived from a Greek word
meaning
(a) to cut up.
(c) functioning part.
(b) to analyze.
(d) to observe death.
2. The most important contribution of
William Harvey was his research on
(a) the continuous circulation of blood.
(b) the microscopic structure of
spermatozoa.
(c) the detailed structure of the kidney.
(d) the striped appearance of skeletal
muscle.
3. Which of the following listings is in
correct chronological order?
(a) Galen, Hippocrates, Harvey,
Vesalius, Aristotle
(b) Hippocrates, Galen, Vesalius,
Aristotle, Harvey
(c) Hippocrates, Aristotle, Galen,
Vesalius, Harvey
(d) Aristotle, Hippocrates, Galen,
Harvey, Vesalius
4. Anatomy was first widely accepted as a
science in ancient
(a) Rome.
(c) China.
(b) Egypt.
(d) Greece.
5. The establishment of sound principles of
medical practice earned this man the title
of father of medicine.
(a) Hippocrates (c) Erasistratus
(b) Aristotle
(d) Galen
6. Which of the four body humors was
believed by Hippocrates to be associated
with the lungs?
(a) black bile
(c) phlegm
(b) yellow bile
(d) blood
7. The anatomical masterpiece De Humani
Corporis Fabrica was the work of
(a) Leonardo.
(c) Vesalius.
(b) Harvey.
(d) Leeuwenhoek.
8. What event of about 1450 helped to
usher in the Renaissance?
(a) the development of the
microscope
(b) an acceptance of the scientific
method
(c) the development of the cell theory
(d) the development of movable type
9. The body organ thought by Aristotle to

be the seat of intelligence was
(a) the liver.
(c) the brain.
(b) the heart.
(d) the intestine.
10. X rays were discovered during the late
nineteenth century by
(a) Roentgen.
(c) Schleiden.
(b) Hooke.
(d) Mller.
Essay Questions
1. Define the terms anatomize, trepanation,
paleopathology, vivisection, and cadaver.
2. Discuss the practical nature of anatomy to
prehistoric people.
3. Why were the techniques of embalming a
corpse, which were perfected in ancient
Egypt, not shared with other cultures or
recorded for future generations?
4. What is acupuncture? What are some of
its uses today?
5. Why is Latin an ideal language from
which to derive anatomical terms? What
is the current trend regarding the use of
proper names (eponyms) in referring to
anatomical structures?
6. Why do you suppose the Hippocratic oath
has survived for over 2,000 years as a
creed for medical practice? What aspects
of the oath are difficult to conform to in
todays society?
7. What is meant by the humoral theory of
body organization? Which great
anatomists were influenced by this
theory? When did it cease to be an
influence on anatomical investigation and
interpretation?
8. Discuss the impact of Galen on the
advancement of anatomy and medicine.
What circumstances permitted the
philosophies of Galen to survive for such
a long period?
9. Briefly discuss the establishment of
anatomy as a science during the
Renaissance.
10. Herophilus popularized anatomy during
his time but was severely criticized by
later anatomists. Why was his work so
controversial?
11. Who invented the microscope? What part
did it play in the advancement of

anatomy? What specialties of anatomical

study have arisen since the introduction
of the microscope?
12. Discuss the impact of the work of Andreas
Vesalius on the science of anatomy.
13. Give some examples of how culture and
religion influenced the science of anatomy.
14. List some techniques currently used to
study anatomy and identify the specialties
within which these techniques are used.
Critical-Thinking Questions
1. Discuss some of the factors that
contributed to a decline in scientific
understanding during the Middle Ages.
How would you account for the
resurgence of interest in the science of
anatomy during the Renaissance?
2. Homeostasis is a physiological term that
was coined by the American physiologist
Walter Cannon in 1932. It refers to the
ability of an organism to maintain the
stability of its internal environment by
adjusting its physiological processes.
Discuss the similarities of the humoral
theory of body organization and the
philosophy of yin and yang as ancient
attempts to explain homeostasis.
3. You learned in this chapter that Galen
relied on dissections of animals other than
human in an attempt to understand
human anatomy. Discuss the value and
limitations of using mammalian
specimens (other than human) in the
laboratory portion of a human anatomy
course. What advantages are gained by
studying human cadavers?
4. Students studying law at European
Universities during the Early Renaissance
were required to take a course in human
anatomy. Considering the breadth of law
practice during those times, explain why
it was important for a lawyer to
understand anatomy.
5. Just as geography describes the
topography for history, anatomy describes
the topography for medicine. Using
specific examples, discuss how discoveries
in anatomy have resulted in advances in
medicine.
CHAPTER 1
Chapter 1

II. Terminology,
Organization, and the
Human Organism
2. Body Organization and

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Body Organization and

Classification and Characteristics of

Humans 23
Body Organization 28
Anatomical Nomenclature 30
Planes of Reference and Descriptive
Terminology 33
Body Regions 35
Body Cavities and Membranes 41
Chapter Summary 46
Clinical Case Study
FIGURE: Radiographic anatomy is

important in assessing trauma to bones and
visceral organs.
A young woman was hit by a car while crossing a street. Upon arrival at the scene, paramedics
found the patient to be a bit dazed but reasonably lucid, complaining of pain in her abdomen
and the left side of her chest. Otherwise, her vital signs were within normal limits. Initial evaluation in the emergency room revealed a very tender abdomen and left chest. The chest radiograph demonstrated a collapsed left lung resulting from air in the pleural space
(pneumothorax). The emergency room physician inserted a drainage tube into the left chest
(into the pleural space) to treat the pneumothorax. Attention was then turned to the abdomen. Because of the finding of tenderness, a peritoneal lavage was performed. This procedure
involves penetrating the abdominal wall and inserting a tube into the peritoneal cavity. Clear
fluid such as sterile water or normal saline is then instilled into the abdomen and siphoned out
again. The fluid used in this procedure is called lavage fluid. A return of lavage fluid containing
blood, fecal matter, or bile indicates injury to an abdominal organ that requires surgery. The return of lavage fluid from this patient was clear. However, the nurse stated that lavage fluid was
draining out of the chest tube.
From what you know about how the various body cavities are organized, do you suppose
this phenomenon could be explained based on normal anatomy? What might have caused it to
occur in our patient? Does the absence of bile, blood, etc., in the peritoneal lavage fluid guarantee that no organ has been ruptured? If it does not, explain why in terms of the relationship of
the various organs to the membranes within the abdomen.

II. Terminology,
Human Organism
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Chapter 2
CLASSIFICATION AND
CHARACTERISTICS OF HUMANS
Body Organization and Anatomical Nomenclature
23
Dorsal hollow
nerve cord
Notochord
Primitive eye
Objective 1
Gut
Classify humans according to the taxonomic
system.
Objective 2
List the characteristics that identify humans as

chordates and as mammals.
Objective 3
Describe the anatomical characteristics that set

humans apart from other primates.
The human organism, or Homo sapiens, as we have named ourselves, is unique in many ways. Our scientific name translates
from the Latin to man the intelligent, and indeed our intelligence is our most distinguishing feature. It has enabled us to
build civilizations, conquer dread diseases, and establish cultures.
We have invented a means of communicating through written
symbols. We record our own history, as well as that of other organisms, and speculate about our future. We continue to devise
ever more ingenious ways for adapting to our changing environment. At the same time, we are so intellectually specialized that
we are not self-sufficient. We need one another as much as we
need the recorded knowledge of the past.
We are constantly challenged to learn more about ourselves. As we continue to make new discoveries about our structure and function, our close relationship to other living organisms
becomes more and more apparent. Often, it is sobering to realize
our biological imperfections and limitations.
We share many characteristics with all living animals. As
human organisms, we breathe, eat and digest food, excrete bodily
wastes, locomote, and reproduce our own kind. We are subject to
disease, injury, pain, aging, mutations, and death. Because we are
composed of organic materials, we will decompose after death as
microorganisms consume our flesh as food. The processes by which
our bodies produce, store, and utilize energy are similar to those
used by all living organisms. The genetic code that regulates our
development is found throughout nature. The fundamental patterns of development of many nonhuman animals also characterize
the formation of the human embryo. Recent genetic mapping of
the human genome confirms that there are fewer than 35,000
genes accounting for all of our physical traits. By far the majority
of these genes are similar to those found in many other organisms.
In the classification, or taxonomic, system established by
biologists to organize the structural and evolutionary relationships of living organisms, each category of classification is re-
taxon: Gk. taxis, order
Pharynx
Pharyngeal
pouches
Limb
bud
Umbilical
cord
Limb bud
Creek
FIGURE 2.1 A schematic diagram of a chordate embryo. The

three diagnostic chordate characteristics are indicated in boldface
type.
ferred to as a taxon. The highest taxon is the kingdom and the

most specific taxon is the species. Humans are species belonging
to the animal kingdom. Phylogeny (fi-loje-ne) is the science that
studies relatedness on the basis of taxonomy.
Phylum Chordata
Human beings belong to the phylum Chordata (fi'lum kor-da'ta),
along with fishes, amphibians, reptiles, birds, and other mammals. All chordates have three structures in common: a notochord (no'to-kord), a dorsal hollow nerve cord, and pharyngeal
(fa-rin'je-al) pouches (fig. 2.1). These chordate characteristics are
well expressed during the embryonic period of development and,
to a certain extent, are present in an adult. The notochord is a
flexible rod of tissue that extends the length of the back of an
embryo. A portion of the notochord persists in the adult as
the nucleus pulposus, located within each intervertebral disc
(fig. 2.2). The dorsal hollow nerve cord is positioned above the
notochord and develops into the brain and spinal cord, which
are supremely functional as the central nervous system in the
adult. Pharyngeal pouches form gill openings in fishes and some
phylogeny: L. phylum, tribe; Gk. logos, study
CHAPTER 2
Humans are biological organisms belonging to the phylum Chordata within the kingdom Animalia and to the family Hominidae
within the class Mammalia and the order Primates.

Unit 2
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Terminology, Organization, and the Human Organism
CHAPTER 2
24
II. Terminology,
Human Organism
L4
L5
S1
(a)
(b)
FIGURE 2.2 (a) A lateral view of the vertebral column showing the intervertebral discs and, to the right, a superior view of an intervertebral
disc showing the nucleus pulposus and (b) a herniated disc (see arrow) compressing a spinal nerve.
amphibians. In other chordates, such as humans, embryonic pharyngeal pouches develop, but only one of the pouches persists,
becoming the middle-ear cavity. The auditory (eustachian) (yoosta'shun) tube, is a persisting connection between the middle-ear
cavity and the pharynx (far'ingks) (throat area).
The function of an intervertebral disc and its nucleus pulposus
is to allow flexibility between vertebrae for movement of the entire spinal column while preventing compression. Spinal nerves exit
between vertebrae, and the discs maintain the spacing to avoid
nerve damage. A slipped disc, resulting from straining the back, is
a misnomer. What actually occurs is a herniation, or rupture, because
of a weakened wall of the nucleus pulposus. This may cause severe
pain as a nerve is compressed.
Class Mammalia
Mammals are chordate animals with hair and mammary glands.
Hair is a thermoregulatory protective covering for most mammals, and mammary glands serve for suckling the young
(fig. 2.3). Other characteristics of mammals include a convoluted
(intricately folded) cerebrum, three auditory ossicles (bones), heterodont dentition (teeth of various shapes), squamosal-dentary
jaw articulation (a joint between the lower jaw and skull), an attached placenta (pla-cen'ta), well-developed facial muscles, a
muscular diaphragm, and a four-chambered heart with a left aortic arch (fig. 2.4).
heterodont: Gk. heteros, other; odontos, tooth

placenta: L. placenta, flat cake
Order Primates
There are several subdivisions of closely related groupings of mammals. These are called orders. Humans, along with lemurs, monkeys, and great apes, belong to the order called Primates. Members
of this order have prehensile hands (fig. 2.5), digits modified for
grasping, and relatively large, well-developed brains (fig. 2.6).
Family Hominidae
Humans are the sole living members of the family Hominidae.
Homo sapiens is included within this family, to which all the varieties or ethnic groups of humans belong (fig. 2.7). Each racial
group has distinguishing features that have been established in
isolated populations over thousands of years. Our classification
pedigree is presented in table 2.1.
Fostered by greater ease of travel and communication, more
frequent contact between diverse cultures has led to a breakdown of some of the traditional barriers to interracial marriage. This
may lead to a mixing of the gene pool, so that distinct ethnic
groups become less evident. Perhaps multiple ethnic ties in everyones pedigree would help reduce cultural hostility and strife.
Primates: L. primas, first

prehensile: L. prehensus, to grasp

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CHAPTER 2
Mammary ridge
Nipple
Accessory nipples
Creek
(a)
(b)
FIGURE 2.3 The mammary ridge and accessory nipples. (a) Mammary glands are positioned along a mammary ridge. (b) In humans, additional nipples (polythelia) occasionally develop elsewhere along the mammary ridge (see arrows).
Convoluted cerebrum
Hair
Three auditory
ossicles
Well-developed
facial muscles
Squamosal-dentary
jaw articulation
Heterodont dentition
Four-chambered
heart with left
aortic arch
Mammary glands
Muscular diaphragm
Placental attachment
for young (within
uterus)
FIGURE 2.4 Mammals have several distinguishing characteristics;

some of these are indicated in the photo with their approximate location within the body.
Characteristics of Humans
As human beings, certain of our anatomical characteristics are so
specialized that they are diagnostic in separating us from other
animals, and even from other closely related mammals. We also
have characteristics that are equally well developed in other animals, but when these function with the human brain, they provide us with remarkable and unique capabilities. Our anatomical
characteristics include the following:
1. A large, well-developed brain. The adult human brain
weighs between 1,350 and 1,400 grams (3 pounds). This
gives us a large brain-to-body-weight ratio. But more important is the development of portions of the brain. Certain extremely specialized regions and structures within the
brain account for emotion, thought, reasoning, memory,
and even precise, coordinated movement.
2. Bipedal locomotion. Because humans stand and walk on
two appendages, our style of locomotion is said to be
bipedal. Upright posture imposes other diagnostic structural
features, such as the sigmoid (S-shaped) curvature of the
bipedal: L. bi, two; pedis, foot

sigmoid: Gk. sigma, shaped like the letter S

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Creek
Tarsier
Aye-aye
Gorilla
Human
FIGURE 2.5 An opposable thumb enables a prehensile grip, which is characteristic of primates.
Cerebrum
Optic lobe
Cerebellum
Cerebrum
Codfish
Frog
Chimpanzee
Cerebellum
Alligator
Goose
Human
Creek
Horse
FIGURE 2.6 The brains of various vertebrates showing the relative size of the cerebrum (shaded pink) to other structures. (The brains are not
drawn to scale. Note that only mammals have a convoluted cerebrum.)

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(a)
(b)
(e)
(c)
(d)
(f)
FIGURE 2.7 The principal races of humans: (a) Mongoloid (Thailand), (b) Caucasoid (Northern Europe), (c) Negroid (Africa), (d) people of Indian subcontinent (Nepal), (e) Capoid (Kalahari bushman), and (f ) Australoid (Ngatatjara man, West Australia).
TABLE 2.1 Classification of Human Beings

Taxon
Designated
Grouping
Kingdom
Animalia
Eucaryotic cells without walls, plastids,

or photosynthetic pigments
Phylum
Chordata
Dorsal hollow nerve cord; notochord;

pharyngeal pouches
Subphylum
Vertebrata
Vertebral column
Class
Mammalia
Mammary glands; hair; convoluted

cerebrum; heterodont dentition
Characteristics
Order
Primates
Well-developed brain; prehensile hands
Family
Hominidae
Large cerebrum, bipedal locomotion
Genus
Homo
Flattened face; prominent chin and nose

with inferiorly positioned nostrils
Species
sapiens
Largest cerebrum
spine, the anatomy of the hips and thighs, and arched feet.
Some of these features may cause clinical problems in older
individuals.
3. An opposable thumb. The human thumb is structurally
adapted for tremendous versatility in grasping objects.
The saddle joint at the base of the thumb allows a wide
range of movement (see fig. 8.13). All primates have opposable thumbs.
4. Well-developed vocal structures. Humans, like no other
animals, have developed articulated speech. The anatomical structure of our vocal organs (larynx, tongue, and lips),
and our well-developed brain have made this possible.
5. Stereoscopic vision. Although this characteristic is well developed in several other animals, it is also keen in humans.
Our eyes are directed forward so that when we focus on an
object, we view it from two angles. Stereoscopic vision gives
us depth perception, or a three-dimensional image.
stereoscopic: Gk. stereos, solid; skopein, to view

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We also differ from other animals in the number and

arrangement of our vertebrae (vertebral formula), the kinds and
number of our teeth (tooth formula), the degree of development
of our facial muscles, and the structural organization of various
body organs.
The human body contains many distinct kinds of cells,

each specialized to perform specific functions. Examples of specialized cells are bone cells, muscle cells, fat cells, blood cells,
liver cells, and nerve cells. The unique structure of each of these
cell types is directly related to its function.
The human characteristics just described account for the

splendor of our cultural achievements. As bipedal animals,
we have our hands free to grasp and manipulate objects with our opposable thumbs. We can store information in our highly developed
brain, make use of it at a later time, and even share our learning
through oral or written communication.
Tissue Level
Knowledge Check
1. What is a chordate? Why are humans considered members
of the phylum Chordata?
2. Why are humans designated as vertebrates, mammals, and
primates? What characteristics distinguish humans from
other primates?
3. Which characteristics of humans are adaptive for social
organization?
Tissues are layers or groups of similar cells that perform a common function. The entire body is composed of only four principal kinds of tissues: epithelial, connective, muscular, and nervous
tissue. An example of a tissue is the muscle within the heart,
whose function it is to pump the blood through the body. The
outer layer of skin (epidermis) is a tissue because it is composed
of similar cells that together serve as a protective shield for the
body. Histology is the science concerned with the microscopic
study of tissues. The characteristic roles of each tissue type are
discussed fully in chapter 4.
Organ Level
BODY ORGANIZATION
Structural and functional levels of organization characterize the
human body, and each of its parts contributes to the total organism.
Objective 4
Identify the components of a cell, tissue, organ,

and system, and explain how these structures relate to one
another in constituting an organism.
Objective 5
Describe the general function of each system.
Cellular Level
The cell is the basic structural and functional component of
life. Humans are multicellular organisms composed of 60 to
100 trillion cells. It is at the microscopic cellular level that
such vital functions of life as metabolism, growth, irritability
(responsiveness to stimuli), repair, and replication are carried on.
Cells are composed of atomsminute particles that
are bound together to form larger particles called molecules
(fig. 2.8). Certain molecules, in turn, are grouped in specific
ways to form small functional structures called organelles
(or''ga-nelz'). Each organelle carries out a specific function
within the cell. A cells nucleus, mitochondria, and endoplasmic reticulum are organelles. The structure of cells and
the functions of the organelles will be examined in detail in
chapter 3.
An organ is an aggregate of two or more tissue types that performs a specific function. Organs occur throughout the body and
vary greatly in size and function. Examples of organs are the
heart, spleen, pancreas, ovary, skin, and even any of the bones
within the body. Each organ usually has one or more primary tissues and several secondary tissues. In the stomach, for example,
the inside epithelial lining is considered the primary tissue because the basic functions of secretion and absorption occur
within this layer. Secondary tissues of the stomach are the connective, nervous, and muscle tissues.
System Level
The systems of the body constitute the next level of structural
organization. A body system consists of various organs that have
similar or related functions. Examples of systems are the circulatory system, nervous system, digestive system, and endocrine system. Certain organs may serve two systems. For example, the
pancreas functions with both the endocrine and digestive systems and the pharynx serves both the respiratory and digestive
systems. All the systems of the body are interrelated and function
together, making up the organism.
tissue: Fr. tissu, woven; from L. texo, to weave

cell: L. cella, small room
organ: Gk. organon, instrument

system: Gk. systema, being together

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Chapter 2
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Increasing complexity
CHAPTER 2
Atom
Molecule
System
Macromolecule
Organelle
Organ
Cell
Organism
Tissue
FIGURE 2.8 The levels of structural organization and complexity within the human body.
Growth is a normal process by which an organism increases
in size as a result of the accretion of cells and tissues similar to
those already present within organs. Growth is an integral part of
development that continues until adulthood. Normal growth depends not only on proper nutrition but on the concerted effect of
several hormones (chemicals produced by endocrine glands), including insulin, growth hormone, and (during adolescence) the
sex hormones. It is through the growth process that each of the
body systems eventually matures (fig. 2.9). Puberty is the developmental transition during the growth process when sexual features become expressed in several of the body systems and the
reproductive organs become functional.
A systematic (systemic) approach to studying anatomy emphasizes the purposes of various organs within a system. For example, the functional role of the digestive system can be best
understood if all of the organs within that system are studied together. In a regional approach, all of the organs and structures in
one particular region are examined at the same time. The regional approach has merit in graduate professional schools (medical, dental, etc.) because the structural relationships of portions
of several systems can be observed simultaneously. Dissections of

cadavers are usually conducted on a regional basis. Trauma or injury usually affects a region of the body, whereas a disease that affects a region may also involve an entire system.
This text uses a systematic approach to anatomy. In the
chapters that follow, you will become acquainted, system by system, with the functional anatomy of the entire body. An
overview of the structure and function of each of the body systems is presented in figure 2.10.
Knowledge Check
4. Construct a diagram to illustrate the levels of structural organization that characterize the body. Which of these levels are microscopic?
5. Why is the skin considered an organ?
6. Which body systems control the functioning of the others?
Which are supportive of the organism? Which serve a
transportive role?

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II. Terminology,
Human Organism
testosterone
and other
androgens
FIGURE 2.9 Developmental changes from childhood to adulthood. Controlled by hormones, puberty is the period of body growth when sexual
characteristics are expressed and the sexual organs become functional.
ANATOMICAL NOMENCLATURE
In order to understand the science of anatomy, students must
master its descriptive terminology.
Objective 6
Explain how anatomical terms are derived.
Objective 7
Describe what is meant by prefixes and suffixes.
Anatomy is a descriptive science. Analyzing anatomical terminology can be a rewarding experience in that one learns something of the character of antiquity in the process. However,
understanding the roots of words is not only of academic interest.
Familiarity with technical terms reinforces the learning process.
Most anatomical terms are derived from Greek or Latin, but
some of the more recent terms are of German and French origin.
As mentioned in chapter 1, some anatomical structures bear the
names of people who discovered or described them. Such terms
are totally nondescriptive; unfortunately, they have little meaning in and of themselves.
Many Greek and Latin terms were coined more than 2,000
years ago. Deciphering the meanings of these terms affords a
glimpse into our medical heritage. Many terms referred to common plants or animals. Thus, the term vermis means worm;
cochlea (kok'le-a), snail shell; cancer, crab; and uvula, little grape.
Even the term muscle comes from the Latin musculus, which
means mouse. Other terms suggest the warlike environment of
ancient Greece and Rome. Thyroid, for example, means shield;

xiphos (zi'fos) means sword; and thorax, breastplate. Sella means
saddle and stapes (sta'pez) means stirrup. Various tools or instruments were referred to in early anatomy. The malleus and anvil
resemble miniatures of a blacksmiths implements, and tympanum refers to a drum.
You will encounter many new terms throughout your
study of anatomy. You can learn these terms more easily if
you know the meaning of their prefixes and suffixes. Use the
glossary of prefixes and suffixes (on the inside front cover) as
an aid in learning new terms. Pronouncing these terms as
you learn them will also help you remember them. A guide
to the singular and plural forms of words is presented in
table 2.2.
The material presented in the remainder of this chapter
provides a basic foundation for anatomy, as well as for all medical
and paramedical fields. Anatomy is a very precise science because of its universally accepted reference language for describing
body parts and locations.
Knowledge Check
7. Explain the statement, Anatomy is a descriptive science.
8. Refer to the glossary of prefixes and suffixes on the front
inside front cover to decipher the terms blastocoel, hypodermic, dermatitis, and orchiectomy.

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CHAPTER 2
Integumentary system
Function: external support
and protection of body
Lymphatic system
Function: body immunity;
absorption of fats; drainage
of tissue fluid
FIGURE 2.10 The body systems.
Skeletal system
Function: internal support and
flexible framework for body
movement; production of
blood cells; storage of
minerals
Endocrine system
Function: secretion of
hormones for
chemical regulation
Muscular system
Function: body movement;
production of body heat
Urinary system
Function: filtration of blood;
maintenance of volume and
chemical composition
of blood; removal of
metabolic wastes from body

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Respiratory system
Function: gaseous exchange
between external environment
and blood
Digestive system
Function: breakdown and
absorption of food materials
FIGURE 2.10 Continued
Nervous system
Function: control and
regulation of all other
systems of the body
Female reproductive system

Function: production of female
sex cells (ova) and female
hormones; receptacle for
sperm from male; site for
fertilization of ovum,
implantation, and development
of embryo and fetus; delivery
of fetus
Circulatory system
Function: transport of
life-sustaining materials to
body cells; removal of
metabolic wastes from cells
Male reproductive system

Function: production of male
sex cells (sperm) and male
hormones; transfer of sperm
to reproductive system of
female

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Chapter 2
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TABLE 2.2 Examples of Singular and

Plural Forms of Body Terminology
Coronal
plane
Examples
-a
-ae
Papilla, papillae; axilla, axillae
-en
-ina
Foramen, foramina; lumen, lumina
-ex
-ices
Cortex, cortices
-is
-es
Testis, testes
-is
-ides
Epididymis, epididymides
-ix
-ices
Matrix, matrices; appendix, appendices
-on
-a
Mitochondrion, mitochondria
-um
-a
Epithelium, epithelia; cilium, cilia; ovum, ova
-us
-i
Humerus, humeri; carpus, carpi; fasciculus,

fasciculi
-us
-ora
Corpus, corpora
-x
-ges
Phalanx, phalanges; pharynx, pharynges
-y
-ies
Artery, arteries; ovary, ovaries
Transverse
plane
PLANES OF REFERENCE
AND DESCRIPTIVE TERMINOLOGY
All of the descriptive planes of reference and terms of direction
used in anatomy are standardized because of their reference to
the body in anatomical position.
FIGURE 2.11 Planes of reference through the body.
Objective 8
Identify the planes of reference used to locate

structures within the body.
Objective 9
Describe the anatomical position.
Objective 10
Define and be able to properly use the

descriptive and directional terms that refer to the body.
Planes of Reference
In order to visualize and study the structural arrangements of various organs, the body may be sectioned (cut) and diagrammed according to three fundamental planes of reference: a sagittal
(saj' -tal) plane, a coronal plane, and a transverse plane
(figs. 2.11 and 2.12). A sagittal plane extends vertically through
the body dividing it into right and left portions. A midsagittal (median) plane is a sagittal plane that passes lengthwise through the
midplane of the body, dividing it equally into right and left halves.
Coronal, or frontal, planes also pass lengthwise and divide the body
coronal: L. corona, crown
into anterior (front) and posterior (back) portions. Transverse

planes, also called horizontal, or cross-sectional, planes, divide the
body into superior (upper) and inferior (lower) portions.
The value of the computerized tomographic X-ray (CT) scan
(see fig. 1.19a) is that it displays an image along a transverse
plane similar to that which could otherwise be obtained only in an actual section through the body. Prior to the development of this technique, the vertical plane of conventional radiographs made it difficult,
if not impossible, to assess the extent of body irregularities.
Descriptive Terminology
Anatomical Position
All terms of direction that describe the relationship of one body
part to another are made in reference to the anatomical position. In the anatomical position, the body is erect, the feet are
parallel to each other and flat on the floor, the eyes are directed
forward, and the arms are at the sides of the body with the palms
of the hands turned forward and the fingers pointed straight
down (fig. 2.13).
CHAPTER 2
Singular Plural
ending
ending
Sagittal
plane

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(a)
(b)
(c)
FIGURE 2.12 The human brain sectioned along (a) a transverse plane, (b) a coronal plane, and (c) a sagittal plane.
Directional Terms
Directional terms are used to locate structures and regions of the
body relative to the anatomical position. A summary of directional terms is presented in table 2.3.
Clinical Procedures
Certain clinical procedures are important in determining
anatomical structure and function in a living individual. The
most common of these are as follows:
Inspection. Visually observing the body to note any clinical symptoms, such as abnormal skin color, swelling, or
rashes. Other observations may include needle marks on
the skin, irregular breathing rates, or abnormal behavior.
Palpation. Applying the fingers with firm pressure to the
surface of the body to feel surface landmarks, lumps, tender
spots, or pulsations.
Percussion. Tapping sharply on various locations on the
thorax or abdomen to detect resonating vibrations as an
aid in locating excess fluids or organ abnormalities.

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Chapter 3
Cytology
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CHAPTER 3
FIGURE 3.26 The stages of mitosis.

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Microscopic Structure of the Body
friction to the skin stimulates mitotic activity and the synthesis

of a fibrous protein, keratin, which results in the formation of a
protective callus.
Cells adapt to potentially injurious stimuli by several specific mechanisms. Hypertrophy (hipertro-fe) refers to an increase in the size of cells resulting from increased synthesis of
protein, nucleic acids, and lipids. Cellular hypertrophy can be either compensatory or hormonal. Compensatory hypertrophy occurs when increased metabolic demands on particular cells result
in an increase in cellular mass. Examples of compensatory hypertrophy include the enlargement of skeletal muscle fibers as a result of exercise and cardiac (heart) muscle fibers or kidney cells
because of an increased work demand. Hypertension (high blood
pressure) causes cardiac cells to hypertrophy because they must
pump blood against raised pressures. After the removal of a diseased kidney, there is a compensatory increase in the size of the
cells of the remaining kidney so that its normal weight is approximately doubled. Examples of hormonal hypertrophy are the increased size of the breasts and smooth muscles of the uterus in a
pregnant woman.
Hyperplasia (hiper-plaze-a) refers to an increase in the
number of cells formed as a result of increased mitotic activity.
The removal of a portion of the liver, for example, leads to regeneration, or hyperplasia, of the remaining liver cells to restore
the loss. But the triggering mechanism for hyperplasia is not
known. In women, a type of hormonally induced hyperplasia occurs in cells of the endometrium of the uterus after menstruation,
which restores this layer to a suitable state for possible implantation of an embryo.
Atrophy (atro-fe) refers to a decrease in the size of cells
and a corresponding decrease in the size of the affected organ.
Atrophy can occur in the cells of any organ and may be classified
as disuse atrophy, disease atrophy, or aging (senile) atrophy.
Metaplasia (meta -pl ze-a) is a specialized cellular change
in which one type of cell transforms into another. Generally, it
involves the change of highly specialized cells into more generalized, protective cells. For example, excessive exposure to inhaled
smoke causes the specialized ciliated columnar epithelial cells
lining the bronchial airways to change into stratified squamous
epithelium, which is more resistant to injury from smoke.
Trauma to Cells
As adaptable as cells are to environmental changes, they are subject to damage from aging and disease. If a trauma causes extensive cellular death, the condition may become life threatening.
A person dies when a vital organ can no longer perform its metabolic role in sustaining the body.
hypertrophy: Gk. hyper, over; trophe, nourishment

hyperplasia: Gk. hyper, over; plasis, a molding
atrophy: Gk. a, without; trophe, nourishment
metaplasia: Gk. meta, between; plasis, a molding
Energy deficit means that more energy is required by a cell

than is available. Cells can tolerate certain mild deficits because
of various reserves stored within the cytoplasm, but a severe or
prolonged deficit will cause cells to die. An energy deficit occurs
when the cells do not have enough glucose or oxygen to allow
for glucose combustion. Examples of energy deficits are low levels
of blood sugar (hypoglycemia) and the impermeability of the cell
membrane to glucose (as in diabetes mellitus). Malnutrition also
may result in an energy deficit. Few cells can tolerate an interruption in oxygen supply. Cells of the brain and the heart have
tremendous oxygen demands, and an interruption of the supply
to these organs can cause death in a matter of minutes.
Physical injury to cells, another type of trauma, occurs in
a variety of ways. High temperature (hyperthermia) is generally
less tolerable to cells than low temperature (hypothermia). Respiratory rate, heart rate, and metabolism accelerate with hyperthermia. Continued hyperthermia causes protein coagulation
within cells, and eventually cellular death. In frostbite, rapid or
prolonged chilling causes cellular injury. In severe frostbite, ice
crystals form and cause the cells to burst.
Burns are particularly significant if they cause damage to
the deeper skin layers, which interferes with the mitotic activity
of cells (see fig. 5.20). Of immediate concern with burns, however, is the devastating effect of fluid loss and infection through
traumatized cell membranes.
Accidental poisoning and suicide through drug overdose
account for large numbers of deaths in the United States and
elsewhere. Drugs and poisons can cause cellular dysfunction by
disrupting DNA replication, RNA transcription, enzyme systems, or cell membrane activity.
Radiation causes a type of cell trauma that is cumulative in
effect. When X rays are administered for therapeutic purposes
(radiotherapy), small doses are focused on a tumorous area over a
course of many days to prevent widespread cellular injury. Some
cells are more sensitive to radiation than others. Immature or mitotically active cells are highly sensitive, whereas cells that are
no longer growing, such as neurons and muscle cells, are not as
vulnerable to radiation injury.
Infectious agents, or pathogens, also cause cellular dysfunction. Viruses and bacteria are the most common pathogens.
Viruses usually invade and destroy cells as they reproduce themselves. Bacteria, on the other hand, do not usually invade cells
but will frequently poison cells with their toxic metabolic wastes.
Medical Genetics
Medical genetics is a branch of medicine concerned with diseases that have a genetic origin. Genetic factors include abnormalities in chromosome number or structure and mutant genes.
Genetic diseases are a diverse group of disorders, including malformed blood cells (sickle-cell anemia), defective blood clotting
(hemophilia), and mental retardation (Down syndrome).
Chromosomal abnormalities occur in approximately 0.6%
of live-birth infants. The majority (70%) are subtle, cause no
problems, and usually go undetected. Structural changes in the
DNA that are passed from parent to offspring by means of sex


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Chapter 3
cells are called mutations (myoo-tashunz). Mutations either
occur naturally or are environmentally induced through chemicals or radiation. Natural mutations are not well understood.
About 12% of all congenital malformations are caused by mutations and probably come about through an interaction of genetic
and environmental factors. Many of these problems can be predicted by knowing the genetic pedigree of prospective parents
and prevented through genetic counseling. Teratology (ter-atolo-je) is the science concerned with developmental defects and
the diagnosis, treatment, and prevention of malformations.
Genetic problems are occasionally caused by having too
few or too many chromosomes. The absence of an entire chromosome is termed monosomy (mono-somme). Embryos with
monosomy usually die. People with Turners syndrome have
only one X chromosome and have a better chance of survival
than those who are missing one of the other chromosomes. Trisomy (tri so-me), a genetic condition in which an extra chromosome is present, occurs more frequently than monosomy. The
best known among the trisomies is Down syndrome.
Cancer
Cancer refers to a complex group of diseases characterized by uncontrolled cell replication. The rapid proliferation of cells results
in the formation of a neoplasm, or new cellular mass. Neoplasms,
frequently called tumors, are classified as benign or malignant
based on their cytological and histological features. Benign neoplasms usually grow slowly and are confined to a particular area.
These types are usually not life threatening unless they grow to
large sizes in vital organs like the brain. Malignant neoplasms
(fig. 3.27) grow rapidly and metastasize (me-tasta-sz) (fragment
and spread) easily through lymphatic or blood vessels. The original malignant neoplasm is called the primary growth and the new
tumors, or metastatic tumors, are called secondary growths.
Cancer cells resemble undifferentiated or primordial cell
types. Generally they do not mature before they divide and are
not capable of maintaining normal cell function. Cancer causes
death when a vital organ regresses because of competition from
cancer cells for space and nutrients. The pain associated with cancer develops when the growing neoplasm affects sensory neurons.
73
Normal cells
(with hairlike cilia)
Cancer cells
FIGURE 3.27 An electron micrograph of cancer cells from the

respiratory tract (59,800).
The various types of cancers are classified on the basis of the
tissue in which they develop. Lymphoma, for example, is a cancer
of lymphoid tissue; osteogenic cancer is a type of bone cancer;
myeloma is cancer of the bone marrow; and sarcoma is a general
term for any cancer arising from cells of connective tissue.
The etiology (cause) of cancers is largely unknown. However, initiating factors, or carcinogens (kar-sino-jenz), such as
viruses, chemicals, or irradiation, may provoke cancer to develop. Cigarette smoking, for example, causes various respiratory
cancers to develop. The tendency to develop other types of cancers has a genetic basis. Some researchers even think that physiological stress can promote certain types of cancerous activity.
Because the causes of cancer are not well understood, emphasis is
placed on early detection with prompt treatment.
Aging
mutation: L. mutare, to change

teratology: Gk. teras, monster; logos, study
Although there are obvious external indicators of aginggraying

and loss of hair, wrinkling of skin, loss of teeth, and decreased
muscle masschanges within cells as a result of aging are not as
apparent and are not well understood. Certain organelles alter
with age. The mitochondria, for example, may change in structure and number, and the Golgi complex may fragment. Also,
lipid vacuoles tend to accumulate in the cytoplasm, and the cytoplasmic food stores that contain glycogen decrease.
Turners syndrome: from Henry H. Turner, American endocrinologist,

18921970
Down syndrome: from John L. H. Down, English physician, 182896
carcinogen: Gk, karkinos, cancer
CHAPTER 3
In an attempt to better understand medical genetics, the

Human Genome Project was launched by Congress in 1988
with the ambitious goal of completely mapping the human genome.
Scientists are currently on the verge of determining the exact sequences of bases with which the 3 billion base pairs are arranged to
form the 50,000 to 100,000 genes in the haploid human genome of a
sperm cell or ovum. Knowing this information will provide the ultimate
reference for diagnosis and treatment of the 4000 genetic diseases
that are known to be directly caused by particular abnormal genes.
Cytology

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The McGrawHill
Companies, 2001
ies, and its deterioration is thought to be associated with such

vascular diseases as arteriosclerosis in aged persons.

Giving a substance that competes for the enzyme alcohol dehydrogenase
can inhibit the reaction that forms the toxic metabolite of ethylene glycol. Thus, infusing a nearly intoxicating dose of alcohol can spare the
kidneys from harm. The ethylene glycol is then excreted harmlessly.
CHAPTER 3
The chromatin and chromosomes within the nucleus

show changes with aging, such as clumping, shrinking, or fragmenting with repeated mitotic divisions. There is strong evidence that certain cell types have a predetermined number of
mitotic divisions that are genetically controlled, thus determining the overall vitality and longevity of an organ. If this is true,
identifying and genetically manipulating the aging gene might
be possible.
Extracellular substances also change with age. Protein
strands of collagen and elastin change in quality and number in
aged tissues. Elastin plays an important role in the walls of arter-
Chapter Summary
Introduction to Cytology (p. 49)
1. Cells are the structural and functional
units of the body. Cellular function is
referred to as metabolism and the study of
cells is referred to as cytology.
2. Cellular function depends on the specific
membranes and organelles characteristic
of each type of cell.
3. All cells have structural modifications
that serve functional purposes.
Cellular Chemistry (pp. 5052)

1. Four elements (oxygen, carbon, hydrogen,
and nitrogen) compose over 95% of the
bodys mass and are linked together to
form inorganic and organic compounds.
2. Water is the most abundant inorganic
compound in cells and is an excellent
solvent.
(a) Water is important in temperature
control and hydrolysis.
(b) Dehydration, a condition in which
fluid loss exceeds fluid intake, may be a
serious problemespecially in infants.
3. Electrolytes are inorganic compounds that
form ions when dissolved in water.
(a) The three classes of electrolytes are
acids, bases, and salts.
(b) Electrolytes are important in
maintaining pH, in conducting
electrical currents, and in regulating
the activity of enzymes.
4. Proteins are organic compounds that may
exist by themselves or be conjugated with
other compounds.
(a) Proteins are important structural
components of the body and are
necessary for cellular growth, repair,
and division.
(b) Enzymes and hormones are examples

of specialized proteins.
5. Carbohydrates are organic compounds
containing carbon, hydrogen, and oxygen,
with a 2:1 ratio of hydrogen to oxygen.
(a) The carbohydrate group includes the
starches and sugars.
(b) Carbohydrates are the most abundant
source of cellular energy.
6. Lipids are organic fats and fat-related
substances.
(a) Lipids are composed primarily of
carbon, hydrogen, and oxygen.
(b) Lipids serve as an important source of
energy, form parts of membranes, and
protect and insulate various parts of
the body.
Cellular Structure (pp. 5264)

1. A cell is composed of a cell membrane,
cytoplasm and organelles, and a nucleus.
2. The cell membrane, composed of
phospholipid and protein molecules,
encloses the contents of the cell and
regulates the passage of substances into
and out of the cell.
(a) The permeability of the cell membrane
depends on its structure, the size of the
molecules, ionic charge, lipid solubility,
and the presence of carrier molecules.
(b) Cell membranes may be specialized
with such structures as microvilli,
sacs, and hair cells.
3. Cytoplasm refers to the material
between the cell membrane and the
nucleus. Nucleoplasm is the material
within the nucleus. Protoplasm is a
collective term for both the cytoplasm
and nucleoplasm.
4. Organelles are specialized components

within the cytoplasm of cells.
(a) Endoplasmic reticulum provides a
framework within the cytoplasm and
forms a site for the attachment of
ribosomes. It functions in the
synthesis of lipids and proteins and in
cellular transport.
(b) Ribosomes are particles of protein
and RNA that function in protein
synthesis. The protein particles
may be used within the cell or
secreted.
(c) The Golgi complex consists of
membranous vesicles that synthesize
glycoproteins and secrete lipids. The
Golgi complex is extensive in
secretory cells, such as those of the
pancreas and salivary glands.
(d) Mitochondria are membranous
sacs that consist of outer and
inner mitochondrial layers and
folded membranous extensions
of the inner layer called cristae.
The mitochondria produce ATP
and are called the powerhouses
of a cell. Mitochondria are lacking
in sperm cells and red blood cells.
(e) Lysosomes are spherical bodies that
contain digestive enzymes. They are
abundant in the phagocytic white
blood cells.
(f) Peroxisomes are enzyme-containing
membranous sacs that are abundant in
the kidneys and liver. Some of the
enzymes in peroxisomes generate
hydrogen peroxide, and one of them,
catalase, breaks down excess hydrogen
peroxide.


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Chapter 3
6. The nucleotides in DNA consist of the sugar
deoxyribose, phosphate, and one of four
nitrogenous bases: adenine, guanine,
cytosine, or thymine. According to the law of
complementary base pairing, bases are
specific in their bonding: adenine bonds with
thymine and guanine bonds with cytosine.
7. RNA contains the sugar ribose (instead of
deoxyribose) and the base uracil (in place
of thymine). The three major forms of
RNA are mRNA, tRNA, and rRNA.
8. The genetic code in mRNA consists of
three bases called codons. Codons bond to
anticodons, which are three bases in tRNA.
9. Each type of tRNA is bonded to a specific
type of amino acid, which the tRNA
brings to the growing polypeptide chain.
Cell Cycle (pp. 6570)

1. The cell cycle consists of growth,
synthesis, and mitosis.
(a) Growth is the increase in cellular
mass that results from metabolism.
75
Synthesis is the production of DNA

and RNA to regulate cellular activity.
Mitosis is the splitting of the cells
nucleus and cytoplasm that results in
the formation of two diploid cells.
(b) Mitosis permits an increase in
the number of cells (body growth)
and allows for the replacement
of damaged, diseased, or
worn-out cells.
2. A DNA molecule is in the shape of a
double helix. The structural unit of the
molecule is a nucleotide, which consists
of deoxyribose (sugar), phosphate, and a
nitrogenous base.
3. Cell division consists of a division of the
chromosomes (mitosis) and a division of
the cytoplasm (cytokinesis). The stages of
mitosis include prophase, metaphase,
anaphase, and telophase.
Review Activities
Objective Questions
1. Inorganic compounds that form ions
when dissociated in water are
(a) hydrolites.
(d) ionizers.
(b) metabolites. (e) nucleic acids.
(c) electrolytes.
2. The four elements that compose over
95% of the body are
(a) oxygen, potassium, hydrogen, carbon.
(b) carbon, sodium, nitrogen, oxygen.
(c) potassium, sodium, magnesium,
oxygen.
(d) carbon, oxygen, nitrogen, hydrogen.
(e) oxygen, carbon, hydrogen, sulfur.
3. Which organelle contains strong
hydrolytic enzymes?
(a) the lysosome
(b) the Golgi complex
(c) the ribosome
(d) the vacuole
(e) the mitochondrion
4. Ciliated cells occur in
(a) the trachea. (c) the bronchioles.
(b) the ductus
(d) the uterine tubes.
deferens.
(e) all of the above.
5. Osmosis deals with the movement of
(a) gases.
(c) oxygen only.
(b) water only.
(d) both a and c.
6. The phase of mitosis in which the

chromosomes line up at the equator
(equatorial plane) of the cell is called
(a) interphase.
(d) anaphase.
(b) prophase.
(e) telophase.
(c) metaphase.
chromatids separate is called
(a) interphase.
(d) anaphase.
(b) prophase.
(e) telophase.
(c) metaphase.
8. The organelle that combines protein with
carbohydrates and packages them within
vesicles for secretion is
(a) the Golgi complex.
(b) the rough endoplasmic reticulum.
(c) the smooth endoplasmic reticulum.
(d) the ribosome.
9. The enlarged skeletal muscle fibers that
result from an increased work demand
serve to illustrate
(a) disuse atrophy.
(b) compensatory hypertrophy.
(c) metaplasia.
(d) inertia.
10. Regeneration of liver cells is an example of
(a) compensatory hypertrophy.
(b) hyperplasia.
(c) metaplasia.
(d) hypertrophy.
11. Which of the following statements about
DNA is false?
(a) It is located in the nucleus.
(b) It is double-stranded.
(c) The bases adenine and thymine can
bond together.
(d) The bases guanine and adenine can
bond together.
RNA is true?
(a) It is made in the nucleus.
(b) It contains the sugar deoxyribose.
(c) It is a complementary copy of the
entire DNA molecule.
(d) It is double-stranded.
Essay Questions
1. Explain why a knowledge of cellular
anatomy is necessary for understanding
tissue and organ function within the
body. How is the study of cells important
for the understanding of body dysfunction
and disease?
2. Why is water a good fluid medium
of the cell?
CHAPTER 3
(g) The centrosome is the dense area of

cytoplasm near the nucleus that
contains the centrioles. The paired
centrioles play an important role in
cell division.
(h) Vacuoles are membranous sacs that
function as storage chambers.
(i) Fibrils and microtubules provide
support in the form of a cytoskeleton.
(j) Cilia and flagella are projections of
the cell that have the same basic
structure and that function in
producing movement.
5. The cell nucleus is enclosed in a nuclear
membrane that controls the movement of
substances between the nucleoplasm and
the cytoplasm.
(a) The nucleoli are small bodies of
protein and RNA within the nucleus
that produce ribosomes.
(b) Chromatin is a coiled fiber of protein
and DNA that shortens to form
chromosomes during cell reproduction.
Cytology

CHAPTER 3
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3. How are proteins, carbohydrates, and

lipids similar? How are they different?
What are enzymes and hormones?
4. Describe the cell membrane. List the
various kinds of movement through the cell
membrane and give an example of each.
5. Describe, diagram, and list the functions
of the following:
(a) endoplasmic reticulum,
(b) ribosome,
(c) mitochondrion,
(d) Golgi complex,
(e) centrioles, and
(f) cilia.
6. Define inorganic compound and organic
compound and give examples of each.
7. Define the terms protoplasm, cytoplasm,
and nucleoplasm. Describe the position of
the membranes associated with each of
these substances.
8. Describe the structure of the nucleus and
the functions of its parts.
9. What is a nucleotide? How does it relate
to the overall structure of a DNA
molecule?
10. Explain the relationship between DNA,
chromosomes, chromatids, and genes.
11. Describe how RNA is produced and list
the different forms of RNA.
12. Explain how one DNA strand can serve
as a template for the synthesis of another
DNA strand.
13. Distinguish between mitosis and

cytokinesis. Describe the major events of
mitosis and discuss the significance of the
mitotic process.
14. Give examples of factors that contribute
cellular hypertrophy, hyperplasia, atrophy,
and metaplasia.
15. Explain how cells respond to
(a) energy deficit,
(b) hyperthermia,
(c) burns,
(d) radiation, and
(e) pathogens.
16. Define the following genetic terms:
teratology, monosomy, trisomy, and
mutation.
17. In what ways do cells of a neoplasm differ
from normal cells. How may a malignant
neoplasm cause death?
18. Discuss the cellular and extracellular
changes that accompany aging.
1. How is the structural organization of its
individual cells essential to a multicellular
organism?
2. Construct a table comparing the structure
and function of several kinds of cells.
Indicate which organelles would be of
particular importance to each kind of cell.
3. Define medical genetics and give some
examples of genetic diseases. Is spending

the billions of dollars required to

complete the Human Genome Project
justified? Why or why not?
4. The brain is protected to some extent by
the blood-brain barriera membrane
between circulating blood and the brain
that keeps certain damaging substances
from reaching brain tissue. However, the
brain is still subject to trauma that can
cause it to swell, much like an ankle
swells with a sprain. Because the cranium
is a cavity of fixed size, brain edema
(swelling) can rapidly lead to coma and
death. Knowing what you do about
movement of water across a membrane,
can you explain why mannitol, a type of
sugar that does not cross the blood-brain
barrier, is commonly used to treat patients
who have suffered head trauma?
5. Your friend knows that you have just
reviewed cellular chemistry, and so he asks
for your opinion about his new diet. In an
attempt to eliminate the lipid content in
adipose tissue and thus lose weight, he has
completely eliminated fat from his diet.
He feels that he is now free to eat as much
food as he likes, provided it consists only
of carbohydrates and protein. Is your
friends logic flawed? Would you advise
him to stick with this diet?


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Histology
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4
Definition and Classification of
Tissues 78
Developmental Exposition:
The Tissues 79
Epithelial Tissue 79
Connective Tissue 89
Muscle Tissue 99
Nervous Tissue 100
CLINICAL CONSIDERATIONS 101

Chapter Summary 103
Clinical Case Study

As a medical student, you are rotating with a gastroenterologist who is performing an upper endoscopy on a patient with long-standing gastroesophageal reflux (heartburn). During the procedure, the doctor quizzes you on what type of cells line the esophagus. What is your answer? He
takes a biopsy of the lower esophagus just above the stomach. Later, the specimen is fixed, and
you examine it under the microscope. You see a single layer of nonciliated, tall, column-shaped
cells. What type of tissue are you examining?
FIGURE: Knowing the structure and function

of body tissues elucidates how they may adapt
to provide protection to body organs.

78
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DEFINITION AND
Shaft of a
hair within
a hair follicle
Objective 1
histology.
CHAPTER 4
Objective 2

cells and tissues.
Objective 3



(a)
Shaft of hair
emerging from
the exposed
surface of
the skin
(b)


Knowledge Check


of the Body
4. Histology
The McGrawHill
Companies, 2001
Embryoblast
Blastocoele
Amniotic cavity
(c)
Ectoderm
Trophoblast
Endoderm
(b)
Amniotic cavity
(a)
Embryonic
disc
(d)
Ectoderm
Mesoderm
Endoderm
Yolk sac
(e)
Trophoblast
Schenk
EXHIBIT 1 The early stages of embryonic development. (a) Fertilization and the formation of the zygote, (b) the morula at about the
third day, (c) the early blastocyst at the time of implantation between the fifth and seventh day, (d) a blastocyst at 2 weeks, and (e) a blastocyst at 3 weeks showing the three primary germ layers that constitute the embryonic disc.
The Tissues
Human prenatal development is initiated by the fertilization of
an ovulated ovum (egg) from a female by a sperm cell from a
male. The chromosomes within the nucleus of a zygote (zgot)
(fertilized egg) contain all the genetic information necessary for
the differentiation and development of all body structures.
Within 30 hours after fertilization, the zygote undergoes a

mitotic division as it moves through the uterine tube toward the
uterus (see chapter 22). After several more cellular divisions, the
embryonic mass consists of 16 or more cells and is called a
morula (moryoo-la), as shown in exhibit I. Three or 4 days after
conception, the morula enters the uterine cavity where it remains unattached for about 3 days. During this time, the center
of the morula fills with fluid absorbed from the uterine cavity. As
the fluid-filled space develops inside the morula, two distinct
groups of cells form. The single layer of cells forming the outer
zygote: Gk. zygotos, yolked
morula: Gk. morus, mulberry
EXPLANATION
(continued)
EPITHELIAL TISSUE
Objective 5
Discuss the functions of the membranous

epithelia in different locations in the body.
There are two major categories of epithelia: membranous and

glandular. Membranous epithelia are located throughout the body
and form such structures as the outer layer of the skin; the inner
lining of body cavities, tubes, and ducts; and the covering of visceral organs. Glandular epithelia are specialized tissues that form
the secretory portion of glands.
Objective 6
Objective 4
Membranous epithelia always have one free surface exposed to a

body cavity, a lumen (hollow portion of a body tube), or to the
skin surface. Some membranous epithelia are derived from ectoderm, such as the outer layer of the skin; some from mesoderm,
79
Compare and contrast the various types of

membranous epithelia.
epithelium: Gk. epi, upon; thelium, to cover
Define gland and compare and contrast the

various types of glands in the body.
Characteristics of Membranous Epithelia


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(continued)
TABLE 4A Derivatives of the Germ Layers

Ectoderm
Mesoderm
Endoderm
Epidermis of skin and epidermal derivatives:

hair, nails, glands of the skin; linings of oral,
nasal, anal, and vaginal cavities
Muscle: smooth, cardiac, skeletal
Nervous tissue; sense organs
Dermis of skin; dentin of teeth
Epithelium of pharynx, auditory canal, tonsils,

thyroid, parathyroid, thymus, larynx, trachea,
lungs, GI tract, urinary bladder and urethra, and
vagina
Lens of eye; enamel of teeth

Pituitary gland
Epithelium (endothelium) of blood vessels,

lymphatic vessels, body cavities, joint cavities
Adrenal medulla
Internal reproductive organs
Connective tissue: embryonic, mesenchyme,

connective tissue proper, cartilage, bone, blood
Liver and pancreas
Kidneys and ureters

Adrenal cortex
wall is known as the trophoblast, and the inner aggregation of

cells is known as the embryoblast. With further development,
the trophoblast differentiates into a structure that will later form
part of the placenta; the embryoblast will eventually become the
embryo. With the establishment of these two groups of cells, the
morula becomes known as a blastocyst (blasto-sist). Implantation
of the blastocyst in the uterine wall begins between the fifth and
seventh day (see chapter 22).
As the blastocyst completes implantation during the second
week of development, the embryoblast undergoes marked differentiation. A slitlike space called the amniotic (amne-ot-ic) cavity
forms within the embryoblast, adjacent to the trophoblast. The
embryoblast now consists of two layers: an upper ectoderm, which
trophoblast: Gk. trophe, nourishment; blastos, germ

embryoblast: Gk. embryon, to be full, swell; blastos, germ
ectoderm: Gk. ecto, outside; derm, skin
such as the inside lining of blood vessels; and others from endoderm, such as the inside lining of the digestive tract (gastrointestinal, or GI, tract).
Membranous epithelia may be one or several cell layers
thick. The upper surface may be exposed to gases, as in the case
of epithelium in the integumentary and respiratory systems; to
liquids, as in the circulatory and urinary systems; or to semisolids,
as in the GI tract. The deep surface of most membranous epithelia is bound to underlying supportive tissue by a basement membrane, that consists of glycoprotein from the epithelial cells and
a meshwork of collagenous and reticular fibers from the underlying connective tissue. With few exceptions, membranous epithe80
is closer to the amniotic cavity, and a lower endoderm, which

borders the blastocoel (blastocyst cavity). A short time later, a
third layer called the mesoderm forms between the endoderm and
ectoderm. These three layers constitute the primary germ layers.
The primary germ layers are of great significance because
all the cells and tissues of the body are derived from them (see
fig. 22.9). Ectodermal cells form the nervous system; the outer
layer of skin (epidermis), including hair, nails, and skin glands;
and portions of the sensory organs. Mesodermal cells form the
skeleton, muscles, blood, reproductive organs, dermis of the skin,
and connective tissue. Endodermal cells produce the lining of the
GI tract, the digestive organs, the respiratory tract and lungs, and
the urinary bladder and urethra. The derivatives of the primary
germ layers are summarized in table 4A.
endoderm: Gk. endo, within; derm, skin

mesoderm: Gk. meso, middle; derm, skin
lia are avascular (without blood vessels) and must be nourished

by diffusion from underlying connective tissues. Cells that make
up membranous epithelia are tightly packed together, with little
intercellular matrix between them.
Some of the functions of membranous epithelia are quite
specific, but certain generalities can be made. Epithelia that cover
or line surfaces provide protection from pathogens, physical injury,
toxins, and desiccation. Epithelia lining the GI tract function in
absorption. The epithelium of the kidneys provides filtration,
whereas that within the pulmonary alveoli (air sacs) of the lungs
allows for diffusion. Highly specialized neuroepithelium in the taste
buds and in the nasal region has a chemoreceptor function.


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Chapter 4
Histology
81
(c)
(a)
FIGURE 4.2 (a) Simple squamous epithelium lines the lumina of vessels, where it permits diffusion. (b) A photomicrograph of this tissue and
(c) a labeled diagram. Simple squamous epithelia that line the lumina of vessels are referred to as endothelia, and that which cover visceral organs are referred to as mesothelia.
Many membranous epithelia are exposed to friction or

harmful substances from the outside environment. For this reason, epithelial tissues have remarkable regenerative abilities. The
mitotic replacement of the outer layer of skin and the lining of
the GI tract, for example, is a continuous process.
Membranous epithelia are histologically classified by the
number of layers of cells and the shape of the cells along the exposed surface. Epithelial tissues that are composed of a single
layer of cells are called simple; those that are layered are said to
be stratified. Squamous cells are flattened; cuboidal cells are cubeshaped; and columnar cells are taller than they are wide.
Simple Epithelia
Simple epithelial tissue is a single cell layer thick and is located
where diffusion, absorption, filtration, and secretion are principal
functions. The cells of simple epithelial tissue range from thin,
flattened cells to tall, columnar cells. Some of these cells have
cilia that create currents for the movement of materials across
cell surfaces. Others have microvilli that increase the surface
area for absorption.
Simple Squamous Epithelium

Simple squamous (skwamus) epithelium is composed of flattened, irregularly shaped cells that are tightly bound together in
a mosaiclike pattern (fig. 4.2). Each cell contains an oval or
squamous: L. squamosus, scaly
spherical central nucleus. This epithelium is adapted for diffusion

and filtration. It occurs in the pulmonary alveoli within the
lungs (where gaseous exchange occurs), in portions of the kidney
(where blood is filtered), on the inside walls of blood vessels, in
the lining of body cavities, and in the covering of the viscera.
The simple squamous epithelium lining the inner walls of blood
and lymphatic vessels is termed endothelium (endo-thele-um)
(fig. 4.2b). That which covers visceral organs and lines body cavities is called mesothelium (meso-thele-um).
Simple Cuboidal Epithelium

Simple cuboidal epithelium is composed of a single layer of
tightly fitted cube-shaped cells (fig. 4.3). This type of epithelium
is found lining small ducts and tubules that have excretory, secretory, or absorptive functions. It occurs on the surface of the
ovaries, forms a portion of the tubules within the kidney, and
lines the ducts of the salivary glands and pancreas.
Simple Columnar Epithelium

Simple columnar epithelium is composed of tall, columnar
cells (fig. 4.4). The height of the cells varies, depending on the
site and function of the tissue. Each cell contains a single nucleus which is usually located near the basement membrane.
endothelium: Gk. endon, within; thelium, to cover

mesothelium: Gk. meso, middle; thelium, to cover
CHAPTER 4
(b)

82
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Lumen of
renal tubule
Kidney
Basement
membrane
Nucleus
CHAPTER 4
(b)
(c)
Ureter
(a)
FIGURE 4.3 (a) Simple cuboidal epithelium lines the lumina of ducts; for example, in the kidneys, where it permits movement of fluids and
ions. (b) A photomicrograph of this tissue and (c) a labeled diagram.
Liver
Stomach
Gallbladder
Large intestine
Small intestine
(b)
(b)
Lumen of small
intestine
Nucleus
Creek
Basement
membrane
(a)
Goblet cell
Cilia
(c)
FIGURE 4.4 (a) Simple columnar epithelium lines the lumen of the digestive tract, where it permits secretion and absorption. (b) A photomicrograph of this tissue and (c) a labeled diagram.


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Chapter 4
Histology
83
Body of
uterus
Uterine tube
Uterine cavity
Ovary
Lumen of
uterine tube
Cilia
(a)
Cell membrane
Vagina
Nucleus
Basement
membrane
(c)
FIGURE 4.5 (a) Simple ciliated columnar epithelium lines the lumen of the uterine tube, where currents generated by the cilia propel the ovum
(egg cell) toward the uterus. (b) a photomicrograph of this tissue and (c) a labeled diagram.
Specialized unicellular glands called goblet cells are scattered

through this tissue at most locations. Goblet cells secrete a lubricative and protective mucus along the free surfaces of the
cells. Simple columnar epithelium is found lining the inside walls
of the stomach and intestine. In the digestive system, it forms a
highly absorptive surface and also secretes certain digestive
chemicals. Within the stomach, simple columnar epithelium has
a tremendous mitotic rate. It replaces itself every 2 to 3 days.
Simple Ciliated Columnar Epithelium

Simple ciliated columnar epithelium is characterized by the presence of cilia along its free surface (fig. 4.5). By contrast, the simple columnar type is unciliated. Cilia produce wavelike
movements that transport materials through tubes or passageways. This type of epithelium occurs in the female uterine tubes
to move the ovum (egg cell) toward the uterus.
Not only do the cilia function to propel the ovum, but recent
evidence indicates that sperm introduced into the female
vagina during sexual intercourse may be moved along the return currents, or eddies, generated by ciliary movement. This greatly enhances the likelihood of fertilization.
Pseudostratified Ciliated Columnar Epithelium

As the name implies, this type of epithelium has a layered appearance (strata = layers). Actually, it is not multilayered (pseudo
= false), because each cell is in contact with the basement membrane. Not all cells are exposed to the surface, however (fig. 4.6).
The tissue appears to be stratified because the nuclei of the cells

are located at different levels. Numerous goblet cells and a ciliated exposed surface are characteristic of this epithelium. It is
found lining the inside walls of the trachea and the bronchial
tubes; hence, it is frequently called respiratory epithelium. Its function is to remove foreign dust and bacteria entrapped in mucus
from the lower respiratory system.
Coughing and sneezing, or simply clearing the throat, are
protective reflex mechanisms for clearing the respiratory passages of obstruction or of inhaled particles that have been trapped in
the mucus along the ciliated lining. The material that is coughed up
consists of the mucus-entrapped particles.
Stratified Epithelia
Stratified epithelia have two or more layers of cells. In contrast
to the single-layered simple epithelia, they are poorly suited for
absorption and secretion. Stratified epithelia have a primarily
protective function that is enhanced by rapid cell divisions. They
are classified according to the shape of the surface layer of cells,
because the layer in contact with the basement membrane is always cuboidal or columnar in shape.
Stratified Squamous Epithelium

Stratified squamous epithelium is composed of a variable number
of cell layers that are flattest at the surface (fig. 4.7). Mitosis occurs only at the deepest layers (see table 5.2). The mitotic rate
CHAPTER 4
(b)
Paras

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Larynx
Trachea
Bronchiole
Lung
Secondary bronchi
(b)
CHAPTER 4
(b)
(a)
Creek
Cilia
Goblet cell
Nucleus
Bronchus
Basement membrane
Connective tissue
(c)
FIGURE 4.6 (a) Pseudostratified ciliated columnar epithelium lines the lumen of the respiratory tract, where it traps foreign material and moves
it away from the pulmonary alveoli of the lungs. (b) a photomicrograph of this tissue and (c) a labeled diagram.
Uterine tube
Ovary
Uterus
Urinary bladder
Cervix
Rectum
Urethra
Vagina
Anus
Waldrop
(b)
(a)
Stratified
squamous
epithelium
(c)
FIGURE 4.7 Stratified squamous epithelium forms the outer layer of skin and the lining of body openings. In the moistened areas, such as in the
vagina (a), it is nonkeratinized, whereas in the epidermis of the skin it is keratinized. (b) a photomicrograph of this tissue and (c) a labeled diagram.


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Parotid gland
Nuclei
Lumen of
parotid duct
Basement
membrane
(b)
(c)
(a)
FIGURE 4.8 (a) Stratified cuboidal epithelium lines the lumina of large ducts like the parotid duct, which drains saliva from the parotid gland.
(b) a photomicrograph of this tissue and (c) a labeled diagram.
approximates the rate at which cells are sloughed off at the surface.
As the newly produced cells grow in size, they are pushed toward
the surface, where they replace the cells that are sloughed off.
Movement of the epithelial cells away from the supportive basement membrane is accompanied by the production of keratin (keratin) (described below), progressive dehydration, and flattening.
There are two types of stratified squamous epithelial tissues: keratinized and nonkeratinized.
1. Keratinized stratified squamous epithelium contains keratin, a protein that strengthens the tissue. Keratin makes
the epidermis (outer layer) of the skin somewhat waterproof
and protects it from bacterial invasion. The outer layers of
the skin are dead, but glandular secretions keep them soft
(see chapter 5).
2. Nonkeratinized stratified squamous epithelium lines the
oral cavity and pharynx, nasal cavity, vagina, and anal
canal. This type of epithelium, called mucosa (myoo-kosa),
is well adapted to withstand moderate abrasion but not
fluid loss. The cells on the exposed surface are alive and are
always moistened.
Stratified squamous epithelium is the first line of defense
against the entry of living organisms into the body. Stratification,
rapid mitotic activity, and keratinization within the epidermis of the skin
are important protective features. An acidic pH along the surfaces of
this tissue also helps to prevent disease. The pH of the skin is between
4.0 and 6.8. The pH in the oral cavity ranges from 5.8 to 7.1, which
tends to retard the growth of microorganisms. The pH of the anal region is about 6, and the pH along the vaginal lining is 4 or lower.
keratin: Gk. keras, horn
Stratified Cuboidal Epithelium

Stratified cuboidal epithelium usually consists of only two or
three layers of cuboidal cells (fig. 4.8). This type of epithelium is
confined to the linings of the large ducts of sweat glands, salivary
glands, and the pancreas, where its stratification probably provides a more robust lining than would simple epithelium.
Transitional Epithelium
Transitional epithelium is similar to nonkeratinized stratified
squamous epithelium except that the surface cells of the former
are large and round rather than flat, and some may have two nuclei (fig. 4.9). Transitional epithelium is found only in the urinary system, particularly lining the cavity of the urinary bladder
and lining the lumina of the ureters. This tissue is specialized to
permit distension (stretching) of the urinary bladder as it fills
with urine. The inner, exposed cells actually transform from
being rounded when the urinary bladder is empty to being somewhat flattened as it distends with urine.
A summary of membranous epithelial tissue is presented in
table 4.1.
Body Membranes
Body membranes are composed of thin layers of epithelial tissue
and, in certain locations, epithelial tissue coupled with supporting connective tissue. Body membranes cover, separate, and support visceral organs and line body cavities. The two basic types of
body membranes, mucous membranes and serous membranes,
are described in detail in chapter 2 under the heading Body
Cavities and Membranes (see p. 41).
CHAPTER 4
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Ureter
CHAPTER 4
(b)
Urinary
bladder
Lumen of
urinary bladder
(b)
Transitional
epithelium
Urethra
Smooth
muscle
tissue
(c)
(a)
FIGURE 4.9 (a) Transitional epithelium lines the lumina of the ureters, part of the urethra and the cavity of the urinary bladder, where it permits
distention. (b) a photomicrograph of this tissue and (c) a labeled diagram.
TABLE 4.1 Summary of Membranous Epithelial Tissue

Type
Structure and Function
Location
Simple Epithelia
Single layer of cells; function varies with type
Covering visceral organs; linings of body cavities,

tubes, and ducts
Simple squamous epithelium
Single layer of flattened, tightly bound cells;

diffusion and filtration
Capillary walls; pulmonary alveoli of lungs;

covering visceral organs; linings of body cavities
Simple cuboidal epithelium
Single layer of cube-shaped cells; excretion,

secretion, or absorption
Surface of ovaries; linings of renal tubules, salivary

ducts, and pancreatic ducts
Simple columnar epithelium
Single layer of nonciliated, tall, column-shaped cells;

protection, secretion, and absorption
Lining of most of GI tract
Simple ciliated columnar epithelium
Single layer of ciliated, column-shaped cells;

transportive role through ciliary motion
Lining of uterine tubes
Pseudostratified ciliated columnar epithelium
Single layer of ciliated, irregularly shaped cells; many

goblet cells; protection, secretion, ciliary
movement
Lining of respiratory passageways
Two or more layers of cells; function varies with type
Epidermal layer of skin; linings of body openings,

ducts, and urinary bladder
Stratified squamous epithelium (keratinized)
Numerous layers containing keratin, with outer

layers flattened and dead; protection
Epidermis of skin
Stratified squamous epithelium (nonkeratinized)
Numerous layers lacking keratin, with outer layers

moistened and alive; protection and pliability
Linings of oral and nasal cavities, vagina, and anal

canal
Stratified cuboidal epithelium
Usually two layers of cube-shaped cells;

strengthening of luminal walls
Large ducts of sweat glands, salivary glands, and

pancreas
Transitional epithelium
Numerous layers of rounded, nonkeratinized cells;

distension
Walls of ureters, part of urethra, and urinary

bladder


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Secretion
Lumen
Mucus
Liver
Stomach
Cell membrane
Gallbladder
Golgi complex
Small intestine
CHAPTER 4
Large intestine
Rough endoplasmic
reticulum
Creek
(a)
(b)
(c)
FIGURE 4.10 A goblet cell is a unicellular gland that secretes mucus, which lubricates and protects surface linings. (a) Goblet cells are abundant in the columnar epithelium lining the lumen of the small intestine. (b) A photomicrograph of a goblet cell and (c) a labeled diagram.
Glandular Epithelia
As tissues develop in the embryo, tiny invaginations (infoldings)
or evaginations (outfoldings) of membranous epithelia give rise to
specialized secretory structures called exocrine (ekso-krin) glands.
These glands remain connected to the epithelium by ducts, and
their secretions pass through the ducts onto body surfaces or into
body cavities. Exocrine glands should not be confused with endocrine glands, which are ductless, and which secrete their products (hormones) into the blood or surrounding extracellular fluid.
Exocrine glands within the skin include oil (sebaceous) glands,
sweat glands, and mammary glands. Exocrine glands within the
digestive system include the salivary and pancreatic glands.
Exocrine glands are classified according to their structure
and how they discharge their products. Classified according to
structure, there are two types of exocrine glands, unicellular and
multicellular glands.
1. Unicellular glands are single-celled glands, such as goblet
cells (fig. 4.10). They are modified columnar cells that
occur within most epithelial tissues. Goblet cells are found
in the epithelial linings of the respiratory and digestive systems. The mucus secretion of these cells lubricates and protects the surface linings.
2. Multicellular glands, as their name implies, are composed

of both secretory cells and cells that form the walls of the
ducts. Multicellular glands are classified as simple or compound glands. The ducts of the simple glands do not
branch, whereas those of the compound type do (fig. 4.11).
Multicellular glands are also classified according to the
shape of their secretory portion. They are identified as
tubular glands if the secretory portion resembles a tube and
as acinar glands if the secretory portion resembles a flask.
Multicellular glands with a secretory portion that resembles both a tube and a flask are termed tubuloacinar glands.
means by which they release their product (fig. 4.12).
1. Merocrine (mero-krin) glands are those that secrete a watery substance through the cell membrane of the secretory
cells. Salivary glands, pancreatic glands, and certain sweat
glands are of this type.
2. Apocrine (apo-krin) glands are those in which the secretion accumulates on the surface of the secretory cell; then,
a portion of the cell, along with the secretion, is pinched
off to be discharged. Mammary glands are of this type.
merocrine: Gk. meros, part; krinein, to separate

exocrine: Gk. exo, outside; krinein, to separate
apocrine: Gk. apo, off; krinein, to separate

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Duct
Secretory cells
Simple
branched tubular
Simple
coiled tubular
Simple acinar
Simple
branched acinar
CHAPTER 4
Simple tubular
Compound tubular
Compound acinar
Compound tubuloacinar
FIGURE 4.11 Structural classification of multicellular exocrine glands. The ducts of the simple glands either do not branch or have few
branches, whereas those of the compound glands have multiple branches.
Secretion
Disintegrating cell
and its contents
discharged
with secretion
Intact cell
New cell
Merocrine gland
FIGURE 4.12 Examples of multicellular exocrine glands.
Apocrine gland
Cr
ee
k
Pinched-off
portion of cell
discharged
with secretion
Holocrine gland


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TABLE 4.2 Summary of Glandular Epithelial Tissue

Classification of Exocrine Glands by Structure
Type
Example
Lubricate and protect

Protect, cool body, lubricate, aid in digestion, maintain
body homeostasis
Goblet cells of digestive, respiratory, urinary, and

reproductive systems
Sweat glands, digestive glands, liver, mammary glands,
sebaceous glands
A. Simple
1. Tubular
2. Branched tubular
3. Coiled tubular
4. Acinar
5. Branched acinar
Aid in digestion
Protect, aid in digestion
Regulate temperature
Provide additive for spermatozoa
Condition skin
Intestinal glands
Uterine glands, gastric glands
Certain sweat glands
Seminal vesicles of male reproductive system
Sebaceous glands of the skin
B. Compound
1. Tubular
2. Acinar
Lubricate urethra of male, assist body digestion

Provide nourishment for infant, aid in digestion
Bulbourethral glands of male reproductive system, liver

Mammary glands, salivary glands (sublingual and
submandibular)
Salivary gland (parotid), pancreas
I. Unicellular
II. Multicellular
3. Tubuloacinar
Aid in digestion
Classification of Exocrine Glands by Mode of Secretion

Type
Description of Secretion
Example
Merocrine glands
Watery secretion for regulating temperature or enzymes

that promote digestion
Salivary and pancreatic glands, certain sweat glands
Apocrine glands
Portion of secretory cell and secretion are discharged; provides

nourishment for infant, assists in regulating temperature
Mammary glands, certain sweat glands
Holocrine glands
Entire secretory cell with enclosed secretion is discharged;

conditions skin
3. Holocrine (holo-krin) glands are those in which the entire

secretory cell is discharged, along with the secretory product. An example of a holocrine gland is an oil-secreting
(sebaceous) gland of the skin (see chapter 5).
A summary of glandular epithelial tissue is presented in
table 4.2.
Knowledge Check
4. List the functions of simple squamous epithelia.
5. What are the three types of columnar epithelia? What do
they have in common? How are they different?
6. What are the two types of stratified squamous epithelia and
how do they differ?
7. Distinguish between unicellular and multicellular glands.
Explain how multicellular glands are classified according to
their mechanism of secretion.
8. In what ways are mammary glands and certain sweat
glands similar?
holocrine: Gk. holos, whole; krinein, to separate
CONNECTIVE TISSUE
Connective tissue is divided into subtypes according to the matrix
that binds the cells. Connective tissue provides structural and
metabolic support for other tissues and organs of the body.
Objective 7
Describe the general characteristics, locations,

and functions of connective tissue.
Objective 8
Explain the functional relationship between

embryonic and adult connective tissue.
Objective 9
List the various ground substances, fiber types,

and cells that constitute connective tissue and explain their
functions.
Characteristics and Classification

of Connective Tissue
Connective tissue is the most abundant tissue in the body. It supports other tissues or binds them together and provides for the
metabolic needs of all body organs. Certain types of connective
tissue store nutritional substances; other types manufacture protective and regulatory materials.
CHAPTER 4
Function

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Mesenchymal
cell
CHAPTER 4
Matrix
(a)
(b)
FIGURE 4.13 Mesenchyme is a type of embryonic connective tissue that can migrate and give rise to all other kinds of connective tissue.
(a) It is found within an early developing embryo and (b) consists of irregularly shaped cells lying in a jellylike homogeneous matrix.
Although connective tissue varies widely in structure and

function, all types of connective tissue have similarities. With
the exception of mature cartilage, connective tissue is highly vascular and well nourished. It is able to replicate and, by so doing,
is responsible for the repair of body organs. Unlike epithelial
tissue, which is composed of tightly fitted cells, connective tissue
contains considerably more matrix (intercellular material) than
cells. Connective tissue does not occur on free surfaces of body
cavities or on the surface of the body, as does epithelial tissue.
Furthermore, connective tissue is embryonically derived from
mesoderm, whereas epithelial tissue derives from ectoderm,
mesoderm, and endoderm.
The classification of connective tissue is not exact, and
several schemes have been devised. In general, however, the
various types are named according to the kind and arrangement
of the matrix. The following are the basic kinds of connective
tissues:
A. Embryonic connective tissue
B. Connective tissue proper
1. Loose (areolar) connective tissue
2. Dense regular connective tissue
3. Dense irregular connective tissue
4. Elastic connective tissue
5. Reticular connective tissue
6. Adipose tissue
C. Cartilage
1. Hyaline cartilage
2. Fibrocartilage
3. Elastic cartilage
D. Bone tissue
E. Blood (vascular tissue)
Embryonic Connective Tissue

The embryonic period of development, which lasts 6 weeks
(from the start of the third to the end of the eighth week), is
characterized by extensive tissue differentiation and organ formation. At the beginning of the embryonic period, all connective
tissue looks alike and is referred to as mesenchyme (mezen-km).
Mesenchyme is undifferentiated embryonic connective tissue
that is derived from mesoderm. It consists of irregularly shaped
cells surrounded by large amounts of a homogeneous, jellylike
matrix (fig. 4.13). In certain periods of development, mesenchyme migrates to predisposed sites where it interacts with
other tissues to form organs. Once mesenchyme has completed
its embryonic migration to a predetermined destination, it differentiates into all other kinds of connective tissue.
Some mesenchymal-like tissue persists past the embryonic
period in certain sites within the body. Good examples are the
undifferentiated cells that surround blood vessels and form fibroblasts if the vessels are traumatized. Fibroblasts assist in healing wounds (see chapter 5).
Another kind of prenatal connective tissue exists only in
the fetus (the fetal period is from 9 weeks to birth) and is called
mucous connective tissue or Whartons jelly. It gives a turgid consistency to the umbilical cord.
Connective Tissue Proper

Connective tissue proper has a loose, flexible matrix, frequently
called ground substance. The most common cell within connec-
Whartons jelly: from Thomas Wharton, English anatomist, 161473


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Pectoralis
major
muscle
Elastic fiber
Collagenous
fiber
Mast cell
Fibroblast
(b)
(a)
Paras
Fascia
FIGURE 4.14 Loose connective tissue is packing and binding tissue that surrounds muscles (a), nerves, and vessels and binds the skin to
the underlying muscles. (b) A photomicrograph of the tissue and (c) a labeled diagram.
tive tissue proper is called a fibroblast (fibro-blast). Fibroblasts

are large, star-shaped cells that produce collagenous (ko-lajenus), elastic, and reticular (re-tikyoo-lar) fibers. Collagenous
fibers are composed of a protein called collagen (kola-jen); they
are flexible, yet they have tremendous strength. Elastic fibers are
composed of a protein called elastin, which provides certain tissues with elasticity. Collagenous and elastic fibers may be either
sparse and irregularly arranged, as in loose connective tissue, or
tightly packed, as in dense connective tissue. Tissues with loosely
arranged fibers generally form packing material that cushions and
protects various organs, whereas those that are tightly arranged
form the binding and supportive connective tissues of the body.
Resilience in tissues that contain elastic fibers is extremely important for several physical functions of the body. Consider, for
example, that elastic fibers are found in the walls of large arteries and in
the walls of the lower respiratory passageways. As these walls are expanded by blood moving through vessels or by inspired air, the elastic
fibers must first stretch and then recoil. This maintains the pressures of
the fluid or air moving through the lumina, thus ensuring adequate flow
rates and rates of diffusion through capillary and lung surfaces.
Reticular fibers reinforce by branching and joining to

form a delicate lattice or reticulum. Reticular fibers are common
in lymphatic glands, where they form a meshlike center called
the stroma.
Six basic types of connective tissue proper are generally

recognized. These tissues are distinguished by the consistency of
the ground substance and the type and arrangement of the reinforcement fibers.
Loose Connective (Areolar) Tissue

Loose connective tissue is distributed throughout the body as a
binding and packing material. It binds the skin to the underlying
muscles and is highly vascular, providing nutrients to the skin.
Loose connective tissue that binds skin to underlying muscles is
known as fascia (fashe-a). It also surrounds blood vessels and
nerves, where it provides both protection and nourishment. Specialized cells called mast cells are dispersed throughout the loose
connective tissue surrounding blood vessels. Mast cells produce heparin (hepa-rin), an anticoagulant that prevents blood from clotting
within the vessels. They also produce histamine, which is released
during inflammation. Histamine acts as a powerful vasodilator.
The cells of loose connective tissue are predominantly fibroblasts, with collagenous and elastic fibers dispersed throughout the ground substance (fig. 4.14). The irregular arrangement
of this tissue provides flexibility, yet strength, in any direction. It
is this tissue layer, for example, that permits the skin to move
when a part of the body is rubbed.
collagen: Gk, kolla, glue

elastin: Gk. elasticus, to drive
reticular: L. rete, net or netlike
stroma: Gk. stroma, a couch or bed
fascia: L. fascia, a band or girdle

heparin: Gk. hepatos, the liver
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Fibroblast
CHAPTER 4
Tendon of
long head
of biceps brachii m.
Collagenous
fibers
Biceps
brachii m.
(a)
Tendon
of short
head of
biceps
brachii m.
(b)
(c)
Paras
FIGURE 4.15 Dense regular connective tissue forms the strong and highly flexible tendons (a) and ligaments. (b) A photomicrograph of the
tissue and (c) a labeled diagram.
Much of the fluid of the body is found within loose connective

tissue and is called interstitial fluid (tissue fluid). Sometimes
excessive tissue fluid accumulates, causing a swollen condition
called edema (e-dema). Edema is a symptom of numerous dysfunctions or disease processes.
Dense Regular Connective Tissue

Dense regular connective tissue is characterized by large amounts
of densely packed collagenous fibers that run parallel to the direction of force placed on the tissue during body movement. Because this tissue is silvery white in appearance, it is sometimes
called white fibrous connective tissue.
Dense regular connective tissue occurs where strong, flexible support is needed (fig. 4.15). Tendons, which attach muscles
to bones and transfer the forces of muscle contractions, and ligaments, which connect bone to bone across articulations, are
composed of this type of tissue.
Trauma to ligaments, tendons, and muscles are common
sports-related injuries. A strain is an excessive stretch of the
tissue composing the tendon or muscle, with no serious damage. A
sprain is a tearing of the tissue of a ligament and may be slight, moderate, or complete. A complete tear of a major ligament is especially
painful and disabling. Ligamentous tissue does not heal well because it has a poor blood supply. Surgical reconstruction is generally
needed for the treatment of a severed ligament.
tendon: L. tendere, to stretch

ligament: L. ligare, bind
Dense Irregular Connective Tissue

Dense irregular connective tissue is characterized by large
amounts of densely packed collagenous fibers that are interwoven to provide tensile strength in any direction. This tissue
is found in the dermis of the skin and the submucosa of the
GI tract. It also forms the fibrous capsules of organs and joints
(fig. 4.16).
Elastic Connective Tissue

Elastic connective tissue is composed primarily of elastic fibers
that are irregularly arranged and yellowish in color (fig. 4.17).
They can be stretched to one and a half times their original
lengths and will snap back to their former size. Elastic connective tissue is found in the walls of large arteries, in portions of the
larynx, and in the trachea and bronchial tubes of the lungs. It is
also present between the arches of the vertebrae that make up
the vertebral column.
Reticular Connective Tissue

Reticular connective tissue is characterized by a network of reticular fibers woven through a jellylike matrix (fig. 4.18). Certain
specialized cells within reticular tissue are phagocytic (fago-sitik)
(macrophages) and therefore can ingest foreign materials. The
liver, spleen, lymph nodes, and bone marrow contain reticular
connective tissue.


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(b)
(c)
(a)
FIGURE 4.16 Dense irregular connective tissue forms joint capsules (a) that contain synovial fluid for lubricating movable joints. (b) A photomicrograph of the tissue and (c) a labeled diagram.
Tunica
intima
(inner coat)
Elastic fibers
Endothelial cells
Elastic tissue
Fibroblast
(b)
Paras
(c)
(a)
FIGURE 4.17 Elastic connective tissue permits stretching of a large artery (a) as blood flows through. (b) A photomicrograph of the tissue and
(c) a labeled diagram.
Adipose Tissue
Adipose tissue is a specialized type of loose fibrous connective
tissue that contains large quantities of adipose cells, or
adipocytes (ad-po-sts). Adipose cells form from mesenchyme
adipose: L. adiposus, fat
and, for the most part, are formed prenatally and during the
first year of life. Adipose cells store droplets of fat within their
cytoplasm, causing them to swell and forcing their nuclei to
one side (fig. 4.19).
Adipose tissue is found throughout the body but is concentrated around the kidneys, in the hypodermis of the skin,
on the surface of the heart, surrounding joints, and in the
CHAPTER 4
Collagenous
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Reticular
fibers
Reticular cell
Nucleus of
reticular cell
CHAPTER 4
(b)
(c)
Spleen
(a)
Paras
FIGURE 4.18 Reticular connective tissue forms the stroma, or framework, of such organs as the spleen (a), liver, thymus, and lymph nodes.
(b) A photomicrograph of this tissue and (c) a labeled diagram.
Nucleus of
adipose cell
Fat droplet
Dermis
Cytoplasm
(b)
Hypodermis
(c)
Hair
follicle
(a)
Paras
FIGURE 4.19 Adipose tissue is abundant in the hypodermis of the skin (a) and around various internal organs. (b) A photomicrograph of the
breasts of sexually mature females. Fat functions not only as a

food reserve, but also supports and protects various organs. It
is a good insulator against cold because it is a poor conductor
of heat.
Excessive fat can be unhealthy by placing a strain on the heart
and perhaps causing early death. For these reasons, good exercise programs and sensible diets are extremely important. Adipose
tissue can also retain lipid-soluble, environmental pollutants that are
ingested or absorbed through the skin. Dieting eliminates the fat
stored within adipose tissue but not the tissue itself.
The surgical procedure of suction lipectomy may be used to

remove small amounts of adipose tissue from localized body areas
such as the breasts, abdomen, buttocks, and thighs. Suction lipectomy is used for cosmetic purposes rather than as a treatment for
obesity, and the risks for potentially detrimental side effects need to
be seriously considered. Potential candidates should be between 30
and 40 years old and only about 15 to 20 pounds overweight. They
should also have good skin elasticity.
The characteristics, functions, and locations of connective

tissue proper are summarized in table 4.3.


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TABLE 4.3 Summary of Connective Tissue Proper

Location
Loose connective (areolar) tissue
Predominantly fibroblast cells with lesser amounts of

collagen and elastin proteins; binds organs, holds
tissue fluids
Surrounding nerves and vessels, between muscles,

beneath the skin
Dense regular connective tissue
Densely packed collagenous fibers that run parallel to the

direction of force; provides strong, flexible support
Tendons, ligaments
Dense irregular connective tissue
Densely packed collagenous fibers arranged in a tight

interwoven pattern; provides tensile strength in any
direction
Dermis of skin, fibrous capsules of organs and joints,

periosteum of bone
Elastic connective tissue
Predominantly irregularly arranged elastic fibers; supports,

provides framework
Large arteries, lower respiratory tract, between the

arches of vertebrae
Reticular connective tissue
Reticular fibers that form a supportive network; stores,

performs phagocytic function
Lymph nodes, liver, spleen, thymus, bone marrow
Adipose tissue
Adipose cells; protects, stores fat, insulates
Hypodermis of skin, surface of heart, omentum, around

kidneys, back of eyeball, surrounding joints
Cartilage
Cartilage (kart-lij) consists of cartilage cells, or chondrocytes
(kondro-sts), and a semisolid matrix that imparts marked elastic
properties to the tissue. It is a supportive and protective connective tissue that is frequently associated with bone. Cartilage
forms a precursor to one type of bone and persists at the articular
surfaces on the bones of all movable joints.
The chondrocytes within cartilage may occur singly but
are frequently clustered. Chondrocytes occupy cavities, called lacunae
(la-kyoonesingular lacuna), within the matrix. Most cartilage is
surrounded by a dense irregular connective tissue called perichondrium (per-kondre-um). Cartilage at the articular surfaces
of bones (articular cartilage) lacks a perichondrium. Because mature cartilage is avascular, it must receive nutrients through diffusion from the perichondrium and the surrounding tissue. For this
reason, cartilaginous tissue has a slow rate of mitotic activity; if
damaged, it heals with difficulty.
There are three kinds of cartilage: hyaline (hia-ln) cartilage,
fibrocartilage, and elastic cartilage. They are distinguished by the
type and amount of fibers embedded within the matrix.
Hyaline Cartilage
Hyaline cartilage, commonly called gristle, has a homogeneous,
bluish-staining matrix in which the collagenous fibers are so fine
that they can be observed only with an electron microscope.
When viewed through a light microscope, hyaline cartilage has a
clear, glassy appearance (fig. 4.20).
Hyaline cartilage is the most abundant cartilage within the
body. It covers the articular surfaces of bones, supports the tubular trachea and bronchi of the respiratory system, reinforces the
nose, and forms the flexible bridge, called costal cartilage, be-
lacuna: L. lacuna, hole or pit

hyaline: Gk. hyalos, glass
tween the anterior portion of each of the first 10 ribs and the
sternum. Most of the bones of the body form first as hyaline cartilage and later become bone in a process called endochondral
ossification.
Fibrocartilage
Fibrocartilage has a matrix that is reinforced with numerous collagenous fibers (fig. 4.21). It is a durable tissue adapted to withstand tension and compression. It is found at the symphysis
pubis, where the two pelvic bones articulate, and between the
vertebrae as intervertebral discs. It also forms the cartilaginous
wedges within the knee joint, called menisci (see chapter 8).
By the end of the day, the intervertebral discs of the vertebral
column are somewhat compacted. So a person is actually
slightly shorter in the evening than in the morning, following a recuperative rest. Aging, however, brings with it a gradual compression of
the intervertebral discs that is irreversible.
Elastic Cartilage
Elastic cartilage is similar to hyaline cartilage except for the presence of abundant elastic fibers that make elastic cartilage very
flexible without compromising its strength (fig. 4.22). The numerous elastic fibers also give it a yellowish appearance. This tissue is found in the outer ear, portions of the larynx, and in the
auditory canal.
The three types of cartilage are summarized in table 4.4.
Bone Tissue
Bone tissue is the most rigid of all the connective tissues. Unlike
cartilage, bone tissue has a rich vascular supply and is the site of
considerable metabolic activity. The hardness of bone is largely
due to the calcium phosphate (calcium hydroxyapatite) deposited
within the intercellular matrix. Numerous collagenous fibers, also
embedded within the matrix, give bone some flexibility.
CHAPTER 4
Type

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Lacuna
Intercellular
matrix
Chondrocyte
Thyroid
cartilage
Larynx
CHAPTER 4
(b)
(c)
Cricoid
cartilage
Tracheal
cartilages
(a)
Paras
FIGURE 4.20 Hyaline cartilage is the most abundant cartilage in the body. It occurs in places such as the larynx (a), trachea, portions of the
rib cage, and embryonic skeleton. (b) A photomicrograph of the tissue and (c) a labeled diagram.
Lacuna
Chondrocyte
Intercellular
matrix
Collagenous
fibers
(b)
(c)
(a)
FIGURE 4.21 Fibrocartilage is located at the symphysis pubis, within the knee joints, and between the vertebrae as the intervertebral discs
(a). A photomicrograph of the tissue is shown in (b) and a labeled diagram in (c).


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Histology
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Lacuna
Chondrocyte
Elastic fibers
Auricular
cartilage
(b)
Paras
FIGURE 4.22 Elastic cartilage gives support to the outer ear (a), auditory canal, and parts of the larynx. A photomicrograph of the tissue is
shown in (b) and a labeled diagram in (c).
TABLE 4.4 Summary of Cartilage

Type
Location
Hyaline cartilage
Homogeneous matrix with extremely fine collagenous fibers;

provides flexible support, protects, is precursor to bone
Articular surfaces of bones, nose, walls of respiratory passages,

fetal skeleton
Fibrocartilage
Abundant collagenous fibers within matrix; supports, withstands

compression
Symphysis pubis, intervertebral discs, knee joint
Elastic cartilage
Abundant elastic fibers within matrix; supports, provides flexibility
Framework of outer ear, auditory canal, portions of larynx
When bone is placed in a weak acid, the calcium salts dissolve

away and the bone becomes pliable. It retains its basic shape
but can be easily bent and twisted (fig. 4.23). In calcium deficiency
diseases, such as rickets, the bone tissue becomes pliable and
bends under the weight of the body (see fig. 5.11).
Based on porosity, bone tissue is classified as either compact or spongy, and most bones have both types (fig. 4.24). Compact (dense) bone tissue constitutes the hard outer portion of a
bone, and spongy (cancellous) bone tissue constitutes the porous,
highly vascular inner portion. The outer surface of a bone is covered by a connective tissue layer called the periosteum that serves
as a site of attachment for ligaments and tendons, provides protection, and gives durable strength to the bone. Spongy bone tissue makes the bone lighter and provides a space for red bone
marrow, where blood cells are produced.
In compact bone tissue, mature bone cells, called
osteocytes, are arranged in concentric layers around a central
FIGURE 4.23 A bone soaked in a weak acid, such as the acidic
acid in vinegar, demineralizes and becomes flexible.
CHAPTER 4
(a)
(c)

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Cell
membrane
Fluid-filled
vesicle
Fluid
Nucleolus
Nucleus
Cytoplasm
CHAPTER 3
(a) Pinocytosis
Cell
membrane
Vesicle
Particle
Phagocytized
particle
Nucleolus Nucleus
(b) Phagocytosis
FIGURE 3.12 Pinocytosis and phagocytosis compared. (a) During pinocytosis, the cell takes in a minute droplet of fluid from its surroundings.
(b) During phagocytosis, a solid particle is engulfed and ingested through the cell membrane.
Microtubules
(a)
(b)
FIGURE 3.13 (a) An electron micrograph showing microtubules forming a type of cytoskeleton (about 30,000). (b) A diagram of a microtubule showing the precisely arranged globular proteins of which they are composed.


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Flagellar
or ciliary
membrane
Cell
membrane
CHAPTER 3
(a)
Cilia
Debris
Goblet
cell
Flagellum
FIGURE 3.15 An electron micrograph of ciliated cells that line the

lumen of the uterine tube (640).
(b)
(c)
Creek
FIGURE 3.14 (a) Cilia and flagella are similar in the structural
arrangement of their microtubules. (b) A sperm cell (spermatozoon)
has a single flagellum for propulsion. (c) Cilia produce a wavelike
motion to move particles toward the outside of the body.
Cell Nucleus
The spherical nucleus is usually located near the center of the
cell (fig. 3.16). It is the largest structure of the cell and contains
the genetic material that determines cellular structure and controls cellular activity.
Most cells contain a single nucleus. Certain cells, however,
such as skeletal muscle cells, are multinucleated. The long skeletal muscle fibers contain so much cytoplasm that several governing centers are necessary. Other cells, such as mature red blood
cells, lack nuclei. These cells are limited to certain types of
chemical activities and are not capable of cell division.
The nucleus is enclosed by a bilayered nuclear membrane
(nuclear envelope) (fig. 3.16). The narrow space between the
inner and outer layers of the nuclear membrane is called the nucleolemma cisterna (sis-terna). Minute nuclear pores are located along the nuclear membrane. These openings are lined
with proteins that act as selective gates, allowing certain molecules, such as proteins, RNA, and protein-RNA complexes, to
move between the nucleoplasm and the cytoplasm.
Two important structures within the nucleoplasm of the

nucleus determine what a cell will look like and what functions
it will perform:
1. Nucleoli. Nucleoli (noo-kleo-li) are small, nonmembranous spherical bodies composed largely of protein and
RNA. It is thought that they function in the production of
ribosomes. As ribosomes are formed, they migrate through
the nuclear membrane into the cytoplasm.
2. Chromatin. Chromatin (kroma-tin) is a coiled, threadlike mass. It is the genetic material of the cell and consists
principally of protein and DNA molecules. When a cell
begins to divide, the chromatin shortens and thickens into
rod-shaped structures called chromosomes (kromo-somz)
(figs. 3.17 and 3.18). Each chromosome carries thousands
of genes that determine the structure and function of a cell.
Knowledge Check
9. Describe the composition and specializations of the cell
membrane. Discuss the importance of the selective permeability of the cell membrane.
10. Describe the various kinds of movements across the cell
membrane. Which are passive and which are active?
11. Describe the structure and function of the following cytoplasmic organelles: rough endoplasmic reticulum, Golgi
complex, lysosomes, and mitochondria.
12. Distinguish between the nucleus and nucleoli.
13. Distinguish between chromatin and chromosomes.

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Nuclear pore
Nuclear membrane
Nucleolus
Outer layer
Nucleolemma
cisterna
Nucleus
CHAPTER 3
Inner layer
(a)
Chromatin
(b)
FIGURE 3.16 (a) An electron micrograph of the cell nucleus (about 20,000). The nucleus contains a nucleolus and masses of chromatin.
(b) The double-layered nuclear membrane has pores that permit substances to pass between nucleus and cytoplasm.
FIGURE 3.17 A color-enhanced light micrograph showing the full

complement of male chromosomes arranged in numbered homologous pairs.
FIGURE 3.18 The structure of a chromosome after DNA replication, in which it consists of two identical strands, or chromatids.


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Chapter 3
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CELL CYCLE
A cell cycle consists of growth, synthesis, and mitosis. Growth is
the increase in cellular mass resulting from metabolism. Synthesis
is the production of DNA and RNA to regulate cellular activity. Mitosis is the division of the nucleus and cytoplasm of a cell that results in the formation of two daughter cells.
Objective 12
Describe the structure of DNA and RNA
molecules.
Objective 13
Discuss genetic transcription and protein
synthesis.
CHAPTER 3
Objective 14
List the stages of mitosis and discuss the

events of each stage.
Objective 15
Discuss the significance of mitosis.
Cellular replication is one of the principal concepts of biology.

Through the process of cellular division called mitosis (mi-tosis),
a multicellular organism can develop and be maintained. Mitosis
enables body growth and the replacement of damaged, diseased,
or worn-out cells. The process ensures that each daughter cell
will have the same number and kind of chromosomes as the original parent cell.
In an average healthy adult, over 100 billion cells will die and
be mitotically replaced during a 24-hour period. This represents a replacement of about 2% of the body mass each day. Some of the most
mitotically active sites are the outer layer of skin, the bone marrow,
the internal lining of the digestive tract, and the liver.
Before a cell can divide, it must first duplicate its chromosomes so that the genetic traits can be passed to the succeeding
generations of cells. A chromosome consists of a coiled deoxyribonucleic acid (DNA) molecule that is complexed with protein.
As mentioned previously, chromosomes are formed by the shortening and thickening of the chromatin within the nucleus when
the cell begins to divide, at which time they are clearly visible
under the compound microscope. There are 23 pairs of chromosomes in each human body (somatic) cell and approximately
20,000 genes are positioned on each chromosome.
Chromosomes are of varied lengths and shapessome
twisted, some rodlike. During mitosis, they shorten and condense, each pair assuming a characteristic shape (see fig. 3.17).
On the chromosome is a small, buttonlike body called a centromere to which are attached the spindle fibers that direct the
chromosome toward the pole of the cell during mitosis.
Structure of DNA
The DNA molecule is frequently called a double helix because of
its resemblance to a spiral ladder (fig. 3.19). The sides of the
DNA molecule are formed by alternating units of the sugar de-
mitosis: Gk. mitos, thread
FIGURE 3.19 The double-helix structure of DNA. Each strand of

the helix contains only four kinds of organic bases (A, T, C, and G).
oxyribose and phosphoric acid called the phosphate group. The rungs
of the molecule are composed of pairs of nitrogenous bases. The
ends of each nitrogenous base are attached to the deoxyribosephosphate units. There are only four types of nitrogenous bases
in a DNA molecule: adenine (A), thymine (T), cytosine (C),
and guanine (G).
The basic structural units of the DNA molecule are called
nucleotides. Each nucleotide consists of a molecule of deoxyribose, a phosphate group, and one of the four nitrogenous bases.
Thus, there is a nucleotide type for each of the four bases.

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G
T
C
Region of parental DNA helix.

(Both backbones are light.)
C
CHAPTER 3
G
C
C
G
G
Region of replication. Parental
DNA is unzipped and new
nucleotides are pairing with
those in parental strands.
A
A
T
T
A
G
G
G
T
C
T
G
Region of completed replication.

Each double helix is composed
of an old parental strand (light
purple) and a new daughter
strand (dark purple). The two
DNA molecules formed are
identical to the original DNA
helix and to one another.
C
C
G
G
C
G
C
C
T
C
T
A
T
FIGURE 3.20 The replication of DNA. Each new double helix is composed of one old and one new strand. The sequence of bases of each of
the new molecules is identical to that of the parent DNA because of complementary base pairing.
The pairing of the nitrogenous bases of the nucleotides is

highly specific. The molecular configuration of each base is such
that adenine always pairs with thymine and cytosine always pairs
with guanine. The hydrogen bonds between these bases are relatively weak and can be easily split during cellular division
(fig. 3.20). During division, the sequence of bases along the sides
of the DNA molecule serves as a template that determines the
sequence along each new strand.
James Watson and Francis Crick, who devised the doublehelix model, first described their vision of DNA in 1953, in the
journal Nature (see table 1.2). The closing sentence of their brief arti-
cle (a mere 900 words) is a marvel of humility and restraint: It has

not escaped our notice that the specific pairing we have
postulated . . . immediately suggests a possible copying mechanism for the genetic material.
Structure of RNA and RNA Synthesis

In the process of protein synthesis, DNA produces a messenger
molecule of RNA of complementary structure to transport the
genetic information. Like DNA, RNA consists of long chains of
nucleotides joined together by sugar-phosphate bonds. However,


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Chapter 3
Cytology
67
A
G
RNA
C
G
G
T
DNA
C
C
G
C
G
G
C
G
G
G
G
C
G
A
C
U
A
C
T
G
C
G
A
U
C
C
G
C
C
G
G
C
FIGURE 3.21 Differences between the nitrogenous bases and
sugars in DNA and RNA.
G
A
U
A
U
as shown in Fig. 3.21, nucleotides in RNA differ from those in

DNA in the following ways:
A ribonucleotide contains the sugar ribose (instead of
deoxyribose).
The base uracil is present in place of thymine.
RNA is composed of a single polynucleotide strand; it is
not double-stranded like DNA.
RNA is considerably shorter than DNA.
Four types of RNA are produced within the nucleus, each with a
different composition and function:
1. Precursor messenger RNA (pre-mRNA), which is altered within the nucleus (through cutting and splicing) to
form mRNA;
2. Messenger RNA (mRNA), which contains the code for
the synthesis of specific proteins;
3. Transfer RNA (tRNA), which transfers amino acids and
which is needed for decoding the genetic message contained in mRNA; and
4. Ribosomal RNA (rRNA), which forms part of the structure of ribosomes.
The DNA that codes for rRNA synthesis is located in the nucleolus. Pre-mRNA and tRNA synthesis is controlled by DNA located elsewhere in the nucleus.
Genetic TranscriptionRNA Synthesis

During cell division, the chromosomes are inactive packages of
DNA. The genes do not become active until the chromosomes
A
C
G
T
G
U
G
G
G
T
C
FIGURE 3.22 RNA synthesis (genetic transcription). Notice that

only one of the two DNA strands is used to form a single-stranded
molecule of RNA.
unravel. Active DNA directs the metabolism of the cell indirectly through its regulation of RNA and protein synthesis.
One gene codes for one polypeptide chain. Each gene is a
strand of DNA that is several thousand nucleotide pairs long. In
order for the genetic code to be translated for the synthesis of
specific proteins, the DNA code must first be transcribed into an
RNA code (fig. 3.22). This is accomplished by DNA-directed
RNA synthesis, or genetic transcription.
During RNA synthesis, the enzyme RNA polymerase breaks
the weak hydrogen bonds between paired DNA bases. This does
not occur throughout the length of DNA, but only in the regions that are to be transcribed (there are base sequences that
code for start and stop). Double-stranded DNA, therefore,
separates in these regions so that the freed bases can pair with
the complementary RNA nucleotide bases that are freely available in the nucleus.
CHAPTER 3

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G
G
T
C
T
T
C
C
C
DNA
double
helix
C
T
G
G
Transcription
DNA coding strand
Messenger RNA
Translation
CHAPTER 3
Codon 1
Codon 2
Codon 3
Codon 4
Codon 5
Codon 6
Codon 7
Methionine
Glycine
Serine
Isoleucine
Glycine
Alanine
Alanine
Protein
FIGURE 3.23 The genetic code is first transcribed into base triplets (codons) in mRNA and then translated into a specific sequence of amino
acids in a protein.
This pairing of bases follows the law of complementary base

pairing: guanine bonds with cytosine (and vice versa), and adenine bonds with uracil (because uracil in RNA is equivalent to
thymine in DNA). In RNA synthesis, only one of the two freed
strands of DNA serves as a guide (see fig. 3.22). Once an RNA
molecule has been produced, it detaches from the DNA strand
on which it was formed. This process can continue indefinitely,
producing many thousands of RNA copies of the DNA strand
being transcribed. When the gene is no longer to be transcribed,
the separated DNA strands can recoil into their helical form.
In the case of pre-mRNA, the finished molecule is altered
after synthesis. Within the pre-mRNA are noncoding regions
known as introns. The introns are removed through the action of
enzymes, and the coding regions are then spliced together so that
they can direct the synthesis of a specific protein.
Protein Synthesis
Once produced, mRNA leaves the nucleus and enters the cytoplasm, where it attaches to ribosomes. The mRNA passes through
a number of ribosomes to form a polyribosome, or polysome for
short. The association of mRNA with ribosomes is needed for

genetic translationthe production of specific proteins according
to the code contained in the mRNA base sequences.
Functions of Codons and Anticodons

Each mRNA molecule contains several hundred or more nucleotides, arranged in the sequence determined by complementary base pairing with DNA during genetic transcription (RNA
synthesis). Every three bases, or base triplet, is a code word
called a codonfor a specific amino acid. Sample codons and
their amino acid translation are shown in figure 3.23. As
mRNA move through the ribosome, the sequence of codons is
translated into a sequence of specific amino acids within a growing polypeptide chain.
Translation of the codons is accomplished by transfer RNA
(tRNA) and particular enzymes. One end of each tRNA contains the anticodon. The anticodon consists of three nucleotides
that are complementary to a specific codon in MRNA. Enzymes
in the cell cytoplasm join specific amino acids to the ends of
tRNA, so that a tRNA with a given anticodon is always bonded
to one specific amino acid. There are 20 different varieties of


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Chapter 3
Codons
mRNA
5
8
Codons
Next amino acid
7
4
6
Ala
tRNA
tRNA
I
H
tRNA
G
tRNA
F
Gly
Next amino acid
E
tRNA
tRNA
Iso
C
Ser
Gly
A
Gly
Ser
Met
Iso
tRN
Amino acids
in growing
polypeptide
chain
Gly
Met
Ribosome
FIGURE 3.24 The actions of mRNA and tRNA in genetic translation. The three-letter abbreviations for the amino acids in the growing polypeptide chain stand for the amino acids indicated in figure 3.23.
synthetase enzymesone for each type of amino acid. Each synthetase must not only recognize its specific amino acid, it must
also be able to attach this amino acid to the particular tRNA
that has the correct anticodon for that amino acid. Each of the
tRNA molecules in the cytoplasm of a cell is thus bonded to a
specific amino acid and is capable of bonding by its anticodon
base triplet with a specific codon in mRNA.
Formation of a Polypeptide
The anticodons of tRNA bind to the codons of mRNA as the
mRNA moves through the ribosome. Because each tRNA molecule carries a specific amino acid, the joining together of
these amino acids by peptide bonds forms a polypeptide whose
amino acid sequence has been determined by the sequence of
codons in mRNA.
The first and second tRNA bring the first and second
amino acids together, and a peptide bond forms between them.
The first amino acid then detaches from its tRNA, so that a
dipeptide is linked by the second amino acid to the second
tRNA. When the third tRNA binds to the third codon, the
amino acid it brings forms a peptide bond with the second amino
acid (which detaches from its tRNA). A tripeptide is thus attached by the third amino acid to the third tRNA. The polypep-
tide chain lengthens as new amino acids are added to its growing
tip (fig. 3.24). This polypeptide chain is always attached by
means of only one tRNA to the strand of mRNA, and this
tRNA molecule is always the one that has added the latest
amino acid to the growing polypeptide.
As the polypeptide chain becomes longer, interactions between its amino acids cause the chain to twist into a helix (secondary structure) and to fold and bend upon itself (tertiary
structure). At the end of this process, the new protein detaches
from the tRNA as the last amino acid is added.
Cell Cycle and Cell Division

A cell cycle is the series of changes that a cell undergoes from
the time it is formed until it has completed a division and reproduced itself. Interphase is the first period of the cycle, from cell
formation to the start of cell division (fig. 3.25). During interphase, the cell grows, carries on metabolic activities, and prepares itself for division.
Interphase is divided into G1, S, and G2 phases. During
the G1 (first growth) phase, the cell grows rapidly and is metabolically active. The duration of G1 varies considerably in different types of cells. It may last only hours in cells that have rapid
CHAPTER 3
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Anticodons
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Highly specialized cells, such as muscle and nerve cells, do

not replicate after a person is born. If these cells die, as the result of disease, injury, or even disuse, they are not replaced and scar
tissue may form. Nerve cells are especially vulnerable to damage
from oxygen deprivation, alcohol, and various other drugs.
es
is
Knowledge Check
Mitosis
Cy
to
k
in
Tel
oph
ase
Anaphase
ase
aph
Met
h
op
Pr
as
Mitotic Phase
CHAPTER 3
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G2
Final growth and
activity before
mitosis
G1
Centrioles
replicate
DNA replication
14. Explain why the DNA molecule is described as a double

helix.
15. Describe the various forms of RNA, and discuss how RNA
directs protein synthesis.
16. List the phases in the life cycle of a cell and describe the
principal events that occur during each phase.
17. Explain why mitosis is such an important biological process.
Cellular Adaptations
Interphase
FIGURE 3.25 Interphase and the mitotic phase are the two principal divisions of the cell cycle. During the mitotic phase, nuclear division is followed by cytoplasmic division and the formation of two
daughter cells.
division rates, or it may be a matter of days or even years for

other cells. At the end of G1, the centrioles replicate in preparation for their role in cell division. During the S (synthetic)
phase, the DNA in the nucleus of the cell replicates, so that the
two future cells will receive identical copies of the genetic material. During the G2 (second growth) phase, the enzymes and
other proteins needed for the division process are synthesized,
and the cell continues to grow.
The actual division of a cell is referred to as the mitotic
phase, or simply M phase (fig. 3.25). The mitotic phase is further divided into mitosis and cytokinesis. Mitosis is the period
of a cell cycle during which there is nuclear division and the
duplicated chromosomes separate to form two genetically identical daughter nuclei. The process of mitosis takes place in four
successive stages, each stage passing into the next without
sharp structural distinctions. These stages are prophase,
metaphase, anaphase, and telophase (fig. 3.26). Cytokinesis
(sito-knsis) is division of the cytoplasm, which takes place
during telophase.
cytokinesis: Gk. kytos, a hollow; kinesis, movement
Apparently included within cellular specialization of structure

and function is mitotic potential. Certain cells do not require
further division once the organ to which they contribute becomes functional. Others, as part of their specialization, require
continuous mitosis to keep an organ healthy. Thus, in the adult,
it is found that some cells divide continually, some occasionally,
and some not at all. For example, epidermal cells, hemopoietic
cells within bone marrow, and cells that line the lumen of the GI
tract divide continually throughout life. Cells within specialized
organs, such as the liver or kidneys, divide as the need becomes
apparent. Naturally occurring cellular death, disease, or trauma
from surgery or injury may necessitate mitosis in these organs.
Still other cells, such as muscle or nerve cells, lose their mitotic
ability as they become differentiated. Trauma to these cells frequently causes a permanent loss of function.
Although the factors regulating mitosis are unclear, evidence suggests that mitotic ability is genetically controlled and,
for those cells that do divide, even the number of divisions is
predetermined. If this is true, it may be a factor in the aging
process. Physical stress, nutrition, and hormones definitely have
an effect on mitotic activity. It is thought that the replication activity of cells might be controlled through a feedback mechanism
involving the release of a growth-inhibiting substance. Such a substance might slow or inhibit the cell divisions and growth of particular organs once they had amassed a certain number of cells or
had reached a certain size.
Except for cells on exposed surfaces, most cells of the body
are located in a fairly homogeneous environment, where continual adaptation to change is not necessary for survival. However,
cells do have remarkable adaptability and resilience, enabling
them to withstand conditions that might otherwise be lethal.
Prolonged exposure to sunlight, for example, stimulates the synthesis of melanin and tanning of the skin. Likewise, mechanical


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CHAPTER 3
FIGURE 3.26 The stages of mitosis.

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friction to the skin stimulates mitotic activity and the synthesis

of a fibrous protein, keratin, which results in the formation of a
protective callus.
Cells adapt to potentially injurious stimuli by several specific mechanisms. Hypertrophy (hipertro-fe) refers to an increase in the size of cells resulting from increased synthesis of
protein, nucleic acids, and lipids. Cellular hypertrophy can be either compensatory or hormonal. Compensatory hypertrophy occurs when increased metabolic demands on particular cells result
in an increase in cellular mass. Examples of compensatory hypertrophy include the enlargement of skeletal muscle fibers as a result of exercise and cardiac (heart) muscle fibers or kidney cells
because of an increased work demand. Hypertension (high blood
pressure) causes cardiac cells to hypertrophy because they must
pump blood against raised pressures. After the removal of a diseased kidney, there is a compensatory increase in the size of the
cells of the remaining kidney so that its normal weight is approximately doubled. Examples of hormonal hypertrophy are the increased size of the breasts and smooth muscles of the uterus in a
pregnant woman.
Hyperplasia (hiper-plaze-a) refers to an increase in the
number of cells formed as a result of increased mitotic activity.
The removal of a portion of the liver, for example, leads to regeneration, or hyperplasia, of the remaining liver cells to restore
the loss. But the triggering mechanism for hyperplasia is not
known. In women, a type of hormonally induced hyperplasia occurs in cells of the endometrium of the uterus after menstruation,
which restores this layer to a suitable state for possible implantation of an embryo.
Atrophy (atro-fe) refers to a decrease in the size of cells
and a corresponding decrease in the size of the affected organ.
Atrophy can occur in the cells of any organ and may be classified
as disuse atrophy, disease atrophy, or aging (senile) atrophy.
Metaplasia (meta -pl ze-a) is a specialized cellular change
in which one type of cell transforms into another. Generally, it
involves the change of highly specialized cells into more generalized, protective cells. For example, excessive exposure to inhaled
smoke causes the specialized ciliated columnar epithelial cells
lining the bronchial airways to change into stratified squamous
epithelium, which is more resistant to injury from smoke.
Trauma to Cells
As adaptable as cells are to environmental changes, they are subject to damage from aging and disease. If a trauma causes extensive cellular death, the condition may become life threatening.
A person dies when a vital organ can no longer perform its metabolic role in sustaining the body.
hypertrophy: Gk. hyper, over; trophe, nourishment

hyperplasia: Gk. hyper, over; plasis, a molding
metaplasia: Gk. meta, between; plasis, a molding
Energy deficit means that more energy is required by a cell

than is available. Cells can tolerate certain mild deficits because
of various reserves stored within the cytoplasm, but a severe or
prolonged deficit will cause cells to die. An energy deficit occurs
when the cells do not have enough glucose or oxygen to allow
for glucose combustion. Examples of energy deficits are low levels
of blood sugar (hypoglycemia) and the impermeability of the cell
membrane to glucose (as in diabetes mellitus). Malnutrition also
may result in an energy deficit. Few cells can tolerate an interruption in oxygen supply. Cells of the brain and the heart have
tremendous oxygen demands, and an interruption of the supply
to these organs can cause death in a matter of minutes.
Physical injury to cells, another type of trauma, occurs in
a variety of ways. High temperature (hyperthermia) is generally
less tolerable to cells than low temperature (hypothermia). Respiratory rate, heart rate, and metabolism accelerate with hyperthermia. Continued hyperthermia causes protein coagulation
within cells, and eventually cellular death. In frostbite, rapid or
prolonged chilling causes cellular injury. In severe frostbite, ice
crystals form and cause the cells to burst.
Burns are particularly significant if they cause damage to
the deeper skin layers, which interferes with the mitotic activity
of cells (see fig. 5.20). Of immediate concern with burns, however, is the devastating effect of fluid loss and infection through
traumatized cell membranes.
Accidental poisoning and suicide through drug overdose
account for large numbers of deaths in the United States and
elsewhere. Drugs and poisons can cause cellular dysfunction by
disrupting DNA replication, RNA transcription, enzyme systems, or cell membrane activity.
Radiation causes a type of cell trauma that is cumulative in
effect. When X rays are administered for therapeutic purposes
(radiotherapy), small doses are focused on a tumorous area over a
course of many days to prevent widespread cellular injury. Some
cells are more sensitive to radiation than others. Immature or mitotically active cells are highly sensitive, whereas cells that are
no longer growing, such as neurons and muscle cells, are not as
vulnerable to radiation injury.
Infectious agents, or pathogens, also cause cellular dysfunction. Viruses and bacteria are the most common pathogens.
Viruses usually invade and destroy cells as they reproduce themselves. Bacteria, on the other hand, do not usually invade cells
but will frequently poison cells with their toxic metabolic wastes.
Medical Genetics
Medical genetics is a branch of medicine concerned with diseases that have a genetic origin. Genetic factors include abnormalities in chromosome number or structure and mutant genes.
Genetic diseases are a diverse group of disorders, including malformed blood cells (sickle-cell anemia), defective blood clotting
(hemophilia), and mental retardation (Down syndrome).
Chromosomal abnormalities occur in approximately 0.6%
of live-birth infants. The majority (70%) are subtle, cause no
problems, and usually go undetected. Structural changes in the
DNA that are passed from parent to offspring by means of sex


of the Body
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The McGrawHill
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Chapter 3
cells are called mutations (myoo-tashunz). Mutations either
occur naturally or are environmentally induced through chemicals or radiation. Natural mutations are not well understood.
About 12% of all congenital malformations are caused by mutations and probably come about through an interaction of genetic
and environmental factors. Many of these problems can be predicted by knowing the genetic pedigree of prospective parents
and prevented through genetic counseling. Teratology (ter-atolo-je) is the science concerned with developmental defects and
the diagnosis, treatment, and prevention of malformations.
Genetic problems are occasionally caused by having too
few or too many chromosomes. The absence of an entire chromosome is termed monosomy (mono-somme). Embryos with
monosomy usually die. People with Turners syndrome have
only one X chromosome and have a better chance of survival
than those who are missing one of the other chromosomes. Trisomy (tri so-me), a genetic condition in which an extra chromosome is present, occurs more frequently than monosomy. The
best known among the trisomies is Down syndrome.
Cancer
Cancer refers to a complex group of diseases characterized by uncontrolled cell replication. The rapid proliferation of cells results
in the formation of a neoplasm, or new cellular mass. Neoplasms,
frequently called tumors, are classified as benign or malignant
based on their cytological and histological features. Benign neoplasms usually grow slowly and are confined to a particular area.
These types are usually not life threatening unless they grow to
large sizes in vital organs like the brain. Malignant neoplasms
(fig. 3.27) grow rapidly and metastasize (me-tasta-sz) (fragment
and spread) easily through lymphatic or blood vessels. The original malignant neoplasm is called the primary growth and the new
tumors, or metastatic tumors, are called secondary growths.
Cancer cells resemble undifferentiated or primordial cell
types. Generally they do not mature before they divide and are
not capable of maintaining normal cell function. Cancer causes
death when a vital organ regresses because of competition from
cancer cells for space and nutrients. The pain associated with cancer develops when the growing neoplasm affects sensory neurons.
73
Normal cells
(with hairlike cilia)
Cancer cells
FIGURE 3.27 An electron micrograph of cancer cells from the

respiratory tract (59,800).
The various types of cancers are classified on the basis of the
tissue in which they develop. Lymphoma, for example, is a cancer
of lymphoid tissue; osteogenic cancer is a type of bone cancer;
myeloma is cancer of the bone marrow; and sarcoma is a general
term for any cancer arising from cells of connective tissue.
The etiology (cause) of cancers is largely unknown. However, initiating factors, or carcinogens (kar-sino-jenz), such as
viruses, chemicals, or irradiation, may provoke cancer to develop. Cigarette smoking, for example, causes various respiratory
cancers to develop. The tendency to develop other types of cancers has a genetic basis. Some researchers even think that physiological stress can promote certain types of cancerous activity.
Because the causes of cancer are not well understood, emphasis is
placed on early detection with prompt treatment.
Aging
mutation: L. mutare, to change

teratology: Gk. teras, monster; logos, study
Although there are obvious external indicators of aginggraying

and loss of hair, wrinkling of skin, loss of teeth, and decreased
muscle masschanges within cells as a result of aging are not as
apparent and are not well understood. Certain organelles alter
with age. The mitochondria, for example, may change in structure and number, and the Golgi complex may fragment. Also,
lipid vacuoles tend to accumulate in the cytoplasm, and the cytoplasmic food stores that contain glycogen decrease.
Turners syndrome: from Henry H. Turner, American endocrinologist,

18921970
Down syndrome: from John L. H. Down, English physician, 182896
carcinogen: Gk, karkinos, cancer
CHAPTER 3
In an attempt to better understand medical genetics, the

Human Genome Project was launched by Congress in 1988
with the ambitious goal of completely mapping the human genome.
Scientists are currently on the verge of determining the exact sequences of bases with which the 3 billion base pairs are arranged to
form the 50,000 to 100,000 genes in the haploid human genome of a
sperm cell or ovum. Knowing this information will provide the ultimate
reference for diagnosis and treatment of the 4000 genetic diseases
that are known to be directly caused by particular abnormal genes.
Cytology

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ies, and its deterioration is thought to be associated with such

vascular diseases as arteriosclerosis in aged persons.

Giving a substance that competes for the enzyme alcohol dehydrogenase
can inhibit the reaction that forms the toxic metabolite of ethylene glycol. Thus, infusing a nearly intoxicating dose of alcohol can spare the
kidneys from harm. The ethylene glycol is then excreted harmlessly.
CHAPTER 3
The chromatin and chromosomes within the nucleus

show changes with aging, such as clumping, shrinking, or fragmenting with repeated mitotic divisions. There is strong evidence that certain cell types have a predetermined number of
mitotic divisions that are genetically controlled, thus determining the overall vitality and longevity of an organ. If this is true,
identifying and genetically manipulating the aging gene might
be possible.
Extracellular substances also change with age. Protein
strands of collagen and elastin change in quality and number in
aged tissues. Elastin plays an important role in the walls of arter-
Chapter Summary
Introduction to Cytology (p. 49)
1. Cells are the structural and functional
units of the body. Cellular function is
referred to as metabolism and the study of
cells is referred to as cytology.
2. Cellular function depends on the specific
membranes and organelles characteristic
of each type of cell.
3. All cells have structural modifications
that serve functional purposes.
Cellular Chemistry (pp. 5052)

1. Four elements (oxygen, carbon, hydrogen,
and nitrogen) compose over 95% of the
bodys mass and are linked together to
form inorganic and organic compounds.
2. Water is the most abundant inorganic
compound in cells and is an excellent
solvent.
(a) Water is important in temperature
control and hydrolysis.
(b) Dehydration, a condition in which
fluid loss exceeds fluid intake, may be a
serious problemespecially in infants.
3. Electrolytes are inorganic compounds that
form ions when dissolved in water.
(a) The three classes of electrolytes are
acids, bases, and salts.
(b) Electrolytes are important in
maintaining pH, in conducting
electrical currents, and in regulating
the activity of enzymes.
4. Proteins are organic compounds that may
exist by themselves or be conjugated with
other compounds.
(a) Proteins are important structural
components of the body and are
necessary for cellular growth, repair,
and division.
(b) Enzymes and hormones are examples

of specialized proteins.
5. Carbohydrates are organic compounds
containing carbon, hydrogen, and oxygen,
with a 2:1 ratio of hydrogen to oxygen.
(a) The carbohydrate group includes the
starches and sugars.
(b) Carbohydrates are the most abundant
source of cellular energy.
6. Lipids are organic fats and fat-related
substances.
(a) Lipids are composed primarily of
carbon, hydrogen, and oxygen.
(b) Lipids serve as an important source of
energy, form parts of membranes, and
protect and insulate various parts of
the body.
Cellular Structure (pp. 5264)

1. A cell is composed of a cell membrane,
cytoplasm and organelles, and a nucleus.
2. The cell membrane, composed of
phospholipid and protein molecules,
encloses the contents of the cell and
regulates the passage of substances into
and out of the cell.
(a) The permeability of the cell membrane
depends on its structure, the size of the
molecules, ionic charge, lipid solubility,
and the presence of carrier molecules.
(b) Cell membranes may be specialized
with such structures as microvilli,
sacs, and hair cells.
3. Cytoplasm refers to the material
between the cell membrane and the
nucleus. Nucleoplasm is the material
within the nucleus. Protoplasm is a
collective term for both the cytoplasm
and nucleoplasm.
4. Organelles are specialized components

within the cytoplasm of cells.
(a) Endoplasmic reticulum provides a
framework within the cytoplasm and
forms a site for the attachment of
ribosomes. It functions in the
synthesis of lipids and proteins and in
cellular transport.
(b) Ribosomes are particles of protein
and RNA that function in protein
synthesis. The protein particles
may be used within the cell or
secreted.
(c) The Golgi complex consists of
membranous vesicles that synthesize
glycoproteins and secrete lipids. The
Golgi complex is extensive in
secretory cells, such as those of the
pancreas and salivary glands.
(d) Mitochondria are membranous
sacs that consist of outer and
inner mitochondrial layers and
folded membranous extensions
of the inner layer called cristae.
The mitochondria produce ATP
and are called the powerhouses
of a cell. Mitochondria are lacking
in sperm cells and red blood cells.
(e) Lysosomes are spherical bodies that
contain digestive enzymes. They are
abundant in the phagocytic white
blood cells.
(f) Peroxisomes are enzyme-containing
membranous sacs that are abundant in
the kidneys and liver. Some of the
enzymes in peroxisomes generate
hydrogen peroxide, and one of them,
catalase, breaks down excess hydrogen
peroxide.


of the Body
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Chapter 3
6. The nucleotides in DNA consist of the sugar
deoxyribose, phosphate, and one of four
nitrogenous bases: adenine, guanine,
cytosine, or thymine. According to the law of
complementary base pairing, bases are
specific in their bonding: adenine bonds with
thymine and guanine bonds with cytosine.
7. RNA contains the sugar ribose (instead of
deoxyribose) and the base uracil (in place
of thymine). The three major forms of
RNA are mRNA, tRNA, and rRNA.
8. The genetic code in mRNA consists of
three bases called codons. Codons bond to
anticodons, which are three bases in tRNA.
9. Each type of tRNA is bonded to a specific
type of amino acid, which the tRNA
brings to the growing polypeptide chain.
Cell Cycle (pp. 6570)

1. The cell cycle consists of growth,
synthesis, and mitosis.
(a) Growth is the increase in cellular
mass that results from metabolism.
75
Synthesis is the production of DNA

and RNA to regulate cellular activity.
Mitosis is the splitting of the cells
nucleus and cytoplasm that results in
the formation of two diploid cells.
(b) Mitosis permits an increase in
the number of cells (body growth)
and allows for the replacement
of damaged, diseased, or
worn-out cells.
2. A DNA molecule is in the shape of a
double helix. The structural unit of the
molecule is a nucleotide, which consists
of deoxyribose (sugar), phosphate, and a
nitrogenous base.
3. Cell division consists of a division of the
chromosomes (mitosis) and a division of
the cytoplasm (cytokinesis). The stages of
mitosis include prophase, metaphase,
anaphase, and telophase.
Review Activities
Objective Questions
1. Inorganic compounds that form ions
when dissociated in water are
(a) hydrolites.
(d) ionizers.
(b) metabolites. (e) nucleic acids.
(c) electrolytes.
2. The four elements that compose over
95% of the body are
(a) oxygen, potassium, hydrogen, carbon.
(b) carbon, sodium, nitrogen, oxygen.
(c) potassium, sodium, magnesium,
oxygen.
(d) carbon, oxygen, nitrogen, hydrogen.
(e) oxygen, carbon, hydrogen, sulfur.
3. Which organelle contains strong
hydrolytic enzymes?
(a) the lysosome
(b) the Golgi complex
(c) the ribosome
(d) the vacuole
(e) the mitochondrion
4. Ciliated cells occur in
(a) the trachea. (c) the bronchioles.
(b) the ductus
deferens.
(e) all of the above.
5. Osmosis deals with the movement of
(a) gases.
(c) oxygen only.
(b) water only.
(d) both a and c.

chromosomes line up at the equator
(equatorial plane) of the cell is called
(a) interphase.
(d) anaphase.
(b) prophase.
(e) telophase.
(c) metaphase.
chromatids separate is called
(a) interphase.
(d) anaphase.
(b) prophase.
(e) telophase.
(c) metaphase.
8. The organelle that combines protein with
carbohydrates and packages them within
vesicles for secretion is
(a) the Golgi complex.
(b) the rough endoplasmic reticulum.
(c) the smooth endoplasmic reticulum.
(d) the ribosome.
9. The enlarged skeletal muscle fibers that
result from an increased work demand
serve to illustrate
(a) disuse atrophy.
(b) compensatory hypertrophy.
(c) metaplasia.
(d) inertia.
10. Regeneration of liver cells is an example of
(a) compensatory hypertrophy.
(b) hyperplasia.
(c) metaplasia.
(d) hypertrophy.
DNA is false?
(a) It is located in the nucleus.
(b) It is double-stranded.
(c) The bases adenine and thymine can
bond together.
(d) The bases guanine and adenine can
bond together.
RNA is true?
(a) It is made in the nucleus.
(b) It contains the sugar deoxyribose.
(c) It is a complementary copy of the
entire DNA molecule.
(d) It is double-stranded.
Essay Questions
1. Explain why a knowledge of cellular
anatomy is necessary for understanding
tissue and organ function within the
body. How is the study of cells important
for the understanding of body dysfunction
and disease?
2. Why is water a good fluid medium
of the cell?
CHAPTER 3
(g) The centrosome is the dense area of

cytoplasm near the nucleus that
contains the centrioles. The paired
centrioles play an important role in
cell division.
(h) Vacuoles are membranous sacs that
function as storage chambers.
(i) Fibrils and microtubules provide
support in the form of a cytoskeleton.
(j) Cilia and flagella are projections of
the cell that have the same basic
structure and that function in
producing movement.
5. The cell nucleus is enclosed in a nuclear
membrane that controls the movement of
substances between the nucleoplasm and
the cytoplasm.
(a) The nucleoli are small bodies of
protein and RNA within the nucleus
that produce ribosomes.
(b) Chromatin is a coiled fiber of protein
and DNA that shortens to form
chromosomes during cell reproduction.
Cytology

CHAPTER 3
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3. How are proteins, carbohydrates, and

lipids similar? How are they different?
What are enzymes and hormones?
4. Describe the cell membrane. List the
various kinds of movement through the cell
membrane and give an example of each.
5. Describe, diagram, and list the functions
of the following:
(a) endoplasmic reticulum,
(b) ribosome,
(c) mitochondrion,
(d) Golgi complex,
(e) centrioles, and
(f) cilia.
6. Define inorganic compound and organic
compound and give examples of each.
7. Define the terms protoplasm, cytoplasm,
and nucleoplasm. Describe the position of
the membranes associated with each of
these substances.
8. Describe the structure of the nucleus and
the functions of its parts.
9. What is a nucleotide? How does it relate
to the overall structure of a DNA
molecule?
10. Explain the relationship between DNA,
chromosomes, chromatids, and genes.
11. Describe how RNA is produced and list
the different forms of RNA.
12. Explain how one DNA strand can serve
as a template for the synthesis of another
DNA strand.
13. Distinguish between mitosis and

cytokinesis. Describe the major events of
mitosis and discuss the significance of the
mitotic process.
14. Give examples of factors that contribute
cellular hypertrophy, hyperplasia, atrophy,
and metaplasia.
15. Explain how cells respond to
(a) energy deficit,
(b) hyperthermia,
(c) burns,
(d) radiation, and
(e) pathogens.
16. Define the following genetic terms:
teratology, monosomy, trisomy, and
mutation.
17. In what ways do cells of a neoplasm differ
from normal cells. How may a malignant
neoplasm cause death?
18. Discuss the cellular and extracellular
changes that accompany aging.
1. How is the structural organization of its
individual cells essential to a multicellular
organism?
2. Construct a table comparing the structure
and function of several kinds of cells.
Indicate which organelles would be of
particular importance to each kind of cell.
3. Define medical genetics and give some
examples of genetic diseases. Is spending

the billions of dollars required to

complete the Human Genome Project
justified? Why or why not?
4. The brain is protected to some extent by
the blood-brain barriera membrane
between circulating blood and the brain
that keeps certain damaging substances
from reaching brain tissue. However, the
brain is still subject to trauma that can
cause it to swell, much like an ankle
swells with a sprain. Because the cranium
is a cavity of fixed size, brain edema
(swelling) can rapidly lead to coma and
death. Knowing what you do about
movement of water across a membrane,
can you explain why mannitol, a type of
sugar that does not cross the blood-brain
barrier, is commonly used to treat patients
who have suffered head trauma?
5. Your friend knows that you have just
reviewed cellular chemistry, and so he asks
for your opinion about his new diet. In an
attempt to eliminate the lipid content in
adipose tissue and thus lose weight, he has
completely eliminated fat from his diet.
He feels that he is now free to eat as much
food as he likes, provided it consists only
of carbohydrates and protein. Is your
friends logic flawed? Would you advise
him to stick with this diet?


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Histology
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4
Definition and Classification of
Tissues 78
The Tissues 79
Epithelial Tissue 79
Connective Tissue 89
Muscle Tissue 99
Nervous Tissue 100

Chapter Summary 103
Clinical Case Study

As a medical student, you are rotating with a gastroenterologist who is performing an upper endoscopy on a patient with long-standing gastroesophageal reflux (heartburn). During the procedure, the doctor quizzes you on what type of cells line the esophagus. What is your answer? He
takes a biopsy of the lower esophagus just above the stomach. Later, the specimen is fixed, and
you examine it under the microscope. You see a single layer of nonciliated, tall, column-shaped
cells. What type of tissue are you examining?
FIGURE: Knowing the structure and function

of body tissues elucidates how they may adapt
to provide protection to body organs.

78
Unit 3

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4. Histology
The McGrawHill
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DEFINITION AND
Shaft of a
hair within
a hair follicle
Objective 1
histology.
CHAPTER 4
Objective 2

cells and tissues.
Objective 3



(a)
Shaft of hair
emerging from
the exposed
surface of
the skin
(b)


Knowledge Check


of the Body
4. Histology
The McGrawHill
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Embryoblast
Blastocoele
Amniotic cavity
(c)
Ectoderm
Trophoblast
Endoderm
(b)
Amniotic cavity
(a)
Embryonic
disc
(d)
Ectoderm
Mesoderm
Endoderm
Yolk sac
(e)
Trophoblast
Schenk
EXHIBIT 1 The early stages of embryonic development. (a) Fertilization and the formation of the zygote, (b) the morula at about the
third day, (c) the early blastocyst at the time of implantation between the fifth and seventh day, (d) a blastocyst at 2 weeks, and (e) a blastocyst at 3 weeks showing the three primary germ layers that constitute the embryonic disc.
The Tissues
Human prenatal development is initiated by the fertilization of
an ovulated ovum (egg) from a female by a sperm cell from a
male. The chromosomes within the nucleus of a zygote (zgot)
(fertilized egg) contain all the genetic information necessary for
the differentiation and development of all body structures.
Within 30 hours after fertilization, the zygote undergoes a

mitotic division as it moves through the uterine tube toward the
uterus (see chapter 22). After several more cellular divisions, the
embryonic mass consists of 16 or more cells and is called a
morula (moryoo-la), as shown in exhibit I. Three or 4 days after
conception, the morula enters the uterine cavity where it remains unattached for about 3 days. During this time, the center
of the morula fills with fluid absorbed from the uterine cavity. As
the fluid-filled space develops inside the morula, two distinct
groups of cells form. The single layer of cells forming the outer
zygote: Gk. zygotos, yolked
morula: Gk. morus, mulberry
EXPLANATION
(continued)
EPITHELIAL TISSUE
Objective 5
Discuss the functions of the membranous

epithelia in different locations in the body.
There are two major categories of epithelia: membranous and

glandular. Membranous epithelia are located throughout the body
and form such structures as the outer layer of the skin; the inner
lining of body cavities, tubes, and ducts; and the covering of visceral organs. Glandular epithelia are specialized tissues that form
the secretory portion of glands.
Objective 6
Objective 4
Membranous epithelia always have one free surface exposed to a

body cavity, a lumen (hollow portion of a body tube), or to the
skin surface. Some membranous epithelia are derived from ectoderm, such as the outer layer of the skin; some from mesoderm,
79
Compare and contrast the various types of

membranous epithelia.
epithelium: Gk. epi, upon; thelium, to cover
Define gland and compare and contrast the

various types of glands in the body.
Characteristics of Membranous Epithelia


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The McGrawHill
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(continued)
TABLE 4A Derivatives of the Germ Layers

Ectoderm
Mesoderm
Endoderm
Epidermis of skin and epidermal derivatives:

hair, nails, glands of the skin; linings of oral,
nasal, anal, and vaginal cavities
Muscle: smooth, cardiac, skeletal
Epithelium of pharynx, auditory canal, tonsils,

thyroid, parathyroid, thymus, larynx, trachea,
lungs, GI tract, urinary bladder and urethra, and
vagina

Pituitary gland
Epithelium (endothelium) of blood vessels,

lymphatic vessels, body cavities, joint cavities
Adrenal medulla
Connective tissue: embryonic, mesenchyme,

connective tissue proper, cartilage, bone, blood
Liver and pancreas
Kidneys and ureters

Adrenal cortex
wall is known as the trophoblast, and the inner aggregation of

cells is known as the embryoblast. With further development,
the trophoblast differentiates into a structure that will later form
part of the placenta; the embryoblast will eventually become the
embryo. With the establishment of these two groups of cells, the
morula becomes known as a blastocyst (blasto-sist). Implantation
of the blastocyst in the uterine wall begins between the fifth and
seventh day (see chapter 22).
week of development, the embryoblast undergoes marked differentiation. A slitlike space called the amniotic (amne-ot-ic) cavity
forms within the embryoblast, adjacent to the trophoblast. The
embryoblast now consists of two layers: an upper ectoderm, which

embryoblast: Gk. embryon, to be full, swell; blastos, germ
such as the inside lining of blood vessels; and others from endoderm, such as the inside lining of the digestive tract (gastrointestinal, or GI, tract).
Membranous epithelia may be one or several cell layers
thick. The upper surface may be exposed to gases, as in the case
of epithelium in the integumentary and respiratory systems; to
liquids, as in the circulatory and urinary systems; or to semisolids,
as in the GI tract. The deep surface of most membranous epithelia is bound to underlying supportive tissue by a basement membrane, that consists of glycoprotein from the epithelial cells and
a meshwork of collagenous and reticular fibers from the underlying connective tissue. With few exceptions, membranous epithe80
is closer to the amniotic cavity, and a lower endoderm, which

borders the blastocoel (blastocyst cavity). A short time later, a
third layer called the mesoderm forms between the endoderm and
ectoderm. These three layers constitute the primary germ layers.
The primary germ layers are of great significance because
all the cells and tissues of the body are derived from them (see
fig. 22.9). Ectodermal cells form the nervous system; the outer
layer of skin (epidermis), including hair, nails, and skin glands;
and portions of the sensory organs. Mesodermal cells form the
skeleton, muscles, blood, reproductive organs, dermis of the skin,
and connective tissue. Endodermal cells produce the lining of the
GI tract, the digestive organs, the respiratory tract and lungs, and
the urinary bladder and urethra. The derivatives of the primary
germ layers are summarized in table 4A.

lia are avascular (without blood vessels) and must be nourished

by diffusion from underlying connective tissues. Cells that make
up membranous epithelia are tightly packed together, with little
intercellular matrix between them.
Some of the functions of membranous epithelia are quite
specific, but certain generalities can be made. Epithelia that cover
or line surfaces provide protection from pathogens, physical injury,
toxins, and desiccation. Epithelia lining the GI tract function in
absorption. The epithelium of the kidneys provides filtration,
whereas that within the pulmonary alveoli (air sacs) of the lungs
allows for diffusion. Highly specialized neuroepithelium in the taste
buds and in the nasal region has a chemoreceptor function.


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Chapter 4
Histology
81
(c)
(a)
FIGURE 4.2 (a) Simple squamous epithelium lines the lumina of vessels, where it permits diffusion. (b) A photomicrograph of this tissue and
(c) a labeled diagram. Simple squamous epithelia that line the lumina of vessels are referred to as endothelia, and that which cover visceral organs are referred to as mesothelia.
Many membranous epithelia are exposed to friction or

harmful substances from the outside environment. For this reason, epithelial tissues have remarkable regenerative abilities. The
mitotic replacement of the outer layer of skin and the lining of
the GI tract, for example, is a continuous process.
Membranous epithelia are histologically classified by the
number of layers of cells and the shape of the cells along the exposed surface. Epithelial tissues that are composed of a single
layer of cells are called simple; those that are layered are said to
be stratified. Squamous cells are flattened; cuboidal cells are cubeshaped; and columnar cells are taller than they are wide.
Simple Epithelia
Simple epithelial tissue is a single cell layer thick and is located
where diffusion, absorption, filtration, and secretion are principal
functions. The cells of simple epithelial tissue range from thin,
flattened cells to tall, columnar cells. Some of these cells have
cilia that create currents for the movement of materials across
cell surfaces. Others have microvilli that increase the surface
area for absorption.
Simple Squamous Epithelium

Simple squamous (skwamus) epithelium is composed of flattened, irregularly shaped cells that are tightly bound together in
a mosaiclike pattern (fig. 4.2). Each cell contains an oval or
squamous: L. squamosus, scaly
spherical central nucleus. This epithelium is adapted for diffusion

and filtration. It occurs in the pulmonary alveoli within the
lungs (where gaseous exchange occurs), in portions of the kidney
(where blood is filtered), on the inside walls of blood vessels, in
the lining of body cavities, and in the covering of the viscera.
The simple squamous epithelium lining the inner walls of blood
and lymphatic vessels is termed endothelium (endo-thele-um)
(fig. 4.2b). That which covers visceral organs and lines body cavities is called mesothelium (meso-thele-um).
Simple Cuboidal Epithelium

Simple cuboidal epithelium is composed of a single layer of
tightly fitted cube-shaped cells (fig. 4.3). This type of epithelium
is found lining small ducts and tubules that have excretory, secretory, or absorptive functions. It occurs on the surface of the
ovaries, forms a portion of the tubules within the kidney, and
lines the ducts of the salivary glands and pancreas.
Simple Columnar Epithelium

Simple columnar epithelium is composed of tall, columnar
cells (fig. 4.4). The height of the cells varies, depending on the
site and function of the tissue. Each cell contains a single nucleus which is usually located near the basement membrane.
endothelium: Gk. endon, within; thelium, to cover

mesothelium: Gk. meso, middle; thelium, to cover
CHAPTER 4
(b)

82
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Lumen of
renal tubule
Kidney
Basement
membrane
Nucleus
CHAPTER 4
(b)
(c)
Ureter
(a)
FIGURE 4.3 (a) Simple cuboidal epithelium lines the lumina of ducts; for example, in the kidneys, where it permits movement of fluids and
ions. (b) A photomicrograph of this tissue and (c) a labeled diagram.
Liver
Stomach
Gallbladder
Large intestine
Small intestine
(b)
(b)
Lumen of small
intestine
Nucleus
Creek
Basement
membrane
(a)
Goblet cell
Cilia
(c)
FIGURE 4.4 (a) Simple columnar epithelium lines the lumen of the digestive tract, where it permits secretion and absorption. (b) A photomicrograph of this tissue and (c) a labeled diagram.


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Body of
uterus
Uterine tube
Uterine cavity
Ovary
Lumen of
uterine tube
Cilia
(a)
Cell membrane
Vagina
Nucleus
Basement
membrane
(c)
FIGURE 4.5 (a) Simple ciliated columnar epithelium lines the lumen of the uterine tube, where currents generated by the cilia propel the ovum
(egg cell) toward the uterus. (b) a photomicrograph of this tissue and (c) a labeled diagram.
Specialized unicellular glands called goblet cells are scattered

through this tissue at most locations. Goblet cells secrete a lubricative and protective mucus along the free surfaces of the
cells. Simple columnar epithelium is found lining the inside walls
of the stomach and intestine. In the digestive system, it forms a
highly absorptive surface and also secretes certain digestive
chemicals. Within the stomach, simple columnar epithelium has
a tremendous mitotic rate. It replaces itself every 2 to 3 days.
Simple Ciliated Columnar Epithelium

Simple ciliated columnar epithelium is characterized by the presence of cilia along its free surface (fig. 4.5). By contrast, the simple columnar type is unciliated. Cilia produce wavelike
movements that transport materials through tubes or passageways. This type of epithelium occurs in the female uterine tubes
to move the ovum (egg cell) toward the uterus.
Not only do the cilia function to propel the ovum, but recent
evidence indicates that sperm introduced into the female
vagina during sexual intercourse may be moved along the return currents, or eddies, generated by ciliary movement. This greatly enhances the likelihood of fertilization.
Pseudostratified Ciliated Columnar Epithelium

As the name implies, this type of epithelium has a layered appearance (strata = layers). Actually, it is not multilayered (pseudo
= false), because each cell is in contact with the basement membrane. Not all cells are exposed to the surface, however (fig. 4.6).
The tissue appears to be stratified because the nuclei of the cells

are located at different levels. Numerous goblet cells and a ciliated exposed surface are characteristic of this epithelium. It is
found lining the inside walls of the trachea and the bronchial
tubes; hence, it is frequently called respiratory epithelium. Its function is to remove foreign dust and bacteria entrapped in mucus
from the lower respiratory system.
Coughing and sneezing, or simply clearing the throat, are
protective reflex mechanisms for clearing the respiratory passages of obstruction or of inhaled particles that have been trapped in
the mucus along the ciliated lining. The material that is coughed up
consists of the mucus-entrapped particles.
Stratified epithelia have two or more layers of cells. In contrast
to the single-layered simple epithelia, they are poorly suited for
absorption and secretion. Stratified epithelia have a primarily
protective function that is enhanced by rapid cell divisions. They
are classified according to the shape of the surface layer of cells,
because the layer in contact with the basement membrane is always cuboidal or columnar in shape.
Stratified Squamous Epithelium

Stratified squamous epithelium is composed of a variable number
of cell layers that are flattest at the surface (fig. 4.7). Mitosis occurs only at the deepest layers (see table 5.2). The mitotic rate
CHAPTER 4
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Larynx
Trachea
Bronchiole
Lung
Secondary bronchi
(b)
CHAPTER 4
(b)
(a)
Creek
Cilia
Goblet cell
Nucleus
Bronchus
Basement membrane
Connective tissue
(c)
FIGURE 4.6 (a) Pseudostratified ciliated columnar epithelium lines the lumen of the respiratory tract, where it traps foreign material and moves
it away from the pulmonary alveoli of the lungs. (b) a photomicrograph of this tissue and (c) a labeled diagram.
Uterine tube
Ovary
Uterus
Urinary bladder
Cervix
Rectum
Urethra
Vagina
Anus
Waldrop
(b)
(a)
Stratified
squamous
epithelium
(c)
FIGURE 4.7 Stratified squamous epithelium forms the outer layer of skin and the lining of body openings. In the moistened areas, such as in the
vagina (a), it is nonkeratinized, whereas in the epidermis of the skin it is keratinized. (b) a photomicrograph of this tissue and (c) a labeled diagram.


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Parotid gland
Nuclei
Lumen of
parotid duct
Basement
membrane
(b)
(c)
(a)
FIGURE 4.8 (a) Stratified cuboidal epithelium lines the lumina of large ducts like the parotid duct, which drains saliva from the parotid gland.
(b) a photomicrograph of this tissue and (c) a labeled diagram.
approximates the rate at which cells are sloughed off at the surface.
As the newly produced cells grow in size, they are pushed toward
the surface, where they replace the cells that are sloughed off.
Movement of the epithelial cells away from the supportive basement membrane is accompanied by the production of keratin (keratin) (described below), progressive dehydration, and flattening.
There are two types of stratified squamous epithelial tissues: keratinized and nonkeratinized.
1. Keratinized stratified squamous epithelium contains keratin, a protein that strengthens the tissue. Keratin makes
the epidermis (outer layer) of the skin somewhat waterproof
and protects it from bacterial invasion. The outer layers of
the skin are dead, but glandular secretions keep them soft
(see chapter 5).
2. Nonkeratinized stratified squamous epithelium lines the
oral cavity and pharynx, nasal cavity, vagina, and anal
canal. This type of epithelium, called mucosa (myoo-kosa),
is well adapted to withstand moderate abrasion but not
fluid loss. The cells on the exposed surface are alive and are
always moistened.
Stratified squamous epithelium is the first line of defense
against the entry of living organisms into the body. Stratification,
rapid mitotic activity, and keratinization within the epidermis of the skin
are important protective features. An acidic pH along the surfaces of
this tissue also helps to prevent disease. The pH of the skin is between
4.0 and 6.8. The pH in the oral cavity ranges from 5.8 to 7.1, which
tends to retard the growth of microorganisms. The pH of the anal region is about 6, and the pH along the vaginal lining is 4 or lower.
keratin: Gk. keras, horn
Stratified Cuboidal Epithelium

Stratified cuboidal epithelium usually consists of only two or
three layers of cuboidal cells (fig. 4.8). This type of epithelium is
confined to the linings of the large ducts of sweat glands, salivary
glands, and the pancreas, where its stratification probably provides a more robust lining than would simple epithelium.
Transitional Epithelium
Transitional epithelium is similar to nonkeratinized stratified
squamous epithelium except that the surface cells of the former
are large and round rather than flat, and some may have two nuclei (fig. 4.9). Transitional epithelium is found only in the urinary system, particularly lining the cavity of the urinary bladder
and lining the lumina of the ureters. This tissue is specialized to
permit distension (stretching) of the urinary bladder as it fills
with urine. The inner, exposed cells actually transform from
being rounded when the urinary bladder is empty to being somewhat flattened as it distends with urine.
A summary of membranous epithelial tissue is presented in
table 4.1.
Body Membranes
Body membranes are composed of thin layers of epithelial tissue
and, in certain locations, epithelial tissue coupled with supporting connective tissue. Body membranes cover, separate, and support visceral organs and line body cavities. The two basic types of
body membranes, mucous membranes and serous membranes,
are described in detail in chapter 2 under the heading Body
Cavities and Membranes (see p. 41).
CHAPTER 4
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Ureter
CHAPTER 4
(b)
Urinary
bladder
Lumen of
urinary bladder
(b)
Transitional
epithelium
Urethra
Smooth
muscle
tissue
(c)
(a)
FIGURE 4.9 (a) Transitional epithelium lines the lumina of the ureters, part of the urethra and the cavity of the urinary bladder, where it permits
distention. (b) a photomicrograph of this tissue and (c) a labeled diagram.
TABLE 4.1 Summary of Membranous Epithelial Tissue

Type
Location
Simple Epithelia
Single layer of cells; function varies with type
Covering visceral organs; linings of body cavities,

tubes, and ducts
Simple squamous epithelium
Single layer of flattened, tightly bound cells;

diffusion and filtration
Capillary walls; pulmonary alveoli of lungs;

covering visceral organs; linings of body cavities
Simple cuboidal epithelium
Single layer of cube-shaped cells; excretion,

secretion, or absorption
Surface of ovaries; linings of renal tubules, salivary

ducts, and pancreatic ducts
Simple columnar epithelium
Single layer of nonciliated, tall, column-shaped cells;

protection, secretion, and absorption
Lining of most of GI tract
Simple ciliated columnar epithelium
Single layer of ciliated, column-shaped cells;

transportive role through ciliary motion
Lining of uterine tubes
Pseudostratified ciliated columnar epithelium
Single layer of ciliated, irregularly shaped cells; many

goblet cells; protection, secretion, ciliary
movement
Lining of respiratory passageways
Two or more layers of cells; function varies with type
Epidermal layer of skin; linings of body openings,

ducts, and urinary bladder
Stratified squamous epithelium (keratinized)
Numerous layers containing keratin, with outer

layers flattened and dead; protection
Epidermis of skin
Stratified squamous epithelium (nonkeratinized)
Numerous layers lacking keratin, with outer layers

moistened and alive; protection and pliability
Linings of oral and nasal cavities, vagina, and anal

canal
Stratified cuboidal epithelium
Usually two layers of cube-shaped cells;

strengthening of luminal walls
Large ducts of sweat glands, salivary glands, and

pancreas
Transitional epithelium
Numerous layers of rounded, nonkeratinized cells;

distension
Walls of ureters, part of urethra, and urinary

bladder


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Secretion
Lumen
Mucus
Liver
Stomach
Cell membrane
Gallbladder
Golgi complex
Small intestine
CHAPTER 4
Large intestine
Rough endoplasmic
reticulum
Creek
(a)
(b)
(c)
FIGURE 4.10 A goblet cell is a unicellular gland that secretes mucus, which lubricates and protects surface linings. (a) Goblet cells are abundant in the columnar epithelium lining the lumen of the small intestine. (b) A photomicrograph of a goblet cell and (c) a labeled diagram.
Glandular Epithelia
As tissues develop in the embryo, tiny invaginations (infoldings)
or evaginations (outfoldings) of membranous epithelia give rise to
specialized secretory structures called exocrine (ekso-krin) glands.
These glands remain connected to the epithelium by ducts, and
their secretions pass through the ducts onto body surfaces or into
body cavities. Exocrine glands should not be confused with endocrine glands, which are ductless, and which secrete their products (hormones) into the blood or surrounding extracellular fluid.
Exocrine glands within the skin include oil (sebaceous) glands,
sweat glands, and mammary glands. Exocrine glands within the
digestive system include the salivary and pancreatic glands.
Exocrine glands are classified according to their structure
and how they discharge their products. Classified according to
structure, there are two types of exocrine glands, unicellular and
multicellular glands.
1. Unicellular glands are single-celled glands, such as goblet
cells (fig. 4.10). They are modified columnar cells that
occur within most epithelial tissues. Goblet cells are found
in the epithelial linings of the respiratory and digestive systems. The mucus secretion of these cells lubricates and protects the surface linings.
2. Multicellular glands, as their name implies, are composed

of both secretory cells and cells that form the walls of the
ducts. Multicellular glands are classified as simple or compound glands. The ducts of the simple glands do not
branch, whereas those of the compound type do (fig. 4.11).
shape of their secretory portion. They are identified as
tubular glands if the secretory portion resembles a tube and
as acinar glands if the secretory portion resembles a flask.
Multicellular glands with a secretory portion that resembles both a tube and a flask are termed tubuloacinar glands.
means by which they release their product (fig. 4.12).
1. Merocrine (mero-krin) glands are those that secrete a watery substance through the cell membrane of the secretory
cells. Salivary glands, pancreatic glands, and certain sweat
glands are of this type.
2. Apocrine (apo-krin) glands are those in which the secretion accumulates on the surface of the secretory cell; then,
a portion of the cell, along with the secretion, is pinched
off to be discharged. Mammary glands are of this type.
merocrine: Gk. meros, part; krinein, to separate

exocrine: Gk. exo, outside; krinein, to separate
apocrine: Gk. apo, off; krinein, to separate

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Duct
Secretory cells
Simple
branched tubular
Simple
coiled tubular
Simple acinar
Simple
branched acinar
CHAPTER 4
Simple tubular
Compound tubular
Compound acinar
Compound tubuloacinar
FIGURE 4.11 Structural classification of multicellular exocrine glands. The ducts of the simple glands either do not branch or have few
branches, whereas those of the compound glands have multiple branches.
Secretion
Disintegrating cell
and its contents
discharged
with secretion
Intact cell
New cell
Merocrine gland
FIGURE 4.12 Examples of multicellular exocrine glands.
Apocrine gland
Cr
ee
k
Pinched-off
portion of cell
discharged
with secretion
Holocrine gland


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TABLE 4.2 Summary of Glandular Epithelial Tissue

Classification of Exocrine Glands by Structure
Type
Example
Lubricate and protect

Protect, cool body, lubricate, aid in digestion, maintain
body homeostasis
Goblet cells of digestive, respiratory, urinary, and

reproductive systems
Sweat glands, digestive glands, liver, mammary glands,
sebaceous glands
A. Simple
1. Tubular
2. Branched tubular
3. Coiled tubular
4. Acinar
5. Branched acinar
Aid in digestion
Protect, aid in digestion
Regulate temperature
Provide additive for spermatozoa
Condition skin
Intestinal glands
Uterine glands, gastric glands
Certain sweat glands
Seminal vesicles of male reproductive system
B. Compound
1. Tubular
2. Acinar
Lubricate urethra of male, assist body digestion

Provide nourishment for infant, aid in digestion
Bulbourethral glands of male reproductive system, liver

Mammary glands, salivary glands (sublingual and
submandibular)
Salivary gland (parotid), pancreas
I. Unicellular
II. Multicellular
3. Tubuloacinar
Aid in digestion
Classification of Exocrine Glands by Mode of Secretion

Type
Description of Secretion
Example
Merocrine glands
Watery secretion for regulating temperature or enzymes

that promote digestion
Salivary and pancreatic glands, certain sweat glands
Apocrine glands
Portion of secretory cell and secretion are discharged; provides

nourishment for infant, assists in regulating temperature
Mammary glands, certain sweat glands
Holocrine glands
Entire secretory cell with enclosed secretion is discharged;

conditions skin
3. Holocrine (holo-krin) glands are those in which the entire

secretory cell is discharged, along with the secretory product. An example of a holocrine gland is an oil-secreting
(sebaceous) gland of the skin (see chapter 5).
A summary of glandular epithelial tissue is presented in
table 4.2.
Knowledge Check
4. List the functions of simple squamous epithelia.
5. What are the three types of columnar epithelia? What do
they have in common? How are they different?
6. What are the two types of stratified squamous epithelia and
how do they differ?
7. Distinguish between unicellular and multicellular glands.
Explain how multicellular glands are classified according to
their mechanism of secretion.
8. In what ways are mammary glands and certain sweat
glands similar?
holocrine: Gk. holos, whole; krinein, to separate
CONNECTIVE TISSUE
Connective tissue is divided into subtypes according to the matrix
that binds the cells. Connective tissue provides structural and
metabolic support for other tissues and organs of the body.
Objective 7
Describe the general characteristics, locations,

and functions of connective tissue.
Objective 8
Explain the functional relationship between

embryonic and adult connective tissue.
Objective 9
List the various ground substances, fiber types,

and cells that constitute connective tissue and explain their
functions.
Characteristics and Classification

of Connective Tissue
Connective tissue is the most abundant tissue in the body. It supports other tissues or binds them together and provides for the
metabolic needs of all body organs. Certain types of connective
tissue store nutritional substances; other types manufacture protective and regulatory materials.
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Function

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Mesenchymal
cell
CHAPTER 4
Matrix
(a)
(b)
FIGURE 4.13 Mesenchyme is a type of embryonic connective tissue that can migrate and give rise to all other kinds of connective tissue.
(a) It is found within an early developing embryo and (b) consists of irregularly shaped cells lying in a jellylike homogeneous matrix.
Although connective tissue varies widely in structure and

function, all types of connective tissue have similarities. With
the exception of mature cartilage, connective tissue is highly vascular and well nourished. It is able to replicate and, by so doing,
is responsible for the repair of body organs. Unlike epithelial
tissue, which is composed of tightly fitted cells, connective tissue
contains considerably more matrix (intercellular material) than
cells. Connective tissue does not occur on free surfaces of body
cavities or on the surface of the body, as does epithelial tissue.
Furthermore, connective tissue is embryonically derived from
mesoderm, whereas epithelial tissue derives from ectoderm,
mesoderm, and endoderm.
The classification of connective tissue is not exact, and
several schemes have been devised. In general, however, the
various types are named according to the kind and arrangement
of the matrix. The following are the basic kinds of connective
tissues:
A. Embryonic connective tissue
B. Connective tissue proper
1. Loose (areolar) connective tissue
2. Dense regular connective tissue
3. Dense irregular connective tissue
4. Elastic connective tissue
5. Reticular connective tissue
6. Adipose tissue
C. Cartilage
1. Hyaline cartilage
2. Fibrocartilage
3. Elastic cartilage
D. Bone tissue
E. Blood (vascular tissue)
Embryonic Connective Tissue

The embryonic period of development, which lasts 6 weeks
(from the start of the third to the end of the eighth week), is
characterized by extensive tissue differentiation and organ formation. At the beginning of the embryonic period, all connective
tissue looks alike and is referred to as mesenchyme (mezen-km).
Mesenchyme is undifferentiated embryonic connective tissue
that is derived from mesoderm. It consists of irregularly shaped
cells surrounded by large amounts of a homogeneous, jellylike
matrix (fig. 4.13). In certain periods of development, mesenchyme migrates to predisposed sites where it interacts with
other tissues to form organs. Once mesenchyme has completed
its embryonic migration to a predetermined destination, it differentiates into all other kinds of connective tissue.
Some mesenchymal-like tissue persists past the embryonic
period in certain sites within the body. Good examples are the
undifferentiated cells that surround blood vessels and form fibroblasts if the vessels are traumatized. Fibroblasts assist in healing wounds (see chapter 5).
Another kind of prenatal connective tissue exists only in
the fetus (the fetal period is from 9 weeks to birth) and is called
mucous connective tissue or Whartons jelly. It gives a turgid consistency to the umbilical cord.
Connective Tissue Proper

Connective tissue proper has a loose, flexible matrix, frequently
called ground substance. The most common cell within connec-
Whartons jelly: from Thomas Wharton, English anatomist, 161473


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Pectoralis
major
muscle
Elastic fiber
Collagenous
fiber
Mast cell
Fibroblast
(b)
(a)
Paras
Fascia
FIGURE 4.14 Loose connective tissue is packing and binding tissue that surrounds muscles (a), nerves, and vessels and binds the skin to
the underlying muscles. (b) A photomicrograph of the tissue and (c) a labeled diagram.
tive tissue proper is called a fibroblast (fibro-blast). Fibroblasts

are large, star-shaped cells that produce collagenous (ko-lajenus), elastic, and reticular (re-tikyoo-lar) fibers. Collagenous
fibers are composed of a protein called collagen (kola-jen); they
are flexible, yet they have tremendous strength. Elastic fibers are
composed of a protein called elastin, which provides certain tissues with elasticity. Collagenous and elastic fibers may be either
sparse and irregularly arranged, as in loose connective tissue, or
tightly packed, as in dense connective tissue. Tissues with loosely
arranged fibers generally form packing material that cushions and
protects various organs, whereas those that are tightly arranged
form the binding and supportive connective tissues of the body.
Resilience in tissues that contain elastic fibers is extremely important for several physical functions of the body. Consider, for
example, that elastic fibers are found in the walls of large arteries and in
the walls of the lower respiratory passageways. As these walls are expanded by blood moving through vessels or by inspired air, the elastic
fibers must first stretch and then recoil. This maintains the pressures of
the fluid or air moving through the lumina, thus ensuring adequate flow
rates and rates of diffusion through capillary and lung surfaces.
Reticular fibers reinforce by branching and joining to

form a delicate lattice or reticulum. Reticular fibers are common
in lymphatic glands, where they form a meshlike center called
the stroma.
Six basic types of connective tissue proper are generally

recognized. These tissues are distinguished by the consistency of
the ground substance and the type and arrangement of the reinforcement fibers.
Loose Connective (Areolar) Tissue

Loose connective tissue is distributed throughout the body as a
binding and packing material. It binds the skin to the underlying
muscles and is highly vascular, providing nutrients to the skin.
Loose connective tissue that binds skin to underlying muscles is
known as fascia (fashe-a). It also surrounds blood vessels and
nerves, where it provides both protection and nourishment. Specialized cells called mast cells are dispersed throughout the loose
connective tissue surrounding blood vessels. Mast cells produce heparin (hepa-rin), an anticoagulant that prevents blood from clotting
within the vessels. They also produce histamine, which is released
during inflammation. Histamine acts as a powerful vasodilator.
The cells of loose connective tissue are predominantly fibroblasts, with collagenous and elastic fibers dispersed throughout the ground substance (fig. 4.14). The irregular arrangement
of this tissue provides flexibility, yet strength, in any direction. It
is this tissue layer, for example, that permits the skin to move
when a part of the body is rubbed.
collagen: Gk, kolla, glue

elastin: Gk. elasticus, to drive
reticular: L. rete, net or netlike
stroma: Gk. stroma, a couch or bed
fascia: L. fascia, a band or girdle

heparin: Gk. hepatos, the liver
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Fibroblast
CHAPTER 4
Tendon of
long head
of biceps brachii m.
Collagenous
fibers
Biceps
brachii m.
(a)
Tendon
of short
head of
biceps
brachii m.
(b)
(c)
Paras
FIGURE 4.15 Dense regular connective tissue forms the strong and highly flexible tendons (a) and ligaments. (b) A photomicrograph of the
Much of the fluid of the body is found within loose connective

tissue and is called interstitial fluid (tissue fluid). Sometimes
excessive tissue fluid accumulates, causing a swollen condition
called edema (e-dema). Edema is a symptom of numerous dysfunctions or disease processes.
Dense Regular Connective Tissue

Dense regular connective tissue is characterized by large amounts
of densely packed collagenous fibers that run parallel to the direction of force placed on the tissue during body movement. Because this tissue is silvery white in appearance, it is sometimes
called white fibrous connective tissue.
Dense regular connective tissue occurs where strong, flexible support is needed (fig. 4.15). Tendons, which attach muscles
to bones and transfer the forces of muscle contractions, and ligaments, which connect bone to bone across articulations, are
composed of this type of tissue.
Trauma to ligaments, tendons, and muscles are common
sports-related injuries. A strain is an excessive stretch of the
tissue composing the tendon or muscle, with no serious damage. A
sprain is a tearing of the tissue of a ligament and may be slight, moderate, or complete. A complete tear of a major ligament is especially
painful and disabling. Ligamentous tissue does not heal well because it has a poor blood supply. Surgical reconstruction is generally
needed for the treatment of a severed ligament.
tendon: L. tendere, to stretch

ligament: L. ligare, bind
Dense Irregular Connective Tissue

Dense irregular connective tissue is characterized by large
amounts of densely packed collagenous fibers that are interwoven to provide tensile strength in any direction. This tissue
is found in the dermis of the skin and the submucosa of the
GI tract. It also forms the fibrous capsules of organs and joints
(fig. 4.16).
Elastic Connective Tissue

Elastic connective tissue is composed primarily of elastic fibers
that are irregularly arranged and yellowish in color (fig. 4.17).
They can be stretched to one and a half times their original
lengths and will snap back to their former size. Elastic connective tissue is found in the walls of large arteries, in portions of the
larynx, and in the trachea and bronchial tubes of the lungs. It is
also present between the arches of the vertebrae that make up
the vertebral column.
Reticular Connective Tissue

Reticular connective tissue is characterized by a network of reticular fibers woven through a jellylike matrix (fig. 4.18). Certain
specialized cells within reticular tissue are phagocytic (fago-sitik)
(macrophages) and therefore can ingest foreign materials. The
liver, spleen, lymph nodes, and bone marrow contain reticular
connective tissue.


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(b)
(c)
(a)
FIGURE 4.16 Dense irregular connective tissue forms joint capsules (a) that contain synovial fluid for lubricating movable joints. (b) A photomicrograph of the tissue and (c) a labeled diagram.
Tunica
intima
(inner coat)
Elastic fibers
Endothelial cells
Elastic tissue
Fibroblast
(b)
Paras
(c)
(a)
FIGURE 4.17 Elastic connective tissue permits stretching of a large artery (a) as blood flows through. (b) A photomicrograph of the tissue and
(c) a labeled diagram.
Adipose Tissue
Adipose tissue is a specialized type of loose fibrous connective
tissue that contains large quantities of adipose cells, or
adipocytes (ad-po-sts). Adipose cells form from mesenchyme
adipose: L. adiposus, fat
and, for the most part, are formed prenatally and during the
first year of life. Adipose cells store droplets of fat within their
cytoplasm, causing them to swell and forcing their nuclei to
one side (fig. 4.19).
Adipose tissue is found throughout the body but is concentrated around the kidneys, in the hypodermis of the skin,
on the surface of the heart, surrounding joints, and in the
CHAPTER 4
Collagenous
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Reticular
fibers
Reticular cell
Nucleus of
reticular cell
CHAPTER 4
(b)
(c)
Spleen
(a)
Paras
FIGURE 4.18 Reticular connective tissue forms the stroma, or framework, of such organs as the spleen (a), liver, thymus, and lymph nodes.
(b) A photomicrograph of this tissue and (c) a labeled diagram.
Nucleus of
adipose cell
Fat droplet
Dermis
Cytoplasm
(b)
Hypodermis
(c)
Hair
follicle
(a)
Paras
FIGURE 4.19 Adipose tissue is abundant in the hypodermis of the skin (a) and around various internal organs. (b) A photomicrograph of the
breasts of sexually mature females. Fat functions not only as a

food reserve, but also supports and protects various organs. It
is a good insulator against cold because it is a poor conductor
of heat.
Excessive fat can be unhealthy by placing a strain on the heart
and perhaps causing early death. For these reasons, good exercise programs and sensible diets are extremely important. Adipose
tissue can also retain lipid-soluble, environmental pollutants that are
ingested or absorbed through the skin. Dieting eliminates the fat
stored within adipose tissue but not the tissue itself.
The surgical procedure of suction lipectomy may be used to

remove small amounts of adipose tissue from localized body areas
such as the breasts, abdomen, buttocks, and thighs. Suction lipectomy is used for cosmetic purposes rather than as a treatment for
obesity, and the risks for potentially detrimental side effects need to
be seriously considered. Potential candidates should be between 30
and 40 years old and only about 15 to 20 pounds overweight. They
should also have good skin elasticity.
The characteristics, functions, and locations of connective

tissue proper are summarized in table 4.3.


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TABLE 4.3 Summary of Connective Tissue Proper

Location
Loose connective (areolar) tissue
Predominantly fibroblast cells with lesser amounts of

collagen and elastin proteins; binds organs, holds
tissue fluids
Surrounding nerves and vessels, between muscles,

beneath the skin
Dense regular connective tissue
Densely packed collagenous fibers that run parallel to the

direction of force; provides strong, flexible support
Tendons, ligaments
Dense irregular connective tissue
Densely packed collagenous fibers arranged in a tight

interwoven pattern; provides tensile strength in any
direction
Dermis of skin, fibrous capsules of organs and joints,

periosteum of bone
Elastic connective tissue
Predominantly irregularly arranged elastic fibers; supports,

provides framework
Large arteries, lower respiratory tract, between the

arches of vertebrae
Reticular connective tissue
Reticular fibers that form a supportive network; stores,

performs phagocytic function
Lymph nodes, liver, spleen, thymus, bone marrow
Adipose tissue
Adipose cells; protects, stores fat, insulates
Hypodermis of skin, surface of heart, omentum, around

kidneys, back of eyeball, surrounding joints
Cartilage
Cartilage (kart-lij) consists of cartilage cells, or chondrocytes
(kondro-sts), and a semisolid matrix that imparts marked elastic
properties to the tissue. It is a supportive and protective connective tissue that is frequently associated with bone. Cartilage
forms a precursor to one type of bone and persists at the articular
surfaces on the bones of all movable joints.
The chondrocytes within cartilage may occur singly but
are frequently clustered. Chondrocytes occupy cavities, called lacunae
(la-kyoonesingular lacuna), within the matrix. Most cartilage is
surrounded by a dense irregular connective tissue called perichondrium (per-kondre-um). Cartilage at the articular surfaces
of bones (articular cartilage) lacks a perichondrium. Because mature cartilage is avascular, it must receive nutrients through diffusion from the perichondrium and the surrounding tissue. For this
reason, cartilaginous tissue has a slow rate of mitotic activity; if
damaged, it heals with difficulty.
There are three kinds of cartilage: hyaline (hia-ln) cartilage,
fibrocartilage, and elastic cartilage. They are distinguished by the
type and amount of fibers embedded within the matrix.
Hyaline Cartilage
Hyaline cartilage, commonly called gristle, has a homogeneous,
bluish-staining matrix in which the collagenous fibers are so fine
that they can be observed only with an electron microscope.
When viewed through a light microscope, hyaline cartilage has a
clear, glassy appearance (fig. 4.20).
Hyaline cartilage is the most abundant cartilage within the
body. It covers the articular surfaces of bones, supports the tubular trachea and bronchi of the respiratory system, reinforces the
nose, and forms the flexible bridge, called costal cartilage, be-
lacuna: L. lacuna, hole or pit

hyaline: Gk. hyalos, glass
tween the anterior portion of each of the first 10 ribs and the
sternum. Most of the bones of the body form first as hyaline cartilage and later become bone in a process called endochondral
ossification.
Fibrocartilage
Fibrocartilage has a matrix that is reinforced with numerous collagenous fibers (fig. 4.21). It is a durable tissue adapted to withstand tension and compression. It is found at the symphysis
pubis, where the two pelvic bones articulate, and between the
vertebrae as intervertebral discs. It also forms the cartilaginous
wedges within the knee joint, called menisci (see chapter 8).
By the end of the day, the intervertebral discs of the vertebral
column are somewhat compacted. So a person is actually
slightly shorter in the evening than in the morning, following a recuperative rest. Aging, however, brings with it a gradual compression of
the intervertebral discs that is irreversible.
Elastic Cartilage
Elastic cartilage is similar to hyaline cartilage except for the presence of abundant elastic fibers that make elastic cartilage very
flexible without compromising its strength (fig. 4.22). The numerous elastic fibers also give it a yellowish appearance. This tissue is found in the outer ear, portions of the larynx, and in the
auditory canal.
The three types of cartilage are summarized in table 4.4.
Bone Tissue
Bone tissue is the most rigid of all the connective tissues. Unlike
cartilage, bone tissue has a rich vascular supply and is the site of
considerable metabolic activity. The hardness of bone is largely
due to the calcium phosphate (calcium hydroxyapatite) deposited
within the intercellular matrix. Numerous collagenous fibers, also
embedded within the matrix, give bone some flexibility.
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Type

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Lacuna
Intercellular
matrix
Chondrocyte
Thyroid
cartilage
Larynx
CHAPTER 4
(b)
(c)
Cricoid
cartilage
Tracheal
cartilages
(a)
Paras
FIGURE 4.20 Hyaline cartilage is the most abundant cartilage in the body. It occurs in places such as the larynx (a), trachea, portions of the
rib cage, and embryonic skeleton. (b) A photomicrograph of the tissue and (c) a labeled diagram.
Lacuna
Chondrocyte
Intercellular
matrix
Collagenous
fibers
(b)
(c)
(a)
FIGURE 4.21 Fibrocartilage is located at the symphysis pubis, within the knee joints, and between the vertebrae as the intervertebral discs
(a). A photomicrograph of the tissue is shown in (b) and a labeled diagram in (c).


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Lacuna
Chondrocyte
Elastic fibers
Auricular
cartilage
(b)
Paras
FIGURE 4.22 Elastic cartilage gives support to the outer ear (a), auditory canal, and parts of the larynx. A photomicrograph of the tissue is
shown in (b) and a labeled diagram in (c).
TABLE 4.4 Summary of Cartilage

Type
Location
Hyaline cartilage
Homogeneous matrix with extremely fine collagenous fibers;

provides flexible support, protects, is precursor to bone
Articular surfaces of bones, nose, walls of respiratory passages,

fetal skeleton
Fibrocartilage
Abundant collagenous fibers within matrix; supports, withstands

compression
Symphysis pubis, intervertebral discs, knee joint
Elastic cartilage
Abundant elastic fibers within matrix; supports, provides flexibility
Framework of outer ear, auditory canal, portions of larynx
When bone is placed in a weak acid, the calcium salts dissolve

away and the bone becomes pliable. It retains its basic shape
but can be easily bent and twisted (fig. 4.23). In calcium deficiency
diseases, such as rickets, the bone tissue becomes pliable and
bends under the weight of the body (see fig. 5.11).
Based on porosity, bone tissue is classified as either compact or spongy, and most bones have both types (fig. 4.24). Compact (dense) bone tissue constitutes the hard outer portion of a
bone, and spongy (cancellous) bone tissue constitutes the porous,
highly vascular inner portion. The outer surface of a bone is covered by a connective tissue layer called the periosteum that serves
as a site of attachment for ligaments and tendons, provides protection, and gives durable strength to the bone. Spongy bone tissue makes the bone lighter and provides a space for red bone
marrow, where blood cells are produced.
In compact bone tissue, mature bone cells, called
osteocytes, are arranged in concentric layers around a central
FIGURE 4.23 A bone soaked in a weak acid, such as the acidic
acid in vinegar, demineralizes and becomes flexible.
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(a)
(c)

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CHAPTER 4
(b)
Lamellae
(a)
Central canal
Osteocyte
within a
lacuna
Canaliculi
(c)
FIGURE 4.24 Bone (a) consists of compact and spongy tissues. (b) A photomicrograph of compact bone tissue and (c) a labeled diagram.
(haversian) canal, which contains a vascular and nerve supply.

Each osteocyte occupies a cavity called a lacuna. Radiating from
each lacuna are numerous minute canals, or canaliculi, which
traverse the dense matrix of the bone tissue to adjacent lacunae.
Nutrients diffuse through the canaliculi to reach each osteocyte.
The matrix is deposited in concentric layers called lamellae.
Bone tissue is described in detail in chapter 6.
Blood (Vascular Tissue)

Blood, or vascular tissue, is a highly specialized fluid connective
tissue that plays a vital role in maintaining homeostasis. The
cells, or formed elements, of blood are suspended in a liquid
matrix called blood plasma (fig. 4.25). The three types of formed
elements are erythrocytes (red blood cells), leukocytes (white
blood cells), and thrombocytes (platelets). Blood is discussed
fully in chapter 16.
haversian canal: from Clopton Havers, English anatomist, 16501702

erythrocyte: Gk. erythros, red; kytos, hollow (cell)
leukocyte: Gk. leukos, white; kytos, hollow (cell)
thrombocyte: Gk. thrombos, a clot; kytos, hollow (cell)
An injury to a portion of the body may stimulate tissue repair

activity, usually involving connective tissue. A minor scrape or
cut results in platelet and plasma activity of the exposed blood and
the formation of a scab. The epidermis of the skin regenerates beneath the scab. A severe open wound heals through connective tissue granulation. In this process, collagenous fibers form from
surrounding fibroblasts to strengthen the traumatized area. The
healed area is known as a scar.
Knowledge Check
9. List the basic types of connective tissue and describe the
structure, function, and location of each.
10. Which of the previously discussed connective tissues
function to protect body organs? Which type is
phagocytic? Which types bind and support various structures? Which types are associated in some way with
the skin?
11. What is the developmental significance of mesenchyme and
how does it differ functionally from adult connective tissue?
12. Briefly describe reticular fibers, fibroblasts, collagenous
fibers, elastic fibers, and mast cells.


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Erythrocytes
Leukocytes
Thrombocytes (platelets)
Paras
FIGURE 4.25 Blood consists of formed elementserythrocytes (red blood cells), leukocytes (white blood cells), and thrombocytes
(platelets)suspended in a liquid plasma matrix.
MUSCLE TISSUE
Muscle tissue is responsible for the movement of materials
through the body, the movement of one part of the body with respect to another, and for locomotion. Fibers in the three kinds of
muscle tissue are adapted to contract in response to stimuli.
Objective 10
Describe the structure, location, and function

of the three types of muscle tissue.
Muscle tissue is unique in its ability to contract, and thus make

movement possible. The muscle cells, or fibers, are elongated in
the direction of contraction, and movement is accomplished
through the shortening of the fibers in response to a stimulus.
Muscle tissue is derived from mesoderm. There are three types of
muscle tissue in the body: smooth, cardiac, and skeletal muscle tissue (fig. 4.26).
Smooth muscle fibers are long, spindle-shaped cells. They

contain a single nucleus and lack striations. These cells are usually grouped together in flattened sheets, forming the muscular
portion of a wall around a lumen.
Cardiac Muscle
Cardiac muscle tissue makes up most of the wall of the heart.
This tissue is characterized by bifurcating (branching) fibers,
each with a single, centrally positioned nucleus, and by transversely positioned intercalated (in-terka-lat-ed) discs. Intercalated discs help to hold adjacent cells together and transmit the
force of contraction from cell to cell. Like skeletal muscle, cardiac muscle is striated, but unlike skeletal muscle it experiences
rhythmic involuntary contractions. Cardiac muscle is further discussed in chapter 16.
Skeletal Muscle
Smooth Muscle
Smooth muscle tissue is common throughout the body, occurring
in many of the systems. For example, in the wall of the GI tract
it provides the contractile force for the peristaltic movements involved in the mechanical digestion of food. Smooth muscle is
also found in the walls of arteries, the walls of respiratory passages, and in the urinary and reproductive ducts. The contraction of smooth muscle is under autonomic (involuntary) nervous
control, and is discussed in more detail in chapter 13.
Skeletal muscle tissue attaches to the skeleton and is responsible

for voluntary body movements. Each elongated, multinucleated
fiber has distinct transverse striations. Fibers of this muscle tissue
are grouped into parallel fasciculi (bundles) that can be seen
without a microscope in fresh muscle. Both cardiac and skeletal
muscle fibers cannot replicate once tissue formation has been
completed shortly after birth. Skeletal muscle tissue is further
discussed in chapter 9. The three types of muscle tissue are summarized in table 4.5.
CHAPTER 4
Cells
Plasma
Centrifuged
blood sample

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Nucleus
CHAPTER 4
(a) Smooth muscle tissue
Striation
Nucleus
Intercalated
disc
(b) Cardiac muscle tissue
Nucleus
Striation
(c) Skeletal muscle tissue
FIGURE 4.26 Muscle tissue: (a) smooth, (b) cardiac, and (c) skeletal.
TABLE 4.5 Summary of Muscle Tissue

Type
Location
Smooth
Elongated, spindle-shaped
fiber with single nucleus;
involuntary movements
of internal organs
Walls of hollow
internal organs
Cardiac
Branched, striated fiber

with single nucleus
and intercalated discs;
involuntary rhythmic
contraction
Heart wall
Skeletal
Multinucleated, striated,
cylindrical fiber that
occurs in fasciculi;
voluntary movement
of skeletal parts
Associated with
skeleton; spans
joints of skeleton
via tendons
Knowledge Check
13. Describe the general characteristics of muscle tissue.
What is meant by voluntary and involuntary as applied to
muscle tissue?
14. Distinguish between smooth, cardiac, and skeletal muscle
tissue on the bases of structure, location, and function.
NERVOUS TISSUE
Nervous tissue is composed of neurons, which respond to stimuli
and conduct impulses to and from all body organs, and neuroglia,
which functionally support and physically bind neurons.
Objective 11
Describe the basic characteristics and

functions of nervous tissue.
Objective 12
Distinguish between neurons and neuroglia.


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Cell body
Capillary
Axon
Astrocyte
Dendrite
Vascular
processes
CHAPTER 4
(a) Neurons
(b) Neuroglia
FIGURE 4.27 Nervous tissue is found within the brain, spinal cord, nerves, and ganglia. It consists of two principal kinds of cells: (a) neurons
and (b) neuroglia.
Neurons
Neuroglia
Although there are several kinds of neurons (nooronz) in nervous tissue, they all have three principal components: (1) a
cell body, or perikaryon; (2) dendrites; and (3) an axon
(fig. 4.27). Dendrites are branched processes that receive stimuli and conduct nerve impulses toward the cell body. The cell
body, or perikaryon (per -kare-on), contains the nucleus and
specialized organelles and microtubules. The axon is a cytoplasmic extension that conducts nerve impulses away from the
cell body. The term nerve fiber refers to any process extending
from the cell body of a neuron and the myelin sheath that surrounds it (see fig. 11.5).
Neurons derive from ectoderm and are the basic structural and functional units of the nervous system. They are specialized to respond to physical and chemical stimuli, convert
stimuli into nerve impulses, and conduct these impulses to
other neurons, muscle fibers, or glands. Of all the bodys cells,
neurons are probably the most specialized. As with muscle cells,
the number of neurons is established prenatally (before
birth); thereafter, they lack the ability to undergo mitosis,
although under certain circumstances a severed portion can regenerate.
In addition to neurons, nervous tissue contains neuroglia (noorogle-a) (fig. 4.27). Neuroglial cells, sometimes called glial cells,
are about 5 times as abundant as neurons and have limited mitotic abilities. They do not transmit impulses but support and
bind neurons together. Certain neuroglial cells are phagocytic;
others assist in providing sustenance to the neurons.
Neurons and neuroglia are discussed in detail in chapter 11.
neuron: Gk. neuron, sinew or nerve

perikaryon: Gk. peri, around; karyon, nut or kernel
Knowledge Check
15. Compare and contrast neurons and neuroglia in terms of
structure, function, and location.
16. List the structures of a neuron following the sequence of a
nerve impulse passing through the cell.
As stated at the beginning of this chapter, the study of tissues is
extremely important in understanding the structure and function
of organs and body systems. Histology has immense clinical
neuroglia: Gk. neuron, nerve; glia, glue

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importance as well. Many diseases are diagnosed through microscopic examination of tissue sections. Even in performing an autopsy, an examination of various tissues is vital in establishing
the cause of death.
Several sciences are concerned with specific aspects of tissues. Histopathology is the study of diseased tissues. Histochemistry
is concerned with the physiology of tissues as they maintain
homeostasis. Histotechnology explores the ways in which tissues
can be better stained and observed. In all of these disciplines, a
thorough understanding of normal, or healthy, tissues is imperative for recognizing altered, or abnormal, tissues.
CHAPTER 4
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Changes in Tissue Composition

Most diseases alter tissue structure locally, where the disease is
prevalent. Some diseases, however, called general conditions,
cause changes that are far removed from the locus of the disease.
Atrophy (wasting of body tissue), for example, may be limited to
a particular organ where the disease interferes with the metabolism of that organ, but it may also involve an entire limb if nourishment or nerve impulses are impaired. Muscle atrophy can be
caused by a disease of the nervous system like polio, or it can be
the result of a diminished blood supply to a muscle. Senescence
(se-ne sens) atrophy, or simply senescence, is the natural aging of
tissues and organs within the body. Disuse atrophy is a local atrophy that results from the inactivity of a tissue or organ. Muscular dystrophy causes a disuse atrophy that decreases muscle size
and strength because of the loss of sarcoplasm within the muscle.
Necrosis (ne-krosis) is death of cells or tissues within the
living body. It can be recognized by physical changes in the dead
tissues. Necrosis can be caused by severe injury; physical agents
(trauma, heat, radiant energy, chemical poisons); or poor nutrition of tissues. When histologically examined, the necrotic tissue
usually appears opaque, with a whitish or yellowish cast. Gangrene is a massive necrosis of tissue accompanied by an invasion
of microorganisms that live on decaying flesh.
Somatic death is the death of the body as a whole. Following somatic death, tissues undergo irreversible changes, such as
rigor mortis (muscular rigidity), clotting of the blood, and cooling of the body. Postmortem (after death) changes occur under
varying conditions at predictable rates, which is useful in estimating the approximate time of death.
Tissue Analysis
In diagnosing a disease, it is frequently important to examine tissues from a living person histologically. When this is necessary,
a biopsy (biop-se) (removal of a section of living tissue) is
performed. There are several techniques for biopsies. Surgical
removal is usually done on large masses or tumors. Curettage
(kyoo re-tazh) involves cutting and scraping tissue, as may be
necrosis: Gk. nekros, corpse
gangrene: Gk. gangraina, gnaw or eat
done in examining for uterine cancer. In a percutaneous needle

biopsy, a biopsy needle is inserted through a small skin incision
and tissue samples are withdrawn. Both normal and diseased tissues are removed for purposes of comparison.
Preparing tissues for examination is a multistep process.
Fixation is fundamental for all histological preparation. It is the
rapid killing, hardening, and preservation of tissue to maintain
its existing structure. Embedding the tissue in a supporting
medium such as paraffin wax usually follows fixation. The next
step, sectioning the tissue into extremely thin slices, is followed
by mounting the specimen on a slide. Some tissues are fixed by
rapid freezing and then sectioned while frozen, making embedding unnecessary. Frozen sections enable the pathologist to make
a quick diagnosis during a surgical operation. These are done frequently, for example, in cases of suspected breast cancer. Staining is the next step. Hematoxylin and eosin (H & E) stains are
routinely used on all tissue specimens. They give a differential
blue and red color to the basic and acidic structures within the
tissue. Other dyes may be needed to stain for specific structures.
Examination is first done with the unaided eye and then
with a microscope. Practically all histological conditions can be
diagnosed with low magnification (40). Higher magnification is
used to clarify specific details. Further examination may be performed with an electron microscope, which reveals the intricacy
of cellular structure. Histological observation provides the foundation for subsequent diagnosis, prognosis, treatment, and
reevaluation.
Tissue Transplantation
In the last two decades, medical science has made tremendous advancements in tissue transplants. Tissue transplants are necessary
for replacing nonfunctional, damaged, or lost body parts. The
most successful transplant is one where tissue is taken from one
place on a persons body and moved to another place, such as a
skin graft from the thigh to replace burned tissue of the hand.
Transfer of ones own tissue is termed an autograft. Isografts are
transplants between genetically identical individuals, the only example being identical twins. These transplants also have a high
success rate. Allografts, or homotransplants, are grafts between
individuals of the same species but of different genotype, and
xenografts, or heterografts, are grafts between individuals of different species. An example of a xenograft is the transplant of a pig
valve to replace a dysfunctional or diseased human heart valve.
Both allografts and xenografts present the problem of a possible
tissue-rejection reaction. When this occurs, the recipients immune
mechanisms are triggered, and the donors tissue is identified as
foreign and is destroyed. The reaction can be minimized by
matching recipient and donor tissue. Immunosuppressive drugs
also may lessen the rejection rate. These drugs act by interfering
with the recipients immune mechanisms. Unfortunately, immunosuppressive drugs may also lower the recipients resistance to
infections. New techniques involving blood transfusions from
donor to recipient before a transplant are proving successful. In
any event, tissue transplants are an important aspect of medical
research, and significant breakthroughs are on the horizon.


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Chapter 4
The use of fetal tissue transplantation to grow new tissues
in adult patients has shown promise for the treatment of many
clinical problems of tissues or organs. Desired cells are harvested from as many as 15 human fetuses and are then quickly
implanted into the transplant recipient. The fetal cells are allowed to follow a normal course of maturation within the adult
patient, with the hope that a healthy, fully functional body
structure will develop. Because the typical source for fetal tissues is aborted fetuses, this procedure has become a medical
ethical dilemma.
Histology
103

The doctor clarifies this paradox by explaining that some cells in the
body undergo structural changes in response to unusual stimuli. Cigarette smoking, for example, impairs ciliary movement, inhibits function
of alveolar macrophages, and leads to enlargement and proliferation of
mucus-secreting glands in the airways. The net effect of this is chronic
bronchitis.
Similarly, acid reflux into the esophagus can induce a transformation of stratified squamous epithelium to simple columnar epithelium.
This condition is termed Barretts esophagus and is a precursor to
esophageal cancer in 5% of cases.
CHAPTER 4
Chapter Summary
Definition and Classification of Tissues
(p. 78)
1. Tissues are aggregations of similar cells
that perform specific functions. The study
of tissues is called histology.
2. Cells are surrounded and bound together
by an intercellular matrix, the
composition of which varies from solid to
liquid.
3. The four principal types of tissues are
epithelial tissue, connective tissue, muscle
tissue, and nervous tissue.
Epithelial Tissue (pp. 7989)

1. Epithelia are derived from all three germ
layers and may be one or several layers
thick. The lower surface of most
membranous epithelia is supported by a
basement membrane.
2. Simple epithelium consists of a single cell
layer that varies in shape and surface
characteristics. It is located where
diffusion, filtration, and secretion occur.
3. Stratified epithelium consists of two or
more layers of cells and is adapted for
protection.
4. Transitional epithelium lines the urinary

bladder, ureters, and parts of the urethra.
The cells of transitional epithelium
permit distension.
5. Body membranes are composed of thin
layers of epithelial tissue that may be
coupled with supporting connective
tissue. The two basic types are mucous
membranes and serous membranes.
6. Glandular epithelia are derived from
developing epithelial tissue and function
as secretory exocrine glands.
Connective Tissue (pp. 8998)

1. Connective tissues are derived from
mesoderm and, with the exception of
cartilage, are highly vascular.
2. Connective tissue proper contains
fibroblasts, collagenous fibers, and elastic
fibers within a flexible ground substance.
3. Cartilage provides a flexible framework
for many organs. It consists of a semisolid
matrix of chondrocytes and various fibers.
4. Bone tissue consists of osteocytes,
collagenous fibers, and a durable matrix of
mineral salts.
5. Blood consists of formed elements

(erythrocytes, leukocytes, and
thrombocytes) suspended in a fluid
plasma matrix.
Muscle Tissue (pp. 99100)

1. Muscle tissues (smooth, cardiac, and
skeletal) are responsible for the
movement of materials through the body,
the movement of one part of the body
with respect to another, and for
locomotion.
2. Fibers in muscle tissue are adapted to
contract in response to stimuli.
Nervous Tissue (pp. 100101)

1. Neurons are the functional units of the
nervous system. They respond to stimuli
and conduct nerve impulses to and from
all body organs.
2. Neuroglia support and bind neurons.
Some are phagocytic; others provide
sustenance to neurons.
Review Activities
Objective Questions
1. Which of the following is not a principal
type of body tissue?
(a) nervous
(d) muscular
(b) integumentary (e) epithelial
(c) connective
2. Which statement regarding tissues is false?
(a) They are aggregations of similar kinds
of cells that perform specific functions.
(b) All of them are microscopic and are

studied within the science of
histology.
(c) All of them are stationary within the
body at the location of their
developmental origin.
(d) An organ is composed of two or more
tissue types.
3. Connective tissue, muscle, and the dermis

of the skin derive from embryonic
(a) mesoderm.
(c) ectoderm.
(b) endoderm.
4. Which statement regarding epithelia is false?
(a) They are derived from mesoderm,
ectoderm, and endoderm.
(b) They are strengthened by elastic
and collagenous fibers.

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(c) One side is exposed to a lumen, a

body cavity, or to the external
environment.
(d) They have very little intercellular
matrix.
A gastric ulcer of the stomach would
involve
(a) simple cuboidal epithelium.
(b) transitional epithelium.
(c) simple ciliated columnar epithelium.
(d) simple columnar epithelium.
Which structural and secretory
designation describes mammary glands?
(a) acinar, apocrine
(b) tubular, holocrine
(c) tubular, merocrine
(d) acinar, holocrine
Dense regular connective tissue is
found in
(a) blood vessels. (c) tendons.
(b) the spleen.
(d) the wall of the
uterus.
The phagocytic connective tissue found
in the lymph nodes, liver, spleen, and
bone marrow is
(a) reticular.
(c) mesenchyme.
(b) loose fibrous. (d) elastic.
Cartilage is slow in healing following an
injury because
(a) it is located in body areas that are
under constant physical strain.
(b) it is avascular.
(c) its chondrocytes cannot reproduce.
(d) it has a semisolid matrix.
Cardiac muscle tissue has
(a) striations.
(b) intercalated discs.
(c) rhythmic involuntary contractions.
(d) all of the above.
Essay Questions
1. Define tissue. What are the differences
between cells, tissues, glands, and organs?
2. What physiological functions are epithelial
tissues adapted to perform?
3. Identify the epithelial tissue
(a) in the pulmonary alveoli of the lungs,
(b) lining the lumen of the GI tract,
(c) in the outer layer of skin,
(d) lining the cavity of the urinary
bladder,
(e) lining the uterine tube, and
(f) lining the trachea and bronchial
tubes.
Describe the function of the tissue in each
case.
4. Why are both keratinized and
nonkeratinized epithelia found within the
body?
5. Describe how epithelial glands are
classified according to structural
complexity and secretory function.
6. Identify the connective tissue
(a) on the surface of the heart and
surrounding the kidneys,
(b) within the wall of the aorta,
(c) forming the symphysis pubis,
(d) supporting the outer ear,
(e) forming the lymph nodes, and
(f) forming the tendo calcaneus.
Describe the function of the tissue in each
case.
7. Compare and contrast the structure and
location of the following: reticular fibers,
collagenous fibers, elastin, fibroblasts, and
mast cells.
8. What is the relationship between adipose
cells and fat? Discuss the function of fat
and explain the potential danger of
excessive fat.

9. Discuss the mitotic abilities of each of the

four principal tissue types.
10. Define the following terms: atrophy,
necrosis, gangrene, and somatic death.
1. The function of a tissue is actually a
function of its cells. And the function of a
cell is a function of its organelles.
Knowing this, what type of organelles
would be particularly abundant in cardiac
muscle tissue that requires a lot of energy;
in reticular tissue within the liver, where
cellular debris and toxins are ingested;
and in dense regular connective tissue
that consists of tough protein strands?
2. The aorta (a principal blood vessel) has
three layers surrounding its lumen. What
is the predominant tissue in each of the
three layers and what is the adaptive
function of each?
3. Your aunt was recently diagnosed as having
brain cancer. In talking with your aunts
physician, she indicated that the cancer
was actually a neuroglioma, and went on to
say that cancer of neurons and muscle cells
is a rare occurrence. Explain why neuroglial
cells are much more susceptible to cancer
than are neurons or muscle cells.
4. Compare the vascular supply of bones and
ligaments and discuss how this may be
relevant to the clinical course of an ankle
sprain and an ankle fracture.
5. The connective tissue diseases are a group of
disorders most likely caused by an abnormal
immune response to a persons own
connective tissue. The best known of these
is rheumatoid arthritis, in which small joints
of the body become inflamed and the
articulating surfaces erode away. Knowing
where connective tissue is found in the
body, can you predict which organs might be
involved in other connective tissue diseases?

IV. Support and Movement
5. Integumentary System
Integumentary System
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5
The Skin as an Organ 106
Layers of the Skin 106
Functions of the Skin 112
Epidermal Derivatives 115
The Integumentary System 120
Important Clinical Terminology 128
Chapter Summary 129
Clinical Case Study

A 27-year-old male was involved in a gasoline explosion and sustained burns to his face, neck,
chest, and arms. Upon arrival at the emergency room, he complained of intense pain on his face
and neck, both of which exhibited extensive blistering and erythema (redness). These findings
were all curiously absent on the burned chest and arms, which had a pale, waxy appearance.
Examination revealed the skin on the patients chest and arms to be leathery and lacking
sensation. The emergency room physician commented to an observing medical student that
third-degree burns were present on the skin of these regions and that excision of the burn eschar (traumatized tissue) with subsequent skin grafting would be required.
Why would the areas that sustained second-degree burns be red, blistered, and painful,
while the third-degree burns were pale and insensate (without sensation, including pain)? Why
would the chest and arms require skin grafting, but probably not the face and neck?
Hints: Think in terms of functions of the skin and survival of the germinal cells in functioning
skin. Carefully examine figures 5.1 and 5.20.
FIGURE: Immediate medical attention is

essential in an attempt to save a person who
has experienced an extensive and severe
burn. Of major concern is the rapid loss of
body fluids.

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Support and Movement
THE SKIN AS AN ORGAN

The skin (integument) is the largest organ of the body, and together with its accessory organs (hair, glands, and nails), it constitutes the integumentary system. In certain areas of the body, it
has adaptive modifications that accommodate protective or metabolic functions. In its role as a dynamic interface between the continually changing external environment and the bodys internal
environment, the skin helps maintain homeostasis.
Objective 1
Explain why the skin is considered an organ

and a component of the integumentary system.
Objective 2
Describe some common clinical conditions of

the skin that result from nutritional deficiencies or body
dysfunctions.
CHAPTER 5
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We are more aware of and concerned with our integumentary

system than perhaps any other system of our body. One of the
first things we do in the morning is to look in a mirror and see
what we have to do to make our skin and hair presentable. Periodically, we examine our skin for wrinkles and our scalp for gray
hairs as signs of aging. We recognize other people to a large extent by features of their skin.
The appearance of our skin frequently determines the initial impression we make on others. Unfortunately, it may also determine whether or not we succeed in gaining social acceptance.
For example, social rejection as a teenager, imagined or real, can
be directly associated with skin problems such as acne. A persons self-image and consequent social behavior may be closely
associated with his or her physical appearance.
Even clothing styles are somewhat determined by how
much skin we, or the designers, want to expose. But our skin is
much more than a showpiece. It helps regulate certain body
functions and protect certain body structures.
The skin, or integument (in-teg'yoo-ment), and its accessory
structures (hair, glands, and nails) constitute the integumentary
system. Included in this system are the millions of sensory receptors of the skin and its extensive vascular network. The skin is a
dynamic interface between the body and the external environment. It protects the body from the environment even as it allows for communication with the environment.
The skin is an organ, because it consists of several kinds
of tissues that are structurally arranged to function together. It
is the largest organ of the body, covering over 7,600 sq cm
(3,000 sq in.) in the average adult, and accounts for approximately 7% of a persons body weight. The skin is of variable
thickness, averaging 1.5 mm. It is thickest on the parts of the
body exposed to wear and abrasion, such as the soles of the feet
and palms of the hand. In these areas, it is about 6 mm thick. It
is thinnest on the eyelids, external genitalia, and tympanic membrane (eardrum), where it is approximately 0.5 mm thick. Even
integument: L. integumentum, a covering
its appearance and texture varies from the rough, callous skin
covering the elbows and knuckles to the soft, sensitive areas of
the eyelids, nipples, and genitalia.
The general appearance of the skin is clinically important
because it provides clues to certain body dysfunctions. Pale skin
may indicate shock, whereas red, flushed, overwarm skin may indicate fever and infection. A rash may indicate allergies or local
infections. Abnormal textures of the skin may be the result of
glandular or nutritional problems (table 5.1). Even chewed fingernails may be a clue to emotional problems.
Knowledge Check
1. Explain why the skin is considered an organ and why the
skin, together with the integumentary derivatives, is considered a system.
2. Which vitamins and minerals are important for healthy
skin? (See table 5.1.)
3. Describe the appearance of the skin that may accompany
each of the following conditions: allergy; shock; infection;
dry, stiff hair; hyperpigmentation; and general dermatitis.
LAYERS OF THE SKIN

The skin consists of two principal layers. The outer epidermis is
stratified into four or five structural layers, and the thick and
deeper dermis consists of two layers. The hypodermis (subcutaneous tissue) connects the skin to underlying organs.
Objective 3
Describe the histological characteristics of

each layer of the skin.
Objective 4
Summarize the transitional events that occur

within each of the epidermal layers.
Epidermis
The epidermis (ep''-der'mis) is the superficial protective layer of
the skin. Derived from ectoderm, the epidermis is composed of
stratified squamous epithelium that varies in thickness from
0.007 to 0.12 mm. All but the deepest layers are composed of
dead cells. Either four or five layers may be present, depending
on where the epidermis is located (figs. 5.1 and 5.2). The epidermis of the palms and soles has five layers because these areas are
exposed to the most friction. In all other areas of the body, the
epidermis has only four layers. The names and characteristics of
the epidermal layers are as follows.
1. Stratum basale (basal layer). The stratum basale (stra'tum ba-sal'e) consists of a single layer of cells in contact
with the dermis. Four types of cells compose the stratum
stratum: L. stratum, something spread out

basale: Gk. basis, base

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TABLE 5.1 Conditions of the Skin and Associated Structures Indicating Nutritional
Deficiencies or Body Dysfunctions
Condition
Deficiency
Comments
General dermatitis
Zinc
Redness and itching
Scrotal or vulval dermatitis
Riboflavin
Inflammation in genital region
Hyperpigmentation
Vitamin B12, folic acid, or starvation
Dark pigmentation on backs of hands and feet
Dry, stiff, brittle hair
Protein, calories, and other nutrients
Usually occurs in young children or infants
Follicular hyperkeratosis
Vitamin A, unsaturated fatty acids
Rough skin caused by keratotic plugs from hair follicles
Pellagrous dermatitis
Niacin and tryptophan
Lesions on areas exposed to sun
Thickened skin at pressure points
Niacin
Noted at belt area at the hips
Spoon nails
Iron
Thin nails that are concave or spoon-shaped
Dry skin
Water or thyroid hormone
Oily skin (acne)
Dehydration, hypothyroidism, rough skin

Hyperactivity of sebaceous glands
CHAPTER 5
Adipose
tissue
FIGURE 5.1 A diagram of the skin.
Hair
bulb
Hair
follicle

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Epidermis
Papillary
layer
of dermis
Dermis
Reticular
layer
of dermis
FIGURE 5.3 A scanning electron micrograph of the surface of the

skin showing the opening of a sweat gland.
CHAPTER 5
FIGURE 5.2 A light micrograph of the epidermis (25).

basale: keratinocytes (ker''a-tin'o-sts), melanocytes (mel'a-nosts), tactile cells (Merkel cells), and nonpigmented granular
dendrocytes (Langerhans cells). With the exception of tactile
cells, these cells are constantly dividing mitotically and
moving outward to renew the epidermis. It usually takes
between 6 to 8 weeks for the cells to move from the stratum basale to the surface of the skin.
Keratinocytes are specialized cells that produce the protein keratin (ker'a-tin), which toughens and waterproofs
the skin. As keratinocytes are pushed away from the vascular nutrient and oxygen supply of the dermis, their nuclei
degenerate, their cellular content becomes dominated by
keratin, and the process of keratinization is completed. By
the time keratinocytes reach the surface of the skin, they
resemble flat dead scales. They are completely filled with
keratin enclosed in loose cell membranes. Melanocytes are
specialized epithelial cells that synthesize the pigment
melanin (mel'a-nin) which provides a protective barrier to
the ultraviolet radiation in sunlight. Tactile cells are sparse
compared to keratinocytes and melanocytes. These sensory
receptor cells aid in tactile (touch) reception. Nonpigmented granular dendrocytes are scattered throughout the
stratum basale. They are protective macrophagic cells that
ingest bacteria and other foreign debris.
2. Stratum spinosum (spiny layer). The stratum spinosum

(spi-no'sum) contains several layers of cells. The spiny appearance of this layer is due to the spinelike extensions
that arise from the keratinocytes when the tissue is fixed
for microscopic examination. Because there is limited mitosis in the stratum spinosum, this layer and the stratum
basale are collectively referred to as the stratum germinativum (jer-m''na-ti'vum).
3. Stratum granulosum (granular layer). The stratum granulosum (gran''yoo-lo'sum) consists of only three or four flattened layers of cells. These cells contain granules that are
filled with keratohyalin, a chemical precursor to keratin.
4. Stratum lucidum (clear layer). The nuclei, organelles,
and cell membranes are no longer visible in the cells of the
stratum lucidum (loo's-dum), and so histologically this
layer appears clear. It exists only in the lips and in the
thickened skin of the soles and palms.
5. Stratum corneum (hornlike layer). The stratum
corneum (kor'ne-um) is composed of 25 to 30 layers of flattened, scalelike cells. Thousands of these dead cells shed
from the skin surface each day, only to be replaced by new
ones from deeper layers. This surface layer is cornified; it is
the layer that actually protects the skin (fig. 5.3). Cornification, brought on by keratinization, is the drying and flattening of the stratum corneum and is an important
protective adaptation of the skin. Friction at the surface of
keratinocyte: Gk. keras, homlike; kytos, cell
spinosum: L. spina, thorn
melanocyte: Gk. melas, black; kytos, cell

Merkel cells: from F. S. Merkel, German anatomist, 18451919
germinativum: L. germinare, spout or growth

granulosum: L. granum, grain
Langerhans cells: from Paul Langerhans, German anatomist, 18471888

macrophagic: L. makros, large; phagein, to eat
lucidum: L. lucidus, light

corneum: L. corneus, hornlike

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TABLE 5.2 Layers of the Epidermis
Stratum corneum
Consists of many layers of keratinized dead cells
that are flattened and nonnucleated; cornified
Stratum lucidum
A thin, clear layer found only in the epidermis of the
lips, palms, and soles
Stratum granulosum
Composed of one or more layers of granular cells
that contain fibers of keratin and shriveled nuclei
Stratum spinosum
Composed of several layers of cells with centrally
located, large, oval nuclei and spinelike processes;
limited mitosis
the skin stimulates additional mitotic activity in the stratum basale and stratum spinosum, which may result in the
formation of a callus for additional protection.
The specific characteristics of each epidermal layer are described in table 5.2.
Tattooing colors the skin permanently because dyes are injected below the mitotic basal layer of the epidermis into
the underlying dermis. In nonsterile conditions, infectious organisms
may be introduced along with the dye. Small tattoos can be removed
by skin grafting; for large tattoos, mechanical abrasion of the skin is
preferred.
Coloration of the Skin

Normal skin color is the expression of a combination of three
pigments: melanin, carotene, and hemoglobin. Melanin is a brownblack pigment produced in the melanocytes of the stratum basale
(fig. 5.4). All individuals of similar size have approximately the
same number of melanocytes, but the amount of melanin produced and the distribution of the melanin determine racial variations in skin color, such as black, brown, yellow, and white.
Melanin protects the basal layer against the damaging effect of
the ultraviolet (UV) rays of the sun. A gradual exposure to the
sunlight promotes the increased production of melanin within
the melanocytes, and hence tanning of the skin. The skin of a
person with albinism (al'b-niz-em) has the normal number of
FIGURE 5.4 Melanocytes throughout the stratum basale (see

arrow) produce melanin.
melanocytes in the epidermis but lacks the enzyme tyrosinase that

converts the amino acid tyrosine to melanin. Albinism is a hereditary condition.
Other genetic expressions of melanocytes are more common than albinism. Freckles, for example, are caused by aggregated patches of melanin. A lack of melanocytes in localized
CHAPTER 5
Stratum basale
Consists of a single layer of cuboidal cells in contact
with the basement membrane that undergo mitosis;
contains pigment-producing melanocytes

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areas of the skin causes distinct white spots in the condition

called vitiligo (vit--li'go). After the age of 50, brown plaquelike
growths, called seborrheic (seb''o-re'ik) hyperkeratoses, may appear
on the skin, particularly on exposed portions. Commonly called
liver spots, these pigmented patches are benign growths of
pigment-producing melanocytes. Usually no treatment is required, unless for cosmetic purposes.
CHAPTER 5
Excessive exposure to sunlight can cause skin cancer (see

Clinical Considerations and fig. 5.18). In sunlight, the skin absorbs two wavelengths of ultraviolet rays known as UVA and UVB.
The DNA within the basal skin cells may be damaged as the suns
more dangerous UVB rays penetrate the skin. Although it was once
believed that UVA rays were harmless, findings now indicate that excessive exposure to these rays may inhibit the DNA repair process
that follows exposure to UVB. Therefore, individuals who are exposed solely to UVA rays in tanning salons are still in danger of
basal cell carcinoma, because they will later be exposed to UVB
rays of sunlight when they go outdoors.
Carotene (kar'o-ten) is a yellowish pigment found in certain plant products, such as carrots, that tends to accumulate in
cells of the stratum corneum and fatty parts of the dermis. It was
once thought to account for the yellow-tan skin of people of
Asian descent, but this coloration is now known to be caused by
variations in melanin.
Hemoglobin (he''mo-glo'bin) is not a pigment of the skin;
rather, it is the oxygen-binding pigment found in red blood cells.
Oxygenated blood flowing through the dermis gives the skin its
pinkish tones.
Certain physical conditions or diseases cause symptomatic
discoloration of the skin. Cyanosis (si-a-no'sis) is a bluish discoloration of the skin that appears in people with certain cardiovascular or respiratory diseases. People also become cyanotic during an
interruption of breathing. In jaundice, the skin appears yellowish because of an excess of bile pigment in the bloodstream. Jaundice is
usually symptomatic of liver dysfunction and sometimes of liver immaturity, as in a jaundiced newborn. Erythema (er''-the'ma) is a redness of the skin generally due to vascular trauma, such as from a
sunburn.
Surface Patterns
The exposed surface of the skin has recognizable patterns that
are either present at birth or develop later. Fingerprints (friction
ridges) are congenital patterns that are present on the finger and
toe pads, as well as on the palms and soles. The designs formed
by these lines have basic similarities but are not identical in any
two individuals (fig. 5.5). They are formed by the pull of elastic
fibers within the dermis and are well established prenatally. The
ridges of fingerprints function to prevent slippage when grasping
objects. Because they are precise and easy to reproduce, fingerprints are customarily used for identifying individuals.
vitiligo: L. vitiatio, blemish

carotene: L. carota, carrot (referring to orange coloration)
hemoglobin: Gk. haima, blood; globus, globe
cyanosis: Gk. kyanosis, dark blue color
jaundice: L. galbus, yellow
erythema: Gk. erythros, red; haima, blood
(a)
(b)
(c)
(d)
FIGURE 5.5 The four basic fingerprint patterns (a) arch, (b) whorl,
(c) loop, and (d) combination.
Acquired lines include the deep flexion creases on the

palms and the shallow flexion lines that can be seen on the
knuckles and on the surface of other joints. Furrows on the forehead and face are acquired from continual contraction of facial
muscles, such as from smiling or squinting in bright light or
against the wind. Facial lines become more strongly delineated as
a person ages.
The science known as dermatoglyphics is concerned with the
classification and identification of fingerprints. Every individuals prints are unique, including those of identical twins. Fingerprints, however, are not exclusive to humans. All other primates have
fingerprints, and even dogs have a characteristic nose print that is
used for identification in the military canine corps and in certain dog
kennels.
Dermis
The dermis is deeper and thicker than the epidermis (see
fig. 5.1). Elastic and collagenous fibers within the dermis are
arranged in definite patterns, producing lines of tension in the skin
and providing skin tone (fig. 5.6). There are many more elastic
fibers in the dermis of a young person than in an elderly one, and
a decreasing number of elastic fibers is apparently associated with
aging. The extensive network of blood vessels in the dermis provides nourishment to the living portion of the epidermis.
The dermis also contains many sweat glands, oil-secreting
glands, nerve endings, and hair follicles.

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FIGURE 5.7 Stretch marks (lineae albicantes) on the abdomen of

a pregnant woman. Stretch marks generally fade with time but may
leave permanent markings.
Innervation of the Skin
collagenous fibers within the dermis of the skin. Surgical incisions

made parallel to the lines of tension heal more rapidly and create less
scar tissue than those made across the lines of tension.
Layers of the Dermis

The dermis is composed of two layers. The upper layer, called the
stratum papillarosum (papillary layer), is in contact with the epidermis and accounts for about one-fifth of the entire dermis (see
fig. 5.2). Numerous projections, called papillae (pa-pil'e), extend
from the upper portion of the dermis into the epidermis. Papillae
form the base for the friction ridges on the fingers and toes.
The deeper and thicker layer of the dermis is called the
stratum reticularosum (reticular layer). Fibers within this layer
are more dense and regularly arranged to form a tough, flexible
meshwork. It is quite distensible, as is evident in pregnant
women or obese individuals, but it can be stretched too far, causing tearing of the dermis. The repair of a strained dermal area
leaves a white streak called a stretch mark, or linea albicans
(lin'e-a al'b-kanz). Lineae albicantes are frequently found on the
buttocks, thighs, abdomen, and breasts (fig. 5.7).
Vascular Supply of the Skin

Blood vessels within the dermis supply nutrients to the mitotically active stratum basale of the epidermis and to the cellular
structures of the dermis, such as glands and hair follicles. Dermal
blood vessels play an important role in regulating body temperature and blood pressure. Autonomic vasoconstriction or vasodilation responses can either shunt the blood away from the
superficial dermal arterioles or permit it to flow freely throughout
dermal vessels. Fever or shock can be detected by the color and
temperature of the skin. Blushing is the result of involuntary vasodilation of dermal blood vessels.
It is the strong, resilient reticular layer of domestic mammals

that is used in making leather and suede. In the tanning
process, the hide of an animal is treated with various chemicals that
cause the epidermis with its hair and the papillary layer of the dermis
to separate from the underlying reticular layer. The reticular layer is
then softened and treated with protective chemicals before being cut
and assembled into consumer goods.
It is important to maintain good blood circulation in people

who are bedridden to prevent bedsores, or decubitus (dekyoo'b-tus) ulcers. When a person lies in one position for an extended period, the dermal blood flow is restricted where the body
presses against the bed. As a consequence, cells die and open
wounds may develop (fig. 5.8). Changing the position of the patient
frequently and periodically massaging the skin to stimulate blood
flow are good preventive measures against decubitus ulcers.
papilla: L. papula, swelling or pimple
decubitus: L. decumbere, lie down

ulcer: L. ulcus, sore
CHAPTER 5
FIGURE 5.6 Lines of tension are caused by the pull of elastic and
The dermis of the skin has extensive innervation (nerve supply).

Specialized integumentary effectors consist of smooth muscles or
glands within the dermis that respond to motor impulses transmitted from the central nervous system to the skin by autonomic
nerve fibers.
Several types of sensory receptors respond to various tactile
(touch), pressure, temperature, tickle, or pain stimuli. Some are
free nerve endings, some form a network around hair follicles, and
some extend into the papillae of the dermis. Certain areas of the
body, such as the palms, soles, lips, and external genitalia, have a
greater concentration of sensory receptors and are therefore more
sensitive to touch. Chapter 15 includes a detailed discussion of
the structure and function of the various sensory receptors.

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6. How do both the dermis and hypodermis function in

thermoregulation?
7. What two basic types of innervation are found within
the dermis?
FUNCTIONS OF THE SKIN

The skin not only protects the body from pathogens and external
injury, it is a highly dynamic organ that plays a key role in maintaining body homeostasis.
Objective 5
Discuss the role of the skin in the protection of

the body from disease and external injury, the regulation of
body fluids and temperature, absorption, synthesis, sensory
reception, and communication.
CHAPTER 5
Physical Protection
FIGURE 5.8 A bedsore (decubitus ulcer) on the medial surface of
the ankle. Bedsores are most common on skin overlying a bony projection, such as at the hip, ankle, heel, shoulder, or elbow.
Hypodermis
The hypodermis, or subcutaneous tissue, is not actually a part of
the skin, but it binds the dermis to underlying organs. The hypodermis is composed primarily of loose connective tissue and adipose cells interlaced with blood vessels (see fig. 5.1). Collagenous
and elastic fibers reinforce the hypodermisparticularly on the
palms and soles, where the skin is firmly attached to underlying
structures. The amount of adipose tissue in the hypodermis varies
with the region of the body and the sex, age, and nutritional
state of the individual. Females generally have about an 8%
thicker hypodermis than males. This layer functions to store
lipids, insulate and cushion the body, and regulate temperature.
The hypodermis is the site for subcutaneous injections. Using
a hypodermic needle, medicine can be administered to patients who are unconscious or uncooperative, and when oral medications are not practical. Subcutaneous devices to administer
slow-release, low-dosage medications are now available. For example, insulin may be administered in this way to treat some forms of diabetes. Even a subcutaneous birth-control device (Norplant) is
currently being marketed (see fig. 21.26).
Knowledge Check
4. List the layers of the epidermis and dermis and explain how
they differ in structure and function.
5. Describe the sequence of cellular replacement within the epidermis and the processes of keratinization and cornification.
hypodermis: Gk. hypo, under; derma, skin
The skin is a barrier to microorganisms, water, and excessive sunlight (UV light). Oily secretions onto the surface of the skin
form an acidic protective film (pH 4.06.8) that waterproofs the
body and retards the growth of most pathogens. The protein keratin in the epidermis also waterproofs the skin, and the cornified
outer layer (stratum corneum) resists scraping and keeps out microorganisms. As mentioned previously, exposure to UV light
stimulates the melanocytes in the stratum basale to synthesize
melanin, which absorbs and disperses sunlight. In addition, surface friction causes the epidermis to thicken by increasing the
rate of mitosis in the cells of the stratum basale and stratum spinosum, resulting in the formation of a protective callus.
Regardless of skin pigmentation, everyone is susceptible to skin
cancer if his or her exposure to sunlight is sufficiently intense.
There are an estimated 800,000 new cases of skin cancer yearly in the
United States, and approximately 9,300 of these are diagnosed as the
potentially life-threatening melanoma (mel-a-no'ma) (cancer of
melanocytes). Melanomas (see fig. 5.19) are usually termed malignant,
because they may spread rapidly. Sunscreens are advised for people
who must be in direct sunlight for long periods of time.
Hydroregulation
The thickened, keratinized, and cornified epidermis of the skin is
adapted for continuous exposure to the air. In addition, the outer
layers are dead and scalelike, and a protein-polysaccharide basement membrane adheres the stratum basale to the dermis.
Human skin is virtually waterproof, protecting the body from
desiccation (dehydration) on dry land, and even from water absorption when immersed in water.
Thermoregulation
The skin plays a crucial role in the regulation of body temperature. Body heat comes from cellular metabolism, particularly in
muscle cells as they maintain tone or a degree of tension. A nor-

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2. The temperature center in the

hypothalamus receives the message and
triggers various physiologic responses
to produce and conserve heat.
1. Responding to a lowering of the

temperature, cutaneous sensory
receptors send a message
to the brain.
(a) Shivering (involuntary muscle

activity) produces heat.
(b) Capillary constriction shunts
blood away from the exposed
surface of the skin.
(c) Perspiration stops as sweat
glands shut down.
FIGURE 5.9 Temperature regulation involves cutaneous sensory receptors that relay messages of decreased body temperature to the brain.
This triggers a response that can quickly generate up to 5 times the normal rate of body heat production.
mal body temperature of 37 C (98.6 F) is maintained in three
ways, all involving the skin (fig. 5.9):
1. through radiant heat loss from dilated blood vessels,
2. through evaporation of perspiration, and
3. through retention of heat from constricted blood vessels
(fig. 5.10).
The volume of perspiration produced is largely a function
of how much the body is overheated. This volume increases approximately 100 to 150 ml/day for each 1 C elevation in body
temperature. For each hour of hard physical work out-of-doors in
the summertime, a person may produce 1 to 10 L of perspiration.
A serious danger of continued exposure to heat and excessive
water and salt loss is heat exhaustion, characterized by nausea, weakness, dizziness, headache, and a decreased blood pressure. Heat stroke is similar to heat exhaustion, except that in heat
stroke sweating is prevented (for reasons that are not clear) and
body temperature rises. Convulsions, brain damage, and death may
follow.
Excessive heat loss triggers a shivering response in muscles,

which increases cellular metabolism. Not only do skeletal muscles contract, but tiny smooth muscles called arrectores pilorum
(a''rek-to'rez pil-o'rumsingular, arrector pili), which are attached
to hair follicles, are also contracted involuntarily and cause
goose bumps.
When the bodys heat-producing mechanisms cannot keep
pace with heat loss, hypothermia results. A lengthy exposure
to temperatures below 20 C (68 F) and dampness may lead to this
condition. This is why it is so important that a hiker, for example,
FIGURE 5.10 A thermogram of the hand showing differential heat

radiation. Hair and body fat are good insulators. Red and yellow indicate the warmest parts of the body. Blue, green, and white indicate
the coolest.
dress appropriately for the weather conditions, especially on cool,

rainy spring or fall days. The initial symptoms of hypothermia are
numbness, paleness, delirium, and uncontrolled shivering. If the core
temperature falls below 32 C (90 F), the heart loses its ability to
pump blood and will go into fibrillation (erratic contractions). If the
victim is not warmed, extreme drowsiness, coma, and death follow.
CHAPTER 5
3. Responding to a rise in the temperature,

cutaneous sensory receptors send a
feedback message to the brain, which
reverses the physiologic processes that
produced and conserved heat.

CHAPTER 5
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(a)
(b)
FIGURE 5.11 (a) Rickets in a child from a Nepalese village, whose inhabitants live in windowless huts. During the rainy season, which may
last 5 to 6 months, the children are kept indoors. (b) A radiograph of a 10-month-old child with rickets. Rickets develops from an improper diet
and also from lack of the sunlight needed to synthesize vitamin D.
Cutaneous Absorption
Because of the effective protective barriers of the integument already described, cutaneous absorption (absorption through the
skin) is limited. Some gases, such as oxygen and carbon dioxide,
may pass through the skin and enter the blood. Small amounts of
UV light, necessary for synthesis of vitamin D, are absorbed readily. Of clinical consideration is the fact that certain chemicals
such as lipid-soluble toxins and pesticides can easily enter the
body through the skin.
Synthesis
The integumentary system synthesizes melanin and keratin, which
remain in the skin synthesis of vitamin D, which is used elsewhere
in the body and begins in the skin with activation of a precursor
molecule by UV light. The molecule is modified in the liver and
kidneys to produce calcitriol (kal-s-tre'ol), the most active form of
vitamin D. Only small amounts of UV light are necessary for vitamin D synthesis, but these amounts are very important to a growing child. Active vitamin D enters the blood and helps regulate
the metabolism of calcium and phosphorus, which are important
in the development of strong and healthy bones. Rickets is a disease caused by vitamin D deficiency (fig. 5.11).
Sensory Reception
Highly specialized sensory receptors (see chapter 15) that respond to the precise stimuli of heat, cold, pressure, touch, vibra-
tion, and pain are located throughout the dermis. Called cutaneous receptors, these sensory nerve cells are especially abundant in the skin of the face and palms, the fingers, the soles of
the feet, and the genitalia. They are less abundant along the
back and on the back of the neck and are sparse in the skin over
joints, especially the elbow. Generally speaking, the thinner the
skin, the greater the sensitivity.
Communication
Humans are highly social animals, and the integument plays
an important role in communication. Various emotions, such
as anger or embarrassment, may be reflected in changes of
skin color. The contraction of specific facial muscles produces facial expressions that convey an array of emotions, including love, surprise, happiness, sadness, and despair.
Secretions from certain integumentary glands have odors that
frequently elicit subconscious responses from others who detect them.
Knowledge Check
8. List five modifications of the integument that are structurally or functionally protective.
9. Explain how the integument functions to regulate body fluids and temperature.
10. What substances are synthesized in the integument?

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Chapter 5
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EPIDERMAL DERIVATIVES
Hair, nails, and integumentary glands form from the epidermal
layer, and are therefore of ectodermal derivation. Hair and nails
are structural features of the integument and have a limited functional role. By contrast, integumentary glands are extremely important in body defense and maintenance of homeostasis.
Objective 6
Describe the structure of hair and list the three

principal types.
Objective 7
Discuss the structure and function of nails.
Objective 8
Compare and contrast the structure and

function of the three principal kinds of integumentary glands.
Hair
(a)
Hirsutism (her'soo-tiz''em) is a condition of excessive body and

facial hair, especially in women. It may be a genetic expression, as in certain ethnic groups, or occur as the result of a metabolic
disorder, usually endocrine. Hirsutism occurs in some women as they
experience hormonal changes during menopause. Various treatments for hirsutism include hormonal injections and electrolysis to
permanently destroy selected hair follicles.
The primary function of hair is protection, even though

its effectiveness is limited. Hair on the scalp and eyebrows protect against sunlight. The eyelashes and the hair in the nostrils
protect against airborne particles. Hair on the scalp may also
protect against mechanical injury. Some secondary functions
of hair are to distinguish individuals and to serve as a sexual
attractant.
Each hair consists of a diagonally positioned shaft, root,
and bulb (fig. 5.13). The shaft is the visible, but dead, portion of
the hair projecting above the surface of the skin. The bulb is the
enlarged base of the root within the hair follicle. Each hair develops from stratum basale cells within the bulb of the hair,
where nutrients are received from dermal blood vessels. As the
cells divide, they are pushed away from the nutrient supply toward the surface, and cellular death and keratinization occur. In
a healthy person, hair grows at the rate of approximately 1 mm
every 3 days. As the hair becomes longer, however, it enters a
resting period, during which there is minimal growth.
hirsutism: L. hirsutus, shaggy
CHAPTER 5
The presence of hair on the body is one of the distinguishing features of mammals, but its distribution, function, density, and texture varies across mammalian species. Humans are relatively
hairless, with only the scalp, face, pubis, and axillae being
densely haired. Men and women have about the same density of
hair on their bodies, but hair is generally more obvious on men
(fig. 5.12) as a result of male hormones. Certain structures and
regions of the body are hairless, such as the palms, soles, lips, nipples, penis, and parts of the female genitalia.
(b)
FIGURE 5.12 A comparison of the expression of body hair in

males and females.
The life span of a hair varies from 3 to 4 months for an eyelash to 3 to 4 years for a scalp hair. Each hair lost is replaced by a
new hair that grows from the base of the follicle and pushes the
old hair out. Between 10 and 100 hairs are lost daily. Baldness results when hair is lost and not replaced. This condition may be
disease-related, but it is generally inherited and most frequently
occurs in males because of genetic influences combined with the
action of the male sex hormone testosterone (tes-tos'te-ron). No
treatment is effective in reversing genetic baldness; however,
flaps or plugs of skin containing healthy follicles from hairy parts
of the body can be grafted onto hairless regions.
Three layers can be observed in hair that is cut in cross section. The inner medulla (me-dul'a) is composed of loosely
arranged cells separated by numerous air cells. The thick cortex
medulla: L. medulla, marrow
cortex: L. cortex, bark

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CHAPTER 5
(a)
(b)
FIGURE 5.13 The structure of hair and the hair follicle. (a) A photomicrograph (63) of the bulb and root of a hair within a hair follicle. (b) A
scanning electron micrograph (280) of a hair as it extends from a follicle. (c) A diagram of hair, a hair follicle, and sebaceous gland, and an arrector pili muscle.
surrounding the medulla consists of hardened, tightly packed
cells. A cuticle covers the cortex and forms the toughened outer
layer of the hair. Cells of the cuticle have serrated edges that give
a hair a scaly appearance when observed under a dissecting scope.
People exposed to heavy metals, such as lead, mercury, arsenic, or cadmium, will have concentrations of these metals
in their hair that are 10 times as great as those found in their blood
or urine. Because of this, hair samples can be extremely important
in certain diagnostic tests.
Even evidence of certain metabolic diseases or nutritional deficiencies may be detected in hair samples. For example, the hair of
children with cystic fibrosis will be deficient in calcium and display
excessive sodium. There is a deficiency of zinc in the hair of malnourished individuals.
Hair color is determined by the type and amount of pigment

produced in the stratum basale at the base of the hair follicle. Varying amounts of melanin produce hair ranging in color from blond
to brunette to black. The more abundant the melanin, the darker
the hair. A pigment with an iron base (trichosiderin) produces red
hair. Gray or white hair is the result of a lack of pigment production
and air spaces within the layers of the shaft of the hair. The texture
of hair is determined by the cross-sectional shape: straight hair is
round in cross section, wavy hair is oval, and kinky hair is flat.
Sebaceous glands and arrectores pilorum muscles (described previously) are attached to the hair follicle (fig. 5.13c).
The arrectores pilorum muscles are involuntary, responding to
thermal or psychological stimuli. When they contract, the hair is
pulled into a more vertical position, causing goose bumps.
Humans have three distinct kinds of hair:
1. Lanugo. Lanugo (la-noo'go) is a fine, silky fetal hair that
appears during the last trimester of development. It is usually seen only on premature infants.
2. Vellus. Vellus is a short, fine hair that replaces lanugo. It
is especially abundant in children and women just barely
extending from the hair follicules.
3. Terminal hair. Terminal hair is coarse, pigmented (except
in most elderly people), and sometimes curly. Examples are
lanugo: L. lana, wool
cuticle: L. cuticula, small skin
vellus: L. vellus, fleece

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Chapter 5
Hyponychium
Free border
Eponychium
Body of nail
117
Nail matrix
Hidden border
Nail groove
Nail fold
Lunula
Eponychium
Body of nail
Nail bed
Free border
Creek
Hyponychium
Hidden border
Nail root
Epidermis
Dermis
Pulp
Distal phalanx
Developing bone
(b)
(a)
scalp hair, axillary hair, pubic hair, eyebrows, eyelashes, and

hair on the extremities. Angora hair is terminal hair that
grows continuously. It is found on the scalp and on the faces
of mature males. Definitive hair is terminal hair that grows
to a certain length and then stops. It is the most common
type of hair. Eyelashes, eyebrows, pubic, and axillary hair are
examples.
Anthropologists have referred to humans as the naked apes
because of our relative hairlessness. The clothing that we wear
over the exposed surface areas of our bodies functions to insulate
and protect us, just as hair or fur does in other mammals. However,
the nakedness of our skin does lead to some problems. Skin cancer
occurs frequently in humans, particularly in regions of the skin exposed to the sun. Acne, another problem unique to humans, is partly
related to the fact that hair is not present to dissipate the oily secretion from the sebaceous glands.
Nails
The nails on the ends of the fingers and toes are formed from the
compressed outer layer (stratum corneum) of the epidermis. The
hardness of the nail is due to the dense keratin fibrils running parallel between the cells. Both fingernails and toenails protect the
digits, and fingernails also aid in grasping and picking up small
objects.
Each nail consists of a body, free border, and hidden border (fig. 5.14). The platelike body of the nail rests on a nail bed,
which is actually the stratum spinosum of the epidermis. The
body and nail bed appear pinkish because of the underlying vascular tissue. The sides of the nail body are protected by a nail
hyponychium: Gk. hypo, under; onyx, nail
fold, and the furrow between the sides and body is the nail
groove. The free border of the nail extends over a thickened region of the stratum corneum called the hyponychium (hi''ponik'e-um) (quick). The root of the nail is attached at the base.
An eponychium (cuticle) covers the hidden border of the
nail. The eponychium frequently splits, causing a hangnail. The
growth area of the nail is the nail matrix. A small part of the nail
matrix, the lunula (loo'nyoo-la), can be seen as a half-moonshaped area near the eponychium of the nail.
The nail grows by the transformation of the superficial
cells of the nail matrix into nail cells. These harder, transparent
cells are then pushed forward over the strata basale and spinosum
of the nail bed. Fingernails grow at the rate of approximately
1 mm each week. The growth rate of toenails is somewhat slower.
The condition of nails may be indicative of a persons general
health and well-being. Nails should appear pinkish, showing
the rich vascular capillaries beneath the translucent nail. A yellowish
hue may indicate certain glandular dysfunctions or nutritional deficiencies. Split nails may also be caused by nutritional deficiencies. A
prominent bluish tint may indicate improper oxygenation of the blood.
Spoon nails (concave body) may be the result of iron-deficiency anemia, and clubbing at the base of the nail may be caused by lung
cancer. Dirty or ragged nails may indicate poor personal hygiene,
and chewed nails may suggest emotional problems.
Glands
Although they originate in the epidermal layer, all of the glands
of the skin are located in the dermis, where they are physically
supported and receive nutrients. Glands of the skin are referred
lunula: L. lunula, small moon
CHAPTER 5
FIGURE 5.14 The fingertip and the associated structures of the nail. (a) A diagram of a dissected nail, and (b) a photomicrograph of a nail
from a fetus (3.5).

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Sebaceous
gland
CHAPTER 5
Sweat
gland
FIGURE 5.16 A photomicrograph of an eccrine sweat gland

(27). The coiled structure of the ductule portion of the gland (see arrows) accounts for its discontinuous appearance.
shaft of the hair to the surface of the skin, where it lubricates and
waterproofs the stratum corneum and also prevents the hair from
becoming brittle. If the ducts of sebaceous glands become
blocked for some reason, the glands may become infected, resulting in acne. Sex hormones regulate the production and secretion
of sebum, and hyperactivity of sebaceous glands can result in serious acne problems, particularly during teenage years.
Sudoriferous Glands
Commonly called oil glands, sebaceous glands are associated

with hair follicles, because they develop from the follicular epithelium of the hair. They are holocrine glands (see chapter 4)
that secrete sebum (se'bum) onto the shaft of the hair (fig. 5.13).
Sebum, which consists mainly of lipids, is dispersed along the
Commonly called sweat glands, sudoriferous glands excrete perspiration, or sweat, onto the surface of the skin. Perspiration is
composed of water, salts, urea, and uric acid. It serves not only
for evaporative cooling, but also for the excretion of certain
wastes. Sweat glands are most numerous on the palms, soles, axillary and pubic regions, and on the forehead. They are coiled and
tubular (fig. 5.15) and are of two types: eccrine (ek'rin) and apocrine (ap'o-krin) sweat glands.
1. Eccrine sweat glands are widely distributed over the
body, especially on the forehead, back, palms, and soles.
These glands are formed before birth and function in evaporative cooling (figs. 5.15 and 5.16).
2. Apocrine sweat glands are much larger than the eccrine
glands. They are found in the axillary and pubic regions,
where they secrete into hair follicles. Apocrine glands are
not functional until puberty, and their odoriferous secretion is thought to act as a sexual attractant.
sebum: L. sebum, tallow or grease
sudoriferous: L. sudorifer, sweat; ferre, to bear
Apocrine
sweat gland
Eccrine
sweat gland
FIGURE 5.15 Types of skin glands.

to as exocrine, because they are externally secreting glands that
either release their secretions directly or through ducts. The
glands of the skin are of three basic types: sebaceous (se-ba'shus),
sudoriferous (soo''dor-if'er-us), and ceruminous (se-roo'm-nus).
Sebaceous Glands

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13. List the three types of integumentary glands and describe

the structure and function of each.
14. Are skin glands mesodermal or ectodermal in derivation?
Are they epidermal or dermal in functional position?
Rib
Intercostal muscle
Pectoralis major
muscle
Deep connective
tissue
Adipose tissue
Secondary tubule
Lactiferous duct
Lactiferous sinus
The skin is a buffer against the external environment and is

therefore subject to a variety of disease-causing microorganisms
and physical assaults. A few of the many diseases and disorders of
the integumentary system are briefly discussed here.
Ampulla
Lobule
Lobe
Mammary glands, found within the breasts, are specialized

sudoriferous glands that secrete milk during lactation (fig. 5.17)
The breasts of the female reach their greatest development during the childbearing years, under the stimulus of pituitary and
ovarian hormones.
Good routine hygiene is very important for health and social
reasons. Washing away the dried residue of perspiration and
sebum eliminates dirt. Excessive bathing, however, can wash off the
natural sebum and dry the skin, causing it to itch or crack. The commercial lotions used for dry skin are, for the most part, refined and
perfumed lanolin, which is sebum from sheep.
Inflammatory skin disorders are caused by immunologic hypersensitivity, infectious agents, poor circulation, or exposure to environmental assaults such as wind, sunlight, or chemicals. Some
people are allergic to certain foreign proteins and, because of this
inherited predisposition, experience such hypersensitive reactions as asthma, hay fever, hives, drug and food allergies, and
eczema. Lesions, as applied to inflammatory conditions, are defined as more or less circumscribed pathologic changes in the tissue. Some of the more common inflammatory skin disorders and
their usual sites are illustrated in fig. 5.18
There are also a number of infectious diseases of the skin,
which is not surprising considering the highly social and communal animals we are. Most of these diseases can now be prevented,
but too frequently people fail to take appropriate precautionary
measures. Infectious diseases of the skin include childhood viral
infections (measles and chicken pox); bacteria, such as staphylococcus (impetigo); sexually transmitted diseases; leprosy; fungi
(ringworm, athletes foot, candida); and mites (scabies).
Ceruminous Glands
Neoplasms
These specialized glands are found only in the external auditory

canal (ear canal) where they secrete cerumen (se-roo'men), or
earwax. Cerumen is a water and insect repellent, and also keeps
the tympanic membrane (eardrum) pliable. Excessive amounts of
cerumen may interfere with hearing.
Both benign and malignant neoplastic conditions or diseases are

common in the skin. Pigmented moles (nevi), for example, are a
type of benign neoplastic growth of melanocytes. Dermal cysts
and benign viral infections are also common. Warts are virally
caused abnormal growths of tissue that occur frequently on the
hands and feet. These warts are usually treated effectively with
liquid nitrogen or acid. A different type of wart, called a venereal
wart, occurs in the anogenital region of affected sexual partners.
Risk factors for cervical cancer may be linked to venereal warts,
so they are treated aggressively with chemicals, cryosurgery,
cautery, or laser therapy.
Knowledge Check
11. Draw and label a hair. Indicate which portion is alive and
discuss what causes the cells in a hair to die.
12. Describe the structure and function of nails.
neoplasm: Gk. neo, new; plasma, something formed

cerumen: L. cera, wax
benign: L. benignus, good-natured

malignant: L. malignus, acting from malice
CHAPTER 5
FIGURE 5.17 A sagittal section of a mammary gland within the

human breast.
Inflammatory Conditions (Dermatitis)

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The Integumentary System
Both the ectodermal and mesodermal germ layers (see chapter 4)
function in the formation of the structures of the integumentary
system. The epidermis and the hair, glands, and nails of the skin
develop from the ectodermal germ layer (exhibits I, II, and III).
The dermis develops from a thickened layer of undifferentiated
mesoderm called mesenchyme (mez'en-k m).
By 6 weeks, the ectodermal layer has differentiated into an
outer flattened periderm and an inner cuboidal germinal (basal)
layer in contact with the mesenchyme. The periderm eventually
EXHIBIT I The development of the skin.
EXHIBIT II The development of hair and glands.
EXPLANATION
Skin cancer is the most common malignancy in the

United States. As shown in figure 5.19, there are three frequently encountered types. Basal cell carcinoma, the most
common skin cancer, accounts for about 70% of total cases. It
usually occurs where exposure to sunlight is the greateston
the face and arms. This type of cancer arises from cells in the
120
stratum basale. It appears first on the surface of the skin as a

small, shiny bump. As the bump enlarges, it often develops a
central crater that erodes, crusts, and bleeds. Fortunately, there
is little danger that it will spread (metastasize) to other body
areas. These carcinomas are usually treated by excision (surgical removal).

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sloughs off, forming the vernix caseosa (ka''se-o'sa), a cheeselike

protective coat that covers the skin of the fetus.
By 11 weeks, the mesenchymal cells below the germinal
cells have differentiated into the distinct collagenous and elastic
connective tissue fibers of the dermis. The tensile properties of
these fibers cause a buckling of the epidermis and the formation of
dermal papillae. During the early fetal period (about 10 weeks),
specialized neural crest cells called melanoblasts migrate into
the developing dermis and differentiate into melanocytes.
The melanocytes soon migrate to the germinal layer of the epidermis, where they produce the pigment melanin that colors the
epidermis.
Before hair can form, a hair follicle must be present. Each
hair follicle begins to develop at about 12 weeks (exhibit II), as a
mass of germinal cells called a hair bud proliferates into the underlying mesenchyme. As the hair bud becomes club-shaped, it is
referred to as a hair bulb. The hair follicle, which physically supports and provides nourishment to the hair, is derived from specialized mesenchyme called the hair papilla, which is localized
around the hair bulb, and from the epithelial cells of the hair
bulb called the hair matrix. Continuous mitotic activity in the epithelial cells of the hair bulb results in the growth of the hair.
Sebaceous glands and sweat glands are the two principal
types of integumentary glands. Both develop from the germinal
layer of the epidermis (exhibit II). Sebaceous glands develop as
proliferations from the sides of the developing hair follicle. Sweat
glands become coiled as the secretory portion of the developing
gland proliferates into the dermal mesenchyme. Mammary glands
(exhibit III) are modified sweat glands that develop in the skin of
the anterior thoracic region.
EXHIBIT III The development of mammary glands at

(a) 12 weeks, (b) 16 weeks, and (c) about 28 weeks.
Squamous cell carcinoma arises from cells immediately superficial to the stratum basale. Normally, these cells undergo very
little division, but in squamous cell carcinoma they continue to
divide as they produce keratin. The result is usually a firm, red
keratinized tumor, confined to the epidermis. If untreated, however, it may invade the dermis and metastasize. Treatment usually consists of excision and radiation therapy.
Malignant melanoma, the most life-threatening form of

skin cancer, arises from the melanocytes located in the stratum
basale. Often, it begins as a small molelike growth, which enlarges, changes color, becomes ulcerated, and bleeds easily.
Metastasis occurs quickly, and unless treated earlyusually by
widespread excision and radiation therapythis cancer is often
fatal.
121

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CHAPTER 5
122
FIGURE 5.18 Common inflammatory skin disorders and their usual sites of occurrence.
Burns
A burn is an epithelial injury caused by contact with a thermal,
radioactive, chemical, or electrical agent. Burns generally occur
on the skin, but they can involve the linings of the respiratory
and GI tracts. The extent and location of a burn is frequently
less important than the degree to which it disrupts body homeostasis. Burns that have a local effect (local tissue destruction)
are not as serious as those that have a systemic effect. Systemic
effects directly or indirectly involve the entire body and are a
threat to life. Possible systemic effects include body dehydration,
shock, reduced circulation and urine production, and bacterial
infections.
Burns are classified as first degree, second degree, or third

degree, based on their severity (fig. 5.20). In first-degree burns,
the epidermal layers of the skin are damaged and symptoms are
restricted to local effects such as redness, pain, and edema
(swelling). A shedding of the surface layers (desquamation) generally follows in a few days. A sunburn is an example. Seconddegree burns involve both the epidermis and dermis. Blisters appear and recovery is usually complete, although slow. Third-degree burns destroy the entire thickness of the skin and
frequently some of the underlying muscle. The skin appears waxy
or charred and is insensitive to touch. As a result, ulcerating
wounds develop, and the body attempts to heal itself by forming
scar tissue. Skin grafts are frequently used to assist recovery.

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(a) Basal cell carcinoma
(a)
CHAPTER 5
(b) Squamous cell carcinoma
(b)
(c) Malignant melanoma
FIGURE 5.19 Three types of skin cancer.
As a way of estimating the extent of damaged skin suffered

in burned patients, the rule of nines (fig. 5.21) is often applied.
The surface area of the body is divided into regions, each of
which accounts for about 9% (or a multiple of 9%) of the total
skin surface. An estimation of the percentage of surface area
damaged is important in treating with intravenous fluid, which
replaces the fluids lost from tissue damage.
Frostbite
Frostbite is a local destruction of the skin resulting from freezing.
Like burns, frostbite is classified by its degree of severity: first degree, second degree, and third degree. In first-degree frostbite,
the skin will appear cyanotic (bluish) and swollen. Vesicle formation and hyperemia (engorgement with blood) are symptoms
(c)
FIGURE 5.20 The classification of burns, (a) First-degree burns

involve the epidermis and are characterized by redness, pain, and
edemasuch as with a sunburn; (b) second-degree burns involve
the epidermis and dermis and are characterized by intense pain,
redness, and blistering; and (c) third-degree burns destroy the entire
skin and frequently expose the underlying organs. The skin is
charred and numb and does not protect against fluid loss.
of second-degree frostbite. As the affected area is warmed, there

will be further swelling, and the skin will redden and blister. In
third-degree frostbite, there will be severe edema, some bleeding, and numbness followed by intense throbbing pain and
necrosis of the affected tissue. Gangrene will follow untreated
third-degree frostbite.

6. Skeletal System:
Introduction and the Axial
Skeleton
Chapter 6
Periosteum
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Skeletal System: Introduction and the Axial Skeleton
External circumferential lamellae

Arteriole
Central canal
139
Venule
Nerve
Central
canal
Canaliculi
Lamellae
Medullary
cavity
(c)
(a)
Osteocyte
Lacuna
CHAPTER 6
Perforating
fibers
Canaliculi
Blood vessels
Perforating
canals
Central
canal
Trabeculae of
spongy bone
(d)
(b)
FIGURE 6.7 Compact bone tissue. (a) A diagram of the femur showing a cut through the compact bone into the medullary cavity. (b) The
arrangement of the osteons within the diaphysis of the bone. (c) An enlarged view of an osteon showing the osteocytes within lacunae and the
concentric lamellae. (d) An osteocyte within a lacuna.
LA
CA
Canaliculi
Osteocyte
within a
lacuna
Central
canal
Lamella
CA
LA
(a)
(b)
FIGURE 6.8 Bone tissue as seen in (a) a scanning electron micrograph and (b) a photomicrograph. The lacunae (LA) provides spaces for the
osteocytes, which are connected to one another by canaliculi (CA). Note the divisions between the lamellae (see arrows).

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6. Skeletal System:
Skeleton
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(a)
(b)
CHAPTER 6
(c)
(d)
(e)
(f)
FIGURE 6.9 The growth process of a long bone, beginning with (a) the cartilaginous model as it occurs in an embryo at 6 weeks. The bone
develops (be) through intermediate stages to (f ) adult bone.
nutrient vessels and a nerve. Osteocytes within spaces called

lacunae (la-kyoo'ne) are regularly arranged between the lamellae. The lacunae are connected by tiny channels called canaliculi (kan'' a-lik'yu-li), through which nutrients diffuse.
Metabolic activity within bone tissue occurs at the osteon
level. Between osteons there are incomplete remnants of osteons, called interstitial systems. Perforating (Volkmanns)
canals penetrate compact bone, connecting osteons with blood
vessels and nerves.
Knowledge Check
9. Construct a sample table listing the location and function
of each type of cell found within bone tissue
10. Define osteon and sketch the arrangement of osteons
within compact bone tissue.
Volkmanns canal: from Alfred Volkmann, German physiologist, 180077
BONE GROWTH
The development of bone from embryo to adult depends on the orderly processes of cell division, growth, and ongoing remodeling.
Bone growth is influenced by genetics, hormones, and nutrition.
Objective 8
Describe the process of endochondral

ossification as related to bone growth.
In most bone development, a cartilaginous model is gradually replaced by bone tissue during endochondral bone formation (see Developmental Exposition, pp. 141142). As the cartilage model
grows, the chondrocytes (cartilage cells) in the center of the shaft
hypertrophy, and minerals are deposited within the matrix in a
process called calcification (fig. 6.9). Calcification restricts the passage of nutrients to the chondrocytes, causing them to die. At the
same time, some cells of the perichondrium (dense regular connective tissue surrounding cartilage) differentiate into osteoblasts.
These cells secrete osteoid (os'te-oid), the hardened organic

6. Skeletal System:
Skeleton
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The Axial Skeleton
certain bones of the cranium are formed this way. Sesamoid bones
are specialized intramembranous bones that develop in tendons.
The patella is an example of a sesamoid bone.
EXPLANATION
DEVELOPMENT OF THE SKULL
Development of Bone
Bone formation, or ossification, begins at about the fourth week of
embryonic development, but ossification centers cannot be readily observed until about the tenth week (exhibit I). Bone tissue
derives from specialized migratory cells of mesoderm (see
fig. 4.13) known as mesenchyme. Some of the embryonic mesenchymal cells will transform into chondroblasts (kon'dro-blasts)
and develop a cartilage matrix that is later replaced by bone in a
process known as endochondral (en''do-kon'dral) ossification.
Most of the skeleton is formed in this fashionfirst it goes
through a hyaline cartilage stage and then it is ossified as bone.
A smaller number of mesenchymal cells develop into bone
directly, without first going through a cartilage stage. This type
of bone-formation process is referred to as intramembranous
(in''tra-mem'bra-nus) ossification. The clavicles, facial bones, and
chondroblast: Gk. chondros, cartilage; blastos, offspring or germ
The formation of the skull is a complex process that begins during the fourth week of embryonic development and continues
well beyond the birth of the baby. Three aspects of the embryonic skull are involved in this process: the chondrocranium, the
neurocranium, and the viscerocranium (exhibit II). The chondrocranium is the portion of the skull that undergoes endochondral ossification to form the bones supporting the brain. The
neurocranium is the portion of the skull that develops through
membranous ossification to form the bones covering the brain
and facial region. The viscerocranium (splanchnocranium) is the
portion that develops from the embryonic visceral arches to form
the mandible, auditory ossicles, the hyoid bone, and specific
processes of the skull.
chondrocranium: Gk. chondros, cartilage; kranion, skull

viscerocranium: L. viscera, soft parts; Gk. kranion, skull
Parietal
bones
Occipital
bone
Frontal
bones
Temporal
bone
Humerus
Zygomatic
bone
Maxilla
Nasal bone
Mandible
Metacarpal
bones
Phalanges
Carpal bones
Ribs
Radius
Ulna
Chondrocranium
Vertebrae
Clavicle
Scapula
Femur
Tibia
Fibula
Ilium
Sacrum
Coccyx
(a)
Phalanges
Metatarsal bones
Tarsal bones
Creek
(b)
EXHIBIT I Ossification centers of the skeleton of a 10-week-old fetus. (a) The diagram depicts endochondrial ossification in red and intramembranous ossification in a stippled pattern. The cartilaginous portions of the skeleton are shown in gray. (b) The photograph shows
the ossification centers stained with a red indicator dye.
141

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(continued)
EXHIBIT II The embryonic skull at 12 weeks is composed of bony elements from three developmental sources: the chondrocranium
(colored blue-gray), the neurocranium (colored light yellow), and the viscerocranium (colored salmon).
component of bone. As the perichondrium calcifies, it gives rise to

a thin plate of compact bone called the periosteal bone collar.
The periosteal bone collar is surrounded by the periosteum.
A periosteal bud, consisting of osteoblasts and blood vessels, invades the disintegrating center of the cartilage model from
the periosteum. Once in the center, the osteoblasts secrete osteoid, and a primary ossification center is established. Ossification then expands into the deteriorating cartilage. This process is
repeated in both the proximal and distal epiphyses, forming secondary ossification centers where spongy bone develops.
Once the secondary ossification centers have been formed,
bone tissue totally replaces cartilage tissue, except at the articular
ends of the bone and at the epiphyseal plates. An epiphyseal
plate contains five histological zones (fig. 6.10). The reserve zone
(zone of resting cartilage) borders the epiphysis and consists of
small chondrocytes irregularly dispersed throughout the intercellular matrix. The chondrocytes in this zone anchor the epiphyseal plate to the bony epiphysis. The proliferation zone (zone of
proliferating cartilage) consists of larger, regularly arranged chondrocytes that are constantly dividing. The hypertrophic zone (zone
142
Epiphyseal border
Reserve
zone
Proliferation
zone
Chondrocytes
Hypertrophic
zone
Epiphyseal border
Resorption
zone
Bone tissue
Ossification
zone
Red bone
marrow
Diaphyseal border
FIGURE 6.10 A photomicrograph from an epiphyseal plate (63).

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Frontal
bone
Ethmoid bone
Zygomatic bone
Middle nasal concha
CHAPTER 6
Maxilla
Inferior nasal concha

Vomer
FIGURE 6.20 A posterior view of a frontal (coronal) section of the skull.
Frontal bone
Nasal bone
Lacrimal bone
Zygomatic
bone
Maxilla
FIGURE 6.21 Bones of the orbit.

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Ethmoidal
sinus
151
Frontal sinus
Sphenoidal sinus
Frontal
sinus
Ethmoidal sinuses
Sphenoidal
sinus
Maxillary sinus
Maxillary
sinus
(a)
(b)
TABLE 6.5 Bones Forming the Orbit

Region of the Orbit
Contributing Bones
Roof (superior)
Frontal bone; lesser wing of sphenoid bone
Floor (inferior)
Maxilla; zygomatic bone; palatine bone
Lateral wall
Zygomatic bone
Posterior wall
Greater wing of sphenoid bone
Medial wall
Maxilla; lacrimal bone; ethmoid bone
Superior margin
Frontal bone
Lateral margin
Zygomatic bone
Medial margin
Maxilla
2. Tympanic part. The tympanic part of the temporal bone

contains the external acoustic meatus (me-a'tus), or ear
canal, which is posterior to the mandibular fossa. A thin,
pointed styloid process (figs. 6.16, 6.17, and 6.18) projects
inferiorly from the tympanic part.
3. Mastoid part. The mastoid process, a rounded projection
posterior to the external acoustic meatus, accounts for the
mass of the mastoid part. The mastoid foramen (fig. 6.16)
is directly posterior to the mastoid process. The stylomastoid foramen, located between the mastoid and styloid
processes (fig. 6.16), provides the passage for part of the facial nerve.
styloid: Gk. stylos, pillar

mastoid: Gk. mastos, breast
4. Petrous part. The petrous (pet'rus) part can be seen in the

floor of the cranium (figs. 6.19 and 6.23). The structures of
the middle ear and inner ear are housed in this dense part of
the temporal bone. The carotid (ka-rot'id) canal and the
jugular foramen border on the medial side of the petrous
part at the junction of the temporal and occipital bones.
The carotid canal allows blood into the brain via the internal carotid artery, and the jugular foramen lets blood drain
from the brain via the internal jugular vein. Three cranial
nerves also pass through the jugular foramen (see table 6.4).
The mastoid process of the temporal bone can be easily palpated as a bony knob immediately behind the earlobe. This
process contains a number of small air-filled spaces called mastoid
cells that can become infected in mastoiditis, as a result, for example, of a prolonged middle-ear infection.
Occipital Bone
The occipital bone forms the posterior and most of the base of
the skull. It articulates with the parietal bones at the lambdoid
suture. The foramen magnum is the large hole in the occipital
bone through which the spinal cord passes to attach to the brain
stem. On each side of the foramen magnum are the occipital
condyles (fig. 6.16), which articulate with the first vertebra (the
atlas) of the vertebral column. At the anterolateral edge of the
occipital condyle is the hypoglossal canal (fig. 6.17), through
which the hypoglossal nerve passes. A condyloid (kon'd-loid)
canal lies posterior to the occipital condyle (fig. 6.16). The
petrous: Gk. petra, rock

magnum: L. magnum, great
CHAPTER 6
FIGURE 6.22 Radiographs of the skull showing the paranasal sinuses. (a) An anteroposterior view and (b) a right lateral view.

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(b)
FIGURE 6.23 The temporal bone. (a) A lateral view and (b) a medial view.
external occipital protuberance is a prominent posterior projection on the occipital bone that can be felt as a definite bump just
under the skin. The superior nuchal (noo'kal) line is a ridge of
bone extending laterally from the occipital protuberance to the
mastoid part of the temporal bone. Sutural bones are small clusters of irregularly shaped bones that frequently occur along the
lambdoid suture.
Sphenoid Bone
The sphenoid (sfe'noid) bone forms part of the anterior base
of the cranium and can be viewed laterally and inferiorly
(figs. 6.15 and 6.16). This bone has a somewhat mothlike shape
(fig. 6.24). It consists of a body and laterally projecting greater
and lesser wings that form part of the orbit. The wedgelike
body contains the sphenoidal sinuses and a prominent saddlelike depression, the sella turcica (sel'a tur's-ka). Commonly
called Turks saddle, the sella turcica houses the pituitary
gland. A pair of pterygoid (ter'-goid) processes project inferiorly from the sphenoid bone and help form the lateral walls of
the nasal cavity.
Several foramina (figs. 6.16, 6.19, and 6.24) are associated
with the sphenoid bone.
nuchal: Fr. nuque, nape of neck

sphenoid: Gk. sphenoeides, wedgelike
1. The optic canal is a large opening through the lesser wing

into the back of the orbit that provides passage for the
optic nerve and the ophthalmic artery.
2. The superior orbital fissure is a triangular opening between the wings of the sphenoid bone that provides passage for the ophthalmic nerve, a branch of the trigeminal
nerve, and for the oculomotor, trochlear, and abducens
nerves.
3. The foramen ovale is an opening at the base of the lateral
pterygoid plate, through which the mandibular nerve passes.
4. The foramen spinosum is a small opening at the posterior
angle of the sphenoid bone that provides passage for the
middle meningeal vessels.
5. The foramen lacerum (las'er-um) is an opening between
the sphenoid and the petrous part of the temporal bone,
through which the internal carotid artery and the
meningeal branch of the ascending pharyngeal artery pass.
6. The foramen rotundum is an opening just posterior to the
superior orbital fissure, at the junction of the anterior and
medial portions of the sphenoid bone. The maxillary nerve
passes through this foramen.
Located on the inferior side of the cranium, the sphenoid bone
would seem to be well protected from trauma. Actually, just the
opposite is trueand in fact the sphenoid is the most frequently fractured bone of the cranium. It has several broad, thin, platelike extensions that are perforated by numerous foramina. A blow to almost any
portion of the skull causes the buoyed, fluid-filled brain to rebound
against the vulnerable sphenoid bone, often causing it to fracture.

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CHAPTER 6
FIGURE 6.24 The sphenoid bone. (a) A superior view and (b) a posterior view.
Ethmoid Bone
The ethmoid bone is located in the anterior portion of the floor
of the cranium between the orbits, where it forms the roof of
the nasal cavity (figs. 6.17, 6.20, and 6.25). An inferior projection of the ethmoid bone, called the perpendicular plate, forms
the superior part of the nasal septum that separates the nasal
cavity into two chambers. Each chamber of the nasal cavity is
referred to as a nasal fossa. Flanking the perpendicular plate on
each side is a large but delicate mass of bone riddled with ethmoidal air cells, collectively constituting the ethmoid sinus. A
spine of the perpendicular plate, the crista galli (kris'ta gal'e),
projects superiorly into the cranial cavity and serves as an attachment for the meninges covering the brain. On both lateral
walls of the nasal cavity are two scroll-shaped plates of the ethmoid bone, the superior and middle nasal conchae (kong'ke
singular, concha) (fig. 6.26), also known as turbinates. At right
angles to the perpendicular plate, within the floor of the cranium, is the cribriform (krib'r-form) plate, which has numerous cribriform foramina for the passage of olfactory nerves
from the epithelial lining of the nasal cavity. The bones of the
nasal cavity are summarized in table 6.6.
Creek
Cribriform plate
Cribriform foramina
Orbital plate
Ethmoidal air cells

ethmoid: Gk. ethmos, sieve
crista galli: L. crista, crest; galli, cocks comb
conchae: L. conchae, shells
cribriform: L. cribrum, sieve; forma, like
Crista galli
Superior nasal concha
Middle nasal concha
Perpendicular plate
FIGURE 6.25 An anterior view of the ethmoid bone.

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FIGURE 6.26 The lateral wall of the nasal cavity.
TABLE 6.6 Bones That Enclose

the Nasal Cavity
Region of
Nasal Cavity
Contributing Bones
Roof (superior)
Ethmoid bone (cribriform plate); frontal bone
Floor (inferior)
Maxilla; palatine bone
Lateral wall
Maxilla; palatine bone
Nasal septum (medial)
Ethmoid bone (perpendicular plate); vomer;

nasal bone
Bridge
Nasal bone
Conchae
Ethmoid bone (superior and middle conchae);

inferior nasal concha
The moist, warm vascular lining within the nasal cavity is susceptible to infections, particularly if a person is not in good
health. Infections of the nasal cavity can spread to several surrounding areas. The paranasal sinuses connect to the nasal cavity and
are especially prone to infection. The eyes may become reddened
and swollen during a nasal infection because of the connection of
the nasolacrimal duct, through which tears drain from the anterior
surface of the eye to the nasal cavity. Organisms may spread via
the auditory tube from the nasopharynx to the middle ear. With prolonged nasal infections, organisms may even ascend to the
meninges covering the brain via the sheaths of the olfactory nerves
and pass through the cribriform plate to cause meningitis.
Facial Bones
The 14 bones of the skull not in contact with the brain are
called facial bones. These bones, together with certain cranial
bones (frontal bone and portions of the ethmoid and temporal
bones), give shape and individuality to the face. Facial bones also
support the teeth and provide attachments for various muscles
that move the jaw and cause facial expressions. With the exceptions of the vomer and mandible, all of the facial bones are
paired. The articulated facial bones are illustrated in figures 6.14
through 6.21.
Maxilla
The two maxillae (mak-sil'e) unite at the midline to form the
upper jaw, which supports the upper teeth. Incisors, canines
(cuspids), premolars, and molars are anchored in dental alveoli,
(tooth sockets), within the alveolar (al-ve'o-lar) process of the
maxilla (fig. 6.27). The palatine (pal'a-tn) process, a horizontal
plate of the maxilla, forms the greater portion of the hard palate
(pal'it), or roof of the mouth. The incisive foramen (fig. 6.16) is
located in the anterior region of the hard palate, behind the in-
incisor: L. incidere, to cut

canine: L. canis, dog
molar: L. mola, millstone
alveolus: L. alveus, little cavity

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(b)
FIGURE 6.27 The maxilla. (a) A lateral view and (b) a medial view.
If the two palatine processes fail to join during early prenatal

development (about 12 weeks), a cleft palate results. A cleft
palate may be accompanied by a cleft lip lateral to the midline.
These conditions can be surgically treated with excellent cosmetic results. An immediate problem, however, is that a baby
with a cleft palate may have a difficult time nursing because it is unable to create the necessary suction within the oral cavity to swallow
effectively.
Palatine Bone
The L-shaped palatine bones form the posterior third of the hard
palate, a part of the orbits, and a part of the nasal cavity. The
horizontal plates of the palatines contribute to the formation of
the hard palate (fig. 6.28). At the posterior angle of the hard
palate is the large greater palatine foramen that provides passage
for the greater palatine nerve and descending palatine vessels
(fig. 6.16). Two or more smaller lesser palatine foramina are positioned posterior to the greater palatine foramen. Branches of
the lesser palatine nerve pass through these openings.
Zygomatic Bone
The two zygomatic bones (cheekbones) form the lateral contours
of the face. A posteriorly extending temporal process of this bone
unites with the zygomatic process of the temporal bone to form the
zygomatic arch (fig. 6.16). The zygomatic bone also forms the lateral margin of the orbit. A small zygomaticofacial (zi''go-mat''kofa'shal) foramen, located on the anterolateral surface of this bone,
allows passage of the zygomatic nerves and vessels.
Lacrimal Bone
The thin lacrimal bones form the anterior part of the medial wall
of each orbit (fig. 6.21). These are the smallest of the facial
bones. Each one has a lacrimal sulcusa groove that helps form
the nasolacrimal canal. This opening permits the tears of the eye
to drain into the nasal cavity.
Nasal Bone
The small, rectangular nasal bones (fig. 6.14) join at the midline
to form the bridge of the nose. The nasal bones support the flexible cartilaginous plates, which are a part of the framework of the
nose. Fractures of the nasal bones or fragmentation of the associated cartilages are common facial injuries.
Inferior Nasal Concha

The two inferior nasal conchae are fragile, scroll-like bones that
project horizontally and medially from the lateral walls of the
nasal cavity (figs. 6.14 and 6.20). They extend into the nasal
cavity just below the superior and middle nasal conchae, which
are part of the ethmoid bone (see fig. 6.25). The inferior nasal
conchae are the largest of the three paired conchae, and, like the
other two, are covered with a mucous membrane to warm,
moisten, and cleanse inhaled air.
CHAPTER 6
cisors. An infraorbital foramen is located under each orbit and

serves as a passageway for the infraorbital nerve and artery to the
nose (figs. 6.14, 6.15, 6.21, and 6.27). A final opening within the
maxilla is the inferior orbital fissure. It is located between the
maxilla and the greater wing of the sphenoid (fig. 6.14) and is
the external opening for the maxillary nerve of the trigeminal
nerve and infraorbital vessels. The large maxillary sinus located
within the maxilla is one of the four paranasal sinuses (figs. 6.20
and 6.22).

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CHAPTER 6
FIGURE 6.28 The palatine bone. (a) A medial view and (b) the
two palatine bones viewed posteriorly. The two palatine bones form
the posterior portion of the hard palate.
Condylar process
Coronoid process
Mandibular notch
Mandibular foramen
Ramus of
mandible
Creek
Oblique line
Mental protuberance
Masseteric
tuberosity
Angle of mandible
Head of condylar process

Neck of condylar process
Coronoid process
Body of mandible
Mandibular margin
(a)
Mental foramen
Mandibular foramen
Pterygoid
tuberosity
Angle of mandible
Mental spine
(b)
FIGURE 6.29 The mandible. (a) A lateral view and (b) a posterior view.

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Vomer
The vomer (vo'mer) is a thin, flattened bone that forms the
lower part of the nasal septum (figs. 6.16, 6.17, and 6.20). Along
with the perpendicular plate of the ethmoid bone, it supports the
layer of septal cartilage that forms most of the anterior and inferior parts of the nasal septum.
Mandible
Dentists use bony landmarks of the facial region to locate the

nerves that traverse the foramina in order to inject anesthetics.
For example, the trigeminal nerve is composed of three large nerves,
the lower two of which convey sensations from the teeth, gums, and
jaws. The mandibular teeth can be desensitized by an injection near
the mandibular foramen called a third-division, or lower, nerve block.
An injection near the foramen rotundum of the skull, called a seconddivision nerve block, desensitizes all of the upper teeth on one side
of the maxilla.
Hyoid Bone
The single hyoid bone is a unique part of the skeleton in that it
does not attach directly to any other bone. It is located in the
neck region, below the mandible, where it is suspended from the
styloid process of the temporal bone by the stylohyoid muscles
and ligaments. The hyoid bone has a body, two lesser cornua
vomer: L. vomer, plowshare
mandible: L. mandere, to chew
CHAPTER 6
The mandible (jawbone) is the largest, strongest bone in the

face. It is attached to the skull by paired temporomandibular
joints (see fig. 8.23), and is the only movable bone of the skull.
The horseshoe-shaped front and horizontal lateral sides of the
mandible are referred to as the body (fig. 6.29). Extending vertically from the posterior part of the body are two rami (ra'mi
singular, ramus). At the superior margin of each ramus is a
knoblike condylar process, which articulates with the mandibular fossa of the temporal bone, and a pointed coronoid process
for the attachment of the temporalis muscle. The depressed area
between these two processes is called the mandibular notch. The
angle of the mandible is where the horizontal body and vertical
ramus meet at the corner of the jaw.
Two sets of foramina are associated with the mandible:
the mental foramen, on the anterolateral aspect of the body of
the mandible below the first molar, and the mandibular foramen, on the medial surface of the ramus. The mental nerve and
vessels pass through the mental foramen, and the inferior alveolar nerve and vessels are transmitted through the mandibular
foramen. Several muscles that close the jaw extend from the
skull to the mandible (see chapter 9). The mandible of an adult
supports 16 teeth within dental alveoli, which occlude with the
16 teeth of the maxilla.
FIGURE 6.30 An anterior view of the hyoid bone.

(kor'nyoo-asingular, cornu) extending anteriorly, and two
greater cornua (fig. 6.30), which project posteriorly to the stylohyoid ligaments.
The hyoid bone supports the tongue and provides attachment for some of its muscles (see fig. 9.18). It may be palpated by
placing a thumb and a finger on either side of the upper neck
under the lateral portions of the mandible and firmly squeezing
medially. This bone is carefully examined in an autopsy when
strangulation is suspected, because it is frequently fractured during strangulation.
Auditory Ossicles
Three small paired bones, called auditory ossicles, are located
within the middle-ear cavities in the petrous part of the temporal
bones (fig. 6.31). From outer to inner, these bones are the
malleus (hammer), incus (anvil), and stapes (stirrup). As
described in chapter 15, their movements transmit sound impulses through the middle-ear cavity (see p. 518).
ramus: L. ramus, branch

condylar: L. condylus, knucklelike
malleus: L. malleus, hammer
coronoid: Gk. korone, like a crows beak

cornu: L. cornu, horn
incus: L. incus, anvil

stapes: L. stapes, stirrup

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FIGURE 6.31 The three auditory ossicles within the middle-ear

cavity.
Knowledge Check
13. State which facial and cranial bones of the skull are paired
and which are unpaired. Also, indicate at least two structural features associated with each bone of the skull.
14. Describe the location of each bone of the skull and indicate the sutures that join these bones.
15. What is the function of each of the following: sella turcica,
foramen magnum, petrous part of the temporal bone, crista
galli, and nasal conchae?
16. Which facial bones support the teeth?
VERTEBRAL COLUMN
The vertebral column consists of a series of irregular bones called
vertebrae, separated from each other by fibrocartilaginous intervertebral discs. Vertebrae enclose and protect the spinal cord,
FIGURE 6.32 The vertebral column of an adult has four curves

named according to the region in which they occur. The bodies of the
vertebrae are separated by intervertebral discs, which allow flexibility.
support the skull and allow for its movement, articulate with the
rib cage, and provide for the attachment of trunk muscles. The
intervertebral discs lend flexibility to the vertebral column and
absorb vertical shock.

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curves until it begins sitting upright and walking. (Note the differences in the curves between the sexes.)
Objective 12
Identify the bones of the five regions of the

vertebral column and describe the characteristic curves of
each region.
Objective 13
Describe the structure of a typical vertebra.
The vertebral column (backbone) and the spinal cord of the

nervous system constitute the spinal column. The vertebral column has four functions:
1. to support the head and upper extremities while permitting
freedom of movement;
2. to enable bipedalism;
3. to provide attachment for various muscles, ribs, and visceral organs; and
4. to protect the spinal cord and permit passage of the
spinal nerves.
The vertebral column is typically composed of 33 individual vertebrae, some of which are fused. There are 7 cervical,
12 thoracic, 5 lumbar, 3 to 5 fused sacral, and 4 or 5 fused coccygeal (kok-sij'e-al) vertebrae; thus, the adult vertebral column is
composed of a total of 26 movable parts. Vertebrae are separated
by fibrocartilaginous intervertebral discs and are secured to each
other by interlocking processes and binding ligaments. This
structural arrangement permits only limited movement between
adjacent vertebrae but extensive movement for the vertebral column as a whole. Between the vertebrae are openings called intervertebral foramina that allow passage of spinal nerves.
When viewed from the side, four curvatures of the vertebral column can be identified (fig. 6.32). The cervical, thoracic,
and lumbar curves are identified by the type of vertebrae they
include. The pelvic curve (sacral curve) is formed by the shape

of the sacrum and coccyx (kok'siks). The curves of the vertebral
column play an important functional role in increasing the
strength and maintaining the balance of the upper part of the
body; they also make possible a bipedal stance.
The four vertebral curves are not present in an infant. The
cervical curve begins to develop at about 3 months as the baby
begins holding up its head, and it becomes more pronounced as
the baby learns to sit up (fig. 6.33). The lumbar curve develops
as a child begins to walk. The thoracic and pelvic curves are
called primary curves because they retain the shape of the fetus.
The cervical and lumbar curves are called secondary curves because they are modifications of the fetal shape.
General Structure of Vertebrae

Vertebrae are similar in their general structure from one region
to another. A typical vertebra consists of an anterior drumshaped body, which is in contact with intervertebral discs
above and below (fig. 6.34). The vertebral arch is attached to
the posterior surface of the body and is composed of two supporting pedicles (ped'-kulz) and two arched laminae (lam'-ne).
The space formed by the vertebral arch and body is the vertebral foramen, through which the spinal cord passes. Between
the pedicles of adjacent vertebrae are the intervertebral foramina, through which spinal nerves emerge as they branch off the
spinal cord.
pedicle: L. pediculus, small foot

lamina: L. lamina, thin layer
CHAPTER 6
FIGURE 6.33 The development of the vertebral curves. An infant is born with the two primary curves but does not develop the secondary

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Body
Transverse
foramen
Superior
articular
process
Occipital
condyle
of skull
Vertebral
foramen
Atlas
Pedicle
Vertebral arch
Lamina
Axis
Spinous process
(note that it is bifid)
Body of C3
(b)
Intervertebral
disc between
C5 and C6
Anterior arch
of atlas
CHAPTER 6
Spinous
process
of C7
(a)
Atlas
Axis
Superior articular
facet (articulates
with occipital condyle)
Dens of axis
Body of
axis
Transverse
foramina
Transverse
processes
Posterior arch
of atlas
Spinous process
of axis
(c)
FIGURE 6.34 Cervical vertebrae. (a) A radiograph of the cervical region, (b) a superior view of a typical cervical vertebra, and (c) the articulated atlas and axis.
Seven processes arise from the vertebral arch of a typical

vertebrae: the spinous process, two transverse processes, two
superior articular processes, and two inferior articular
processes (fig. 6.35). The spinous process and transverse
processes serve for muscle attachment and the superior and inferior articular processes limit twisting of the vertebral column.
The spinous process protrudes posteriorly and inferiorly from the
vertebral arch. The transverse processes extend laterally from
each side of a vertebra at the point where the lamina and pedicle
join. The superior articular processes of a vertebra interlock with
the inferior articular processes of the bone above.
A laminectomy is the surgical removal of the spinous
processes and their supporting vertebral laminae in a particular region of the vertebral column. A laminectomy may be performed
to relieve pressure on the spinal cord or nerve root caused by a
blood clot, a tumor, or a herniated (ruptured) disc. It may also be performed on a cadaver to expose the spinal cord and its surrounding
meninges.
Regional Characteristics of Vertebrae

Cervical Vertebrae
The seven cervical vertebrae form a flexible framework for the
neck and support the head. The bone tissue of cervical vertebrae
is more dense than that found in the other vertebral regions,
and, except for those in the coccygeal region, the cervical vertebrae are smallest. Cervical vertebrae are distinguished by the
presence of a transverse foramen in each transverse process
(fig. 6.34). The vertebral arteries and veins pass through this
opening as they contribute to the blood flow associated with the
brain. Cervical vertebrae C2C6 generally have a bifid, or
notched, spinous process. The bifid spinous processes increase
the surface area for attachment of the strong nuchal ligament that
attaches to the back of the skull. The first cervical vertebra has
no spinous process, and the process of C7 is not bifid and is larger
than those of the other cervical vertebrae.

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Creek
Facet for
head of rib
Transverse process
Body
Body
T1
Demifacet for
head of rib
Demifacet for
head of rib
Vertebral
foramen
Pedicle
Facet for
tubercle of rib
Superior
articular
process
T2T8
Demifacet for
head of rib
Inferior articular
process
Spinous
process
Transverse
process
Facet for
tubercle
of rib
Facet for
head of rib
Superior articular
process
Pedicle
Lamina
Spinous
process
(b)
CHAPTER 6
T9
Intervertebral
foramen
Intervertebral
disc
T10
Facet for
head of rib
(a)
FIGURE 6.35 Thoracic vertebrae. Representative vertebrae in (a) a lateral view and (b) a superior view.
The atlas is the first cervical vertebra (sometimes called cervical 1 or C1). The atlas lacks a body, but it does have a short,
rounded spinous process called the posterior tubercle. It also has
cupped superior articular surfaces that articulate with the oval occipital condyles of the skull. This atlanto-occipital joint supports the
skull and permits the nodding of the head in a yes movement.
The axis is the second cervical vertebra (C2). It has a peglike dens (odontoid process) for rotation with the atlas in turning
the head from side to side, as in a no movement.
Whiplash is a common term for any injury to the neck. Muscle,
bone, or ligament injury in this portion of the spinal column is
relatively common in individuals involved in automobile accidents
and sports injuries. Joint dislocation occurs commonly between the
fourth and fifth or fifth and sixth cervical vertebrae, where neck movement is greatest. Bilateral dislocations are particularly dangerous because of the probability of spinal cord injury. Compression fractures
of the first three cervical vertebrae are common and follow abrupt
forced flexion of the neck. Fractures of this type may be extremely
painful because of pinched spinal nerves.
Thoracic Vertebrae
Twelve thoracic vertebrae articulate with the ribs to form the
posterior anchor of the rib cage. Thoracic vertebrae are larger
than cervical vertebrae and increase in size from superior (T1) to
inferior (T12). Each thoracic vertebra has a long spinous process,
which slopes obliquely downward, and facets (fovea) for articulation with the ribs (fig. 6.35).
Lumbar Vertebrae
The five lumbar vertebrae are easily identified by their heavy
bodies and thick, blunt spinous processes (fig. 6.36) for attachment of powerful back muscles. They are the largest vertebrae of
atlas: from Gk. mythology, Atlasthe Titan who supported the heavens
axis: L. axis, axle
odontoid: Gk. odontos, tooth
lumbar: L. lumbus, loin

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L3
Transverse
process
L4
Spinous
process
L5
Superior
articular
process
CHAPTER 6
Intervertebral
disc
Sacrum
Coccyx
(a)
FIGURE 6.36 Lumbar vertebrae. (a) A radiograph, (b) a superior view, and (c) a lateral view.
the vertebral column. Their articular processes are also distinctive in that the facets of the superior pair are directed medially
instead of posteriorly and the facets of the inferior pair are directed laterally instead of anteriorly.
Sacrum
The wedge-shaped sacrum provides a strong foundation for the
pelvic girdle. It consists of four or five sacral vertebrae (fig. 6.37)
that become fused after age 26. The sacrum has an extensive auricular surface on each lateral side for the formation of a slightly
movable sacroiliac (sak''ro-il'e-ak) joint with the ilium of the
hip. A median sacral crest is formed along the posterior surface
by the fusion of the spinous processes. Posterior sacral foramina
on either side of the crest allow for the passage of nerves from
the spinal cord. The sacral canal is the tubular cavity within the
sacrum: L. sacris, sacred
sacrum that is continuous with the vertebral canal. Paired superior articular processes, which articulate with the fifth lumbar
vertebra, arise from the roughened sacral tuberosity along the
posterior surface.
The smooth anterior surface of the sacrum forms the posterior surface of the pelvic cavity. It has four transverse lines denoting the fusion of the vertebral bodies. At the ends of these
lines are the paired pelvic foramina (anterior sacral foramina).
The superior border of the anterior surface of the sacrum, called
the sacral promontory (prom'on-tor''e), is an important obstetric
landmark for pelvic measurements.
Coccyx
The triangular coccyx (tailbone) is composed of three to five fused
coccygeal vertebrae. The first vertebra of the fused coccyx has two
coccyx: Gk. kokkyx, like a cuckoos beak

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CHAPTER 6
FIGURE 6.37 The sacrum and coccyx. (a) An anterior view and (b) a posterior view.
long coccygeal cornua, which are attached by ligaments to the

sacrum (fig. 6.37). Lateral to the cornua are the transverse processes.
Distinct losses in height occur during middle and old age.
Between the ages of 50 and 55, there is a decrease of 0.5 to
2.0 cm (0.25 to 0.75 in.) because of compression and shrinkage of the
intervertebral discs. Elderly individuals may suffer a further loss of
height because of osteoporosis (see Clinical Considerations at the
end of this chapter).
TABLE 6.7 Regions of the Vertebral

Column
Region
Number of
Bones
Cervical
Transverse foramina; superior

facets of atlas articulate
with occipital condyle;
dens of axis; spinous
processes of third through
sixth vertebrae are generally
bifid
Thoracic
12
Long spinous processes that

slope obliquely downward;
facets for articulation
with ribs
Lumbar
Large bodies; prominent

transverse processes; short,
thick spinous processes
Sacrum
4 or 5 fused
vertebrae
Extensive auricular surface;

median sacral crest;
posterior sacral foramina;
sacral promontory; sacral
canal
Coccyx
3 to 5 fused
vertebrae
Small and triangular; coccygeal

cornua
The regions of the vertebral column are summarized in

table 6.7.
When a person sits, the coccyx flexes anteriorly, acting as
a shock absorber. An abrupt fall on the coccyx, however,
may cause a painful subperiosteal bruising, fracture, or fracturedislocation of the sacrococcygeal joint. An especially difficult childbirth can even injure the coccyx of the mother. Coccygeal trauma is
painful and may require months to heal.
Knowledge Check
17. Which are the primary curves of the vertebral column and
which are the secondary curves? Describe the characteristic
curves of each region.
18. What is the function of the transverse foramina of the cervical vertebrae?
19. Describe the diagnostic differences between a thoracic and
a lumbar vertebra. Which structures are similar and could
therefore be characteristic of a typical vertebra?
Diagnostic Features

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CHAPTER 6
Costal cartilages
FIGURE 6.38 The rib cage.
RIB CAGE
The cone-shaped, flexible rib cage consists of the thoracic vertebrae, 12 paired ribs, costal cartilages, and the sternum. It encloses
and protects the thoracic viscera and is directly involved in the
mechanics of breathing.
Objective 14
Identify the parts of the rib cage and compare

and contrast the various types of ribs.
The sternum (breastbone), ribs, costal cartilages, and the previously described thoracic vertebrae form the rib cage (fig. 6.38).
The rib cage is anteroposteriorly compressed and more narrow
superiorly than inferiorly. It supports the pectoral girdle and
upper extremities, protects and supports the thoracic and upper
abdominal viscera, and plays a major role in breathing (see
fig. 9.21). Certain bones of the rib cage contain active sites in
the bone marrow for the production of red blood cells.
Sternum
The sternum is an elongated, flattened bony plate consisting of
three separate bones: the upper manubrium, (ma-noo'bre-um),
the central body, and the lower xiphoid (zif'oid; zi'foid) process.
The xiphoid process is often cartilaginous. On the lateral sides of
the sternum are costal notches where the costal cartilages attach.
A jugular notch is formed at the superior end of the manubrium,
and a clavicular notch for articulation with the clavicle is present on both sides of the sternal notch. The manubrium articulates with the costal cartilages of the first and second ribs. The
body of the sternum attaches to the costal cartilages of the sec-
sternum: Gk. sternon, chest

manubrium: L. manubrium, a handle
xiphoid: Gk. xiphos, sword
costal: L. costa, rib

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Chapter 6
Articular surfaces
of head of rib
165
Head
Neck
Tubercle
Articular surface
of tubercle
Body
ee
Superior surface
Angle
Cr
Costochondral
joint
Costal groove
Internal surface
Inferior surface
Costal cartilage
FIGURE 6.39 The structure of a rib.
Ribs
Embedded in the muscles of the body wall are 12 pairs of ribs,
each pair attached posteriorly to a thoracic vertebra. Anteriorly,
the first seven pairs are anchored to the sternum by individual
costal cartilages; these ribs are called true ribs. The remaining
five pairs (ribs 8, 9, 10, 11, and 12) are termed false ribs. Because
the last two pairs of false ribs do not attach to the sternum at all,
they are referred to as floating ribs.
Although the ribs vary structurally, each of the first 10
pairs has a head and a tubercle for articulation with a vertebra.
The last two have a head but no tubercle. In addition, each of
the 12 pairs has a neck, angle, and body (fig. 6.39). The head
angle of Louis: from Pierre C. A. Louis, French physician, 17871872
projects posteriorly and articulates with the body of a thoracic

vertebra (fig. 6.40). The tubercle is a knoblike process, just lateral to the head. It articulates with the facet on the transverse
process of a thoracic vertebra. The neck is the constricted area
between the head and the tubercle. The body is the curved main
part of the rib. Along the inner surface of the body is a depressed
canal called the costal groove that protects the costal vessels and
nerve. Spaces between the ribs are called intercostal spaces and
are occupied by the intercostal muscles.
Fractures of the ribs are relatively common, and most frequently occur between ribs 3 and 10. The first two pairs of ribs
are protected by the clavicles; the last two pairs move freely and will
give with an impact. Little can be done to assist the healing of broken
ribs other than binding them tightly to limit movement.
Knowledge Check
20. Describe the rib cage and list its functions. What determines whether a rib is true, false, or floating?
21. Distinguish between the costal margin and costal angle.
Each bone is a dynamic living organ that is influenced by hormones, diet, aging, and disease. Because the development of
bone is genetically controlled, congenital abnormalities may
occur. The hardness of bones gives them strength, yet they lack
the resiliency to avoid fracture when they undergo severe
CHAPTER 6
ond through the tenth ribs. The xiphoid process does not attach
to ribs but is an attachment for abdominal muscles. The costal
cartilages of the eighth, ninth, and tenth ribs fuse to form the
costal margin of the rib cage. A costal angle is formed where the
two costal margins come together at the xiphoid process. The
sternal angle (angle of Louis) may be palpated as an elevation
between the manubrium and body of the sternum at the level of
the second rib (fig. 6.38). The costal angle, costal margins, and
sternal angle are important surface landmarks of the thorax and
abdomen (see figs. 10.19 and 10.20).

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Body of
vertebra
Radiate ligament
Articular facet
for tubercle
of rib
Rib
Costotransverse
ligament
Cr
ee
166
Transverse
process
Lateral
costotransverse
ligament
Spinous
process
CHAPTER 6
FIGURE 6.40 Articulation of a rib with a thoracic vertebra as seen in a superior view.
trauma. (Fractures are discussed in chapter 7 and joint injuries

are discussed in chapter 8.) All of these aspects of bone make for
some important and interesting clinical considerations.
Developmental Disorders
Congenital malformations account for several types of skeletal
deformities. Certain bones may fail to form during osteogenesis,
or they may form abnormally. Cleft palate and cleft lip are malformations of the palate and face. They vary in severity and seem
to involve both genetic and environmental factors. Spina bifida
(sp' na bif'-da) is a congenital defect of the vertebral column resulting from a failure of the laminae of the vertebrae to fuse,
leaving the spinal cord exposed (fig. 6.41). The lumbar area is
most likely to be affected, and frequently only a single vertebra is
involved.
Nutritional and Hormonal Disorders

Several bone disorders result from nutritional deficiencies or
from excessive or deficient amounts of the hormones that regulate bone development and growth. Vitamin D has a tremendous
influence on bone structure and function. When there is a deficiency of this vitamin, the body is unable to metabolize calcium
and phosphorus. Vitamin D deficiency in children causes rickets. The bones of a child with rickets remain soft and structurally
weak, and bend under the weight of the body (see fig. 5.11).
A vitamin D deficiency in the adult causes the bones to
give up stored calcium and phosphorus. This demineralization results in a condition called osteomalacia (os''te-o-ma-la'sha). Osteomalacia occurs most often in malnourished women who have
repeated pregnancies and who experience relatively little exposure to sunlight. It is marked by increasing softness of the bones,
so that they become flexible and thus cause deformities.
FIGURE 6.41 In spina bifida, failure of the vertebral arches to fuse

permits a herniation of the meninges that cover the spinal cord
through the vertebral column. This results in a condition called
meningomyelocele.
The consequences of endocrine disorders are described in

chapter 14. Because hormones exert a strong influence on bone
development, however, a few endocrine disorders will be briefly
mentioned here. Hypersecretion of growth hormone from the pituitary gland leads to gigantism in young people if it begins before ossification of their epiphyseal plates. In adults, it leads to
acromegaly (ak''ro-meg'a-le), which is characterized by hypertrophy of the bones of the face, hands, and feet. In a child, growth
hormone deficiency results in slowed bone growtha condition
called dwarfism.
Pagets disease, a bone disorder that affects mainly older
adults, occurs more frequently in males than in females. It is
characterized by disorganized metabolic processes within bone
Pagets disease: from Sir James Paget, English surgeon, 181499

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167
FIGURE 6.42 A bone scan of the legs of a patient suffering from

arthritis in the left knee joint. In a bone scan, an image of an arthritic
joint shows up lighter than most of a normal joint.
Neoplasms of Bone
Malignant bone tumors are three times more common than benign
tumors. Pain is the usual symptom of either type of osseous neoplasm, although benign tumors may not have accompanying pain.
Two types of benign bone tumors are osteomas, which are
the more frequent and which often involve the skull, and osteoid
osteomas, which are painful neoplasms of the long bones, usually
in children.
Osteogenic sarcoma (sar-ko'ma) is the most virulent type
of bone cancer. It frequently metastasizes through the blood to
the lungs. This disease usually originates in the long bones and is
accompanied by aching and persistent pain.
A bone scan (fig. 6.42) is a diagnostic procedure frequently
done on a person who has had a malignancy elsewhere in the
body that may have metastasized to the bone. The patient receiving a bone scan may be injected with a radioactive substance
that accumulates more rapidly in malignant tissue than in normal tissue. Entire body radiographs show malignant bone areas as
intensely dark dots.
CHAPTER 6
tissue. The activity of osteoblasts and osteoclasts becomes irregular, resulting in thick bony deposits in some areas of the skeleton and fragile, thin bones in other areas. The vertebral
column, pelvis, femur, and skull are most often involved, and
become increasingly painful and deformed. Bowed leg bones,
abnormal curvature of the spine, and enlargement of the skull
may develop. The cause of Pagets disease is currently not
known.
FIGURE 6.43 A geriatric skull. Note the loss of teeth and the degeneration of bone, particularly in the facial region.
Aging of the Skeletal System

Senescence affects the skeletal system by decreasing skeletal mass
and density and increasing porosity and erosion (fig. 6.43). Bones
become more brittle and susceptible to fracture. Articulating surfaces also deteriorate, contributing to arthritic conditions.
Arthritic diseases are second to heart disease as the most common debilitation in the elderly.
Osteoporosis (os''te-o-po-ro'sis) is a weakening of the
bones, primarily as a result of calcium loss. The causes of osteoporosis include aging, inactivity, poor diet, and an imbalance in
hormones or other chemicals in the blood. It is most common in
older women because low levels of estrogens after menopause
lead to increased bone resorption, and the formation of new
bone is not sufficient to keep pace. People with osteoporosis are
prone to bone fracture, particularly at the pelvic girdle and vertebrae, as the bones become too brittle to support the weight of the
body. Complications of hip fractures often lead to permanent disability, and vertebral compression fractures may produce a permanent curved deformity of the spine.
Although there is no known cure for osteoporosis, good
eating habits and a regular program of exercise, established at an
early age and continued throughout adulthood, can minimize its

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effects. Treatment in women through dietary calcium, exercise,

and estrogens has had limited positive results. In addition, a drug
called alendronate (Fosamax), approved by the FDA in 1995, has
been shown to be effective in managing osteoporosis. This drug
works without hormones to block osteoclast activity, making it
useful for women who choose not to be treated with estrogen replacement therapy.

The height (overall length) of the vertebral column is equal to the
sum of the thicknesses of the vertebrae plus the sum of the thicknesses
of the intervertebral discs. The body of a vertebra consists of outer
CHAPTER 6
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A 50-year-old male is brought to the emergency room by his spouse because he is acting strangely. She states that he has been
ill for several days, but in the past 24 to
48 hours, he has become forgetful, less communicative, and sleepy. He had been complaining of a headache, which was localized
to the front of his head and face, and a fever
for several days. His wife reports a history of
sinus infections. Physical examination reveals a fever, frontal tenderness, and altered
mental status. You order a head CT. Two
representative images are shown here.
QUESTIONS
1. What abnormalities do you observe
in the CT?
2. Where is the fluid in the right subdural
space (see arrow) coming from?
3. What is the diagnosis?
4. What is your suggested course of
treatment?
compact bone and inner spongy bone. An intervertebral disc consists

of a fibrocartilage sheath called the anulus fibrosus and a mucoid center
portion called the nucleus pulposus. The intervertebral discs generally
change their anatomical configuration as one ages. In early adulthood,
the nucleus pulposus is spongy and moist. With advanced age, however, it desiccates, resulting in a flattening of the intervertebral disc.
Collectively, the intervertebral discs account for 25% of the height of
the vertebral column. As they flatten with age, there is a gradual decrease in a persons overall height. Height loss may also result from undetectable compression fractures of the vertebral bodies, which are
common in elderly people. This phenomenon, however, is considered
pathological and is not an aspect of the normal aging process. In a person with osteoporosis, there is often a marked decrease in height and
perhaps more serious clinical problems as well, such as compression of
spinal nerves.

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169

A 53-year-old male is injured in an automobile accident. He was wearing his seat
belt when the accident occurred. The
paramedics stabilize his spine and bring
him to the emergency room. He is complaining of low back pain. You examine
the patient and note that he has de- Q U E S T I O N S

creased strength and sensation in both 1. What is the patients injury?
legs, as well as tenderness in the small 2. Why does the patient have neurological
of his back. A radiograph of the lumbar
symptoms in his legs?
spine as well as a CT scan are obtained.
3. How does the seat belt contribute to
this injury?
CHAPTER 6
Important Clinical Terminology

achondroplasia (a-kon''dro-pla'ze-a) A
genetic defect that inhibits formation
of cartilaginous bone during fetal
development.
craniotomy Surgical cutting into the
cranium to provide access to the brain.
epiphysiolysis (ep''-fiz''e-ol'-sis) A
separation of the epiphysis from the diaphysis
of a growing long bone.
laminectomy The surgical removal of the
posterior arch of a vertebra, usually to repair
a herniated intervertebral disc.
orthopedics The branch of medicine

concerned with the diagnosis and treatment
of trauma, diseases, and abnormalities
involving the skeletal and muscular systems.
osteitis (os-te- tis) An inflammation of bone
tissue.

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osteoblastoma (os''te-o-blas-to'ma) A benign

tumor produced from bone-forming cells,
most frequently in the vertebrae of young
children.
osteochondritis (os''te-o-kon-dri'tis) An
inflammation of bone and cartilage tissues.
osteomyelitis (os''te-o-mi''e-ltis) An
inflammation of bone marrow caused by
bacteria or fungi.
osteonecrosis (os''te-o-ne kro'sis) The death of
bone tissue, usually caused by obstructed
arteries.
osteopathology The study of bone diseases.

osteosarcoma A malignant tumor of bone
tissue.
osteotomy (os''te-ot'o-me) The cutting of a
bone, usually by means of a saw or a chisel.
Chapter Summary
Organization of the Skeletal System
(pp. 132134)
CHAPTER 6
1. The axial skeleton consists of the skull,

auditory ossicles, hyoid bone, vertebral
column, and rib cage.
2. The appendicular skeleton consists of the
bones within the pectoral girdle, upper
extremities, pelvic girdle, and lower
extremities.
Functions of the Skeletal System

(pp. 134135)
1. The mechanical functions of bones
include the support and protection of
softer body tissues and organs. In addition,
certain bones function as levers during
body movement.
2. The metabolic functions of bones include
hemopoiesis and storage of fat and
minerals.
Bone Structure (pp. 135138)

1. Bone structure includes the shape and
surface features of each bone, along with
gross internal components.
2. Bones may be structurally classified as
long, short, flat, or irregular.
3. The surface features of bones are classified
as articulating surfaces, nonarticulating
prominences, and depressions and
openings.
4. A typical long bone has a diaphysis, or
shaft, filled with marrow in the medullary
cavity; epiphyses; epiphyseal plates for
linear growth; and a covering of
periosteum for appositional growth and
the attachments of ligaments and
tendons.
Bone Tissue (pp. 138140)

1. Compact bone is the dense outer portion;
spongy bone is the porous, vascular inner
portion.
2. The five types of bone cells are osteogenic
cells, in contact with the endosteum and
periosteum; osteoblasts (bone-forming
cells); osteocytes (mature bone cells);
osteoclasts (bone-destroying cells); and
bone-lining cells, along the surface of
most bones.
3. In compact bone, the lamellae of osteons
are the layers of inorganic matrix
surrounding a central canal. Osteocytes
are mature bone cells, located within
capsules called lacunae.
Bone Growth (pp. 140144)

1. Bone growth is an orderly process
determined by genetics, diet, and
hormones.
2. Most bones develop through
endochondral ossification.
3. Bone remodeling is a continual process
that involves osteoclasts in bone
resorption and osteoblasts in the formation
of new bone tissue.
Skull (pp. 144158)

1. The eight cranial bones include the
frontal (1), parietals (2), temporals (2),
occipital (1), sphenoid (1),
and ethmoid (1).
(a) The cranium encloses and protects the
brain and provides for the attachment
of muscles.
(b) Sutures are fibrous joints between
cranial bones.
2. The 14 facial bones include the nasals (2),

maxillae (2), zygomatics (2), mandible (1),
lacrimals (2), palatines (2), inferior nasal
conchae (2), and vomer (1).
(a) The facial bones form the basic shape
of the face, support the teeth, and
provide for the attachment of the
facial muscles.
(b) The hyoid bone is located in the neck,
between the mandible and the larynx.
(c) The auditory ossicles (malleus, incus,
and stapes) are located within each
middle-ear chamber of the petrous
part of the temporal bone.
3. The paranasal sinuses include the
sphenoidal and ethmoidal sinuses in the
sphenoid and ethmoid bones respectively in
the cranial region and the frontal and
maxillary sinuses in the frontal and maxillary
bones respectively in the facial region.
Vertebral Column (pp. 158163)

1. The vertebral column consists of 7 cervical,
12 thoracic, 5 lumbar, 4 or 5 fused sacral,
and 3 to 5 fused coccygeal vertebrae.
2. Cervical vertebrae have transverse
foramina; thoracic vertebrae have facets
and demifacets for articulation with ribs;
lumbar vertebrae have large bodies; sacral
vertebrae are triangularly fused and
contribute to the pelvic girdle; and the
coccygeal vertebrae form a small
triangular bone.
Rib Cage (pp. 164165)

1. The sternum consists of a manubrium,
body, and xiphoid process.
2. There are seven pairs of true ribs and five
pairs of false ribs. The inferior two pairs of
false ribs (pairs 11 and 12) are called
floating ribs.
Review Activities
Objective Questions
1. A bone is considered to be
(a) a tissue.
(b) a cell.
(c) an organ.
(d) a system.
2. Which of the following statements is

false?
(a) Bones are important in the synthesis
of vitamin D.
(b) Bones and teeth contain about 99%
of the bodys calcium.
(c) Red bone marrow is the primary site

for hemopoiesis.
(d) Most bones develop through
endochondral ossification.

6. Skeletal System:
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Chapter 6
(c) thoracic vertebrae.

(d) cervical vertebrae.
10. The bone disorder that frequently
develops in elderly people, particularly if
they experience prolonged inactivity,
malnutrition, or a hormone imbalance is
(a) osteitis.
(b) osteonecrosis.
(c) osteoporosis. (d) osteomalacia.
Essay Questions
1. What are the functions of the skeletal
system? Do the individual bones of the
skeleton carry out these functions equally?
Explain.
2. Explain why there are approximately
270 bones in an infant but 206 bones in a
mature adult.
3. List the bones of the skull that are paired.
Which are unpaired? Identify the bones of
the skull that can be palpated.
4. Describe the development of the skull.
What are the fontanels, where are they
located, and what are their functions?
5. Which facial bones contain foramina? What
structures pass through these openings?
6. Distinguish between the axial and
appendicular skeletons. Describe where
these two components articulate.
7. List four types of bones based on shape
and give an example of each type.
8. Diagram a typical long bone. Label the
epiphyses, diaphysis, epiphyseal plates,
medullary cavity, nutrient foramina,
periosteum, and articular cartilages.
9. List the bones that form the cranial
cavity, the orbit, and the nasal cavity.
Describe the location of the paranasal
sinuses, the mastoidal sinus, and the
inner-ear cavity.
10. Describe how bones grow in length and
width. How are these processes similar,
and how do they differ? Explain how
radiographs can be used to determine
normal bone growth.
11. Explain the process of endochondral
ossification of a long bone. Why is it
important that a balance be maintained
between osteoblast activity and osteoclast
activity?
12. Describe the curvature of the vertebral
column. What do the terms primary curves
and secondary curves refer to?
13. List two or more characteristics by which
vertebrae from each of the five regions of
the vertebral column can be identified.

171
14. Identify the bones that form the rib cage.

What functional role do the bones and
the costal cartilages have in respiration?
15. Describe the nature of bones in each of
the following age periods: newborn,
teenager, adult, and elderly.
1. Many people think that the bones in our
bodies are deadunderstandable
considering that we associate bones with
graveyards, Halloween, and leftover
turkey from a Thanksgiving dinner. Your
kid brother is convinced of this. What
information could you use to try to get
him to change his mind?
2. The sensory organs involved with sight,
smell, and hearing are protected by bone.
Describe the locations of each of these
sensory organs and list the associated
bones that provide protection.
3. Explain why a proper balance of vitamins,
hormones, and minerals is essential in
maintaining healthy bone tissue. Give
examples of diseases or skeletal conditions
that may occur in the event of an
imbalance of any of these three essential
substances.
4. The most common surgical approach to a
pituitary gland tumor is through the nasal
cavity. With the knowledge that the
pituitary gland is supported by the sella
turcica of the braincase, list the bones
that would be involved in the removal of
the tumor.
5. The contour of a childs head is distinctly
different from that of an adult. Which
skull bones exhibit the greatest amount of
change as a child grows to adulthood?
6. You read in National Geographic that a
team of archaeologists recently completed
an examination of 18 skeletons from
people buried under tons of volcanic ash
1,200 years ago. By analyzing the bones,
the scientists were able to determine the
sex, physical health (including a partial
medical history), approximate age, and
even the general profession of each of the
18 individuals. How could the
examination of a preserved skeleton yield
such a vast array of information?
CHAPTER 6
3. Match each of the following foramina

with the bone in which it occurs.
1. foramen rotundum
2. mental foramen
3. carotid canal
4. cribriform foramina
5. foramen magnum
(a) ethmoid bone
(b) occipital bone
(c) sphenoid bone
(d) mandible
(e) temporal bone
4. With respect to the hard palate, which
of the following statements is false?
(a) It is composed of two maxillae and
two palatine bones.
(b) It separates the oral cavity from the
nasal cavity.
(c) The mandible articulates with the
posterolateral angles of the hard
palate.
(d) The median palatine suture, incisive
foramen, and greater palatine
foramina are three of its structural
features.
5. The location of the sella turcica is
immediately
(a) superior to the sphenoidal sinus.
(b) inferior to the frontal sinus.
(c) medial to the petrous parts of the
temporal bones.
(d) superior to the perpendicular plate
of the ethmoid bone.
6. Specialized bone cells that enzymatically
reabsorb bone tissue are
(a) osteoblasts.
(b) osteocytes.
(c) osteons.
(d) osteoclasts.
7. The mandibular fossa is located in which
structural part of the temporal bone?
(a) the squamous part
(b) the tympanic part
(c) the mastoid part
(d) the petrous part
8. The crista galli is a structural feature of
which bone?
(a) the sphenoid bone
(b) the ethmoid bone
(c) the palatine bone
(d) the temporal bone
9. Transverse foramina are
characteristic of
(a) lumbar vertebrae.
(b) sacral vertebrae.
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VI. Maintenance of the

Body
17. Respiratory System
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Chapter 17
Respiratory System
627
(a)
FIGURE 17.29 A pneumothorax of the right lung. The right side of

the thorax appears uniformly dark because it is filled with air; the
spaces between the ribs are also greater than those on the left, because the ribs are released from the elastic tension of the lungs. The
left lung appears denser (less dark) because of shunting of blood
from the right to the left lung.
(b)
Trauma or Injury
Humans are especially susceptible to epistaxes (ep''-stak'sez)
(nosebleeds) because the prominent nose can be easily bumped
and because the nasal mucosa has extensive vascularity for
warming the inspired air. Epistaxes may also be caused by high
blood pressure or diseases such as leukemia.
When air enters the pleural cavity surrounding either lung,
the condition is referred to as a pneumothorax (fig. 17.29). A
pneumothorax can result from an external injury, such as a stabbing, bullet wound, or penetrating fractured rib, or it can be the
result of internal conditions. A severely diseased lung, as in emphysema, can create a pneumothorax as the wall of the lung deteriorates along with the visceral pleura and permits air to enter
the pleural cavity.
If you are alone and choking, use whatever is available to

apply force just below your diaphragm. Press into a table or a
sink, or use your own fist.
CHAPTER 17
FIGURE 17.28 (a) An infant with a unilateral cleft lip and palate.
(b) The same child following corrective surgery.
Choking on a foreign object is a common serious trauma to

the respiratory system. More than eight Americans choke to
death each day on food lodged in their trachea. A simple procedure termed the abdominal thrust (Heimlich) maneuver can
save the life of a person who is choking (fig. 17.30). The abdominal thrust maneuver is performed as follows:
A. If the victim is standing or sitting:
1. Stand behind the victim or the victims chair and wrap
your arms around his or her waist.
2. Grasp your fist with your other hand and place the fist
against the victims abdomen, slightly above the navel
and below the rib cage.
3. Press your fist into the victims abdomen with a quick
upward thrust.
4. Repeat several times if necessary.
B. If the victim is lying down:
1. Position the victim on his or her back.
2. Face the victim, and kneel on his or her hips.
3. With one of your hands on top of the other, place the
heel of your bottom hand on the abdomen, slightly
above the navel and below the rib cage.
4. Press into the victims abdomen with a quick upward
thrust.

628
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Maintenance of the Body
FIGURE 17.30 The abdominal thrust (Heimlich) maneuver.
People saved from drowning and victims of shock frequently experience apnea (cessation of breathing) and will soon
die if not revived by artificial respiration. The accepted treatment for reviving a person who has stopped breathing is illustrated in figure 17.31.
Common Respiratory Disorders

CHAPTER 17
The McGrawHill
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A cough is the most common symptom of respiratory disorders.

Acute problems may be accompanied by dyspnea or wheezing.
Respiratory or circulatory problems may cause cyanosis (si'a-no'sis), a blue discoloration of the skin resulting from blood with a
low oxygen content.
Although the common cold is the most widespread of all
respiratory diseases, there is still no cure for this ailmentonly
medications that offer symptomatic relief. Colds occur repeatedly
because acquired immunity for one virus does not protect against
other viruses. Cold viruses generally incite acute inflammation in
the respiratory mucosa, causing flow of mucus, sometimes accompanied by fever and/or headache.
Nearly all of the structures and regions of the respiratory
passageways can become infected and inflamed. Influenza is a
viral disease that causes an inflammatory condition of the
upper respiratory tract. Influenza can be epidemic, but fortu-
nately vaccines are available. Sinusitis is an inflammation of

the paranasal sinuses. Sinusitis can be quite painful if the
drainage ducts from the sinuses into the nasal cavity become
blocked. Tonsillitis may involve any or all of the tonsils and
frequently follows other lingering diseases of the oral or pharyngeal regions. Chronic tonsillitis sometimes requires a tonsillectomy. Laryngitis is inflammation of the larynx, which often
produces a hoarse voice and limits the ability to talk. Tracheobronchitis and bronchitis are infections of the regions for
which they are named. Severe inflammation in these areas can
cause smaller respiratory tubules to collapse, blocking the passage of air.
Diseases of the lungs are likewise common and usually serious. Pneumonia is an acute infection and inflammation of lung
tissue accompanied by exudation (accumulation of fluid). It is
usually caused by bacteria, most commonly by the pneumococcus
bacterium. Viral pneumonia is caused by a number of different
viruses. Tuberculosis is an inflammatory disease of the lungs
contracted by inhaling air sneezed or coughed by someone who is
carrying active tuberculosis bacteria. Tuberculosis softens lung
tissue, which eventually becomes ulcerated. Asthma is a disease
that affects people who are allergic to certain inhaled antigens. It
causes a swelling and blocking of lower respiratory tubes, often
accompanied by the formation of mucus plugs. Pleurisy (ploor'se) is an inflammation of the pleura and is usually secondary to
some other respiratory disease. Inspiration may become painful,
and fluid may collect within the pleural cavity. Emphysema
(em''f -se'ma) is a disease that causes the breakdown of the pulmonary alveoli, thus increasing the size of air spaces and decreasing the surface area (fig. 17.32). It is a frequent cause of death
among heavy cigarette smokers.
Cancer in the respiratory system is known to be caused by
the repeated inhalation of irritating substances, such as cigarette
smoke. Cancers of the lip, larynx, and lungs (fig. 17.33) are especially common in smokers over the age of 50.
Disorders of Respiratory Control

A variety of disease processes can result in cessation of breathing
during sleep, or sleep apnea. Sudden infant death syndrome
(SIDS) is an especially tragic form of sleep apnea that claims the
lives of about 10,000 babies annually in the United States. Victims of this condition are apparently healthy 2-to-5-month-old
babies who die in their sleep without apparent reasonhence,
the laypersons term, crib death. These deaths seem to be
caused by failure of the respiratory control mechanisms in the
brain stem and/or by failure of the carotid bodies to be stimulated
by reduced arterial oxygen.
tuberculosis: L. tuberculum, diminutive of tuber, swelling

cyanosis: Gk. kyanosis, dark-blue color
influenza: L. influentia, a flowing in
asthma: Gk. asthma, panting

emphysema: Gk. emphysan, blow up, inflate


Body
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Chapter 17
Respiratory System
629
CHAPTER 17

Unit 6
646

Body
18. Digestive System
The McGrawHill
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Accessory salivary
gland
Tongue
Accessory salivary
gland
Lingual frenulum
Opening of
submandibular
duct
Parotid gland
Sublingual ducts
Parotid duct
Masseter muscle
Sublingual
gland
Submandibular
gland
Submandibular
duct
Mandible (cut)
FIGURE 18.11 The salivary glands.
CHAPTER 18
Mucous cells
Intralobular
parotid duct
Lumen of
submandibular
intralobular duct
Seromucous
acini
(a)
Serous cells
(b)
Mucous cells
Serous cells
(c)
Intralobular
sublingual duct
FIGURE 18.12 The histology of the salivary glands. (a) The parotid gland, (b) the submandibular gland, and (c) the sublingual gland.


Body
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Chapter 18
Digestive System
647
TABLE 18.4 Major Salivary Glands

Gland
Location
Duct
Entry into Oral Cavity
Type of Secretion
Parotid gland
Anterior and inferior to auricle;

subcutaneous over masseter
muscle
Parotid (Stensens) duct
Lateral to upper second molar
Watery serous fluid, salts,

and enzyme
Submandibular gland
Inferior to the base of the tongue
Submandibular (Whartons)
duct
Papilla lateral to lingual

frenulum
Watery serous fluid with

some mucus
Sublingual gland
Anterior to submandibular gland

under the tongue
Several small sublingual ducts

(Rivinus ducts)
Ducts along the base of the

tongue
Mostly thick, stringy

mucus; salts; and enzyme
(salivary amylase)
CHAPTER 18
FIGURE 18.13 A posterior view of the constrictor muscles of the pharynx. The right side has been cut away to illustrate the interior structures
in the pharynx.
and the lumen is lined with a mucous membrane containing

stratified squamous epithelium. The pharynx is divided into
three regions: the nasopharynx, posterior to the nasal cavity; the
oropharynx, posterior to the oral cavity; and the laryngopharynx,
at the level of the larynx (see fig. 17.3).
The external circular layer of pharyngeal muscles, called
constrictors (fig. 18.13), compresses the lumen of the pharynx
involuntarily during swallowing. The superior constrictor mus-
cle attaches to bony processes of the skull and mandible and encircles the upper portion of the pharynx. The middle constrictor
muscle arises from the hyoid bone and stylohyoid ligament and
encircles the middle portion of the pharynx. The inferior constrictor muscle arises from the cartilages of the larynx and encircles the lower portion of the pharynx. During breathing, the
lower portion of the inferior constrictor muscle is contracted,
preventing air from entering the esophagus.

648
Unit 6

Body
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The motor and most of the sensory innervation to the

pharynx is via the pharyngeal plexus, situated chiefly on the
middle constrictor muscle. It is formed by the pharyngeal
branches of the glossopharyngeal and vagus nerves, together with
a deep sympathetic branch from the superior cervical ganglion.
The pharynx is served principally by ascending pharyngeal
arteries, which branch from the external carotid arteries. The
pharynx is also served by small branches from the inferior thyroid
arteries, which arise from the thyrocervical trunk. Venous return
is via the internal jugular veins.
Adventitia
Tunica
muscularis
Submucosa
Mucosa
Lumen
Knowledge Check
8. Define the terms heterodont and diphyodont. Which of the
four kinds of teeth is not found in deciduous dentition?
9. Where are the enamel, dentin, cementum, and pulp of a
tooth located? Explain how a tooth is anchored into its
socket.
10. Describe the location of the parotid, submandibular, and sublingual ducts and state where they empty into the oral cavity.
ESOPHAGUS AND STOMACH

A bolus of food is passed from the esophagus to the stomach,
where it is churned and mixed with gastric secretions. The chyme
thus produced is sent past the pyloric sphincter of the stomach to
the duodenum.
Objective 9
Describe the steps in deglutition.
Objective 10
Describe the location, gross structure, and

functions of the stomach.
Objective 11
CHAPTER 18
Describe the histological structure of the

esophagus and stomach. List the cell types in the gastric
mucosa, along with their secretory products.
Esophagus
The esophagus is that portion of the GI tract that connects the
pharynx to the stomach (see figs. 18.1 and 18.15). It is a collapsible tubular organ, approximately 25 cm (10 in.) long, originating
at the larynx and lying posterior to the trachea.
The esophagus is located within the mediastinum of the
thorax and passes through the diaphragm just above the opening
into the stomach. The opening through the diaphragm is called
the esophageal hiatus (e-sof''a-je'al hi-a'tus) The esophagus
is lined with a nonkeratinized stratified squamous epithelium
(fig. 18.14); its walls contain either skeletal or smooth muscle, depending on the location. The upper third of the esophagus contains skeletal muscle; the middle third, a combination of skeletal
and smooth muscle; and the terminal portion, smooth muscle only.
esophagus: Gk. oisein, to carry; phagema, food
FIGURE 18.14 The histology of the esophagus seen in cross

section.
The lower esophageal (gastroesophageal) sphincter is a

slight thickening of the circular muscle fibers at the junction of
the esophagus and the stomach. After food or fluid pass into the
stomach, this sphincter constricts to prevent the stomach contents from regurgitating into the esophagus. There is a normal
tendency for this to occur because the thoracic pressure is lower
than the abdominal pressure as a result of the air-filled lungs.
The lower esophageal sphincter is not a well-defined sphincter muscle comparable to others located elsewhere along the
GI tract, and it does at times permit the acidic contents of the stomach to enter the esophagus. This can create a burning sensation
commonly called heartburn, although the heart is not involved. In infants under a year of age, the lower esophageal sphincter may function erratically, causing them to spit up following meals. Certain
mammals, such as rodents, have a true lower esophageal sphincter
and cannot regurgitate, which is why poison grains are effective in
killing mice and rats.
Swallowing Mechanisms
Swallowing, or deglutition (de''gloo-tish'un), is the complex mechanical and physiological act of moving food or fluid from the
oral cavity to the stomach. For descriptive purposes, deglutition
is divided into three stages.
The first deglutitory stage is voluntary and follows mastication, if food is involved. During this stage, the mouth is closed
and breathing is temporarily interrupted. A bolus is formed as
the tongue is elevated against the transverse palatine folds
(palatal rugae) of the hard palate (see fig. 18.5) through contraction of the mylohyoid and styloglossus muscles and the intrinsic
muscles of the tongue.
The second stage of deglutition is the passage of the bolus
through the pharynx. The events of this stage are involuntary
and are elicited by stimulation of sensory receptors located at the
opening of the oropharynx. Pressure of the tongue against the
transverse palatine folds seals off the nasopharynx from the oral
cavity, creates a pressure, and forces the bolus into the oropharynx. The soft palate and pendulant palatine uvula are elevated to
close the nasopharynx as the bolus passes. The hyoid bone and


Body
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Chapter 18
Digestive System
649
Esophagus
Esophagus
Fundus
Cardia
Peristaltic
contraction
of muscularis
layer of
esophagus
Lesser
curvature
Swallowed bolus
entering stomach
Greater
curvature
Duodenum
Pylorus
Body
Stomach
(a)
FIGURE 18.15 An anteroposterior radiograph of the esophagus

showing peristaltic contraction and movement of a bolus into the
stomach.
the larynx are also elevated. Elevation of the larynx against the
epiglottis seals the glottis so that food or fluid is less likely to
enter the trachea. Sequential contraction of the constrictor muscles of the pharynx moves the bolus through the pharynx to the
esophagus. This stage is completed in just a second or less.
The third stage, the entry and passage of food through the
esophagus, is also involuntary. The bolus is moved through the
esophagus by peristalsis (fig. 18.15). In the case of fluids, the entire process of deglutition takes place in slightly more than a second; for a typical bolus, the time frame is 5 to 8 seconds.
The stomachthe most distensible part of the GI tractis located in the upper left abdominal quadrant, immediately below
the diaphragm. Typically J-shaped when empty, the stomach is
continuous with the esophagus superiorly and empties into the
duodenal portion of the small intestine inferiorly (fig. 18.16). In
the stomach, which serves as a holding organ for ingested food,
the food is mechanically churned with gastric secretions to form
a pasty material called chyme (km). Once formed, chyme is
moved from the stomach to the small intestine.
The stomach is divided into four regions: the cardia, fundus, body, and pylorus (fig. 18.17). The cardia is the narrow
upper region immediately below the lower esophageal sphincter.
The fundus is the dome-shaped portion to the left of and in di-
chyme: L. chymus, juice

cardia: Gk. kardia, heart (upper portion, nearer the heart)
(b)
FIGURE 18.16 The stomach. (a) As seen from a cadaver, the

stomach is a J-shaped organ. (b) A radiograph of the stomach is of
clinical value for detecting ulcers, constrictions, or ingested objects.
A patient swallows radiopaque barium, which coats the lining of the
stomach and duodenum. These structures and certain abnormalities
may then show up in the radiograph.
rect contact with the diaphragm. The body is the large central
portion, and the pylorus is the funnel-shaped terminal portion.
The pyloric sphincter is the modified circular muscle at the end
of the pylorus, where it joins the small intestine. Pylorus is a
fundus: L. fundus, bottom

pylorus: Gk. pyloros, gatekeeper
CHAPTER 18
Stomach

650
Unit 6

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Gastric
folds
CHAPTER 18
FIGURE 18.17 The major regions and structures of the stomach.

Greek word meaning gatekeeper, and this junction is just that,
regulating the movement of chyme into the small intestine and
prohibiting backflow.
The stomach has two surfaces and two borders. The
broadly rounded surfaces are referred to as the anterior and posterior surfaces. The medial concave border is the lesser curvature (fig. 18.17), and the lateral convex border is the greater
curvature. The lesser omentum extends between the lesser curvature and the liver, and the greater omentum is attached to the
greater curvature (see fig. 18.3d).
The wall of the stomach is composed of the same four tunics found in other regions of the GI tract, with two principal
modifications: (1) an extra oblique muscle layer is present in the
muscularis, and (2) the mucosa is thrown into numerous longitudinal folds, called gastric folds or gastric rugae, which permit
stomach distension. The mucosa is further characterized by
the presence of microscopic gastric pits and gastric glands
(figs. 18.18 and 18.19).
There are five types of cells in the gastric glands that secrete specific products.
Goblet cells secrete protective mucus.
Parietal cells secrete hydrochloric acid (HCl).
Principal cells (chief cells) secrete pepsinogen, an inactive
form of the protein-digesting enzyme pepsin.
Columnar
epithelium of
mucosal ridge
Gastric pit
Gastric glands
with principal and
parietal cells
Lamina propria
FIGURE 18.18 The histology of the mucosa of the stomach.


Body
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Chapter 18
Digestive System
651
Gastric pits
(a)
Mucous
cell
Mucosa
Gastric
gland
Parietal
cell
Submucosa
Principal
cell
(b)
FIGURE 18.19 Gastric pits and gastric glands of the stomach mucosa. (a) Gastric pits are the openings of the gastric glands. (b) Gastric
glands consist of mucous cells, principal cells, and parietal cells, each type producing a specific secretion.
Argentaffin (ar-jent'a-fin) cells secrete serotonin, histamine, and autocrine regulators.
Endocrine cells (G cells) secrete the hormone gastrin into
the blood.
The stomach is sensitive to emotional stress. Mucus, secreted

by mucous cells of the stomach, is important in preventing hydrochloric acid and the digestive enzyme pepsin from eroding the
stomach wall. Peptic ulcers may be caused by an increase in cellular
secretion or by insufficient secretions of protective mucus. Another
protective feature is the rapid mitotic activity of the columnar epithelium of the stomach. The entire lining of the stomach is usually replaced every few days. Nevertheless, in the United States
approximately 10% of the male population and 4% of the female population develop peptic ulcers.
Regulation of gastric activity is autonomic. The sympathetic neurons arise from the celiac plexus, and the parasympathetic neurons arise from the vagus nerves. Parasympathetic
neurons synapse in the myenteric plexus between the muscular
layers and in the submucosal plexus in the submucosa. Parasymargentaffin: L. argentum, silver; affinis, attraction (become colored with
silver stain)
Phase
Response
Cephalic phase
Sight, taste, small or mental stimuli evoke

parasympathetic response via vagus nerves; 50150
ml of gastric juice is secreted
Gastric phase
Food in the stomach stretches the mucosa, and

chemical breakdown of protein stimulates the release
of gastrin; gastrin stimulates the production of
600750 ml of gastric juice
Intestinal phase
Chyme entering the duodenum stimulates intestinal

cells to release intestinal gastrin; intestinal gastrin
stimulates the production of additional small
quantities of gastric juice
pathetic impulses promote gastric activity, the phases of which

are presented in table 18.5.
Vomiting is the reflexive response of emptying the stomach
through the esophagus, pharynx, and oral cavity. This action is
controlled by the vomiting center of the medulla oblongata. Stimuli within the GI tract, especially the duodenum, may activate the
vomiting center, as may nauseating odors or sights, motion sickness, or body stress. Various drugs called emetics can also stimulate
a vomiting reflex. The mechanics of vomiting are as follows:
(1) strong, sustained contractions of the upper small intestine, followed by a contraction of the pyloric sphincter; (2) relaxation of
CHAPTER 18
In addition to these products, the gastric mucosa (probably

the parietal cells) secretes intrinsic factor, a polypeptide that is required for absorption of vitamin B12 in the small intestine. Continued gastric activity when the stomach is empty causes hunger
sensations known as hunger pangs. Eating fills the stomach resulting in hunger satiety, or a perception of fullness.
TABLE 18.5 Phases of Gastric Secretion

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the lower esophageal sphincter and contraction of the pyloric

portion of the stomach; (3) a shallow inspiration and closure of
the glottis; and (4) compression of the stomach against the liver
by contraction of the diaphragm and the abdominal muscles.
This reflexive sequence causes a forceful ejection of vomit. The
feeling of nausea is caused by stimuli in the vomiting center and
may or may not cause vomiting.
The only function of the stomach that appears to be essential
for life is the secretion of intrinsic factor. This polypeptide is
needed for the intestinal absorption of vitamin B12, which is required
for maturation of red blood cells in the bone marrow. Following surgical removal of the stomach (gastrectomy), a patient has to receive vitamin B12 (together with intrinsic factor) orally or through injections,
so that he or she will not develop pernicious anemia.
Knowledge Check
11. Describe the three stages of deglutition with reference to
the structures involved.
12. Describe the structure and function of the lower
esophageal sphincter.
13. List the functions of the stomach. What is the function of
the gastric folds?
14. Describe the modifications of the stomach that aid in mechanical and chemical digestion.
SMALL INTESTINE
The small intestine, consisting of the duodenum, jejunum, and
ileum, is the site where digestion is completed and nutrients are
absorbed. The surface area of the intestinal wall is increased by
plicae circulares, intestinal villi, and microvilli.
Objective 12
Describe the location and regions of the small

intestine and the way in which it is supported.
CHAPTER 18
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Objective 13
List the functions of the small intestine and

describe the structural adaptations through which these
functions are accomplished.
Objective 14
Describe the movements that occur within the

small intestine.
The small intestine is that portion of the GI tract between the

pyloric sphincter of the stomach and the ileocecal valve that
opens into the large intestine. It is positioned in the central and
lower portions of the abdominal cavity and is supported, except
for the first portion, by the mesentery (fig. 18.20). The fanshaped mesentery permits movement of the small intestine but
leaves little chance for it to become twisted or kinked. Enclosed
within the mesentery are blood vessels, nerves, and lymphatic
vessels that supply the intestinal wall.
The small intestine is approximately 3 m (12 ft) long and
2.4 cm (1 in.) wide in a living person, but it will measure nearly
twice this length in a cadaver when the muscular wall is relaxed.
It is called the small intestine because of its relatively small diameter compared to that of the large intestine. The small intestine is the bodys major digestive organ and the primary site of
nutrient absorption. Its digestive enzymes, along with those of
the salivary glands, gastric glands, and pancreas, are summarized
in table 18.6.
The small intestine is innervated by the superior mesenteric plexus. The branches of the plexus contain sensory fibers,
postganglionic sympathetic fibers, and preganglionic parasympathetic fibers. The arterial blood supply to the small intestine is
through the superior mesenteric artery and branches from the
celiac trunk and the inferior mesenteric artery. The venous
drainage is through the superior mesenteric vein. This vein
unites with the splenic vein to form the hepatic portal vein,
which carries nutrient-rich blood to the liver (see fig. 16.41).
Regions of the Small Intestine

On the basis of function and histological structure, the small intestine is divided into three regions.
1. The duodenum (doo''o-de'num, doo-od'e-num) is a relatively fixed, C-shaped tubular organ, measuring approximately 25 cm (10 in.) from the pyloric sphincter of the
stomach to the duodenojejunal (doo-od''e-no''je-joo'nal)
flexure. Except for a short portion near the stomach, the
duodenum is retroperitoneal. Its concave surface faces to
the left, where it receives bile secretions from the liver and
gallbladder through the common bile duct and pancreatic
secretions through the pancreatic duct of the pancreas
(fig. 18.21). These two ducts unite to form a common
entry into the duodenum called the hepatopancreatic ampulla (ampulla of Vater), which pierces the duodenal wall
and drains into the duodenum from an elevation called the
duodenal papilla. It is here that bile and pancreatic juice
enter the small intestine. The duodenal papilla can be
opened or closed by the action of the sphincter of ampulla
(of Oddi). The duodenum differs histologically from the
rest of the small intestine by the presence of duodenal
(Brunners) glands in the submucosa (fig. 18.22). These
compound tubuloalveolar glands secrete mucus and are
most numerous near the superior end of the duodenum.
2. The jejunum (je-joo'num), which extends from the duodenum to the ilium, is approximately 1 m (3 ft) long. It has
a slightly larger lumen and more internal folds than does
the ileum, but its histological structure is similar to that of
the ileum.
duodenum: L. duodeni, 12 each (length of 12 fingers breadth)

ampulla of Vater: from Abraham Vater, German anatomist, 16841751
sphincter of Oddi: from Rugger Oddi, Italian physician, 18641913
Brunners glands: from Johann C. Brunner, Swiss anatomist, 16531727


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Duodenum
Jejunum
Intestinal
v
Ileum
Tunica
muscularis
FIGURE 18.20 The small intestine in relation to the stomach and a part of the large intestine. (a) The regions and mesenteric attachment. (b)
A section of the intestinal wall showing the mucosa and submucosa folded into structures called plicae circulares. (The regions of the small intestine are labeled in boldface type.)
Structural Modifications
of the Small Intestine
The products of digestion are absorbed across the epithelial lining of the intestinal mucosa. Absorption occurs primarily in the
jejunum, although some also occurs in the duodenum and ileum.
Peyers patches: from Johann K. Peyer, Swiss anatomist, 16531712
Absorption occurs at a rapid rate as a result of four specializations

that increase the intestinal surface area.
The three meter length of the small intestine.
The plicae (pli'se) circulares are large macroscopic folds of
mucosa (see fig. 18.20).
The intestinal villi (vil'e) are fingerlike macroscopic folds
of the mucosa that project into the lumen of the small intestine (fig. 18.23).
The microvilli are microscopic projections formed by the
folding of each epithelial cell membrane. In a light microscope, the microvilli display a somewhat vague brush
plica: L. plicatus, folded

villus: L. villosus, shaggy
CHAPTER 18
3. The ileum (il'e-um)not to be confused with the ilium of

the os coxaemakes up the remaining 2 m (67 ft) of the
small intestine. The terminal portion of the ileum empties
into the medial side of the cecum through the ileocecal
valve. Lymph nodules, called mesenteric (Peyers) patches,
are abundant in the walls of the ileum.

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TABLE 18.6 Digestive Enzymes

Enzyme
Source
Digestive Action
Salivary glands
Begins carbohydrate digestion by converting starch and glycogen to disaccharides
Gastric glands
Begins the digestion of nearly all types of proteins
Peptidase
Intestinal glands
Converts proteins into amino acids
Sucrase
Intestinal glands
Converts disaccharides into monosaccharides
Lipase
Intestinal glands
Converts fats into fatty acids and glycerol
Amylase
Intestinal glands
Converts starch and glycogen into disaccharides
Nuclease
Intestinal glands
Converts nucleic acids into nucleotides
Enterokinase
Intestinal glands
Activates trypsin
Salivary enzyme
Amylase
Gastric juice enzyme
Pepsin
Intestinal juice enzymes
Maltase
Lactase
Pancreatic juice enzymes

Amylase
Pancreas
Lipase
Pancreas
Peptidases
Pancreas
Convert proteins or partially digested proteins into amino acids
Pancreas
Trypsin
Chymotrypsin
Carboxypeptidase
Nuclease
Common bile duct
CHAPTER 18
Accessory pancreatic duct
Pancreatic duct
Duodenum
Sphincter
of ampulla
Pancreas
Hepatopancreatic ampulla
Duodenal papilla
FIGURE 18.21 A section of the duodenum showing the location of entry of the common bile duct and the pancreatic duct.


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Digestive System
655
Simple
columnar
epithelium
Intestinal villus
(lined with simple
columnar
epithelium)
Lacteal
Lamina propria
Intestinal
villus
Capillary
network
Goblet
cells
Muscularis
mucosa
Intestinal
crypt
Duodenal glands
Lymph
vessel
FIGURE 18.22 The histology of the duodenum.
Arteriole
border on the edges of the columnar epithelium

(fig. 18.24). The terms brush border and microvilli are
often used interchangeably in describing the small intestine.
Venule
FIGURE 18.23 The structure of an intestinal villus and intestinal

crypt.
CHAPTER 18
The intestinal villi are covered with columnar epithelial

cells, among which are interspersed the mucus-secreting goblet
cells. The lamina propria, which forms the connective tissue core
of each intestinal villus, contains numerous lymphocytes, blood
capillaries, and a lymphatic vessel called the lacteal (lak'te-al)
(fig. 18.23). Absorbed monosaccharides and amino acids enter
the blood capillaries; absorbed fatty acids and cholesterols enter
the lacteals. Intestinal villi are considered the functional units of
the digestive system because absorption through these structures
is how digested molecules enter the blood or lymph.
Epithelial cells at the tips of the intestinal villi are continuously shed and are replaced by cells that are pushed up from the
bases of the intestinal villi. The epithelium at the base of the intestinal villi invaginates downward at various points to form narrow pouches that open through pores into the intestinal lumen.
These structures are called the intestinal crypts (crypts of
Lieberkhn) (see fig. 18.23).
Waldrop
Mechanical Activities
Contractions of the longitudinal and circular muscles of the
small intestine produce three distinct types of movement: rhythmic segmentation, pendular movements, and peristalsis.
lacteal: L. lacteus, milk
crypts of Lieberkhn: from Johann N. Lieberkhn, German anatomist, 171156
FIGURE 18.24 An electron photomicrograph of microvilli

(arrow) at the exposed surface of a columnar epithelial cell in the
small intestine.

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Rhythmic segmentations are local contractions of the circular muscular layer. They occur at the rate of about 12 to 16 per
minute in regions containing chyme. Rhythmic segmentations
churn the chyme with digestive juices and bring it into contact
with the mucosa. During these contractions, the vigorous motion
of the intestinal villi stirs the chyme and facilitates absorption.
Pendular movements primarily occur in the longitudinal
muscle layer. In this motion, a constrictive wave moves along a
segment of the small intestine and then reverses and moves in
the opposite direction, moving the chyme back and forth. Pendular movements also mix the chyme but do not seem to have a
particular frequency.
Peristalsis (per''-stal'sis) is responsible for the propulsive
movement of the chyme through the small intestine. These
wavelike contractions are usually weak and relatively short, occurring at a frequency of about 15 to 18 per minute. Chyme requires 3 to 10 hours to travel the length of the small intestine.
Both muscle layers are involved in peristalsis.
The sounds of digestive peristalsis can be easily heard
through a stethoscope placed at various abdominal locations.
These sounds can be detected even through clothing. The sounds,
mostly clicks and gurgles, occur at a frequency of 5 to 30 per minute.
Knowledge Check
15. Describe the small intestine. Where is it located? How is it
subdivided? How is it supported?
16. What are the primary functions of the small intestine?
17. List four structural modifications of the small intestine that
increase its absorptive surface area.
18. Describe the movements of the small intestine. Which
movements are produced by the circular layer of the tunica
muscularis?
19. Which region of the small intestine is the longest? Which
is the shortest? How long does it take a portion of chyme to
move through the small intestine?
CHAPTER 18
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LARGE INTESTINE
The large intestine receives undigested food from the small intestine, absorbs water and electrolytes from chyme, and passes
feces out of the GI tract.
Objective 15
Identify the regions of the large intestine and

describe its gross and histological structure.
orly on the right side to a point just below the liver; it then
crosses to the left, descends into the pelvis, and terminates at the
anus. A specialized portion of the mesentery, the mesocolon,
supports the transverse portion of the large intestine along the
posterior abdominal wall.
The large intestine has little or no digestive function, but
it does absorb water and electrolytes from the remaining chyme.
In addition, the large intestine functions to form, store, and
expel feces from the body.
Regions and Structures

of the Large Intestine
The large intestine is structurally divided into the cecum, colon,
rectum, and anal canal (figs. 18.25 and 18.26). The cecum
(se'kum) is a dilated pouch positioned slightly below the ileocecal
valve. The ileocecal valve is a fold of mucous membrane at the
junction of the small intestine and large intestine that prohibits
the backflow of chyme. A fingerlike projection called the appendix is attached to the inferior medial margin of the cecum. The
8 cm (3 in.) appendix contains an abundance of lymphatic tissue
(fig. 18.27) that may serve to resist infection. Although the appendix serves no digestive function, it is thought to be a vestigial
remnant of an organ that was functional in human ancestors.
A common disorder of the large intestine is inflammation of the
appendix, or appendicitis. Wastes that accumulate in the appendix cannot be moved easily by peristalsis, because the appendix
has only one opening. Although the symptoms of appendicitis are
quite variable, they often include a high white blood cell count, localized pain in the lower right quadrant, and loss of appetite. Vomiting
may or may not occur. Rupture of the appendix (a burst appendix)
spreads infectious material throughout the peritoneal cavity, resulting
in peritonitis.
The superior portion of the cecum is continuous with the

colon, which consists of ascending, transverse, descending, and
sigmoid portions (fig. 18.25). The ascending colon extends superiorly from the cecum along the right abdominal wall to the inferior surface of the liver. Here the colon bends sharply to the left
at the hepatic flexure (right colic flexure) and continues across
the upper abdominal cavity as the transverse colon. At the left
abdominal wall, another right-angle bend called the splenic flexure (left colic flexure) marks the beginning of the descending
colon. From the splenic flexure, the descending colon extends
inferiorly along the left abdominal wall to the pelvic region. The
colon then angles medially from the brim of the pelvis to form an
S-shaped bend, known as the sigmoid colon.
Objective 16
Describe the functions of the large intestine

and explain how defecation is accomplished.
The large intestine averages 1.5 m (5 ft) in length and 6.5 cm

(2.5 in.) in diameter. It is called the large intestine because of
its relatively large diameter compared to that of the small intestine. The large intestine begins at the end of the ileum in the
lower right quadrant of the abdomen. From there, it leads superi-
cecum: L. caecum, blind pouch

appendix: L. appendix, attachment
colon: Gk. kolon, member of the whole
sigmoid: Gk. sigmoeides, shaped like a sigma,


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Digestive System
657
Transverse colon
Hepatic flexure
Splenic flexure
Mesocolon
Taeniae coli
Ascending colon
Descending colon
Epiploic appendage
Ileum
Ileocecal valve
Orifice of appendix
Haustrum
Cecum
Appendix
Sigmoid colon
Rectum
Loechel
Anal canal
FIGURE 18.25 The large intestine.
A hemorrhoid (hem'o-rod) is a mass of varicose veins in the

anal area caused, in part, by difficulty in defecating. Hemorrhoids, in reference to the condition in which such masses occur, are
also called piles. A first-degree hemorrhoid is contained within the
anal canal. A second-degree hemorrhoid prolapses, or extends out-
rectum: L. rectum, straight tube

anus: L. anus, ring
ward during defecation. A third-degree hemorrhoid remains prolapsed through the anal orifice. Rubber band constriction is a common medical treatment for a prolapsed hemorrhoid. In this technique,
a rubber band is tied around the hemorrhoid, constricting its blood
supply, so that the tissue dries and falls off. In a relatively new treatment, infrared photocoagulation, a high-energy light beam coagulates the hemorrhoid.
Although the large intestine consists of the same tunics as

the small intestine, there are some structural differences. The large
intestine lacks intestinal villi but does have numerous goblet cells
in the mucosal layer (fig. 18.29). The longitudinal muscle layer of the
muscularis forms three distinct muscle bands called taeniae coli
(te'ne-e ko'li) that run the length of the large intestine. A series of
bulges in the walls of the large intestine form sacculations, or haustra
(haws'tra), along its entire length (see figs. 18.25 and 18.26).
taenia: L. tainia, a ribbon

haustrum: L. haustrum, bucket or scoop
CHAPTER 18
The terminal 20 cm (7.5 in.) of the GI tract is the rectum,

and the last 2 to 3 cm of the rectum is referred to as the anal
canal (fig. 18.28). The rectum lies anterior to the sacrum, where
it is firmly attached by peritoneum. The anus is the external
opening of the anal canal. Two sphincter muscles guard the anal
opening: the internal anal sphincter, which is composed of
smooth muscle fibers, and the external anal sphincter, composed of skeletal muscle. The mucous membrane of the anal
canal is arranged in highly vascular, longitudinal folds called
anal columns.

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Rectum
Levator ani
muscle
Hemorrhoidal
vessels
Anal canal
External anal
sphincter
Anal columns
Internal anal
sphincter
Anus
Loechel
FIGURE 18.28 The anal canal.
Serosa
FIGURE 18.26 An anteroposterior radiograph after a barium
Tunica
muscularis
enema showing the regions, flexures, and the haustra of the large
intestine.
Submucosa
Mucosa
Center of
lymphatic
nodule
CHAPTER 18
Intestinal
glands
Columnar
epithelium with
goblet cells
(a)
Lumen
Tunica
muscularis
Submucosa
Serosa
FIGURE 18.27 The histology of the appendix shown in cross section.
(b)
FIGURE 18.29 The histology of the rectum. (a) A photomicrograph of the tunics. (b) A scanning electron photomicrograph of the
mucosa from a section of the colon. The arrow indicates the opening
of a goblet cell into the intestinal lumen.
(From R.G. Kessel and R.H. Kardon. Tissues and Organs: A Text-Atlas of Scanning
Electron Microscopy, 1979 W.H. Freeman and Company.)


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Chapter 18
Digestive System
659
TABLE 18.7 Mechanical Activity in the GI Tract

Region
Type of Motility
Frequency
Stimulus
Result
Oral cavity
Mastication
Variable
Initiated voluntarily; proceeds

reflexively
Pulverization: mixing with

saliva
Oral cavity and

pharynx
Deglutition
Maximum of 20 per min
Initiated voluntarily; reflexively

controlled by swallowing center
Clears oral cavity of food
Esophagus
Peristalsis
Depends on frequency of swallowing
Initiated by swallowing
Movement through the

esophagus
Stomach
Small intestine
Large intestine
Receptive relaxation
Matches frequency of swallowing
Unknown
Permits filling of stomach
Tonic contraction
1520 per min
Autonomic plexuses
Mixing and churning
Peristalsis
12 per min
Autonomic plexuses
Evacuation of stomach
Hunger contractions
3 per min
Low blood sugar level
Feeding
Peristalsis
1518 per min
Autonomic plexuses
Transfer through intestine
Rhythmic segmentation
1216 per min
Autonomic plexuses
Mixing
Pendular movements
Variable
Autonomic plexuses
Mixing
Peristalsis
312 per min
Autonomic plexuses
Transport
Mass movements
23 per day
Stretch
Fills sigmoid colon
Haustral churning
312 per min
Autonomic plexuses
Mixing
Defecation
Variable: 1 per day to 3 per week
Reflex triggered by rectal distension
Defecation
Mechanical Activities of the Large Intestine

Chyme enters the large intestine through the ileocecal valve. About
15 ml of pasty material enters the cecum with each rhythmic opening of the valve. The ingestion of food intensifies peristalsis of the
ileum and increases the frequency with which the ileocecal valve
opens; this is called the gastroileal reflex. Material entering the
large intestine accumulates in the cecum and ascending colon.
Three types of movements occur throughout the large intestine: peristalsis, haustral churning, and mass movement. Peristaltic movements of the colon are similar to those of the small
intestine, although they are usually more sluggish in the colon.
In haustral churning, a relaxed haustrum fills with food residues
until a point of distension is reached that stimulates the muscuepiploic: Gk. epiplein, to float on
laris to contract. Besides moving the material to the next haustrum, this contraction churns the material and exposes it to the
mucosa, where water and electrolytes are absorbed. As a result of
water absorption, the material becomes solid or semisolid, and is
now known as feces (fe'sez). Mass movement is a very strong
peristaltic wave, involving the action of the taeniae coli, which
moves the fecal material toward the rectum. Mass movements
generally occur only two or three times a day, usually during or
shortly after a meal. This response to eating, called the gastrocolic reflex, can best be observed in infants who have a bowel
movement during or shortly after feeding.
As material passes through the large intestine, Na+ K+, and
water are absorbed. It has been estimated that an average volume of
850 ml of water per day is absorbed across the mucosa of the colon.
The fecal material that is left then passes to the rectum, leading to
an increase in rectal pressure and the urge to defecate. If the urge to
defecate is denied, feces are prevented from entering the anal canal
by the external anal sphincter. In this case, the feces remain in the
rectum and may even back up into the sigmoid colon.
The defecation reflex normally occurs when the rectal
pressure rises to a particular level that is determined largely by
habit. At this point, the internal anal sphincter relaxes to admit
feces into the anal canal.
During the act of defecation, the longitudinal rectal muscles contract to increase rectal pressure, and the internal and external anal sphincter muscles relax. Excretion is aided by
contractions of abdominal and pelvic skeletal muscles, which
raise the intraabdominal pressure and help push the feces from
the rectum through the anal canal and out the anus.
The various mechanical activities of the GI tract are summarized in table 18.7.
CHAPTER 18
Finally, the large intestine has small but numerous fat-filled

pouches called epiploic (ep- -plo'ik) appendages (see
fig. 18.25) that are attached superficially to the taeniae coli.
The sympathetic innervation of the large intestine arises
from superior and inferior mesenteric plexuses, as well as from
the celiac plexus. The parasympathetic innervation arises from
the paired pelvic splanchnic and vagus nerves. Sensory fibers
from the large intestine respond to bowel pressure and signal the
need to defecate. Blood is supplied to the large intestine by
branches from the superior mesenteric and inferior mesenteric
arteries. Venous blood is returned through the superior and inferior mesenteric veins, which in turn drain into the hepatic portal
vein that enters the liver.

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Visceral peritoneum
Parietal peritoneum
Spinal cord
Vertebra
Plane of
section
Right kidney
Left kidney
Rib
Spleen
Small intestine
Waldrop
Liver
Large intestine
Inferior
vena cava
Stomach
Peritoneal cavity
Gallbladder
Aorta
Pancreas
FIGURE 18.30 A cross section of the abdomen showing the relative position of the liver to other abdominal organs.
Constipation occurs when fecal material accumulates because

of longer than normal periods between defecations. The
slower rate of elimination allows more time for water absorption, so
that the waste products become harder. Although uncomfortable and
sometimes painful, this condition is usually not serious. Diarrhea occurs when waste material passes too quickly through the colon, so
that insufficient time is allowed for water absorption. Excessive diarrhea can result in dangerous levels of dehydration and electrolyte imbalance, particularly in infants because of their small body size.
CHAPTER 18
Knowledge Check
20. Identify the four principal regions of the large intestine and
describe the functions of the colon.
21. Describe the haustra and the taeniae coli and explain their
role in the movements of the large intestine.
22. Describe the location of the rectum, anal canal, and
anal sphincter muscles and explain how defecation is
accomplished.
LIVER, GALLBLADDER,
AND PANCREAS
The liver, consisting of four lobes, processes nutrients and secretes bile, which is stored and concentrated in the gallbladder
prior to discharge into the duodenum. The pancreas, consisting of
endocrine (islet) cells and exocrine (acini) cells, secretes important hormones into the blood and essential digestive enzymes into
the duodenum.
Objective 17
Describe the location, structure, and functions
of the liver.
Objective 18
Describe the location of the gallbladder and

trace the flow of bile through the systems of ducts into the
duodenum.
Objective 19

of the pancreas.
Three accessory digestive organs in the abdominal cavity aid in

the chemical breakdown of food. These are the liver, gallbladder
and pancreas. The liver and pancreas function as exocrine glands
in this process because their secretions are transported to the
lumen of the GI tract via ducts.
Liver
The liver is the largest internal organ of the body, weighing about
1.3 kg (3.54.0 lbs) in an adult. It is positioned immediately beneath the diaphragm in the epigastric and right hypochondriac
regions (see fig. 2.15 and table 2.4) of the abdomen (fig. 18.30).
Its reddish brown color is due to its great vascularity.
The liver has four lobes and two supporting ligaments. Anteriorly, the right lobe is separated from the smaller left lobe by
the falciform ligament (fig. 18.31). Inferiorly, the caudate
falciform: L. falcis, sickle; forma, form


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Gallbladder
Inferior vena cava
Digestive System
661
Quadrate lobe
Right lobe
Coronary ligament
Ligamentum teres
Left lobe
Cystic
duct
Hepatic
duct
Hepatic
artery
Right lobe
Left lobe
Falciform
ligament
Hepatic
portal
vein
Ligamentum teres
Inferior vena cava
Gallbladder
Quadrate lobe
Falciform ligament
Caudate lobe
Common bile duct
Hepatic artery
Left lobe
Right lobe
Common bile
duct
Inferior vena
cava
Left
triangular
ligament
Hepatic portal
Caudate lobe
vein
Ligamentum
venosum
(c)
(The lobes of the liver are labeled in boldface type.)
(kaw'dt) lobe is positioned near the inferior vena cava, and the
quadrate lobe is adjacent to the gallbladder. The falciform ligament attaches the liver to the anterior abdominal wall and the
diaphragm. The ligamentum teres (round ligament) extends
from the falciform ligament to the umbilicus. This ligament is
the remnant of the umbilical vein of the fetus (see table 16.6).
Although the liver is the largest internal organ, it is, in a
sense, only one to two cells thick. This is because the liver cells,
or hepatocytes, form hepatic plates that are one to two cells
thick and separated from each other by large capillary spaces
called liver (hepatic) sinusoids (fig. 18.32). The sinusoids are
lined with phagocytic Kupffer (koop'fer) cells, but the large intercellular gaps between adjacent Kupffer cells make these sinusoids more highly permeable than other capillaries. The plate
structure of the liver and the high permeability of the sinusoids
allow each hepatocyte to be in direct contact with the blood.
The hepatic plates are arranged to form functional units
called liver lobules (fig. 18.33). In the middle of each lobule is a
central vein, and at the periphery of each lobule are branches of
the hepatic portal vein and of the hepatic artery, which open
into the spaces between hepatic plates. Portal venous blood,
containing molecules absorbed in the GI tract, thus mixes with
hepatic: Gk. hepatos, liver
Kupffer cells: from Karl Wilhelm von Kupffer, Bavarian anatomist and
embryologist, 18291902
CHAPTER 18
FIGURE 18.31 The liver and gallbladder. (a) An anterior view and (b) a rotated inferior view, and (c) a photograph of a rotated inferior view.

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Branch of hepatic portal vein

Hepatic triad
Biliary ductule
Branch of hepatic artery
Hepatic plate
Creek
Hepatic plate
Central vein
Liver (hepatic) sinusoid

Intralobular biliary ductule
Liver (hepatic)
sinusoids
(a)
Branch of hepatic
portal vein
Branch of
hepatic artery
Biliary ductule
Biliary canaliculi
Liver (hepatic)
sinusoid
(c)
Connective tissue
sheath
(b)
CHAPTER 18
FIGURE 18.32 A liver lobule and the histology of the liver. (a) A cross section of a liver lobule and (b) a longitudinal section. Blood enters a
liver lobule through the vessels in a hepatic triad, passes through hepatic sinusoids, and leaves the lobule through a central vein. The central
veins converge to form hepatic veins that transport venous blood from the liver. (c) A photomicrograph of a liver lobule in cross section.
arterial blood as the blood flows within the liver sinusoids from the
periphery of the lobule to the central vein (fig. 18.33). The central
veins of different liver lobules converge to form the hepatic vein,
which carries blood from the liver to the inferior vena cava.
The liver lobules have numerous functions, including synthesis, storage, and release of vitamins; synthesis, storage, and release of glycogen; synthesis of blood proteins; phagocytosis of old
red blood cells and certain bacteria; removal of toxic substances;
and production of bile. Bile is stored in the gallbladder and is
eventually secreted into the duodenum for the emulsification
(breaking down into smaller particles) and absorption of fats.
Bile is produced by the hepatocytes and secreted into tiny
channels called bile canaliculi (kan''a-lik'yu-li) located within each
hepatic plate (fig. 18.33). These bile canaliculi are drained at the
periphery of each lobule by bile ducts, which in turn drain into hepatic ducts that carry bile away from the liver. Because blood travels in the sinusoids and bile travels in the opposite direction within
the hepatic plates, blood and bile do not mix in the liver lobules.
The liver receives parasympathetic innervation from the
vagus nerves and sympathetic innervation from thoracolumbar
nerves through the celiac ganglia.
Gallbladder
The gallbladder is a saclike organ attached to the inferior surface
of the liver (figs. 18.31 and 18.34). This organ stores and concentrates bile. A sphincter valve at the neck of the gallbladder allows
a storage capacity of about 35 to 50 ml. The inner mucosal layer
of the gallbladder is thrown into folds similar to the gastric folds
of the stomach. When the gallbladder fills with bile, it expands
to the size and shape of a small pear. Bile is a yellowish green
fluid containing bile salts, bilirubin (a product resulting from the
breakdown of blood), cholesterol, and other compounds. Contraction of the muscularis ejects bile from the gallbladder.
Bile is continuously produced by the liver and drains
through the hepatic ducts and common bile duct to the duodenum. When the small intestine is empty of food, the sphincter of
ampulla (see fig. 18.21) constricts, and bile is forced up the cystic
duct to the gallbladder for storage.
The gallbladder is supplied with blood from the cystic
artery, which branches from the right hepatic artery. Venous
blood is returned through the cystic vein, which empties into the
hepatic portal vein. Autonomic innervation of the gallbladder is


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Digestive System
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Branch of portal vein

Liver (hepatic)
sinusoids
Bile canaliculus
Biliary ductule
Loechel
Central vein
Hepatic plate
FIGURE 18.33 The flow of blood and bile in a liver lobule. Blood flows within sinusoids from branches of the hepatic portal vein to the central
vein (from the periphery to the center of a lobule). Bile flows within hepatic plates from the center of a lobule to biliary ductules at the periphery.
similar to that of the liver; both receive parasympathetic innervation from the vagus nerves and sympathetic innervation from
thoracolumbar nerves through the celiac ganglia.
Pancreas
The soft, lobulated pancreas is known as a mixed gland because it
has both exocrine and endocrine functions. The endocrine function is performed by clusters of cells called the pancreatic islets
(islets of Langerhans). The islet cells secrete the hormones insulin and glucagon into the blood. As an exocrine gland, the pancreas secretes pancreatic juice through the pancreatic duct
(fig. 18.36), which empties into the duodenum.
FIGURE 18.34 The pancreatic duct joins the bile duct to empty its
secretions through the duodenal papilla into the duodenum. The release of bile and pancreatic juice into the duodenum is controlled by
the sphincter of ampulla (see fig. 18.21).
pancreas: Gk. pan, all; kreas, flesh

islets of Langerhans: from Paul Langerhans, German anatomist, 184788
CHAPTER 18
A common clinical problem of the gallbladder is the development of gallstones. Bile is composed of various salts, pigments, and cholesterols that become concentrated as water is
removed. Cholesterols normally remain in solution, but under certain
conditions they precipitate to form solid crystals. Large crystals may
block the bile duct and have to be surgically removed. The radiograph in figure 18.35 shows gallstones in position, and the photograph shows removed gallstones.

664
Unit 6

Body
The McGrawHill
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(a)
(b)
FIGURE 18.35 (a) A radiograph of a gallbladder that contains gallstones (biliary calculi). (b) Following surgical removal of the gallbladder
(cholecystectomy), it has been cut open to reveal its gallstones. (Note their size relative to that of a dime.)
CHAPTER 18
Pancreatic
acinar cells
Pancreatic
islet cells
Pancreatic
duct
Tail of pancreas
Body of pancreas
Head of pancreas
FIGURE 18.36 The pancreas is both an exocrine and an endocrine gland. Pancreatic juicethe exocrine productis secreted by acinar
cells into the pancreatic duct. Scattered islands of cells, called pancreatic islets (islets of Langerhans), secrete the hormones insulin and
glucagon into the blood.


Body
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The Digestive System
EXPLANATION
The entire digestive system develops from modifications of an
elongated tubular structure called the primitive gut. These modifications are initiated during the fourth week of embryonic development. The primitive gut is composed solely of endoderm and
for descriptive purposes can be divided into three regions: the
foregut, midgut, and hindgut (exhibit I).
Foregut
The stomodeum (sto''mo-de'um), or oral pit, is not part of the
foregut but an invagination of ectoderm that breaks through a thin
oral membrane to become continuous with the foregut and form
part of the oral cavity, or mouth. Structures in the mouth, therefore, are ectodermal in origin. The esophagus, pharynx, stomach, a
stomodeum: Gk. stoma, mouth; hodaios, on the way to
EXHIBIT I A sagittal section of a 5-week-old embryo showing the development of the digestive system and its association with the extraembryonic membranes and organs.
665


Body
The McGrawHill
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EXHIBIT II Progressive stages of development of the foregut to form the stomach, duodenum, liver, gallbladder, and pancreas:
(a) 4 weeks, (b) 5 weeks, (c) 6 weeks, and (d) 7 weeks.
portion of the duodenum, the pancreas, liver, and gallbladder are

the organs that develop from the foregut (exhibit II). Along the GI
tract, only the inside epithelial lining of the lumen is derived from
the endoderm of the primitive gut. The vascular portion and
smooth muscle layers are formed from mesoderm that develops
from the surrounding splanchnic mesenchyme.
The stomach first appears as an elongated dilation of the
foregut. The dorsal border of the stomach undergoes more rapid
growth than the ventral border, forming a distinct curvature. The
caudal portion of the foregut and the cranial portion of the
midgut form the duodenum. The liver and pancreas arise from
the wall of the duodenum as small hepatic and pancreatic buds,
respectively. The hepatic bud experiences incredible growth to
form the gallbladder, associated ducts, and the various lobes of
the liver (exhibit II). By the sixth week, the liver is carrying out
hemopoiesis (the formation of blood cells). By the ninth week,
666
the liver has developed to the point where it represents 10% of

the total weight of the fetus.
The pancreas develops from dorsal and ventral pancreatic
buds of endodermal cells. As the duodenum grows, it rotates
clockwise, and the two pancreatic buds fuse (exhibit II).
Midgut
During the fourth week of the embryonic stage (exhibit I), the
midgut is continuous with the yolk sac. By the fifth week, the
midgut has formed a ventral U-shaped midgut loop, which projects into the umbilical cord (exhibit III). As development continues, the anterior limb of the midgut loop coils to form most of
the small intestine. The posterior limb of the midgut loop expands to form the large intestine and a portion of the small intestine. A cecal diverticulum appears during the fifth week.

VII. Reproduction and

Development
21. Female Reproductive

System
The McGrawHill
Companies, 2001
Chapter 21
(a)
Female Reproductive System
749
(b)
FIGURE 21.25 (a) In mammography, the breast is placed alternately on a metal plate and radiographed from above and from the side. (b) A
mammogram of a patient with carcinoma of the breast. (Note the presence of a neoplasm indicated with an arrow.)
Methods of Contraception
CHAPTER 21
The rhythm method of birth control continues to be used by many

people, but the popularity of this technique has declined as more
successful methods of contraception have been introduced
(fig. 21.26). In the rhythm method, an attempt is made to predict the day of the womans ovulation and restrict coitus to safe
times of the cycle that allow no chance for fertilization to occur.
The day of ovulation can be determined by a rise in basal body
temperature or by a change in mucous discharge from the vagina.
This technique has one of the highest failure rates of any of the
widely used birth control methods, largely because womens cycles are often irregular.
More popular methods of birth control include sterilization,
oral contraceptives, intrauterine devices (IUDs), and barrier
methodsincluding condoms for the male and female and diaphragms for the female. All of these techniques are effective, but
they vary with respect to safety, side effects, and degree of efficacy.
Sterilization techniques include vasectomy for the male and
tubal ligation for the female. In the latter technique (which accounts for over 60% of sterilization procedures performed in the
United States), the uterine tubes are cut and tied. This is analogous to the procedure performed on the ductus deferentia in a
vasectomy. It prevents fertilization of the ovulated ovum.
About 10 million women in the United States and 60 million women in the world are currently using oral steroid contraceptives (the Pill). These contraceptives usually consist of a
synthetic estrogen combined with a synthetic progesterone in
the form of pills that are taken once each day for 3 weeks after
the last day of a menstrual period. This procedure causes an immediate increase in blood levels of ovarian steroids (from the
pill), which is maintained for the normal duration of a monthly
cycle. As a result of negative feedback inhibition of gonadotrophin
secretion, ovulation never occurs. The entire cycle is like a false
luteal phase, with high levels of progesterone and estrogen and
low levels of gonadotrophins.
Because the contraceptive pills contain ovarian steroid
hormones, the endometrium proliferates and becomes secretory,
just as it does during a normal cycle. In order to prevent an abnormal growth of the endometrium, women stop taking the pill
after 3 weeks. This causes estrogen and progesterone levels to
fall, and permits menstruation to occur. The contraceptive pill is
an extremely effective method of birth control, but it does have
potentially serious side effectsincluding an increased incidence
of thromboembolism and cardiovascular disorders. It has been
pointed out, however, that the mortality risk associated with
contraceptive pills is still much lower than the risk of death from
complications of pregnancyor from automobile accidents.

CHAPTER 21
750
Unit 7

Development

System
The McGrawHill
Companies, 2001
Reproduction and Development
(a)
(b)
(d)
(e)
(f)
(g)
(c)
(h)
FIGURE 21.26 Various types of birth control devices. (a) IUD, (b) contraceptive sponge, (c) diaphragm, (d ) birth control pills, (e) vaginal
spermicides, (f ) condom, (g) female condom, and (h) Norplant.


Development

System
The McGrawHill
Companies, 2001
Chapter 21
Another way in which to deliver hormonal contraceptives
to a womans body is by means of a subdermal implant. Implants
are 2-in. rods filled with a hormonal contraceptive drug. They
are implanted just under the skin, usually on the upper arm,
through a tiny incision. The contraceptive hormone gradually
leaches out through the walls of the rods and enters the bloodstream, preventing pregnancy for at least 5 years.
Intrauterine devices (IUDs) do not prevent ovulation,
but instead prevent implantation of the blastocyst in the uterine
wall in the event that fertilization occurs. The mechanisms by
which their contraceptive effects are produced are not well understood but appear to involve their ability to cause inflammatory reactions in the uterus. Uterine perforations are the
foremost complication associated with the use of IUDs. Because
of the potential problems with IUDs, their use has diminished.
Barrier contraceptivescondoms, diaphragms, and cervical capsare only slightly less effective than hormonal contraceptives or IUDs, but they do not have serious side effects.
Barrier contraceptives are most effective when they are used in
conjunction with spermicidal (sperm-killing) foams and gels.
Many couples avoid them, however, because they detract from
the spontaneity of sexual intercourse. Latex condoms offer an additional benefit; they provide some protection against sexually
transmitted diseases, including AIDS.
Several new chemical contraceptives are being developed
that should be soon available. These include hormonal patches,
hormonal IUDs, and injectible contraceptives. A hormonal patch
is intended to be placed on the skin directly over an ovary. De-
751
signed to resist moisture, the patch releases a weeks worth of estrogen and progestin. A hormonal IUD is a T-shaped structure
that fits into the uterine cavity where it slowly releases progestin
up to a five-year period. Improved monthly injectible contraceptives have fewer side effects than Depro-Provera, which has been
available since 1992.

An ectopic pregnancy is any pregnancy that implants outside the uterine
cavity. This is most likely to occur in the uterine tube (see fig. 21.24).
The events leading up to our patients problem are as follows: An oocyte
is extruded from the ovary during ovulation and received into the uterine tube. Soon after it enters the tube, the oocyte is fertilized, creating a
zygote. Up to this point, the events are no different from those that
occur in a normal pregnancy. In the case of tubal pregnancy, however,
the transporting ability of the uterine tube fails, causing the conceptus
to be retained in the tube. Implantation then occurs within tissues that
are not well suited for that purpose. For example, the uterine tube does
not expand well to accommodate a growing embryo, nor does it possess
the necessary epithelium and glandular structures as does the endometrium of the uterus. The result is overexpansion and erosion of the
tubal wall, leading to rupture and hemorrhage. Because the uterine tube
is basically exposed to the peritoneal cavity, blood from the site of rupture can flow into and collect in the rectouterine pouch (pouch of Douglas or Douglas cul-de-sac), which lies posterior to the vaginal fornix.
There, it is easily aspirated by a needle placed through the vaginal wall.
QUESTIONS
1. These adnexal masses are consistent
with endometriomas, foci of
endometrial tissue outside the uterus on
the ovaries. How would these masses

cause the patients symptoms?
2. How does endometrial tissue get to the
ovaries?
3. Endometrial implants can be found
throughout the peritoneal space from
direct spread and have even been
reported in the pleural space.
Remembering that these implants
cause irritation of, as well as, scarring
around the organs on which they
are implanted, what symptoms would
the patient have if implants were found
on the following organs: (a) rectum,
(b) urinary bladder, (c) stomach, or
(d) diaphragm?
A
U
CHAPTER 21
A 32-year-old female comes to you because

she and her husband have been unsuccessfully attempting to become pregnant for
more than a year. In questioning her, you
find that she has been suffering from vague
pelvic pain during her periods for many
years. She also reports occasional pain during intercourse over that same time. On
pelvic exam, you find bilateral tender adnexal masses, flanking a normal-sized uterus.
An MRI of the pelvis confirms these findings. [uterus (U), adnexal masses (A)]

752
Unit 7

Development

System
The McGrawHill
Companies, 2001
Chapter Summary
Introduction to the Female Reproductive
System (pp. 726728)
1. The reproductive period of a female is the
period between puberty (about age 12)
and menopause (about age 50). In the
course of this age span, cyclic ovulation
and menstruation patterns occur in
nonpregnant females.
2. The functions of the female reproductive
system are to produce ova; secrete sex
hormones; receive sperm from the male;
provide sites for fertilization,
implantation, and development of the
embryo and fetus; facilitate parturition;
and secrete milk from the mammary
glands.
3. The female reproductive system consists
of (a) primary sex organsthe ovaries;
(b) secondary sex organsthose that are
essential for sexual reproduction,
characterized by latent development; and
(c) secondary sex characteristics
features that are sexual attractants,
expressed after puberty.
Structure and Function of the Ovaries

(pp. 728732)
CHAPTER 21
1. The ovaries are supported by the

mesovarium, which extends from the
broad ligament, and by the ovarian and
suspensory ligaments.
2. The ovarian follicles within the ovarian
cortex undergo cyclic changes.
(a) Primary oocytes, arrested at prophase
I of meiosis, are contained within
primordial follicles.
(b) Upon stimulation by gonadotropic
hormones, some of the primordial
follicles enlarge to become primary
follicles.
(c) When a follicle develops a fluid-filled
antrum, it is called a secondary
follicle.
(d) Generally only one follicle continues
to grow to become a vesicular ovarian
follicle.
(e) The vesicular ovarian follicle

contains a secondary oocyte, arrested
at metaphase II of meiosis.
(f) In the process of ovulation, the
vesicular ovarian follicle ruptures and
releases its secondary oocyte, which
becomes a zygote upon fertilization.
(g) After ovulation, the empty follicle
becomes a corpus luteum.
Secondary Sex Organs (pp. 732738)

1. The uterine tube, which conveys ova
from the ovary to the uterus, provides a
site for fertilization.
(a) The open-ended portion of each
uterine tube is expanded; its margin
bears fimbriae that extend over the
lateral surface of the ovary.
(b) Movement of an ovum is aided by
ciliated cells that line the lumen and
by peristaltic contractions in the wall
of the uterine tube.
2. The uterus is supported by the broad
ligaments, uterosacral ligaments, cardinal
ligaments, and round ligaments. The
regions of uterus are the fundus, body, and
cervix. The cervical canal opens into the
vagina at the uterine ostium.
(a) The endometrium consists of a
stratum basale and a stratum
functionale; the superficial stratum
functionale is shed during
menstruation.
(b) The myometrium produces the
muscular contractions needed for
labor and parturition.
3. The vagina serves to receive the erect
penis during coitus, to convey menses to
the outside during menstruation, and to
transport the fetus during parturition.
(a) The vaginal wall is composed of an
inner mucosa, a middle muscularis,
and an outer fibrous layer.
(b) The vaginal orifice may be partially
covered by a thin membranous
hymen.
4. The external genitalia, or vulva, include

the mons pubis, labia majora and minora,
clitoris, vaginal vestibule, vestibular
bulbs, and vestibular glands.
5. Impulses through parasympathetic nerves
stimulate erectile tissues in the clitoris
and vestibular bulbs; in orgasm, muscular
contraction occurs in the perineum,
uterus, and uterine tubes.
Mammary Glands (pp. 738740)

1. Mammary glands, located within the
breasts, are modified sweat glands.
(a) Each mammary gland is composed of
15 to 20 lobes; the lobes are
subdivided into lobules that contain
mammary alveoli.
(b) During lactation, the mammary
alveoli secrete milk. The milk passes
through mammary ducts, lactiferous
ducts, and lactiferous sinuses and is
discharged through the nipple.
2. The nipple is a cylindrical projection near
the center of the breast, surrounded by
the circular pigmented areola.
Ovulation and Menstruation

(pp. 740742)
1. Ovulation and menstruation are
reproductive cyclic events that are
regulated by hormones secreted by the
hypothalamus, the anterior pituitary, and
the ovaries.
2. The menstrual cycle is divided into
menstrual, proliferative, and secretory
phases.
3. The principal hormones that regulate
ovulation and menstruation are estrogen,
progesterone, follicle-stimulating
hormone (FSH), and luteinizing
hormone (LH).
Review Activities
Objective Questions
1. The cervix is a portion of
(a) the vulva.
(b) the vagina.
(c) the uterus.
2. Fertilization normally occurs in

(a) the ovary.
(b) the uterine tube.
(c) the uterus.
(d) the vagina.
3. The secretory phase of the endometrium

corresponds to which of the following
ovarian phases?
(a) the follicular phase
(b) ovulation


Development

System
The McGrawHill
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Chapter 21
4.
5.
6.
7.
8.
9.
(c) the luteal phase

(d) the menstrual phase
Which of the following statements about
oogenesis is true?
(a) Oogonia, like spermatogonia, form
continuously during postnatal life.
(b) Primary oocytes are haploid.
(c) Meiosis is completed prior to
ovulation.
(d) During ovulation, a secondary oocyte
is released from a vesicular ovarian
follicle.
The function of the mesovarium is
(a) movement of the sperm to the ova.
(b) nourishment of the ovarian walls.
(c) muscular contraction of the uterus.
(d) suspension of the ovary.
Which of the following layers is shed as
menses?
(a) the perimetrial layer
(b) the fibrous layer
(c) the functionalis layer
(d) the menstrual layer
The transverse folds in the mucosal layer
of the vagina are called
(a) perineal folds. (c) fornices.
(b) vaginal rugae. (d) labia gyri.
Which of the following is not a part of the
vulva?
(a) the mons pubis
(b) the clitoris
(c) the vaginal vestibule
(d) the vagina
(e) the labia minora
The paramesonephric (mllerian) ducts
give rise to
(a) the uterine tubes.
(b) the uterus.
(c) the pudendum.
(d) both a and b.
(e) both b and c.
10. In a female, the homologue of the male

scrotum is/are
(a) the labia majora.
(b) the labia minora.
(c) the clitoris.
(d) the vestibule.
Essay Questions
1. Define puberty, ovulation, menstruation,
and menopause.
2. Describe the follicular changes within the
ovarian cortex during the events that
precede and follow ovulation.
3. Define oogenesis. When is the process
initiated and when is it completed?
4. Describe the gross and histologic structure
of the uterus and comment on the
significance of the two endometrial layers.
5. Identify the secondary sex organs and
explain their functions.
6. Summarize the events of a menstrual
cycle and explain the roles of estrogen
and progesterone.
7. List the functions of the vagina and
describe its structure.
8. Distinguish between the labia majora and
labia minora, between the pudendal cleft
and vaginal vestibule, and between the
vestibular bulbs and vestibular glands.
9. List the events that cause an erection of
the female genitalia, and explain the
process of orgasm.
10. Describe the structure and position of
the mammary glands in the breasts.
11. List the homologous reproductive organs
of the male and female reproductive
systems and note the undifferentiated
structures from which they develop.
12. Define gynecology, obstetrics, speculum,
laparoscopy, breast self-examination, and
Pap smear.
753
13. Distinguish between normal, primary, and

secondary amenorrhea. What causes
each?
14. List the various kinds of uterine
neoplasms. How is cancer of the uterus
generally detected?
15. Distinguish between dysplasia,
fibroadenoma, and carcinoma of the
breast.
1. Why is the hormonal regulation of the
reproductive system much more complex
in females than it is in males?
2. Your 14-year-old daughter is serious about
her gymnastics lessons and exercises
rigorously for at least 2 hours every day.
She has not yet had her first menstrual
period and is beginning to get worried.
How would you reassure her?
3. A wide variety of commercial douche
preparations and scented aerosols can be
purchased in drug stores and supermarkets
as feminine hygiene products. Why
should women avoid using them?
4. Both sterility and impotence are sexual
dysfunctions in men. Only one of these is
associated with women, however.
Explain.
5. Contraceptive pills trick the brain into
thinking youre pregnant. Explain what
is meant by this statement.
6. With respect to homologous structures,
explain why men have nipples? Why do
mammary glands normally function only in
females?
7. If a woman is on estrogen replacement
therapy following menopause, why is it
important to monitor her uterine lining?

CHAPTER 21


Development
22
22. Developmental
Anatomy, Postnatal
Growth, and Inheritance
The McGrawHill
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Developmental Anatomy,
Postnatal Growth,
and Inheritance
Fertilization 755
Preembryonic Period 757
Embryonic Period 762
Fetal Period 772
Labor and Parturition 775
Periods of Postnatal Growth 775
Inheritance 782

Genetic Disorders
of Clinical Importance 790
Chapter Summary 791
Clinical Case Study

A 27-year-old woman gave birth to twin boys, followed by an apparently single placenta. After
examining the two infants, the pediatrician informed the mother that one of them had a cleft
palate but that the other was normal. She added that cleft palate could be hereditary and asked
if any family members had the problem. The mother said that she knew of none. Further examination of the placenta revealed two amnions and only one chorion.
Does the presence of two amnions and one chorion indicate monozygotic or dizygotic
twins? How would you account for the fact that one baby has a cleft palate, whereas the other
does not?
Hints: Read the section on multiple pregnancy at the end of the chapter and carefully study
figure 22.37. Can any conclusions be drawn regarding genetic similarities between the two infants? Note that identical twins are always the same gender.
FIGURE: A cleft lip and an accompanying

cleft palate are congenital disorders in which
one or two sides of the upper lip and hard
palate fail to fuse. Cleft lips and cleft palates
can be treated very effectively with surgery.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
FERTILIZATION
Upon fertilization of an ovum by a spermatozoon in the uterine tube,
meiotic development is completed and a diploid zygote is formed.
Objective 1
Define fertilization, capacitation, and

morphogenesis.
Objective 2
Describe the changes that occur in the

spermatozoon and ovum prior to, during, and immediately
following fertilization.
Fertilization refers to the penetration of an ovum (egg) by a spermatozoon (fig. 22.1), with the subsequent union of their genetic
material. It is this event that determines a persons gender (see
p. 716) and biological inheritance. Fertilization cannot occur,
however, unless certain conditions are met. First of all, an ovum
must be present in the uterine tubeit can be there for at most
24 hours before it becomes incapable of undergoing fertilization.
Second, large numbers of spermatozoa must be ejaculated to ensure fertilization. Although a recent study has shown that sperm
The McGrawHill
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Developmental Anatomy, Postnatal Growth, and Inheritance
755
cells can remain viable 5 days after ejaculation, this still leaves a
window of fertility of only 6 days each monththe day of ovulation and the 5 days leading up to it.
As described in chapter 21, a woman usually ovulates one
ovum a month, totaling about 400 during her reproductive years.
Each ovulation releases an ovum that is actually a secondary
oocyte arrested at metaphase of the second meiotic division. An
ovum is surrounded by a thin layer of protein and polysaccharides, called the zona pellucida, and a layer of granulosa cells,
called the corona radiata (fig. 22.1). These layers provide a protective shield around the ovum as it enters the uterine tube.
During coitus, a male ejaculates between 100 million and
500 million spermatozoa into the females vagina. This tremendous number is needed because of the high fatality rateonly
about 100 sperm cells survive to encounter the ovum in the uterine tube. In addition to deformed sperm cellsup to 20% in the
zona pellucida: L. zone, a girdle; pellis, skin

corona radiata: Gk. korone, crown; radiata, radiate
Corona radiata
First polar body
Zona pellucida
Second meiotic spindle
Oocyte membrane
Cytoplasm
(a)
(c)
Nucleus
containing
chromosome
Acrosome
containing
enzymes
Perforations
in acrosome wall
CHAPTER 22
1
3
(b)
Creek
FIGURE 22.1 The process of fertilization. (a,b) As the head of the sperm encounters the corona radiata of the oocyte (2), digestive enzymes
are released from the acrosome (3, 4), clearing a path to the oocyte membrane. When membrane of the sperm contacts the oocyte membrane
(5), the membranes become continuous, and the nucleus and other contents of the sperm move into the egg cytoplasm of the oocyte. (c) A
scanning electron micrograph of a sperm cell bound to the surface of an oocyte.

756
Unit 7

Development
22. Developmental
Anatomy, Postnatal
Cell
membrane
Acrosomal
cap
Nucleus
Basal
plate
FIGURE 22.2 A transmission electron micrograph showing the

head of a human spermatozoon with its nucleus and acrosome.
average fertile maleanother 25% will perish as soon as they contact

the vaginas acidic environment. Still others are destroyed by the
womans immune cells, which recognize them as foreign cells. Even if
they manage to reach the uterine ostium (see fig. 21.12), many sperm
cells will get stuck there and never make it through the uterus.
If it finally encounters an ovum, the spermatozoon must
penetrate the protective corona radiata and zona pellucida for
fertilization to occur. To do this, the head of each sperm cell is
capped by an organelle called an acrosome (ak'ro-som) (figs. 22.1
and 22.2). The acrosome contains a trypsinlike proteindigesting enzyme and hyaluronidase (hi''a-loo-ron'-das), which
digests hyaluronic acid, an important constituent of connective
tissue. When a spermatozoon meets an ovum in the uterine
tube, an acrosomal reaction allows the spermatozoon to penetrate the corona radiata and the zona pellucida. A spermatozoon that comes along relatively lateafter many others
have undergone acrosomal reactions to expose the ovum
membraneis more likely to be the one that finally achieves
penetration of the egg.
CHAPTER 22
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Experiments confirm that freshly ejaculated spermatozoa are

infertile and must be in the female reproductive tract for at
least 7 hours before they can fertilize a secondary oocyte. Their
membranes must become fragile enough to permit the release of the
acrosomal enzymesa process called capacitation. During in vitro
fertilization, capacitation is induced artificially by treating the ejaculate with a solution of gamma globulin, free serum, follicular fluid,
dextran, serum dialysate, and adrenal gland extract to chemically
mimic the conditions of the female reproductive tract.
As soon as a spermatozoon penetrates the zona pellucida, a rapid chemical change in this layer prevents other
spermatozoa from attaching to it. Therefore, only one spermatozoon is permitted to fertilize an oocyte. With the penetration of a single spermatozoon through its cell membrane, the
oocyte is stimulated to complete its second meiotic division
(fig. 22.3). Like the first meiotic division, the second produces one cell that contains most of the cytoplasm and one
polar body. The cell containing the cytoplasm is the mature
ovum, and the second polar body, like the first, ultimately
fragments and disintegrates.
At fertilization, the head of the sperm cell enters the cytoplasm of the much larger ovum. Within 12 hours, the nuclear
membranes in both the sperm cell and ovum disappear, and the
haploid number of chromosomes (23) in the ovum is joined by
the haploid number of chromosomes from the spermatozoon. A
fertilized egg, or zygote (zi'got), containing the diploid number of
chromosomes (46) is thus formed.
Within hours after conception, the structure of the body
begins to form from this single fertilized egg, culminating some
38 weeks later with the birth of a baby. The transformation involved in the growth and differentiation of cells and tissues is
known as morphogenesis (mor''fo-jen'-sis), and it is through
this awesome process that the organs and systems of the body
are established in a functional relationship. Moreover, there are
sensitive periods of morphogenesis for each organ and system,
during which genetic or environmental factors may affect normal development.
Prenatal development can be divided into a preembryonic
period, which is initiated by the fertilization of an ovum; an embryonic period, during which the bodys organ systems are
formed; and a fetal period, which culminates in parturition
(childbirth).
Knowledge Check
1. Explain why capacitation and the acrosomal reaction of
spermatozoa are necessary to accomplish fertilization of a
secondary oocyte.
2. Discuss the changes that occur in a spermatozoon from the
time of ejaculation to the time of fertilization. What
changes occur in a secondary oocyte following ovulation to
the time of fertilization?
3. Define morphogenesis. When does this process begin? What
does it accomplish?
haploid: Gk. haplous, single; L. ploideus, multiple in form

acrosome: Gk. akron, extremity; soma, body
capacitation: L. capacitas, capable of
zygote: Gk. zygotos, yolked, joined

diploid: Gk. diplous, double; L. ploideus, multiple in form
morphogenesis: Gk. morphe, form; genesis, beginning


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
Primary oocyte
Secondary oocyte
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Secondary oocyte
at metaphase II
First
polar
body
First
polar
body
First
meiotic
division
Spindle
apparatus
Ovulation
First polar
body
757
Second
polar body
Fertilization
Nucleus
of ovum
Sperm cell
inside ovum
Chromosomes
Zygote
Nuclear membrane
disappearing
Sperm cell
nucleus
FIGURE 22.3 A secondary oocyte, arrested at metaphase II of meiosis, is released at ovulation. If this cell is fertilized, it will become a mature
ovum, complete its second meiotic division, and produce a second polar body. The fertilized ovum is known as a zygote.
PREEMBRYONIC PERIOD
The events of the 2-week preembryonic period include fertilization, transportation of the zygote through the uterine tube, mitotic divisions, implantation, and the formation of primordial
embryonic tissue.
Objective 3
Describe the events of preembryonic

development that result in the formation of the blastocyst.
Objective 4
Discuss the role of the trophoblast in the

implantation and development of the placenta.
Objective 5
Explain how the primary germ layers develop

and list the structures produced by each layer.
cells. The morula floats freely in the uterine cavity for about
3 days. During this time, the center of the morula fills with fluid
passing in from the uterine cavity. As the fluid-filled space develops within the morula, two distinct groups of cells form, and the
structure becomes known as a blastocyst (blas'to-sist) (fig. 22.4).
The hollow, fluid-filled center of the blastocyst is called the blastocyst cavity. The blastocyst is composed of an outer layer of
cells, known as the trophoblast, and an inner aggregation of cells,
called the embryoblast (internal cell mass) (see fig. 22.6). With
further development, the trophoblast differentiates into a structure called the chorion (kor'e-on), which will later become a portion of the placenta. The embryoblast will become the embryo. A
diagrammatic summary of the ovarian cycle, fertilization, and the
morphogenic events of the first week is presented in figure 22.5.
Objective 6
Define gestation and explain how the

parturition date is determined.
Cleavage and Formation of the Blastocyst
morula: L. morus, mulberry
The process of implantation begins between the fifth and seventh day following fertilization. This is the process by which the
blastocyst embeds itself into the endometrium of the uterine wall
(fig. 22.6a). Implantation is made possible by the secretion of
proteolytic enzymes by the trophoblast, which digest a portion of
the endometrium. The blastula sinks into the depression, and endometrial cells move back to cover the defect in the wall. At the
same time, the part of the uterine wall below the implanting
blastocyst: Gk. blastos, germ; kystis, bladder

chorion: Gk. chorion, membrane
implantation: L. im, in; planto, to plant
CHAPTER 22
Within 30 hours following fertilization, the zygote undergoes a

mitotic division called cleavage. This first division results in the
formation of two identical daughter cells called blastomeres
(fig. 22.4). Additional cleavages occur as the structure passes
down the uterine tube and enters the uterus on about the third
day. It is now composed of a ball of 16 or more cells called a
morula (mor'yu-la). Although the morula has undergone several
mitotic divisions, it is not much larger than the zygote because no
additional nutrients necessary for growth have been entering the
Implantation

758
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Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
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Polar body
Zona pellucida
Blastomere
2-cell stage
4-cell stage
8-cell stage
Embryoblast
Degenerating
zona pellucida
Blastocyst cavity
Trophoblast
Morula
Early blastocyst
Late blastocyst
FIGURE 22.4 Sequential illustrations from the first cleavage of the zygote to the formation of the blastocyst. (Note the deterioration of the zona
pellucida in the early blastocyst.)
Cleavage
(30 hours)
Zygote
Egg nucleus
Morula
(72 hours)
8-cell
stage
4-cell
stage
2-cell
stage
Sperm nucleus
Sperm
cells
Ovary
CHAPTER 22
Fertilization
Blastocyst
(4 days)
Corpus
luteum
Maturing
follicle
Ovulation
Secondary
oocyte
Implanted
blastocyst
(6 days)
FIGURE 22.5 A diagram of the ovarian cycle, fertilization, and the events of the first week. Implantation of the blastocyst begins between the
fifth and seventh day, and is generally completed by the tenth day.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
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759
Blastocoel
Uterine cavity
Trophoblast
Embryonic endoderm
Embryoblast
Cytotrophoblasts
Blastocyst
cavity
Syncytiotrophoblasts
Embryoblast
Endometrium
Trophoblast
Endometrial
epithelium
Endometrial
capillary
Endometrial
gland
Glandular
secretion
Endometrial
storma
(a)
Creek
(b)
FIGURE 22.6 The blastocyst adheres to the endometrium on about the sixth day as seen in (a), a photomicrograph. By the seventh day (b),
specialized syncytiotrophoblasts from the trophoblast have begun to invade the endometrium. The syncytiotrophoblasts secrete human chorionic
gonadotropin (hCG) to sustain pregnancy and will eventually participate in the formation of the placenta for embryonic and fetal sustenance.
All pregnancy tests assay for the presence of hCG in the

blood or urine because this hormone is secreted only by the
blastocyst. Because there is no other source of hCG, the presence of
this hormone confirms a pregnancy. Modern pregnancy tests detect
the presence of hCG by use of antibodies against hCG or by the use
of cellular receptor proteins for hCG.
Human chorionic
gonadotropin
Estrogen
Progesterone
FIGURE 22.7 Human chorionic gonadotropin (hCG) is secreted

by syncytiotrophoblasts during the first trimester of pregnancy. This
pituitary-like hormone maintains the mothers corpus luteum for the
first 512 weeks of pregnancy. The placenta then assumes the role of
estrogen and progesterone production, and the corpus luteum degenerates.
CHAPTER 22
blastocyst thickens, and specialized cells of the trophoblast

produce fingerlike projections, called syncytiotrophoblasts
(sin-sit''e-o-trof'o-blasts), into the thickened area. The syncytiotrophoblasts arise from a specific portion of the trophoblast called the cytotrophoblast (fig. 22.6b), located next to
the embryoblast.
The blastocyst saves itself from being aborted by secreting
a hormone that indirectly prevents menstruation. Even before
the sixth day when implantation begins, the syncytiotrophoblasts
secrete human chorionic gonadotropin (kor''e-on-ik go-nad-otro'pin) (hCG). This hormone is identical to LH in its effects,
and therefore is able to maintain the corpus luteum past the time
when it would otherwise regress. The secretion of estrogen and
progesterone is maintained, and menstruation is normally prevented (fig. 22.7).
The secretion of hCG declines by the tenth week of pregnancy. Actually, this hormone is required only for the first 5 to
6 weeks of pregnancy because the placenta itself becomes an active steroid-secreting gland by this time. At the fifth to sixth
week, the mothers corpus luteum begins to regress (even in the
presence of hCG), but the placenta secretes more than sufficient
amounts of steroids to maintain the endometrium and prevent
menstruation.

760
Unit 7

Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
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Chorion
Ectoderm
Germ
layers
Amnion
Mesoderm
Amniotic cavity
Endoderm
Body stalk
(umbilical cord)
Yolk sac
of embryo
Chorionic villi
FIGURE 22.8 The completion of implantation occurs as the primary germ layers develop at the end of the second week.
TABLE 22.1 Summary of Preembryonic Development

Morphogenic Stage
Time Period
Principal Events
Zygote
24 to 30 hours following
ovulation
Egg is fertilized; zygote has 23 pairs of chromosomes (diploid) from haploid sperm and haploid
egg and is genetically unique
Cleavage
30 hours to third day
Mitotic divisions produce increased number of cells
Morula
Third to fourth day
Solid ball-like structure forms, composed of 16 or more cells
Blastocyst
Fifth day to end of second week
Hollow ball-like structure forms, a single layer thick; embryoblast and trophoblast form;
implantation occurs; embryonic disc forms, followed by primary germ layers
CHAPTER 22
Formation of the Germ Layers

week of development, the embryoblast undergoes marked differentiation. A slitlike space called the amniotic (am''ne-ot'ik)
cavity forms between the embryoblast and the trophoblast
(fig. 22.8). The embryoblast flattens into the embryonic disc (see
fig. 22.10), which consists of two layers: an upper ectoderm, which
is closer to the amniotic cavity, and a lower endoderm, which borders the blastocyst cavity. A short time later, a third layer called
the mesoderm forms between the endoderm and ectoderm. These
three layers constitute the primary germ layers (fig. 22.8). Once
they are formed, at the end of the second week, the preembryonic
period is completed and the embryonic period begins.
The primary germ layers are especially significant because all of

the cells and tissues of the body are derived from them. Ectodermal
cells form the nervous system; the outer layer of skin (epidermis), including hair, nails, and skin glands; and portions of the sensory organs. Mesodermal cells form the skeleton, muscles, blood,
reproductive organs, dermis of the skin, and connective tissue. Endodermal cells produce the lining of the GI tract, the digestive organs,
the respiratory tract and lungs, and the urinary bladder and urethra.
The events of the preembryonic period are summarized in
table 22.1. Refer to figure 22.9 for an illustration and a listing of the organs and body systems that derive from each of the primary germ layers.
The period of prenatal development is referred to as gestation.
Normal gestation for humans is 9 months. Knowing this and the
pattern of menstruation make it possible to determine the babys delivery date. In a typical reproductive cycle, a woman ovulates 14 days
prior to the onset of the next menstruation and is fertile for approximately 20 to 24 hours following ovulation. Adding 9 months, or
38 weeks, to the time of ovulation gives one the estimated delivery date.

gestation: L. gestatus, to bear


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
Amnion
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Digestive tract
Amniotic fluid
Skin
Heart
Spinal cord
Chorion
Body stalk with

umbilical vessels
Brain
Nels
on
Yolk sac
Endoderm
Mesoderm
Ectoderm
FIGURE 22.9 The body systems and the primary germ layers from which they develop.
CHAPTER 22
Epidermis of skin and epidermal

derivatives: hair, nails, glands
of the skin; linings of oral, nasal,
anal, and vaginal cavities
Pituitary gland
Adrenal medulla
Muscle: smooth, cardiac,

and skeletal
Connective tissue: embryonic,
connective tissue proper,
cartilage, bone, blood
Epithelium of blood vessels,
lymphatic vessels, body
cavities, joint cavities
Kidneys and ureters
Adrenal cortex
Epithelium of pharynx, external
acoustic canal, tonsils, thyroid,
parathyroid, thymus, larynx,
trachea, lungs, GI tract, urinary
bladder and urethra, and vagina
Liver and pancreas

762
Unit 7

Development
22. Developmental
Anatomy, Postnatal
Knowledge Check
4. List the structural characteristics of a zygote, morula, and
blastocyst. Approximately when do each of these stages of
the preembryonic period of development occur?
5. Discuss the process of implantation and describe the physiological events that ensure pregnancy.
6. Describe the development of the placenta.
7. List the major structures that derive from each germ layer.
8. Define gestation. What is the average gestation time for humans? How is the parturition date determined?
EMBRYONIC PERIOD
The events of the 6-week embryonic period include the differentiation of the germ layers into specific body organs and the formation
of the placenta, the umbilical cord, and the extraembryonic membranes. Through these morphogenic events, the needs of the embryo are met.
Objective 7
Define embryo and describe the major events

of the embryonic period of development.
Objective 8
List the embryonic needs that must be met to

avoid a spontaneous abortion.
Objective 9
Describe the structure and function of each of

the extraembryonic membranes.
Objective 10
Describe the development and functions of

the placenta and the umbilical cord.
During the embryonic periodfrom the beginning of the third

week to the end of the eighth weekthe developing organism is
correctly called an embryo. It is at this period that all of the
body tissues and organs form, as well as the placenta, umbilical
cord, and extraembryonic membranes. The term conceptus refers
to the embryo, or to the fetus later on, and all of the extraembryonic structuresthe products of conception.
CHAPTER 22
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Embryology is the study of the sequential changes in an organism as the various tissues, organs, and systems develop.
Chick embryos are frequently studied because of the easy access
through the shell and their rapid development. Mice and pig embryos
are also extensively studied as mammalian models. Genetic manipulation, induction of drugs, exposure to disease, radioactive tagging or
dyeing of developing tissues, and X-ray treatments are some of the
commonly conducted experiments that provide information that can
be applied to human development and birth defects.
During the preembryonic period of cell division and differentiation, the developing structure is self-sustaining. The embryo, however, must derive sustenance from the mother. For
morphogenesis to continue, certain immediate needs must be
met. These needs include (1) formation of a vascular association
between the uterus of the mother and the embryo so that nutrients and oxygen can be provided and metabolic wastes and car-
bon dioxide can be removed; (2) establishment of a constant,

protective environment around the embryo that is conducive to
development; (3) establishment of a structural foundation for
embryonic morphogenesis along a longitudinal axis; (4) provision for structural support for the embryo, both internally and
externally; and (5) coordination of the morphogenic events
through genetic expression. If these needs are not met, a spontaneous abortion will generally occur.
The first and second of these needs are provided for by extraembryonic structures; the last three are provided for intraembryonically. The extraembryonic membranes, the placenta, and
the umbilical cord will be considered separately, prior to a discussion of the development of the embryo.
Serious developmental defects usually cause the embryo to
be naturally aborted. About 25% of early aborted embryos
have chromosomal abnormalities. Other abortions may be caused by
environmental factors, such as infectious agents or teratogenic
drugs (drugs that cause birth defects). In addition, an implanted embryo is regarded as foreign tissue by the immune system of the
mother, and is rejected and aborted unless maternal immune responses are suppressed.
Extraembryonic Membranes
At the same time that the internal organs of the embryo are
being formed, a complex system of extraembryonic membranes
is also developing (fig. 22.10). The extraembryonic membranes
are the amnion, yolk sac, allantois, and chorion. These membranes
are responsible for the protection, respiration, excretion, and
nutrition of the embryo and subsequent fetus. At parturition,
the placenta, umbilical cord, and extraembryonic membranes
separate from the fetus and are expelled from the uterus as the
afterbirth.
Amnion
The amnion (am'ne-on) is a thin membrane, derived from ectoderm and mesoderm. It loosely envelops the embryo, forming an
amniotic sac that is filled with amniotic fluid (fig. 22.11b). In
later fetal development, the amnion expands to come in contact
with the chorion. The development of the amnion is initiated
early in the embryonic period, at which time its margin is attached to the free edge of the embryonic disc (see fig. 22.10). As
the amniotic sac enlarges during the late embryonic period (at
about 8 weeks), the amnion gradually sheaths the developing
umbilical cord with an epithelial covering (fig. 22.12).
The buoyant amniotic fluid performs several functions for
the embryo and subsequent fetus.
It ensures symmetrical structural development and growth.
It cushions and protects by absorbing jolts that the mother
may receive.
It helps maintain consistent pressure and temperature.
It helps eliminate metabolic wastes.
It permits freedom of fetal movement, which is important
for skeletomuscular development and blood flow.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
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763
Amniotic
fluid
Embryonic
disc
Amnion
Allantois
Implanted
embryo
Yolk sac
Body of
uterus
Chorion
(a)
Amnion
(b) Connecting
stalk
(a)
Umbilical cord
Placenta
Yolk sac
Allantois
Chorion
frondosum
(c)
Villi of chorion
frondosum
Creek
FIGURE 22.10 The formation of the extraembryonic membranes
Chorion
Amnion
during a single week of rapid embryonic development. (a) At

3 weeks, (b) at 312 weeks, and (c) at 4 weeks.
Amniotic sac
containing
amniotic fluid
Amniotic fluid is formed initially as an isotonic fluid absorbed from the maternal blood in the endometrium surrounding
the developing embryo. Later, the volume is increased and the
concentration changed by urine excreted from the fetus into the
amniotic sac. Amniotic fluid also contains cells that are sloughed
off from the fetus, placenta, and amniotic sac. Because all of these
cells are derived from the same fertilized egg, all have the same
genetic composition. Many genetic abnormalities can be detected
by aspirating a sample of this fluid and examining the cells obtained, in a procedure called amniocentesis (am''ne-o-sen-te'sis).
Amniotic fluid is normally swallowed by the fetus and absorbed in the GI tract. The fluid enters the fetal blood, and the
waste products it contains enter the maternal blood in the plaDown syndrome: from John L. H. Down, English physician, 182896
(b)
Umbilical blood vessels
FIGURE 22.11 An implanted embryo at approximately 412 weeks.

(a) The interior of a uterus showing the implantation site. (b) The developing embryo, extraembryonic membranes, and placenta.
centa. Prior to delivery, the amnion is naturally or surgically ruptured, and the amniotic fluid (bag of waters) is released.
As the fetus grows, the amount of amniotic fluid increases.
It is also continually absorbed and renewed. For the near-term
baby, almost 8 liters of fluid are completely replaced each day.
Yolk Sac
The yolk sac is established during the end of the second week as
cells from the trophoblast form a thin exocoelomic (ek''so-selo'mik) membrane. Unlike the yolk sac of many vertebrates, the
CHAPTER 22
Amniocentesis (fig. 22.13) is usually performed at the fourteenth or fifteenth week of pregnancy, when the amniotic sac
contains 175225 ml of fluid. Genetic diseases, such as Down
syndrome, or trisomy 21 (in which there are three instead of two
number21 chromosomes), can be detected by examining chromosomes. Diseases such as Tay-Sachs disease, in which there is a defective enzyme involved in formation of myelin sheaths, can be
detected by biochemical techniques from these fetal cells.
Yolk sac

764
Unit 7

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22. Developmental
Anatomy, Postnatal
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Decidua
basalis
Chorion
frondosum
Maternal
vein
Umbilical
cord
Chorion
Maternal
artery
Amnion
Amniotic sac containing

amniotic fluid
Placenta
FIGURE 22.12 The embryo, extraembryonic membranes, and placenta at approximately 7 weeks of development. Blood from the embryo is
CHAPTER 22
carried to and from the chorion frondosum by the umbilical arteries and vein. The maternal tissue between the chorionic villi is known as the decidua basalis; this tissue, together with the villi, form the functioning placenta.
human yolk sac contains no nutritive yolk, but it is still an essential structure during early embryonic development. Attached to
the underside of the embryonic disc (see figs. 22.10 and 22.11), it
produces blood for the embryo until the liver forms during the
sixth week. A portion of the yolk sac is also involved in the formation of the primitive gut. In addition, primordial germ cells
form in the wall of the yolk sac. During the fourth week, they
migrate to the developing gonads, where they become the primitive germ cells (spermatogonia or oogonia).
The stalk of the yolk sac usually detaches from the gut by
the sixth week. Following this, the yolk sac gradually shrinks as
pregnancy advances. Eventually, it becomes very small and
serves no additional function.
blood cells and gives rise to the fetal umbilical arteries and vein.
It also contributes to the development of the urinary bladder.
The extraembryonic portion of the allantois degenerates
during the second month. The intraembryonic portion involutes
to form a thick urinary tube called the urachus (yoo'ra-kus) (see
p. 690). After birth, the urachus becomes a fibrous cord called
the median umbilical ligament that attaches to the urinary bladder.
Chorion
The allantois forms during the third week as a small outpouching, or diverticulum, near the base of the yolk sac (see
fig. 22.10). It remains small but is involved in the formation of
The chorion (kor'e-on) is the outermost extraembryonic membrane. It contributes to the formation of the placenta as small fingerlike extensions, called chorionic villi, penetrate deeply into the
uterine tissue (see fig. 22.10). Initially, the entire surface of the
chorion is covered with villi. But those chorionic villi on the surface toward the uterine cavity gradually degenerate, producing a
smooth, bare area known as the smooth chorion. As this occurs,
the chorionic villi associated with the uterine wall rapidly increase
in number and branch out. This portion of the chorion is known
allantois: Gk. allanto, sausage; iodos, resemblance
chorion: Gk. chorion, external fetal membrane
Allantois


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
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Two
chorions
787
Two
amnions
Implantation
of blastocysts
Two
zygotes
Two-cell
stage
Blastocysts
Separate
chorionic sacs
Separate placentas
(a)
Two
chorions
fused
(b)
Two
amnions
Fused placentas
Cre
ek
Fused chorionic sacs
(c)
FIGURE 22.36 The formation of dizygotic twins. Twins of this type are fraternal rather than identical and may have (a) separate or (b) fused
placentas. (c) A photograph of fraternal twins at 11 weeks.
Fetoscopy (fig. 22.39) goes beyond amniocentesis by allowing direct examination of the fetus. Using fetoscopy, physicians
scan the uterus with pulsed sound waves to locate fetal structures,
the umbilical cord, and the placenta. Skin samples are taken from
the head of the fetus and blood samples extracted from the placenta. The principal advantage of fetoscopy is that external features of the fetus (such as fingers, eyes, ears, mouth, and genitals)
can be carefully observed. Fetoscopy is also used to determine
several diseases, including hemophilia, thalassemia, and sicklecell anemia cases, 40% of which are missed by amniocentesis.
Most hospitals are now equipped with instruments that
monitor fetal heart rate and uterine contractions during labor.
These instruments can detect any complication that may arise
during the delivery. The procedure is called Electronic Monitoring
amniocentesis: Gk. amnion, lamb (fetal membrane); kentesis, puncture
fetoscopy: L. fetus, offspring; skopein, to view
CHAPTER 22
sonography employs a mechanical vibration of high frequency to

produce a safe, high-resolution (sharp) image of fetal structure
(fig. 22.38). Ultrasonic imaging is a reliable way to determine
pregnancy as early as 6 weeks after ovulation. It can also be used
to determine fetal weight, length, and position, as well as to diagnose multiple fetuses.
Amniocentesis is a technique used to obtain a small sample of amniotic fluid so that it can be assessed genetically and
biochemically (see fig. 22.13). A wide-bore needle is inserted
into the amniotic sac with guidance by ultrasound, and 510 ml
of amniotic fluid is withdrawn with a syringe. Amniocentesis is
most often performed to determine fetal maturity, but it can also
help predict serious disorders such as Down syndrome or
Gauchers disease (a metabolic disorder).

788
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Development
22. Developmental
Anatomy, Postnatal
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One placenta
One chorionic sac

Two amniotic sacs
Implantation
of blastocyst
Zygote
Two-cell
stage
Blastocyst with two

embryoblasts
(a)
Single
placenta
(b)
C re
ek
Anastomosis of
placental vessels
FIGURE 22.37 The formation of monozygotic twins. Twins of this type develop from a single zygote and are identical. Such twins have two
amnions but only one chorion, and they share a common placenta.
Amniotic fluid
Placenta
Left cerebral
hemisphere
Syringe
withdraws
blood
Endoscope (guided
to exact location by
pulsed sound waves)
Orbit of eye
Left hand
Uterine wall
CHAPTER 22
Thorax
FIGURE 22.38 A color-enhanced ultrasonogram of a fetus during

the third trimester. The left hand is raised, as if waving to the viewer.
Placenta
Fiber optics
Needle punctures fetal
vein on placenta
FIGURE 22.39 Fetoscopy.


Development
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Anatomy, Postnatal
Chapter 22
The McGrawHill
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789
Fetal monitor
Amplifier
Cardiotachometer
Amplifier
Fetal
heart rate
Uterine
activity
Pressure transducer
FIGURE 22.40 Monitoring the fetal heart rate and uterine contractions using an FHRUC device.
of Fetal Heart Rate and Uterine Contractions (FHR-UC Monitoring). The extent of stress to the fetus from uterine contractions
can be determined through FHR-UC monitoring (fig. 22.40).
Long, arduous deliveries are taxing to both the mother and fetus.
If the babys health and vitality are presumed to be in danger because of a difficult delivery, the physician may decide to perform
a cesarean section.
Congenital Defects
Major developmental problems called congenital malformations
occur in approximately 2% of all newborn infants. They may be
caused by genetic inheritance, mutation (genetic change), or environmental factors. About 15% of neonatal deaths are attributed to congenital malformations. The branch of developmental
biology concerned with abnormal development and congenital

Two amnions and one chorion in all but very unusual cases prove the
twins to be monozygotic. The two infants, therefore, are genetically
identical. Thus, when considering genetic disorders such as cleft palate,
one would expect a high degree of concordance (both twins of a
monozygotic pair exhibit a particular anomaly). Many such defects however can be present in only one twina consequence of nongenetic factors, such as intrauterine environment. An example would be
inadequate blood supply to only one twin, resulting in a defect in that
twin but not in the other.
teratology: Gk. teras, monster; logos, study of
CHAPTER 22
congenital: L. congenitus, born with
malformations is called teratology (tar''a-tol'o-je). Many congenital problems have been discussed in previous chapters, in connection with the body system in which they occur.

790
Unit 7

Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
Companies, 2001

A 31-year-old female comes to your office
for her annual gynecologic exam. She reports no menstrual abnormalities. On
physical exam, you note a mass in her left
adnexa (accessory component of the main
organ). Because of this finding, you order a
CT scan to further evaluate this area. (B =
urinary bladder.)
QUESTIONS
1. What is the heterogeneous mass
(arrow) seen to the left of the urinary
bladder (B)?
2. What organ and cell line does this mass
arise from?
3. What is the significance of this mass?
Genetic Disorders of Clinical Importance
CHAPTER 22
cystic fibrosis An autosomal recessive disorder

characterized by the formation of thick mucus
in the lungs and pancreas that interferes with
normal breathing and digestion.
familial cretinism An autosomal recessive
disorder characterized by a lack of thyroid
secretion because of a defect in the iodine
transport mechanism. Untreated children are
dwarfed, sterile, and may be mentally retarded.
galactosemia (ga-lak''te-se'me-a) An autosomal
recessive disorder characterized by an
inability to metabolize galactose, a
component of milk sugar. Patients with this
disorder have cataracts, damaged livers, and
mental retardation.
gout An autosomal dominant disorder
characterized by an accumulation of uric acid
in the blood and tissue resulting from an
abnormal metabolism of purines.
hepatic porphyria (por-fer'e-a) An autosomal
dominant disorder characterized by painful GI
disorders and neurologic disturbances resulting
from an abnormal metabolism of porphyrins.
Huntingtons chorea: from George Huntington,

American physician, 18501916
Marfans syndrome: from Antoine Bernard-Jean
Marfan, French physician 18581942
hereditary hemochromatosis (he''mo-kro''mato'sis) A sex-influenced autosomal dominant

disorder characterized by an accumulation of
iron in the pancreas, liver, and heart,
resulting in diabetes, cirrhosis, and heart
failure.
hereditary leukomelanopathy (loo''ko-mel''anop'a-the) An autosomal recessive disorder
characterized by decreased pigmentation in
the skin, hair, and eyes and abnormal white
blood cells. People with this condition are
generally susceptible to infections and early
deaths.
Huntingtons chorea An autosomal dominant
disorder characterized by uncontrolled
twitching of skeletal muscles and the
deterioration of mental capacities. A latent
expression of this disorder allows the mutant
gene to be passed to children before
symptoms develop.
Marfans syndrome An autosomal dominant
disorder characterized by tremendous growth
of the extremities, extreme looseness of the
joints, dislocation of the lenses of the eyes,

and congenital cardiovascular defects.
phenylketonuria (fen''il-ket''n-oor'e-a)
(PKU) An autosomal recessive disorder
characterized by an inability to metabolize
the amino acid phenylalanine. It is
accompanied by brain and nerve damage and
mental retardation. (Newborns are routinely
tested for PKU; those that are affected are
placed on a diet low in phenylalanine.)
pseudohypertrophic muscular dystrophy A
sex-linked recessive disorder characterized by
progressive muscle atrophy. It usually begins
during childhood and causes death in
adolescence.
retinitis pigmentosa A sex-linked recessive
disorder characterized by progressive atrophy
of the retina and eventual blindness.
Tay-Sachs disease An autosomal recessive
disorder characterized by a deterioration of
physical and mental abilities, early blindness,
and early death. It has a disproportionately high
incidence in Jews of Eastern European origin.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
The McGrawHill
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791
Chapter Summary
Fertilization (pp. 755757)
1. Upon fertilization of a secondary oocyte
by a spermatozoon in the uterine tube,
meiotic development is completed and a
diploid zygote is formed.
2. Morphogenesis is the sequential
formation of body structures during the
prenatal period of human life. The
prenatal period lasts 38 weeks and is
divided into a preembryonic, an
embryonic, and a fetal period.
3. A capacitated spermatozoon digests its
way through the zona pellucida and
corona radiata of the secondary oocyte to
complete the fertilization process and
formation of a zygote.
2.
Preembryonic Period (pp. 757762)
Embryonic Period (pp. 762771)

1. The events of the 6-week embryonic
period include the differentiation of the
germ layers into specific body organs and
3.
4.
5.
Fetal Period (pp. 772774)

1. A small amount of tissue differentiation and
organ development occurs during the fetal
period, but for the most part fetal development
is primarily limited to body growth.
2. Between weeks 9 and 12, ossification

centers appear, the genitalia are formed,
and the digestive, urinary, respiratory, and
muscle systems show functional activity.
3. Between weeks 13 and 16, facial features
are formed and the fetal heartbeat can be
detected with a stethoscope.
4. During the 17-to-20-week period,
quickening can be felt by the mother, and
vernix caseosa and lanugo cover the skin
of the fetus.
substantial weight gain occurs and the
fetal skin becomes wrinkled and pinkish.
6. Toward the end of the 26-to-29-week
period, the eyes have opened, the gonads
have descended in a male, and the fetus is
developed to the extent that it might
survive if born prematurely.
7. At 38 weeks, the fetus is full-term; the
normal gestation is 266 days.
Labor and Parturition (p. 775)

1. Labor and parturition are the culmination
of gestation and require the action of
oxytocin, secreted by the posterior
pituitary, and prostaglandins, produced in
the uterus.
2. Labor is divided into dilation, expulsion,
and placental stages.
Periods of Postnatal Growth

(pp. 775782)
1. The course of human life after birth is
seen in terms of physical and
physiological changes and the attainment
of maturity in the neonatal period,
infancy, childhood, adolescence, and
adulthood.
2. The neonatal period, extending from birth to
the end of the fourth week, is characterized
by major physiological changes.
(a) The most critical need of the newborn
is to establish adequate respiratory and
heart rates. A normal respiratory rate
is 30 to 40 respirations per minute,
and a normal heart rate ranges from
120 to 160 beats per minute.
(b) The four reflexes in the newborn
critical to survival are the suckling
reflex, the rooting reflex, the crying
reflex, and the breathing reflex.
3. Infancy, extending from 4 weeks through
the second year, is characterized by
tremendous growth, increased
coordination, and mental development.
(a) By 2 years, most infants weigh about
4 times their birth weight and average
between 32 and 36 inches in length.
CHAPTER 22
1. Cleavage of the zygote is initiated within

30 hours and continues until a morula
forms; the morula enters the uterine
cavity on about the third day.
2. A hollow, fluid-filled space forms within
the morula, at which point it is called a
blastocyst.
3. Implantation begins between the fifth and
seventh day and is enabled by the
secretion of enzymes that digest a portion
of the endometrium.
(a) During implantation, the trophoblast
cells secrete human chorionic
gonadotropin (hCG), which prevents
the breakdown of the endometrium
and menstruation.
(b) The secretion of hCG declines by the
tenth week as the developed placenta
secretes steroids that maintain the
endometrium.
4. The embryoblast of the implanted
blastocyst flattens into the embryonic
disc, from which the primary germ layers
of the embryo develop.
(a) Ectoderm gives rise to the nervous
system, the epidermis of the skin and
epidermal derivatives, and to portions
of the sensory organs.
(b) Mesoderm gives rise to bones,
muscles, blood, reproductive organs,
the dermis of the skin, and
connective tissue.
(c) Endoderm gives rise to linings of the
GI tract, digestive organs, the
respiratory tract and lungs, and the
urinary bladder and urethra.
the formation of the placenta, the

umbilical cord, and the extraembryonic
membranes. These events make it possible
for morphogenesis to continue.
The extraembryonic membranes include
the amnion, yolk sac, allantois, and
chorion.
(a) The amnion is a thin membrane
surrounding the embryo. It contains
amniotic fluid that cushions and
protects the embryo.
(b) The yolk sac produces blood for the
embryo.
(c) The allantois also produces blood for
the embryo and gives rise to the
umbilical arteries and vein.
(d) The chorion participates in the
formation of the placenta.
The placenta, formed from both maternal
and embryonic tissue, has a transport role
in providing for the metabolic needs of
the fetus and in removing its waste.
(a) The placenta produces steroid and
polypeptide hormones.
(b) Nicotine, drugs, alcohol, and viruses
can cross the placenta to the fetus.
The umbilical cord, containing two
umbilical arteries and one umbilical vein,
is formed as the amnion envelops the
tissues on the underside of the embryo.
From the third to the eighth week, the
structure of all the body organs, except
the genitalia, becomes apparent.
(a) During the third week, the primitive
node forms from the primitive line,
which later gives rise to the
notochord and intraembryonic
mesoderm.
(b) By the end of the fourth week, the
heart is beating; the primordial tissues
of the eyes, brain, spinal cord, lungs,
and digestive organs are properly
positioned; and the superior and
inferior limb buds are recognizable.
(c) At the end of the fifth week, the
sense organs are formed in the
enlarged head and the appendages
have developed with digital primordia
evident.
(d) During the seventh and eighth weeks,
the body organs are formed, except
for the genitalia, and the embryo
appears distinctly human.

614
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Body
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Thyroid cartilage
Cricoid cartilage
Trachea
Clavicle
Apex of left lung
Scapula
Superior lobe
of left lung
Superior lobe
of right lung
Sternum
Middle lobe
of right lung
Inferior lobe
of right lung
Cardiac impression
Inferior lobe
of left lung
Base of
left lung
Costal cartilage
Creek
FIGURE 17.17 The position of the lungs within the rib cage.
CHAPTER 17
distance between each alveolar duct and its terminal pulmonary

alveoli is only about 0.5 mm, these units together compose most
of the mass of the lungs.
Lungs
The large, spongy lungs are paired organs within the thoracic cavity (fig. 17.17). Each lung extends from the diaphragm to a point
just above the clavicle, and its surfaces are bordered by the ribs to
the front and back. The lungs are separated from one another by the
heart and other structures of the mediastinum (me''de-a-sti'num)
which is the area between the lungs. All structures of the respiratory system beyond the principal bronchi, including the bronchial
tree and the pulmonary alveoli, are contained within the lungs.
Each lung has four surfaces that match the contour of the
thoracic cavity. The mediastinal (medial) surface of the lung is
slightly concave and contains a vertical slit, the hilum through
which pulmonary vessels, nerves, and bronchi pass (fig. 17.18).

The inferior surface of the lung, called the base of the lung, is
concave as it fits over the convex dome of the diaphragm. The
superior surface, called the apex (cupola) of the lung, extends
above the level of the clavicle. Finally, the broad, rounded surface in contact with the membranes covering the ribs is called
the costal surface of the lung.
Although the right and left lungs are basically similar, they
are not identical. The left lung is somewhat smaller than the
right and has a cardiac impression on its medial surface to accommodate the heart. The left lung is subdivided into a superior
lobe and an inferior lobe by a single fissure. The right lung is
subdivided by two fissures into three lobes: superior, middle, and
inferior lobes (see figs. 17.1, 17.17, and 17.18). Each lobe of the
lung is divided into many small lobules, which in turn contain
the pulmonary alveoli. Lobular divisions of the lungs make up
specific bronchial segments. Each bronchial segment has its own
mediastinum: L. mediastinus, intermediate

hilum: L. hilum, a trifle (little significance)
cupola: L. cupula, diminutive of cupa, dome or tub


Body
The McGrawHill
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Chapter 17
Respiratory System
Apex of lung
Apex of lung
Superior lobe
Superior lobe
615
Oblique fissure
Pulmonary artery
Bronchus
Bronchus
Pulmonary artery
Pulmonary vein
Pulmonary veins
Middle lobe
Inferior lobe
Inferior lobe
Cardiac impression
Oblique fissure
Aortic impression
Pulmonary ligament
Diaphragmatic surface
Diaphragmatic surface
FIGURE 17.18 Lungs from a cadaver. (a) A medial view of the right lung and (b) a medial view of the left lung.
blood supply and if diseased it can be surgically isolated. The

right lung contains 10 bronchial segments and the left lung contains 8 (fig. 17.19).
The lungs of a newborn are pink but may become discolored
in an adult as a result of smoking or air pollution. Smoking not
only discolors the lungs, it may also cause deterioration of the pulmonary alveoli. Emphysema (em''f -se'ma) and lung cancer (see figs.
17.32 and 17.33) are diseases that are linked to smoking. If a person
moves to a less polluted environment or gives up smoking, the lungs
will get pinker and function more efficiently, unless they have been
permanently damaged by disease.
Pleurae
pleura: Gk. pleura, side or rib

pulmonary: Gk. pleumon, lung
Knowledge Check
10. Describe the structure of the respiratory division of the
lungs and explain how this structure aids in gas exchange.
11. Compare the structure of the right and left lungs.
12. Describe the location of the mediastinum and list the organs it contains.
13. Describe the arrangement of the visceral pleura, parietal
pleura, and pleural cavity. Comment on the functional significance of the compartmentalization of the thoracic cavity.
CHAPTER 17
Pleurae (ploor'e) are serous membranes surrounding the lungs and

lining the thoracic cavity (figs. 17.20 and 17.21). The visceral
pleura adheres to the outer surface of the lung and extends into
each of the interlobar fissures. The parietal pleura lines the thoracic walls and the thoracic surface of the diaphragm. A continuation of the parietal pleura and between the lungs forms the
boundary of the mediastinum. Between the visceral and parietal
pleurae is the slitlike pleural cavity. It contains a lubricating fluid
that allows the membranes to slide past each other easily during
respiration. An inferiorly extending reflection of the pleural layers
around the roots of each lung is called the pulmonary ligament.
The pulmonary ligaments help support the lungs.
The moistened serous membranes of the visceral and parietal pleurae are normally flush against each other like two wet
pieces of glass, and therefore the lungs are stuck to the thoracic
wall. The pleural cavity (intrapleural space) between the two

moistened membranes contains only a thin layer of fluid secreted
by the serous membranes. The pleural cavity in a healthy, living
person is thus potential rather than real; it can become real only
in abnormal situations when air enters the intrapleural space. Because the lungs normally remain in contact with the thoracic
wall, they get larger and smaller along with the thoracic cavity
during respiratory movements.
The thoracic cavity has four distinct compartments: a
pleural cavity surrounds each lung; the pericardial cavity surrounds the heart; and the mediastinum contains the esophagus,
thoracic duct, major vessels, various nerves, and portions of the
respiratory tract. This compartmentalization has protective value
in that infections are usually confined to one compartment.
Also, damage to one organ usually will not involve another. For
example, pleurisy, an inflamed pleura, is generally confined to
one side; and a penetrating injury to the thoracic cavity, such as
a knife wound, may cause one lung to collapse but not the other.
A summary of the major structures of the respiratory system is presented in table 17.1.

616
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Body
The McGrawHill
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Anterior view of bronchopulmonary segments

of the lungs
Apicoposterior
Apical
Posterior
Anterior
Superior
lingular
Medial
Lateral
Inferior
lingular
Anterior basal
Medial basal
Posterior basal
Lateral basal
Right lung
Left lung
Trachea
Carina
Left principal bronchus
Right principal bronchus
Apicoposterior
Apical
CHAPTER 17
Serving
superior lobe
Fused into
single element
Anterior
Posterior
Anterior
Lobar
(secondary)
bronchi
Serving
middle lobe
Lateral
Medial
Serving
inferior lobe
Superior
Anterior basal
Medial basal
Lateral basal
Posterior basal
Segmental (tertiary) bronchi
Superior
Inferior
Lingular
Superior
Anterior basal
Medial basal
Lateral basal
Posterior basal
Posterior view of bronchopulmonary segments

of the lungs
Apicoposterior
Apical
Posterior
Anterior
Superior
Superior
lingular
Anterior
Lateral
Posterior basal
Lateral basal
Creek
Left lung
FIGURE 17.19 Lobes, lobules, and bronchopulmonary segments of the lungs.
Right lung
Serving
inferior lobe
Serving
superior lobe


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Intercostal
muscles
Parietal pleura
Pleural cavity
Apex of lung
Rib
Visceral pleura
Trachea
Lung
Thymus
Superior lobe
of right lung
Superior lobe
of left lung
Horizontal fissure
Cardiac notch
Oblique fissure
Oblique fissure
Middle lobe
of right lung
Inferior lobe
of left lung
Inferior lobe
of right lung
Base of lung
Diaphragm
Heart (in mediastinum)

Creek
FIGURE 17.20 The position of the lungs and associated pleurae.
Left lung
Thoracic aorta
FIGURE 17.21 A cross section of the thoracic cavity showing the mediastinum and pleural membranes.
CHAPTER 17
Left pulmonary
artery
Trachea

Unit 6
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TABLE 17.1 Major Structures of the Respiratory System

Structure
Description
Function
Nose
Primary passageway for air entering the respiratory system; consists

of jutting external portion and internal nasal cavity
Warms, moistens, and filters inhaled air as it is conducted to the

pharynx
Paranasal sinuses
Air spaces in the maxillary, frontal, sphenoid, and ethmoid bones
Produce mucus; provide sound resonance; lighten the skull
Pharynx
Tubular organ connecting oral and nasal cavities to the larynx
Serves as passageway for air entering the larynx and for food
entering the esophagus
Larynx
Voice box; short passageway that connects the pharynx to the

trachea
Serves as passageway for air; produces sound; prevents foreign

materials from entering the trachea
Trachea
Flexible tubular connection between the larynx and bronchial

tree
Serves as passageway for air; pseudostratified ciliated columnar

epithelium cleanses the air
Bronchial tree
Bronchi and branching bronchioles in the lung; tubular

connection between the trachea and pulmonary alveoli
Serves as passageway for air; continued cleansing of inhaled air
Pulmonary alveoli
Microscopic membranous air sacs within the lungs
Functional units of respiration; site of gaseous exchange between

the respiratory and circulatory systems
Lungs
Major organs of the respiratory system; located in the thoracic

cavity and surrounded by pleural cavities
Contain bronchial trees, pulmonary alveoli, and associated

pulmonary vessels
Pleurae
Serous membranes covering the lungs and lining the thoracic

cavity
Compartmentalize, protect, and lubricate the lungs
MECHANICS OF BREATHING
Normal quiet inspiration is achieved by muscle contraction, and
quiet expiration results from muscle relaxation and elastic recoil. A
deeper inspiration and expiration can be forced by contractions of
the accessory respiratory muscles. The amount of air inspired and
expired can be measured to test pulmonary function.
Objective 12
CHAPTER 17
Identify and describe the actions of the

muscles involved in both quiet and forced inspiration.
Objective 13
Describe how quiet expiration occurs and

identify and describe the actions of the muscles involved in
forced expiration.
Objective 14
List the various lung volumes and capacities

and describe how pulmonary function tests help in the
diagnosis of lung disorders.
Breathing, or pulmonary ventilation, requires that the thorax be

flexible in order to function as a bellows during the ventilation
cycle. Breathing consists of two phases, inspiration and expiration.
Inspiration (inhaling) and expiration (exhaling) are accomplished by alternately increasing and decreasing the volume of
the thoracic cavity (fig. 17.22). Breathing in takes place when
the air pressure within the lungs is lower than the atmospheric
pressure; breathing out takes place when the air pressure within
the lungs is greater than the atmospheric pressure.
Pressure gradients change as the size of the thoracic cavity

changes. Not only must the thorax be flexible, it also must be
sufficiently rigid to protect the vital organs it contains. In addition, it must provide extensive attachment surfaces for many
short, powerful muscles. These requirements are met through the
structure and composition of the rib cage. The rib cage is pliable
because the ribs are separated from one another and because
most ribs (upper 10 of the 12 pairs) are attached to the sternum
by resilient costal cartilage. The vertebral attachment likewise
allows for considerable mobility. The structure of the rib cage
and associated cartilage provides continuous elastic tension so
that an expanded thorax will return passively to its resting position when relaxed.
Inspiration
The overall size of the thoracic cavity increases with inspiration
(fig. 17.23). During relaxed inspiration, the muscles of importance are the diaphragm, the external intercostal muscles, and the
interchondral portion of the internal intercostal muscles
(fig. 17.24). Contraction of the dome-shaped diaphragm causes it
to flatten, lowering its dome. This increases the vertical dimen-
diaphragm: Gk. dia, across; phragma, fence


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Chapter 17
At rest
Respiratory System
619
Inspiration
Expiration
FIGURE 17.22 The mechanics of pulmonary ventilation. At rest (a), the atmospheric pressure at sea level and within the plural cavities is 760
mmHg. During inspiration (b), the diaphragm contracts, causing a decrease in intrapleural pressure and consequent inflation of the lungs. During
expiration (c), the diaphragm recoils, causing an increase in intrapleural pressure and consequent deflation of the lungs.
(b)
FIGURE 17.23 A change in lung volume, as shown by radiographs, during expiration (a) and inspiration (b). The increase in
lung volume during full inspiration is shown by comparison with the
lung volume in full expiration (dashed lines).
Expiration
For the most part, expiration is a passive process that occurs as the
muscles of inspiration are relaxed and the rib cage returns to its
original position. The lungs recoil during expiration as elastic fibers
within the lung tissue shorten and the pulmonary alveoli draw together. Lowering of the surface tension in the pulmonary alveoli,
CHAPTER 17
(a)
sion of the thoracic cavity. A simultaneous contraction of the

external intercostal muscles and interchondral portion of the internal intercostal muscles increases the diameter of the thorax.
The scalenes and sternocleidomastoid muscles are involved in
deep inspiration or forced breathing. When these muscles are contracted, the ribs are elevated. At the same time, the upper rib cage
is stabilized so that the external intercostal muscles become more
effective.
The expanded thoracic cavity decreases the air pressure
within the pleural cavities to below that of the atmosphere. It is
this pressure difference that causes the lungs to become inflated.

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FIGURE 17.24 The muscles of respiration. The principal muscles of inspiration are shown on the right side of the trunk and the principal mus-
CHAPTER 17
cles of forced expiration are shown on the left side. For the most part, expiration is passive.
which brings on recoiling, is due to a lipoprotein substance called

surfactant produced by type II alveolar cells. Surfactant is extremely
important in reducing the surface tension in the pulmonary alveoli
by becoming interspersed between water molecules, thereby reducing the attracting forces of hydrogen bonds. A deficiency in surfactant in premature infants can cause respiratory distress syndrome
(RDS) or, as it is commonly called, hyaline membrane disease.
Even under normal conditions, the first breath of life is a difficult one because the newborn must overcome large surface
tension forces in order to inflate its partially collapsed pulmonary
alveoli. The transpulmonary pressure required for the first breath is
15 to 20 times that required for subsequent breaths, and an infant
with respiratory distress syndrome must duplicate this effort with
every breath. Fortunately, many babies with this condition can be
saved by mechanical ventilators that keep them alive long enough for
their lungs to mature and manufacture sufficient surfactant.
During forced expiration, such as coughing or sneezing,

contraction of the interosseous portion of the internal intercostal
muscles causes the rib cage to be depressed. The abdominal muscles may also aid expiration because, when contracted, they force
abdominal organs up against the diaphragm and further decrease
the volume of the thorax. Thus, intrapulmonary pressure can rise
to 20 or 30 mmHg above the atmospheric pressure.
The events that occur during inspiration and expiration

are summarized in table 17.2.
Respiratory Volumes and Capacities

The respiratory system is somewhat inefficient because the air
enters and exits at the same place, through either the nose or the
mouth. Consequently, there is an incomplete exchange of gas
during each ventilatory cycle, and approximately five-sixths of
the air present in the lungs still remains when the next inspiration begins.
The amount of air breathed in a given time and the degree of difficulty in breathing are important indicators of a persons respiratory status. The amount of air exchanged during
pulmonary ventilation varies from person to person according
to age, gender, activity level, general health, and individual differences. Respiratory volumes are measured with a spirometer
(fig. 17.25). Any ventilatory abnormalities can then be compared to what is accepted as normal. The normal adult respiratory volumes and capacities are presented in table 17.3 and
figure 17.26.


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Chapter 17
Respiratory System
621
TABLE 17.2 Pulmonary Ventilation:

Events of Inspiration and Expiration*
Nerve Stimulus
Event
Inspiration
Phrenic nerves
The diaphragm contracts, moving inferiorly,

which measures the volume of the thorax.
The diaphragm is the principal muscle
involved in quiet inspiration.
Intercostal nerves
Contraction of the external intercostal muscles

and the interchondral portion of the internal
intercostal muscles elevates the ribs, thus
increasing the capacity of the thoracic cavity.
Accessory, cervical,
and thoracic nerves
Forced inspiration is accomplished through

contraction of the scalenes and sternocleidomastoid muscles, which increases the
dimension of the thoracic cavity anteroposteriorly. Pulmonary ventilation during forced
inspiration usually occurs through the mouth
rather than through the nose.
As the dimension of the thoracic cavity
increases, the pressure within the pleural
cavities decreases; the lungs inflate because
the atmospheric pressure is greater than the
intraplural pressure.
Expiration
Nerve stimuli to the inspiratory muscles cease
and the muscles relax.
Intercostal and lower

spinal nerves
The rib cage and lungs recoil as air is forced out

of the lungs because of the increased pressure.
Forced expiration occurs when the interosseus
portion of the internal intercostal and
abdominal muscles are contracted.
FIGURE 17.25 A spirometer. With the exception of the residual

volume, which is measured using special techniques, this instrument
can determine respiratory air volumes.
TABLE 17.3 Respiratory Volumes

and Capacities of Healthy Adult Males
Volume
Quanity
of Air
Tidal volume (TV)
500 cc
Volume moved in or out of the

lungs during quiet breathing
Inspiratory reserve
volume (IRV)
3,000 cc
Volume that can be inhaled during

forced breathing in addition to
tidal volume
Expiratory reserve
volume (ERV)
1,000 cc
Volume that can be exhaled during

forced breathing in addition to
tidal volume
Vital capacity (VC)
4,500 cc
Maximum amount of air that can be

exhaled after taking the deepest
breath possible: VC = TV + IRV
+ ERV
Residual volume
(RV)
1,500 cc
Volume that cannot be exhaled
Total lung
capacity (TLC)
6,000 cc
Total volume of air that the lungs

can hold: TLC = VC + RV
*Some of the events during inspiration and expiration may occur

simultaneously.
Nonrespiratory Air Movements

Air movements through the respiratory system that are not associated with pulmonary ventilation are termed nonrespiratory
movements. Such movements accompany emotional displays such
as laughing, sighing, crying, or yawning, or they may function to
expel foreign matter from the respiratory tract, as in coughing
and sneezing. Nonrespiratory movements are generally reflexive.
Some of them, however, can be voluntarily initiated. These types
dyspnea: Gk. dys, bad; pnoe, breathing
of air movements and the reflexive mechanisms involved are

summarized in table 17.5.
Knowledge Check
14. Describe the actions of the diaphragm and intercostal muscles during relaxed inspiration.
15. Describe how forced inspiration and forced expiration are
produced.
16. Define the terms tidal volume and vital capacity.
17. Indicate the respiratory volumes being used during a
sneeze, a deep inspiration prior to jumping into a swimming pool, maximum ventilation while running, and quiet
breathing while sleeping.
CHAPTER 17
People with pulmonary disorders frequently complain of dyspnea (disp'ne-a), a subjective feeling of shortness of breath.
Dyspnea may occur even when ventilation is normal, however, and
may not occur even during exercise, when the total volume of air
movement is very high. Some of the terms used to describe ventilation are defined in table 17.4.
Description

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Functional residual
capacity
Inspiratory
capacity
622

Body
FIGURE 17.26 Respiratory volumes and capacities.
TABLE 17.4 Ventilation Terminology
CHAPTER 17
Term
Definition
Air spaces
Alveolar ducts, alveolar sacs, and pulmonary alveoli
Airways
Structures that conduct air from the mouth and nose to the
respiratory bronchioles
Alveolar
ventilation
Removal and replacement of gas in pulmonary alveoli; equal

to the tidal volume minus the volume of dead space times
the breathing rate
Anatomical dead
space
Volume of the conducting airways to the zone where gas

exchange occurs
Apnea
Cessation of breathing
Dyspnea
Unpleasant subjective feeling of difficult or labored breathing
Eupnea
Normal, comfortable breathing at rest
Term
Definition
Hyperventilation
Abnormally rapid, deep breathing; results in

abnormally low alveolar CO2
Hypoventilation
Abnormally slow, shallow breathing; results in

abnormally high alveolar CO2
Orthopnea
Inability to breathe comfortably while lying

down
Physiological
dead space
Combination of anatomical dead space and

underventilated pulmonary alveoli that do not
contribute normally to blood-gas exchange
Pneumothorax
Presence of gas in the pleural cavity (the space

between the visceral and parietal pleural
membranes) that may cause lung collapse
TABLE 17.5 Nonrespiratory Air Movements

Air Movement
Mechanism
Comments
Coughing
Deep inspiration followed by a closure of the glottis. The forceful

expiration that results abruptly opens the glottis, sending a blast
of air through the upper respiratory tract.
Reflexive or voluntary. Stimulus may be foreign material

irritating the larynx or trachea.
Sneezing
Similar to a cough, except that the forceful expired air is directed primarily
through the nasal cavity. The eyelids close reflexively during a sneeze.
Reflexive response to irritating stimulus of the nasal mucosa.

Sneezing clears the upper respiratory passages.
Sighing
Deep, prolonged inspiration followed by a rapid, forceful expiration.
Reflexive or voluntary, usually in response to boredom or

sadness.
Yawning
Deep inspiration through a widely opened mouth. The inspired air

is usually held for a short period before sudden expiration.
Usually reflexive in response to drowsiness, fatigue, or

boredom, but exact stimulus-receptor cause is unknown.
Laughing
Deep inspiration followed by a rapid convulsive expiration. Air

movements are accompanied by expressive facial distortions.
Reflexive; may be voluntary to express emotions.
Crying
Similar to laughing, but the glottis remains open during entire

expiration and different facial muscles are involved.
Somewhat reflexive but under voluntary control.
Hiccuping
Spasmodic contraction of the diaphragm while the glottis is closed,

producing a sharp inspiratory sound.
Reflexive; serves no known function.


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Chapter 17
Respiratory System
623
REGULATION OF BREATHING
The rhythm of breathing is controlled by centers in the brain stem.
These centers are influenced by higher brain function and regulated by sensory input that makes breathing responsive to the
changing respiratory needs of the body.
Objective 15
Describe the functions of the pneumotaxic,

apneustic, and rhythmicity centers in the brain stem.
Objective 16
Identify the chemoreceptors and describe

their pathway of innervation.
Sensory nerve fibers

(in glossopharyngeal nerve)
Carotid body
Carotid sinus
Common carotid
artery
Sensory nerve fibers

(in vagus nerve)
Aortic bodies
Ascending
aorta
Heart
FIGURE 17.27 The peripheral chemoreceptors (aortic and carotid

bodies) regulate the brain stem respiratory centers by means of sensory nerve stimulation.
Knowledge Check
18. State where in the brain the three respiratory areas are located. Which of these three areas is responsible for autonomic rhythmic breathing?
19. Describe the locations of the peripheral chemoreceptors
and identify the two paired cranial nerves that carry sensory impulses from these sites to the respiratory centers
within the brain stem.
CHAPTER 17
Pulmonary ventilation is primarily an involuntary, rhythmic action so effective that it continues to function even when a person is unconscious. In order for the neural control center of
respiration to function effectively, it must possess monitoring,
stimulating, and inhibiting properties so that the body can respond appropriately to increased or decreased metabolic needs.
In addition, the center must be connected to the cerebrum to receive the voluntary impulses from a person who wants to change
the rate of respiration.
The three respiratory centers of the brain are the rhythmicity, apneustic, and pneumotaxic areas (see fig. 11.28). The rhythmicity area, located in the medulla oblongata, contains two
aggregations of nerve cell bodies that form the inspiratory and expiratory portions. Nerve impulses from the inspiratory portion
travel through the phrenic and intercostal nerves and stimulate the
diaphragm and intercostal muscles. Impulses from the expiratory
portion stimulate the muscles of expiration. These two portions act
reciprocally; that is, when one is stimulated, the other is inhibited.
Stretch receptors in the visceral pleura of the lungs provide feedback through the vagus nerves to stimulate the expiratory portion.
The apneustic (ap-noo'stik) and pneumotaxic (noo''motak'sik) areas are located in the pons. These areas influence the
activity of the rhythmicity area. The apneustic center promotes
inspiration and the pneumotaxic center inhibits the activity of
inspiratory neurons.
The brain stem respiratory centers produce rhythmic
breathing even in the absence of other neural input. This intrinsic respiratory pattern, however, is modified by input from higher
brain centers and by input from receptors sensitive to the chemical composition of the blood. The influence of higher brain centers is evidenced by the fact that you can voluntarily
hypoventilate (as in breath holding) or hyperventilate. The inability to hold your breath for more than a short period is due to
reflex breathing in response to input from the chemoreceptors.
Two groups of chemoreceptors respond to changes in blood
chemistry. These are the central chemoreceptors in the medulla
oblongata and the peripheral chemoreceptors. The peripheral
chemoreceptors include the aortic bodies, located in the aortic
arch, and the carotid bodies, located at the junctions of the internal and external carotid arteries (fig. 17.27). These peripheral
chemoreceptors control breathing indirectly via sensory neurons
to the medulla oblongata. The aortic bodies send sensory information to the medulla oblongata in the vagus nerves; the carotid
bodies stimulate sensory fibers in the glossopharyngeal nerve.


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The Respiratory System
EXPLANATION
The development of the respiratory system is initiated early in
embryonic development and involves both ectoderm and endoderm. Although all of the structures of the respiratory system develop simultaneously, we will consider the upper and
lower systems separately because of the different germ layers
involved.
Development of the Upper Respiratory System

Cephalization (sef''a-l-za'shun) is the evolutionary tendency toward structural and functional differentiation of the cephalic, or
head, end of an organism from the rest of the body. In humans,
cephalization is apparent early in development. One of the initial events is the formation of the nasal cavity at 3 1/2 to 4
weeks of embryonic life. A region of thickened ectoderm called
the olfactory (nasal) placode (plak'od) appears on the front inferior part of the head (exhibit I). The placode invaginates to
form the olfactory pit, which extends posteriorly to connect
with the foregut. The foregut, derived of endoderm, later develops into the pharynx.
The mouth, or oral cavity, develops at the same time as the
nasal cavity, and for a short time there is a thin oronasal membrane separating the two cavities. This membrane ruptures dur-
cephalization: Gk. kephale, head
624
ing the seventh week, and a single large oronasal cavity forms.
Shortly thereafter, tissue plates of mesoderm begin to grow horizontally across the cavity. At approximately the same time, a vertical plate develops inferiorly from the roof of the nasal cavity.
These plates have completed their formation by 3 months of development. The vertical plate forms the nasal septum, and the
horizontal plates form the hard palate.
A cleft palate forms when the horizontal plates fail to meet
in the midline. This condition can be corrected surgically
(see fig. 17.28). The more immediate and serious problem
facing an infant with a cleft palate is that it may be unable
to create enough suction to nurse properly.
Development of the Lower Respiratory System

The lower respiratory system begins as a diverticulum, or outpouching, from the ventral surface of endoderm along the
lower pharyngeal region (exhibit II). This diverticulum, which
forms during the fourth week of development, is referred to as
the laryngotracheal (la-ring''go-tra'ke-al) bud. As the bud
grows, the proximal portion forms the trachea and the distal
portion bifurcates (splits) into a right and left principal
bronchus.
The buds continue to elongate and split until the entire
tubular network within the lower respiratory tract is formed
(exhibit II). As the terminal portion forms air sacs, called pulmonary alveoli, at about 8 weeks of development, the supporting lung tissue begins to form. The complete structure of
the lungs, however, is not fully developed until about 26
weeks of fetal development. Premature infants born prior to
this time therefore require special artificial respiratory equipment to live.


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EXHIBIT I The development of the upper respiratory system. (a) An anterior view of the developing head of an embryo at 4 weeks showing the position of a transverse cut depicted in (a1), (a2), and (a3). (a2) Development at 5 weeks and (a3) at 5 1/2 weeks. (b) An anterior view
of the developing head of an embryo at 6 weeks showing the position of a sagittal cut depicted in (b1), (b2), (b3), and (b4) at 14 weeks.
625


Body
Right principal bronchus
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Left principal bronchus
EXHIBIT II The development of the lower respiratory tract. Developments in (a) and (b) occur during the first month. Those in (c), (d),
and (e) occur during the second month.
The respiratory system is particularly vulnerable to infectious diseases simply because many pathogens are airborne; humans are
highly social, and the warm, moistened environment along the
respiratory tract allows pathogens to thrive. Injury and trauma
are also frequent problems. Protruding noses are subject to fractures; the large, spongy lungs are easily penetrated by broken ribs;
and portions of the respiratory tract may become occluded because they also have a digestive function.
Developmental Problems
of the Respiratory System
Birth defects, inherited disorders, and premature births commonly cause problems in the respiratory system of infants. A
cleft palate is a developmental deformity of the hard palate of
the mouth. An opening persists between the oral and nasal cavi626
ties that makes it difficult, if not impossible, for the infant to

nurse. A cleft palate may be hereditary or a complication of some
disease (e.g., German measles) contracted by the mother during
pregnancy. A cleft lip is a genetically based developmental disorder in which the two sides of the upper lip fail to fuse. Cleft
palates and cleft lips can be treated very effectively with cosmetic surgery (fig. 17.28).
As mentioned earlier, hyaline membrane disease is a fairly
common respiratory disorder that accounts for about one-third of
neonatal deaths. This condition results from the deficient production of surfactant. Cystic fibrosis is a genetic disorder that affects the respiratory system, as well as other systems of the body,
and accounts for approximately 1 childhood death in 20. The effect of this disease on the respiratory system is usually a persistent
inflammation and infection of the respiratory tract.
Pulmonary alveoli are not developed sufficiently to sustain life until after week 20 of gestation. Thus, extrauterine life
prior to that time is extremely difficult even with life-supporting devices.


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Chapter 17
Respiratory System
627
(a)
FIGURE 17.29 A pneumothorax of the right lung. The right side of

the thorax appears uniformly dark because it is filled with air; the
spaces between the ribs are also greater than those on the left, because the ribs are released from the elastic tension of the lungs. The
left lung appears denser (less dark) because of shunting of blood
from the right to the left lung.
(b)
Trauma or Injury
Humans are especially susceptible to epistaxes (ep''-stak'sez)
(nosebleeds) because the prominent nose can be easily bumped
and because the nasal mucosa has extensive vascularity for
warming the inspired air. Epistaxes may also be caused by high
blood pressure or diseases such as leukemia.
When air enters the pleural cavity surrounding either lung,
the condition is referred to as a pneumothorax (fig. 17.29). A
pneumothorax can result from an external injury, such as a stabbing, bullet wound, or penetrating fractured rib, or it can be the
result of internal conditions. A severely diseased lung, as in emphysema, can create a pneumothorax as the wall of the lung deteriorates along with the visceral pleura and permits air to enter
the pleural cavity.
If you are alone and choking, use whatever is available to

apply force just below your diaphragm. Press into a table or a
sink, or use your own fist.
CHAPTER 17
FIGURE 17.28 (a) An infant with a unilateral cleft lip and palate.
(b) The same child following corrective surgery.
Choking on a foreign object is a common serious trauma to

the respiratory system. More than eight Americans choke to
death each day on food lodged in their trachea. A simple procedure termed the abdominal thrust (Heimlich) maneuver can
save the life of a person who is choking (fig. 17.30). The abdominal thrust maneuver is performed as follows:
A. If the victim is standing or sitting:
1. Stand behind the victim or the victims chair and wrap
your arms around his or her waist.
2. Grasp your fist with your other hand and place the fist
against the victims abdomen, slightly above the navel
and below the rib cage.
3. Press your fist into the victims abdomen with a quick
upward thrust.
B. If the victim is lying down:
1. Position the victim on his or her back.
2. Face the victim, and kneel on his or her hips.
3. With one of your hands on top of the other, place the
heel of your bottom hand on the abdomen, slightly
above the navel and below the rib cage.
4. Press into the victims abdomen with a quick upward
thrust.

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FIGURE 17.30 The abdominal thrust (Heimlich) maneuver.
People saved from drowning and victims of shock frequently experience apnea (cessation of breathing) and will soon
die if not revived by artificial respiration. The accepted treatment for reviving a person who has stopped breathing is illustrated in figure 17.31.
Common Respiratory Disorders

CHAPTER 17
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A cough is the most common symptom of respiratory disorders.

Acute problems may be accompanied by dyspnea or wheezing.
Respiratory or circulatory problems may cause cyanosis (si'a-no'sis), a blue discoloration of the skin resulting from blood with a
low oxygen content.
Although the common cold is the most widespread of all
respiratory diseases, there is still no cure for this ailmentonly
medications that offer symptomatic relief. Colds occur repeatedly
because acquired immunity for one virus does not protect against
other viruses. Cold viruses generally incite acute inflammation in
the respiratory mucosa, causing flow of mucus, sometimes accompanied by fever and/or headache.
Nearly all of the structures and regions of the respiratory
passageways can become infected and inflamed. Influenza is a
viral disease that causes an inflammatory condition of the
upper respiratory tract. Influenza can be epidemic, but fortu-
nately vaccines are available. Sinusitis is an inflammation of

the paranasal sinuses. Sinusitis can be quite painful if the
drainage ducts from the sinuses into the nasal cavity become
blocked. Tonsillitis may involve any or all of the tonsils and
frequently follows other lingering diseases of the oral or pharyngeal regions. Chronic tonsillitis sometimes requires a tonsillectomy. Laryngitis is inflammation of the larynx, which often
produces a hoarse voice and limits the ability to talk. Tracheobronchitis and bronchitis are infections of the regions for
which they are named. Severe inflammation in these areas can
cause smaller respiratory tubules to collapse, blocking the passage of air.
Diseases of the lungs are likewise common and usually serious. Pneumonia is an acute infection and inflammation of lung
tissue accompanied by exudation (accumulation of fluid). It is
usually caused by bacteria, most commonly by the pneumococcus
bacterium. Viral pneumonia is caused by a number of different
viruses. Tuberculosis is an inflammatory disease of the lungs
contracted by inhaling air sneezed or coughed by someone who is
carrying active tuberculosis bacteria. Tuberculosis softens lung
tissue, which eventually becomes ulcerated. Asthma is a disease
that affects people who are allergic to certain inhaled antigens. It
causes a swelling and blocking of lower respiratory tubes, often
accompanied by the formation of mucus plugs. Pleurisy (ploor'se) is an inflammation of the pleura and is usually secondary to
some other respiratory disease. Inspiration may become painful,
and fluid may collect within the pleural cavity. Emphysema
(em''f -se'ma) is a disease that causes the breakdown of the pulmonary alveoli, thus increasing the size of air spaces and decreasing the surface area (fig. 17.32). It is a frequent cause of death
among heavy cigarette smokers.
Cancer in the respiratory system is known to be caused by
the repeated inhalation of irritating substances, such as cigarette
smoke. Cancers of the lip, larynx, and lungs (fig. 17.33) are especially common in smokers over the age of 50.
Disorders of Respiratory Control

A variety of disease processes can result in cessation of breathing
during sleep, or sleep apnea. Sudden infant death syndrome
(SIDS) is an especially tragic form of sleep apnea that claims the
lives of about 10,000 babies annually in the United States. Victims of this condition are apparently healthy 2-to-5-month-old
babies who die in their sleep without apparent reasonhence,
the laypersons term, crib death. These deaths seem to be
caused by failure of the respiratory control mechanisms in the
brain stem and/or by failure of the carotid bodies to be stimulated
by reduced arterial oxygen.
tuberculosis: L. tuberculum, diminutive of tuber, swelling

cyanosis: Gk. kyanosis, dark-blue color
influenza: L. influentia, a flowing in
asthma: Gk. asthma, panting

emphysema: Gk. emphysan, blow up, inflate


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Respiratory System
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CHAPTER 17

Unit 6
646

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Accessory salivary
gland
Tongue
Accessory salivary
gland
Lingual frenulum
Opening of
submandibular
duct
Parotid gland
Sublingual ducts
Parotid duct
Masseter muscle
Sublingual
gland
Submandibular
gland
Submandibular
duct
Mandible (cut)
FIGURE 18.11 The salivary glands.
CHAPTER 18
Mucous cells
Intralobular
parotid duct
Lumen of
submandibular
intralobular duct
Seromucous
acini
(a)
Serous cells
(b)
Mucous cells
Serous cells
(c)
Intralobular
sublingual duct
FIGURE 18.12 The histology of the salivary glands. (a) The parotid gland, (b) the submandibular gland, and (c) the sublingual gland.


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Chapter 18
Digestive System
647
TABLE 18.4 Major Salivary Glands

Gland
Location
Duct
Entry into Oral Cavity
Type of Secretion
Parotid gland
Anterior and inferior to auricle;

subcutaneous over masseter
muscle
Parotid (Stensens) duct
Lateral to upper second molar
Watery serous fluid, salts,

and enzyme
Submandibular gland
Inferior to the base of the tongue
Submandibular (Whartons)
duct
Papilla lateral to lingual

frenulum
Watery serous fluid with

some mucus
Sublingual gland
Anterior to submandibular gland

under the tongue
Several small sublingual ducts

(Rivinus ducts)
Ducts along the base of the

tongue
Mostly thick, stringy

mucus; salts; and enzyme
(salivary amylase)
CHAPTER 18
FIGURE 18.13 A posterior view of the constrictor muscles of the pharynx. The right side has been cut away to illustrate the interior structures
in the pharynx.
and the lumen is lined with a mucous membrane containing

stratified squamous epithelium. The pharynx is divided into
three regions: the nasopharynx, posterior to the nasal cavity; the
oropharynx, posterior to the oral cavity; and the laryngopharynx,
at the level of the larynx (see fig. 17.3).
The external circular layer of pharyngeal muscles, called
constrictors (fig. 18.13), compresses the lumen of the pharynx
involuntarily during swallowing. The superior constrictor mus-
cle attaches to bony processes of the skull and mandible and encircles the upper portion of the pharynx. The middle constrictor
muscle arises from the hyoid bone and stylohyoid ligament and
encircles the middle portion of the pharynx. The inferior constrictor muscle arises from the cartilages of the larynx and encircles the lower portion of the pharynx. During breathing, the
lower portion of the inferior constrictor muscle is contracted,
preventing air from entering the esophagus.

648
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The motor and most of the sensory innervation to the

pharynx is via the pharyngeal plexus, situated chiefly on the
middle constrictor muscle. It is formed by the pharyngeal
branches of the glossopharyngeal and vagus nerves, together with
a deep sympathetic branch from the superior cervical ganglion.
The pharynx is served principally by ascending pharyngeal
arteries, which branch from the external carotid arteries. The
pharynx is also served by small branches from the inferior thyroid
arteries, which arise from the thyrocervical trunk. Venous return
is via the internal jugular veins.
Adventitia
Tunica
muscularis
Submucosa
Mucosa
Lumen
Knowledge Check
8. Define the terms heterodont and diphyodont. Which of the
four kinds of teeth is not found in deciduous dentition?
9. Where are the enamel, dentin, cementum, and pulp of a
tooth located? Explain how a tooth is anchored into its
socket.
10. Describe the location of the parotid, submandibular, and sublingual ducts and state where they empty into the oral cavity.
ESOPHAGUS AND STOMACH

A bolus of food is passed from the esophagus to the stomach,
where it is churned and mixed with gastric secretions. The chyme
thus produced is sent past the pyloric sphincter of the stomach to
the duodenum.
Objective 9
Describe the steps in deglutition.
Objective 10
Describe the location, gross structure, and

functions of the stomach.
Objective 11
CHAPTER 18
Describe the histological structure of the

esophagus and stomach. List the cell types in the gastric
mucosa, along with their secretory products.
Esophagus
The esophagus is that portion of the GI tract that connects the
pharynx to the stomach (see figs. 18.1 and 18.15). It is a collapsible tubular organ, approximately 25 cm (10 in.) long, originating
at the larynx and lying posterior to the trachea.
The esophagus is located within the mediastinum of the
thorax and passes through the diaphragm just above the opening
into the stomach. The opening through the diaphragm is called
the esophageal hiatus (e-sof''a-je'al hi-a'tus) The esophagus
is lined with a nonkeratinized stratified squamous epithelium
(fig. 18.14); its walls contain either skeletal or smooth muscle, depending on the location. The upper third of the esophagus contains skeletal muscle; the middle third, a combination of skeletal
and smooth muscle; and the terminal portion, smooth muscle only.
esophagus: Gk. oisein, to carry; phagema, food
FIGURE 18.14 The histology of the esophagus seen in cross

section.
The lower esophageal (gastroesophageal) sphincter is a

slight thickening of the circular muscle fibers at the junction of
the esophagus and the stomach. After food or fluid pass into the
stomach, this sphincter constricts to prevent the stomach contents from regurgitating into the esophagus. There is a normal
tendency for this to occur because the thoracic pressure is lower
than the abdominal pressure as a result of the air-filled lungs.
The lower esophageal sphincter is not a well-defined sphincter muscle comparable to others located elsewhere along the
GI tract, and it does at times permit the acidic contents of the stomach to enter the esophagus. This can create a burning sensation
commonly called heartburn, although the heart is not involved. In infants under a year of age, the lower esophageal sphincter may function erratically, causing them to spit up following meals. Certain
mammals, such as rodents, have a true lower esophageal sphincter
and cannot regurgitate, which is why poison grains are effective in
killing mice and rats.
Swallowing Mechanisms
Swallowing, or deglutition (de''gloo-tish'un), is the complex mechanical and physiological act of moving food or fluid from the
oral cavity to the stomach. For descriptive purposes, deglutition
is divided into three stages.
The first deglutitory stage is voluntary and follows mastication, if food is involved. During this stage, the mouth is closed
and breathing is temporarily interrupted. A bolus is formed as
the tongue is elevated against the transverse palatine folds
(palatal rugae) of the hard palate (see fig. 18.5) through contraction of the mylohyoid and styloglossus muscles and the intrinsic
muscles of the tongue.
The second stage of deglutition is the passage of the bolus
through the pharynx. The events of this stage are involuntary
and are elicited by stimulation of sensory receptors located at the
opening of the oropharynx. Pressure of the tongue against the
transverse palatine folds seals off the nasopharynx from the oral
cavity, creates a pressure, and forces the bolus into the oropharynx. The soft palate and pendulant palatine uvula are elevated to
close the nasopharynx as the bolus passes. The hyoid bone and


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Digestive System
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Esophagus
Esophagus
Fundus
Cardia
Peristaltic
contraction
of muscularis
layer of
esophagus
Lesser
curvature
Swallowed bolus
entering stomach
Greater
curvature
Duodenum
Pylorus
Body
Stomach
(a)
FIGURE 18.15 An anteroposterior radiograph of the esophagus

showing peristaltic contraction and movement of a bolus into the
stomach.
the larynx are also elevated. Elevation of the larynx against the
epiglottis seals the glottis so that food or fluid is less likely to
enter the trachea. Sequential contraction of the constrictor muscles of the pharynx moves the bolus through the pharynx to the
esophagus. This stage is completed in just a second or less.
The third stage, the entry and passage of food through the
esophagus, is also involuntary. The bolus is moved through the
esophagus by peristalsis (fig. 18.15). In the case of fluids, the entire process of deglutition takes place in slightly more than a second; for a typical bolus, the time frame is 5 to 8 seconds.
The stomachthe most distensible part of the GI tractis located in the upper left abdominal quadrant, immediately below
the diaphragm. Typically J-shaped when empty, the stomach is
continuous with the esophagus superiorly and empties into the
duodenal portion of the small intestine inferiorly (fig. 18.16). In
the stomach, which serves as a holding organ for ingested food,
the food is mechanically churned with gastric secretions to form
a pasty material called chyme (km). Once formed, chyme is
moved from the stomach to the small intestine.
The stomach is divided into four regions: the cardia, fundus, body, and pylorus (fig. 18.17). The cardia is the narrow
upper region immediately below the lower esophageal sphincter.
The fundus is the dome-shaped portion to the left of and in di-
chyme: L. chymus, juice

cardia: Gk. kardia, heart (upper portion, nearer the heart)
(b)
FIGURE 18.16 The stomach. (a) As seen from a cadaver, the

stomach is a J-shaped organ. (b) A radiograph of the stomach is of
clinical value for detecting ulcers, constrictions, or ingested objects.
A patient swallows radiopaque barium, which coats the lining of the
stomach and duodenum. These structures and certain abnormalities
may then show up in the radiograph.
rect contact with the diaphragm. The body is the large central
portion, and the pylorus is the funnel-shaped terminal portion.
The pyloric sphincter is the modified circular muscle at the end
of the pylorus, where it joins the small intestine. Pylorus is a
fundus: L. fundus, bottom

pylorus: Gk. pyloros, gatekeeper
CHAPTER 18
Stomach

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Gastric
folds
CHAPTER 18
FIGURE 18.17 The major regions and structures of the stomach.

Greek word meaning gatekeeper, and this junction is just that,
regulating the movement of chyme into the small intestine and
prohibiting backflow.
The stomach has two surfaces and two borders. The
broadly rounded surfaces are referred to as the anterior and posterior surfaces. The medial concave border is the lesser curvature (fig. 18.17), and the lateral convex border is the greater
curvature. The lesser omentum extends between the lesser curvature and the liver, and the greater omentum is attached to the
greater curvature (see fig. 18.3d).
The wall of the stomach is composed of the same four tunics found in other regions of the GI tract, with two principal
modifications: (1) an extra oblique muscle layer is present in the
muscularis, and (2) the mucosa is thrown into numerous longitudinal folds, called gastric folds or gastric rugae, which permit
stomach distension. The mucosa is further characterized by
the presence of microscopic gastric pits and gastric glands
(figs. 18.18 and 18.19).
There are five types of cells in the gastric glands that secrete specific products.
Goblet cells secrete protective mucus.
Parietal cells secrete hydrochloric acid (HCl).
Principal cells (chief cells) secrete pepsinogen, an inactive
form of the protein-digesting enzyme pepsin.
Columnar
epithelium of
mucosal ridge
Gastric pit
Gastric glands
with principal and
parietal cells
Lamina propria
FIGURE 18.18 The histology of the mucosa of the stomach.


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Chapter 18
Digestive System
651
Gastric pits
(a)
Mucous
cell
Mucosa
Gastric
gland
Parietal
cell
Submucosa
Principal
cell
(b)
FIGURE 18.19 Gastric pits and gastric glands of the stomach mucosa. (a) Gastric pits are the openings of the gastric glands. (b) Gastric
glands consist of mucous cells, principal cells, and parietal cells, each type producing a specific secretion.
Argentaffin (ar-jent'a-fin) cells secrete serotonin, histamine, and autocrine regulators.
Endocrine cells (G cells) secrete the hormone gastrin into
the blood.
The stomach is sensitive to emotional stress. Mucus, secreted

by mucous cells of the stomach, is important in preventing hydrochloric acid and the digestive enzyme pepsin from eroding the
stomach wall. Peptic ulcers may be caused by an increase in cellular
secretion or by insufficient secretions of protective mucus. Another
protective feature is the rapid mitotic activity of the columnar epithelium of the stomach. The entire lining of the stomach is usually replaced every few days. Nevertheless, in the United States
approximately 10% of the male population and 4% of the female population develop peptic ulcers.
Regulation of gastric activity is autonomic. The sympathetic neurons arise from the celiac plexus, and the parasympathetic neurons arise from the vagus nerves. Parasympathetic
neurons synapse in the myenteric plexus between the muscular
layers and in the submucosal plexus in the submucosa. Parasymargentaffin: L. argentum, silver; affinis, attraction (become colored with
silver stain)
Phase
Response
Cephalic phase
Sight, taste, small or mental stimuli evoke

parasympathetic response via vagus nerves; 50150
ml of gastric juice is secreted
Gastric phase
Food in the stomach stretches the mucosa, and

chemical breakdown of protein stimulates the release
of gastrin; gastrin stimulates the production of
600750 ml of gastric juice
Intestinal phase
Chyme entering the duodenum stimulates intestinal

cells to release intestinal gastrin; intestinal gastrin
stimulates the production of additional small
quantities of gastric juice
pathetic impulses promote gastric activity, the phases of which

are presented in table 18.5.
Vomiting is the reflexive response of emptying the stomach
through the esophagus, pharynx, and oral cavity. This action is
controlled by the vomiting center of the medulla oblongata. Stimuli within the GI tract, especially the duodenum, may activate the
vomiting center, as may nauseating odors or sights, motion sickness, or body stress. Various drugs called emetics can also stimulate
a vomiting reflex. The mechanics of vomiting are as follows:
(1) strong, sustained contractions of the upper small intestine, followed by a contraction of the pyloric sphincter; (2) relaxation of
CHAPTER 18
In addition to these products, the gastric mucosa (probably

the parietal cells) secretes intrinsic factor, a polypeptide that is required for absorption of vitamin B12 in the small intestine. Continued gastric activity when the stomach is empty causes hunger
sensations known as hunger pangs. Eating fills the stomach resulting in hunger satiety, or a perception of fullness.
TABLE 18.5 Phases of Gastric Secretion

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the lower esophageal sphincter and contraction of the pyloric

portion of the stomach; (3) a shallow inspiration and closure of
the glottis; and (4) compression of the stomach against the liver
by contraction of the diaphragm and the abdominal muscles.
This reflexive sequence causes a forceful ejection of vomit. The
feeling of nausea is caused by stimuli in the vomiting center and
may or may not cause vomiting.
The only function of the stomach that appears to be essential
for life is the secretion of intrinsic factor. This polypeptide is
needed for the intestinal absorption of vitamin B12, which is required
for maturation of red blood cells in the bone marrow. Following surgical removal of the stomach (gastrectomy), a patient has to receive vitamin B12 (together with intrinsic factor) orally or through injections,
so that he or she will not develop pernicious anemia.
Knowledge Check
11. Describe the three stages of deglutition with reference to
the structures involved.
12. Describe the structure and function of the lower
esophageal sphincter.
13. List the functions of the stomach. What is the function of
the gastric folds?
14. Describe the modifications of the stomach that aid in mechanical and chemical digestion.
SMALL INTESTINE
The small intestine, consisting of the duodenum, jejunum, and
ileum, is the site where digestion is completed and nutrients are
absorbed. The surface area of the intestinal wall is increased by
plicae circulares, intestinal villi, and microvilli.
Objective 12
Describe the location and regions of the small

intestine and the way in which it is supported.
CHAPTER 18
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Objective 13
List the functions of the small intestine and

describe the structural adaptations through which these
functions are accomplished.
Objective 14
Describe the movements that occur within the

small intestine.
The small intestine is that portion of the GI tract between the

pyloric sphincter of the stomach and the ileocecal valve that
opens into the large intestine. It is positioned in the central and
lower portions of the abdominal cavity and is supported, except
for the first portion, by the mesentery (fig. 18.20). The fanshaped mesentery permits movement of the small intestine but
leaves little chance for it to become twisted or kinked. Enclosed
within the mesentery are blood vessels, nerves, and lymphatic
vessels that supply the intestinal wall.
The small intestine is approximately 3 m (12 ft) long and
2.4 cm (1 in.) wide in a living person, but it will measure nearly
twice this length in a cadaver when the muscular wall is relaxed.
It is called the small intestine because of its relatively small diameter compared to that of the large intestine. The small intestine is the bodys major digestive organ and the primary site of
nutrient absorption. Its digestive enzymes, along with those of
the salivary glands, gastric glands, and pancreas, are summarized
in table 18.6.
The small intestine is innervated by the superior mesenteric plexus. The branches of the plexus contain sensory fibers,
postganglionic sympathetic fibers, and preganglionic parasympathetic fibers. The arterial blood supply to the small intestine is
through the superior mesenteric artery and branches from the
celiac trunk and the inferior mesenteric artery. The venous
drainage is through the superior mesenteric vein. This vein
unites with the splenic vein to form the hepatic portal vein,
which carries nutrient-rich blood to the liver (see fig. 16.41).
Regions of the Small Intestine

On the basis of function and histological structure, the small intestine is divided into three regions.
1. The duodenum (doo''o-de'num, doo-od'e-num) is a relatively fixed, C-shaped tubular organ, measuring approximately 25 cm (10 in.) from the pyloric sphincter of the
stomach to the duodenojejunal (doo-od''e-no''je-joo'nal)
flexure. Except for a short portion near the stomach, the
duodenum is retroperitoneal. Its concave surface faces to
the left, where it receives bile secretions from the liver and
gallbladder through the common bile duct and pancreatic
secretions through the pancreatic duct of the pancreas
(fig. 18.21). These two ducts unite to form a common
entry into the duodenum called the hepatopancreatic ampulla (ampulla of Vater), which pierces the duodenal wall
and drains into the duodenum from an elevation called the
duodenal papilla. It is here that bile and pancreatic juice
enter the small intestine. The duodenal papilla can be
opened or closed by the action of the sphincter of ampulla
(of Oddi). The duodenum differs histologically from the
rest of the small intestine by the presence of duodenal
(Brunners) glands in the submucosa (fig. 18.22). These
compound tubuloalveolar glands secrete mucus and are
most numerous near the superior end of the duodenum.
2. The jejunum (je-joo'num), which extends from the duodenum to the ilium, is approximately 1 m (3 ft) long. It has
a slightly larger lumen and more internal folds than does
the ileum, but its histological structure is similar to that of
the ileum.
duodenum: L. duodeni, 12 each (length of 12 fingers breadth)

ampulla of Vater: from Abraham Vater, German anatomist, 16841751
sphincter of Oddi: from Rugger Oddi, Italian physician, 18641913
Brunners glands: from Johann C. Brunner, Swiss anatomist, 16531727


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Digestive System
653
Duodenum
Jejunum
Intestinal
v
Ileum
Tunica
muscularis
FIGURE 18.20 The small intestine in relation to the stomach and a part of the large intestine. (a) The regions and mesenteric attachment. (b)
A section of the intestinal wall showing the mucosa and submucosa folded into structures called plicae circulares. (The regions of the small intestine are labeled in boldface type.)
Structural Modifications
The products of digestion are absorbed across the epithelial lining of the intestinal mucosa. Absorption occurs primarily in the
jejunum, although some also occurs in the duodenum and ileum.
Peyers patches: from Johann K. Peyer, Swiss anatomist, 16531712
Absorption occurs at a rapid rate as a result of four specializations

that increase the intestinal surface area.
The three meter length of the small intestine.
The plicae (pli'se) circulares are large macroscopic folds of
mucosa (see fig. 18.20).
The intestinal villi (vil'e) are fingerlike macroscopic folds
of the mucosa that project into the lumen of the small intestine (fig. 18.23).
The microvilli are microscopic projections formed by the
folding of each epithelial cell membrane. In a light microscope, the microvilli display a somewhat vague brush
plica: L. plicatus, folded

villus: L. villosus, shaggy
CHAPTER 18
3. The ileum (il'e-um)not to be confused with the ilium of

the os coxaemakes up the remaining 2 m (67 ft) of the
small intestine. The terminal portion of the ileum empties
into the medial side of the cecum through the ileocecal
valve. Lymph nodules, called mesenteric (Peyers) patches,
are abundant in the walls of the ileum.

Unit 6
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TABLE 18.6 Digestive Enzymes

Enzyme
Source
Digestive Action
Salivary glands
Begins carbohydrate digestion by converting starch and glycogen to disaccharides
Gastric glands
Begins the digestion of nearly all types of proteins
Peptidase
Intestinal glands
Converts proteins into amino acids
Sucrase
Intestinal glands
Converts disaccharides into monosaccharides
Lipase
Intestinal glands
Amylase
Intestinal glands
Nuclease
Intestinal glands
Enterokinase
Intestinal glands
Activates trypsin
Salivary enzyme
Amylase
Gastric juice enzyme
Pepsin
Intestinal juice enzymes
Maltase
Lactase
Pancreatic juice enzymes

Amylase
Pancreas
Lipase
Pancreas
Peptidases
Pancreas
Convert proteins or partially digested proteins into amino acids
Pancreas
Trypsin
Chymotrypsin
Carboxypeptidase
Nuclease
Common bile duct
CHAPTER 18
Accessory pancreatic duct
Pancreatic duct
Duodenum
Sphincter
of ampulla
Pancreas
Hepatopancreatic ampulla
Duodenal papilla
FIGURE 18.21 A section of the duodenum showing the location of entry of the common bile duct and the pancreatic duct.


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Chapter 18
Digestive System
655
Simple
columnar
epithelium
Intestinal villus
(lined with simple
columnar
epithelium)
Lacteal
Lamina propria
Intestinal
villus
Capillary
network
Goblet
cells
Muscularis
mucosa
Intestinal
crypt
Duodenal glands
Lymph
vessel
FIGURE 18.22 The histology of the duodenum.
Arteriole
border on the edges of the columnar epithelium

(fig. 18.24). The terms brush border and microvilli are
often used interchangeably in describing the small intestine.
Venule
FIGURE 18.23 The structure of an intestinal villus and intestinal

crypt.
CHAPTER 18
The intestinal villi are covered with columnar epithelial

cells, among which are interspersed the mucus-secreting goblet
cells. The lamina propria, which forms the connective tissue core
of each intestinal villus, contains numerous lymphocytes, blood
capillaries, and a lymphatic vessel called the lacteal (lak'te-al)
(fig. 18.23). Absorbed monosaccharides and amino acids enter
the blood capillaries; absorbed fatty acids and cholesterols enter
the lacteals. Intestinal villi are considered the functional units of
the digestive system because absorption through these structures
is how digested molecules enter the blood or lymph.
Epithelial cells at the tips of the intestinal villi are continuously shed and are replaced by cells that are pushed up from the
bases of the intestinal villi. The epithelium at the base of the intestinal villi invaginates downward at various points to form narrow pouches that open through pores into the intestinal lumen.
These structures are called the intestinal crypts (crypts of
Lieberkhn) (see fig. 18.23).
Waldrop
Mechanical Activities
Contractions of the longitudinal and circular muscles of the
small intestine produce three distinct types of movement: rhythmic segmentation, pendular movements, and peristalsis.
lacteal: L. lacteus, milk
crypts of Lieberkhn: from Johann N. Lieberkhn, German anatomist, 171156
FIGURE 18.24 An electron photomicrograph of microvilli

(arrow) at the exposed surface of a columnar epithelial cell in the
small intestine.

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Rhythmic segmentations are local contractions of the circular muscular layer. They occur at the rate of about 12 to 16 per
minute in regions containing chyme. Rhythmic segmentations
churn the chyme with digestive juices and bring it into contact
with the mucosa. During these contractions, the vigorous motion
of the intestinal villi stirs the chyme and facilitates absorption.
Pendular movements primarily occur in the longitudinal
muscle layer. In this motion, a constrictive wave moves along a
segment of the small intestine and then reverses and moves in
the opposite direction, moving the chyme back and forth. Pendular movements also mix the chyme but do not seem to have a
particular frequency.
Peristalsis (per''-stal'sis) is responsible for the propulsive
movement of the chyme through the small intestine. These
wavelike contractions are usually weak and relatively short, occurring at a frequency of about 15 to 18 per minute. Chyme requires 3 to 10 hours to travel the length of the small intestine.
Both muscle layers are involved in peristalsis.
The sounds of digestive peristalsis can be easily heard
through a stethoscope placed at various abdominal locations.
These sounds can be detected even through clothing. The sounds,
mostly clicks and gurgles, occur at a frequency of 5 to 30 per minute.
Knowledge Check
15. Describe the small intestine. Where is it located? How is it
subdivided? How is it supported?
16. What are the primary functions of the small intestine?
17. List four structural modifications of the small intestine that
increase its absorptive surface area.
18. Describe the movements of the small intestine. Which
movements are produced by the circular layer of the tunica
muscularis?
19. Which region of the small intestine is the longest? Which
is the shortest? How long does it take a portion of chyme to
move through the small intestine?
CHAPTER 18
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LARGE INTESTINE
The large intestine receives undigested food from the small intestine, absorbs water and electrolytes from chyme, and passes
feces out of the GI tract.
Objective 15
Identify the regions of the large intestine and

describe its gross and histological structure.
orly on the right side to a point just below the liver; it then
crosses to the left, descends into the pelvis, and terminates at the
anus. A specialized portion of the mesentery, the mesocolon,
supports the transverse portion of the large intestine along the
posterior abdominal wall.
The large intestine has little or no digestive function, but
it does absorb water and electrolytes from the remaining chyme.
In addition, the large intestine functions to form, store, and
expel feces from the body.
Regions and Structures

of the Large Intestine
The large intestine is structurally divided into the cecum, colon,
rectum, and anal canal (figs. 18.25 and 18.26). The cecum
(se'kum) is a dilated pouch positioned slightly below the ileocecal
valve. The ileocecal valve is a fold of mucous membrane at the
junction of the small intestine and large intestine that prohibits
the backflow of chyme. A fingerlike projection called the appendix is attached to the inferior medial margin of the cecum. The
8 cm (3 in.) appendix contains an abundance of lymphatic tissue
(fig. 18.27) that may serve to resist infection. Although the appendix serves no digestive function, it is thought to be a vestigial
remnant of an organ that was functional in human ancestors.
A common disorder of the large intestine is inflammation of the
appendix, or appendicitis. Wastes that accumulate in the appendix cannot be moved easily by peristalsis, because the appendix
has only one opening. Although the symptoms of appendicitis are
quite variable, they often include a high white blood cell count, localized pain in the lower right quadrant, and loss of appetite. Vomiting
may or may not occur. Rupture of the appendix (a burst appendix)
spreads infectious material throughout the peritoneal cavity, resulting
in peritonitis.
The superior portion of the cecum is continuous with the

colon, which consists of ascending, transverse, descending, and
sigmoid portions (fig. 18.25). The ascending colon extends superiorly from the cecum along the right abdominal wall to the inferior surface of the liver. Here the colon bends sharply to the left
at the hepatic flexure (right colic flexure) and continues across
the upper abdominal cavity as the transverse colon. At the left
abdominal wall, another right-angle bend called the splenic flexure (left colic flexure) marks the beginning of the descending
colon. From the splenic flexure, the descending colon extends
inferiorly along the left abdominal wall to the pelvic region. The
colon then angles medially from the brim of the pelvis to form an
S-shaped bend, known as the sigmoid colon.
Objective 16
Describe the functions of the large intestine

and explain how defecation is accomplished.
The large intestine averages 1.5 m (5 ft) in length and 6.5 cm

(2.5 in.) in diameter. It is called the large intestine because of
its relatively large diameter compared to that of the small intestine. The large intestine begins at the end of the ileum in the
lower right quadrant of the abdomen. From there, it leads superi-
cecum: L. caecum, blind pouch

appendix: L. appendix, attachment
colon: Gk. kolon, member of the whole
sigmoid: Gk. sigmoeides, shaped like a sigma,


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Chapter 18
Digestive System
657
Transverse colon
Hepatic flexure
Splenic flexure
Mesocolon
Taeniae coli
Ascending colon
Descending colon
Epiploic appendage
Ileum
Ileocecal valve
Orifice of appendix
Haustrum
Cecum
Appendix
Sigmoid colon
Rectum
Loechel
Anal canal
FIGURE 18.25 The large intestine.
A hemorrhoid (hem'o-rod) is a mass of varicose veins in the

anal area caused, in part, by difficulty in defecating. Hemorrhoids, in reference to the condition in which such masses occur, are
also called piles. A first-degree hemorrhoid is contained within the
anal canal. A second-degree hemorrhoid prolapses, or extends out-
rectum: L. rectum, straight tube

anus: L. anus, ring
ward during defecation. A third-degree hemorrhoid remains prolapsed through the anal orifice. Rubber band constriction is a common medical treatment for a prolapsed hemorrhoid. In this technique,
a rubber band is tied around the hemorrhoid, constricting its blood
supply, so that the tissue dries and falls off. In a relatively new treatment, infrared photocoagulation, a high-energy light beam coagulates the hemorrhoid.
Although the large intestine consists of the same tunics as

the small intestine, there are some structural differences. The large
intestine lacks intestinal villi but does have numerous goblet cells
in the mucosal layer (fig. 18.29). The longitudinal muscle layer of the
muscularis forms three distinct muscle bands called taeniae coli
(te'ne-e ko'li) that run the length of the large intestine. A series of
bulges in the walls of the large intestine form sacculations, or haustra
(haws'tra), along its entire length (see figs. 18.25 and 18.26).
taenia: L. tainia, a ribbon

haustrum: L. haustrum, bucket or scoop
CHAPTER 18
The terminal 20 cm (7.5 in.) of the GI tract is the rectum,

and the last 2 to 3 cm of the rectum is referred to as the anal
canal (fig. 18.28). The rectum lies anterior to the sacrum, where
it is firmly attached by peritoneum. The anus is the external
opening of the anal canal. Two sphincter muscles guard the anal
opening: the internal anal sphincter, which is composed of
smooth muscle fibers, and the external anal sphincter, composed of skeletal muscle. The mucous membrane of the anal
canal is arranged in highly vascular, longitudinal folds called
anal columns.

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Rectum
Levator ani
muscle
Hemorrhoidal
vessels
Anal canal
External anal
sphincter
Anal columns
Internal anal
sphincter
Anus
Loechel
FIGURE 18.28 The anal canal.
Serosa
FIGURE 18.26 An anteroposterior radiograph after a barium
Tunica
muscularis
enema showing the regions, flexures, and the haustra of the large
intestine.
Submucosa
Mucosa
Center of
lymphatic
nodule
CHAPTER 18
Intestinal
glands
Columnar
epithelium with
goblet cells
(a)
Lumen
Tunica
muscularis
Submucosa
Serosa
FIGURE 18.27 The histology of the appendix shown in cross section.
(b)
FIGURE 18.29 The histology of the rectum. (a) A photomicrograph of the tunics. (b) A scanning electron photomicrograph of the
mucosa from a section of the colon. The arrow indicates the opening
of a goblet cell into the intestinal lumen.
(From R.G. Kessel and R.H. Kardon. Tissues and Organs: A Text-Atlas of Scanning
Electron Microscopy, 1979 W.H. Freeman and Company.)


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Chapter 18
Digestive System
659
TABLE 18.7 Mechanical Activity in the GI Tract

Region
Type of Motility
Frequency
Stimulus
Result
Oral cavity
Mastication
Variable
Initiated voluntarily; proceeds

reflexively
Pulverization: mixing with

saliva
Oral cavity and

pharynx
Deglutition
Maximum of 20 per min
Initiated voluntarily; reflexively

controlled by swallowing center
Clears oral cavity of food
Esophagus
Peristalsis
Depends on frequency of swallowing
Initiated by swallowing
Movement through the

esophagus
Stomach
Small intestine
Large intestine
Receptive relaxation
Matches frequency of swallowing
Unknown
Permits filling of stomach
Tonic contraction
1520 per min
Autonomic plexuses
Mixing and churning
Peristalsis
12 per min
Autonomic plexuses
Evacuation of stomach
Hunger contractions
3 per min
Low blood sugar level
Feeding
Peristalsis
1518 per min
Autonomic plexuses
Transfer through intestine
Rhythmic segmentation
1216 per min
Autonomic plexuses
Mixing
Pendular movements
Variable
Autonomic plexuses
Mixing
Peristalsis
312 per min
Autonomic plexuses
Transport
Mass movements
23 per day
Stretch
Fills sigmoid colon
Haustral churning
312 per min
Autonomic plexuses
Mixing
Defecation
Variable: 1 per day to 3 per week
Reflex triggered by rectal distension
Defecation
Mechanical Activities of the Large Intestine

Chyme enters the large intestine through the ileocecal valve. About
15 ml of pasty material enters the cecum with each rhythmic opening of the valve. The ingestion of food intensifies peristalsis of the
ileum and increases the frequency with which the ileocecal valve
opens; this is called the gastroileal reflex. Material entering the
large intestine accumulates in the cecum and ascending colon.
Three types of movements occur throughout the large intestine: peristalsis, haustral churning, and mass movement. Peristaltic movements of the colon are similar to those of the small
intestine, although they are usually more sluggish in the colon.
In haustral churning, a relaxed haustrum fills with food residues
until a point of distension is reached that stimulates the muscuepiploic: Gk. epiplein, to float on
laris to contract. Besides moving the material to the next haustrum, this contraction churns the material and exposes it to the
mucosa, where water and electrolytes are absorbed. As a result of
water absorption, the material becomes solid or semisolid, and is
now known as feces (fe'sez). Mass movement is a very strong
peristaltic wave, involving the action of the taeniae coli, which
moves the fecal material toward the rectum. Mass movements
generally occur only two or three times a day, usually during or
shortly after a meal. This response to eating, called the gastrocolic reflex, can best be observed in infants who have a bowel
movement during or shortly after feeding.
As material passes through the large intestine, Na+ K+, and
water are absorbed. It has been estimated that an average volume of
850 ml of water per day is absorbed across the mucosa of the colon.
The fecal material that is left then passes to the rectum, leading to
an increase in rectal pressure and the urge to defecate. If the urge to
defecate is denied, feces are prevented from entering the anal canal
by the external anal sphincter. In this case, the feces remain in the
rectum and may even back up into the sigmoid colon.
The defecation reflex normally occurs when the rectal
pressure rises to a particular level that is determined largely by
habit. At this point, the internal anal sphincter relaxes to admit
feces into the anal canal.
During the act of defecation, the longitudinal rectal muscles contract to increase rectal pressure, and the internal and external anal sphincter muscles relax. Excretion is aided by
contractions of abdominal and pelvic skeletal muscles, which
raise the intraabdominal pressure and help push the feces from
the rectum through the anal canal and out the anus.
The various mechanical activities of the GI tract are summarized in table 18.7.
CHAPTER 18
Finally, the large intestine has small but numerous fat-filled

pouches called epiploic (ep- -plo'ik) appendages (see
fig. 18.25) that are attached superficially to the taeniae coli.
The sympathetic innervation of the large intestine arises
from superior and inferior mesenteric plexuses, as well as from
the celiac plexus. The parasympathetic innervation arises from
the paired pelvic splanchnic and vagus nerves. Sensory fibers
from the large intestine respond to bowel pressure and signal the
need to defecate. Blood is supplied to the large intestine by
branches from the superior mesenteric and inferior mesenteric
arteries. Venous blood is returned through the superior and inferior mesenteric veins, which in turn drain into the hepatic portal
vein that enters the liver.

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Visceral peritoneum
Parietal peritoneum
Spinal cord
Vertebra
Plane of
section
Right kidney
Left kidney
Rib
Spleen
Small intestine
Waldrop
Liver
Large intestine
Inferior
vena cava
Stomach
Peritoneal cavity
Gallbladder
Aorta
Pancreas
FIGURE 18.30 A cross section of the abdomen showing the relative position of the liver to other abdominal organs.
Constipation occurs when fecal material accumulates because

of longer than normal periods between defecations. The
slower rate of elimination allows more time for water absorption, so
that the waste products become harder. Although uncomfortable and
sometimes painful, this condition is usually not serious. Diarrhea occurs when waste material passes too quickly through the colon, so
that insufficient time is allowed for water absorption. Excessive diarrhea can result in dangerous levels of dehydration and electrolyte imbalance, particularly in infants because of their small body size.
CHAPTER 18
Knowledge Check
20. Identify the four principal regions of the large intestine and
describe the functions of the colon.
21. Describe the haustra and the taeniae coli and explain their
role in the movements of the large intestine.
22. Describe the location of the rectum, anal canal, and
anal sphincter muscles and explain how defecation is
accomplished.
LIVER, GALLBLADDER,
AND PANCREAS
The liver, consisting of four lobes, processes nutrients and secretes bile, which is stored and concentrated in the gallbladder
prior to discharge into the duodenum. The pancreas, consisting of
endocrine (islet) cells and exocrine (acini) cells, secretes important hormones into the blood and essential digestive enzymes into
the duodenum.
Objective 17
of the liver.
Objective 18
Describe the location of the gallbladder and

trace the flow of bile through the systems of ducts into the
duodenum.
Objective 19

of the pancreas.
Three accessory digestive organs in the abdominal cavity aid in

the chemical breakdown of food. These are the liver, gallbladder
and pancreas. The liver and pancreas function as exocrine glands
in this process because their secretions are transported to the
lumen of the GI tract via ducts.
Liver
The liver is the largest internal organ of the body, weighing about
1.3 kg (3.54.0 lbs) in an adult. It is positioned immediately beneath the diaphragm in the epigastric and right hypochondriac
regions (see fig. 2.15 and table 2.4) of the abdomen (fig. 18.30).
Its reddish brown color is due to its great vascularity.
The liver has four lobes and two supporting ligaments. Anteriorly, the right lobe is separated from the smaller left lobe by
the falciform ligament (fig. 18.31). Inferiorly, the caudate
falciform: L. falcis, sickle; forma, form


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Chapter 18
Gallbladder
Inferior vena cava
Digestive System
661
Quadrate lobe
Right lobe
Coronary ligament
Ligamentum teres
Left lobe
Cystic
duct
Hepatic
duct
Hepatic
artery
Right lobe
Left lobe
Falciform
ligament
Hepatic
portal
vein
Ligamentum teres
Inferior vena cava
Gallbladder
Quadrate lobe
Falciform ligament
Caudate lobe
Common bile duct
Hepatic artery
Left lobe
Right lobe
Common bile
duct
Inferior vena
cava
Left
triangular
ligament
Hepatic portal
Caudate lobe
vein
Ligamentum
venosum
(c)
(The lobes of the liver are labeled in boldface type.)
(kaw'dt) lobe is positioned near the inferior vena cava, and the
quadrate lobe is adjacent to the gallbladder. The falciform ligament attaches the liver to the anterior abdominal wall and the
diaphragm. The ligamentum teres (round ligament) extends
from the falciform ligament to the umbilicus. This ligament is
the remnant of the umbilical vein of the fetus (see table 16.6).
Although the liver is the largest internal organ, it is, in a
sense, only one to two cells thick. This is because the liver cells,
or hepatocytes, form hepatic plates that are one to two cells
thick and separated from each other by large capillary spaces
called liver (hepatic) sinusoids (fig. 18.32). The sinusoids are
lined with phagocytic Kupffer (koop'fer) cells, but the large intercellular gaps between adjacent Kupffer cells make these sinusoids more highly permeable than other capillaries. The plate
structure of the liver and the high permeability of the sinusoids
allow each hepatocyte to be in direct contact with the blood.
The hepatic plates are arranged to form functional units
called liver lobules (fig. 18.33). In the middle of each lobule is a
central vein, and at the periphery of each lobule are branches of
the hepatic portal vein and of the hepatic artery, which open
into the spaces between hepatic plates. Portal venous blood,
containing molecules absorbed in the GI tract, thus mixes with
hepatic: Gk. hepatos, liver
Kupffer cells: from Karl Wilhelm von Kupffer, Bavarian anatomist and
embryologist, 18291902
CHAPTER 18
FIGURE 18.31 The liver and gallbladder. (a) An anterior view and (b) a rotated inferior view, and (c) a photograph of a rotated inferior view.

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Branch of hepatic portal vein

Hepatic triad
Biliary ductule
Hepatic plate
Creek
Hepatic plate
Central vein
Liver (hepatic) sinusoid

Intralobular biliary ductule
Liver (hepatic)
sinusoids
(a)
Branch of hepatic
portal vein
Branch of
hepatic artery
Biliary ductule
Biliary canaliculi
Liver (hepatic)
sinusoid
(c)
Connective tissue
sheath
(b)
CHAPTER 18
FIGURE 18.32 A liver lobule and the histology of the liver. (a) A cross section of a liver lobule and (b) a longitudinal section. Blood enters a
liver lobule through the vessels in a hepatic triad, passes through hepatic sinusoids, and leaves the lobule through a central vein. The central
veins converge to form hepatic veins that transport venous blood from the liver. (c) A photomicrograph of a liver lobule in cross section.
arterial blood as the blood flows within the liver sinusoids from the
periphery of the lobule to the central vein (fig. 18.33). The central
veins of different liver lobules converge to form the hepatic vein,
which carries blood from the liver to the inferior vena cava.
The liver lobules have numerous functions, including synthesis, storage, and release of vitamins; synthesis, storage, and release of glycogen; synthesis of blood proteins; phagocytosis of old
red blood cells and certain bacteria; removal of toxic substances;
and production of bile. Bile is stored in the gallbladder and is
eventually secreted into the duodenum for the emulsification
(breaking down into smaller particles) and absorption of fats.
Bile is produced by the hepatocytes and secreted into tiny
channels called bile canaliculi (kan''a-lik'yu-li) located within each
hepatic plate (fig. 18.33). These bile canaliculi are drained at the
periphery of each lobule by bile ducts, which in turn drain into hepatic ducts that carry bile away from the liver. Because blood travels in the sinusoids and bile travels in the opposite direction within
the hepatic plates, blood and bile do not mix in the liver lobules.
The liver receives parasympathetic innervation from the
vagus nerves and sympathetic innervation from thoracolumbar
nerves through the celiac ganglia.
Gallbladder
The gallbladder is a saclike organ attached to the inferior surface
of the liver (figs. 18.31 and 18.34). This organ stores and concentrates bile. A sphincter valve at the neck of the gallbladder allows
a storage capacity of about 35 to 50 ml. The inner mucosal layer
of the gallbladder is thrown into folds similar to the gastric folds
of the stomach. When the gallbladder fills with bile, it expands
to the size and shape of a small pear. Bile is a yellowish green
fluid containing bile salts, bilirubin (a product resulting from the
breakdown of blood), cholesterol, and other compounds. Contraction of the muscularis ejects bile from the gallbladder.
Bile is continuously produced by the liver and drains
through the hepatic ducts and common bile duct to the duodenum. When the small intestine is empty of food, the sphincter of
ampulla (see fig. 18.21) constricts, and bile is forced up the cystic
duct to the gallbladder for storage.
The gallbladder is supplied with blood from the cystic
artery, which branches from the right hepatic artery. Venous
blood is returned through the cystic vein, which empties into the
hepatic portal vein. Autonomic innervation of the gallbladder is


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Chapter 18
Digestive System
663
Branch of portal vein

Liver (hepatic)
sinusoids
Bile canaliculus
Biliary ductule
Loechel
Central vein
Hepatic plate
FIGURE 18.33 The flow of blood and bile in a liver lobule. Blood flows within sinusoids from branches of the hepatic portal vein to the central
vein (from the periphery to the center of a lobule). Bile flows within hepatic plates from the center of a lobule to biliary ductules at the periphery.
similar to that of the liver; both receive parasympathetic innervation from the vagus nerves and sympathetic innervation from
thoracolumbar nerves through the celiac ganglia.
Pancreas
The soft, lobulated pancreas is known as a mixed gland because it
has both exocrine and endocrine functions. The endocrine function is performed by clusters of cells called the pancreatic islets
(islets of Langerhans). The islet cells secrete the hormones insulin and glucagon into the blood. As an exocrine gland, the pancreas secretes pancreatic juice through the pancreatic duct
(fig. 18.36), which empties into the duodenum.
FIGURE 18.34 The pancreatic duct joins the bile duct to empty its
secretions through the duodenal papilla into the duodenum. The release of bile and pancreatic juice into the duodenum is controlled by
the sphincter of ampulla (see fig. 18.21).
pancreas: Gk. pan, all; kreas, flesh

islets of Langerhans: from Paul Langerhans, German anatomist, 184788
CHAPTER 18
A common clinical problem of the gallbladder is the development of gallstones. Bile is composed of various salts, pigments, and cholesterols that become concentrated as water is
removed. Cholesterols normally remain in solution, but under certain
conditions they precipitate to form solid crystals. Large crystals may
block the bile duct and have to be surgically removed. The radiograph in figure 18.35 shows gallstones in position, and the photograph shows removed gallstones.

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(a)
(b)
FIGURE 18.35 (a) A radiograph of a gallbladder that contains gallstones (biliary calculi). (b) Following surgical removal of the gallbladder
(cholecystectomy), it has been cut open to reveal its gallstones. (Note their size relative to that of a dime.)
CHAPTER 18
Pancreatic
acinar cells
Pancreatic
islet cells
Pancreatic
duct
Tail of pancreas
Body of pancreas
Head of pancreas
FIGURE 18.36 The pancreas is both an exocrine and an endocrine gland. Pancreatic juicethe exocrine productis secreted by acinar
cells into the pancreatic duct. Scattered islands of cells, called pancreatic islets (islets of Langerhans), secrete the hormones insulin and
glucagon into the blood.


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The Digestive System
EXPLANATION
The entire digestive system develops from modifications of an
elongated tubular structure called the primitive gut. These modifications are initiated during the fourth week of embryonic development. The primitive gut is composed solely of endoderm and
for descriptive purposes can be divided into three regions: the
foregut, midgut, and hindgut (exhibit I).
Foregut
The stomodeum (sto''mo-de'um), or oral pit, is not part of the
foregut but an invagination of ectoderm that breaks through a thin
oral membrane to become continuous with the foregut and form
part of the oral cavity, or mouth. Structures in the mouth, therefore, are ectodermal in origin. The esophagus, pharynx, stomach, a
stomodeum: Gk. stoma, mouth; hodaios, on the way to
EXHIBIT I A sagittal section of a 5-week-old embryo showing the development of the digestive system and its association with the extraembryonic membranes and organs.
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EXHIBIT II Progressive stages of development of the foregut to form the stomach, duodenum, liver, gallbladder, and pancreas:
(a) 4 weeks, (b) 5 weeks, (c) 6 weeks, and (d) 7 weeks.
portion of the duodenum, the pancreas, liver, and gallbladder are

the organs that develop from the foregut (exhibit II). Along the GI
tract, only the inside epithelial lining of the lumen is derived from
the endoderm of the primitive gut. The vascular portion and
smooth muscle layers are formed from mesoderm that develops
from the surrounding splanchnic mesenchyme.
The stomach first appears as an elongated dilation of the
foregut. The dorsal border of the stomach undergoes more rapid
growth than the ventral border, forming a distinct curvature. The
caudal portion of the foregut and the cranial portion of the
midgut form the duodenum. The liver and pancreas arise from
the wall of the duodenum as small hepatic and pancreatic buds,
respectively. The hepatic bud experiences incredible growth to
form the gallbladder, associated ducts, and the various lobes of
the liver (exhibit II). By the sixth week, the liver is carrying out
hemopoiesis (the formation of blood cells). By the ninth week,
666
the liver has developed to the point where it represents 10% of

the total weight of the fetus.
The pancreas develops from dorsal and ventral pancreatic
buds of endodermal cells. As the duodenum grows, it rotates
clockwise, and the two pancreatic buds fuse (exhibit II).
Midgut
During the fourth week of the embryonic stage (exhibit I), the
midgut is continuous with the yolk sac. By the fifth week, the
midgut has formed a ventral U-shaped midgut loop, which projects into the umbilical cord (exhibit III). As development continues, the anterior limb of the midgut loop coils to form most of
the small intestine. The posterior limb of the midgut loop expands to form the large intestine and a portion of the small intestine. A cecal diverticulum appears during the fifth week.


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(a)
(b)
(c)
(d)
(e)
EXHIBIT III Progressive stages of the development of the midgut to form the distal portion of the small intestine and the proximal portion of the large intestine: (a) 5 weeks, (b) 6 weeks, (c) 7 weeks, (d) 10 weeks, and (e) 18 weeks.
During the tenth week, the intestines are drawn up into

the abdominal cavity, and further differentiation and rotation
occur. The cecal diverticulum continues to develop, forming the
cecum and appendix. The remainder of the midgut gives rise to
the ascending colon and hepatic flexure (exhibit III).
Hindgut
The hindgut extends from the midgut to the cloacal (klo-a'kal)
membrane (exhibit IV). The proctodeum (prok''t o-de'um), or
anal pit, is a depression in the anal region formed from an invagination of ectoderm that contributes to the cloacal membrane.
proctodeum: Gk. proktos, anus; hodaios, on the way to
The allantois (a-lan''to-is), which receives urinary wastes from

the fetus, connects to the hindgut at a region called the cloaca, as
seen in exhibit IV. A band of mesenchymal cells called the
urorectal septum grows caudally between the fourth and seventh
week until a complete partition separates the cloaca into a dorsal
anal canal and a ventral urogenital sinus. With the completion
of the urorectal septum, the cloacal membrane is divided into an
anterior urogenital membrane and a posterior anal membrane.
Toward the end of the seventh week, the anal membrane
perforates and forms the anal opening, which is lined with ectodermal cells. About this time, the urogenital membrane ruptures
to provide further development of the urinary and reproductive
systems.
cloaca: L. cloaca, sewer
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EXHIBIT IV The progressive development of the hindgut illustrating the developmental separation of the digestive system from the urogenital system. (a) An anterolateral view of an embryo at 4 weeks showing the position of a sagittal cut depicted in (a1), (b), and (c). (a1) At
4 weeks, the hindgut, cloaca, and allantois are connected. (b) At 6 weeks, the connections between the gut and extraembryonic structures
are greatly diminished. (c) By 7 weeks, structural and functional separation between the digestive system and the urogenital system is almost complete.
The pancreas is positioned horizontally along the posterior

abdominal wall, adjacent to the greater curvature of the stomach.
It measures about 12.5 cm (6 in.) in length and is approximately
2.5 cm (1 in.) thick. It has an expanded head, positioned near the
duodenum; a centrally located body; and a tapering tail, positioned near the spleen. All but a portion of the head is retroperitoneal. Within the lobules of the pancreas are the exocrine
secretory units, called pancreatic acini (as'-ni), and the endocrine

secretory units, called pancreatic islet cells. Each acinus consists
of a single layer of epithelial acinar cells surrounding a lumen into
which the constituents of pancreatic juice are secreted.
The pancreas is innervated by branches of the celiac
plexus. The glandular portion of the pancreas receives parasymacinus: L. acinus, grape
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Chapter 18
Digestive System
669
pathetic innervation, whereas the pancreatic blood vessels receive sympathetic innervation. The pancreas is supplied with
blood by the pancreatic branch of the splenic artery, which arises
from the celiac artery, and by the pancreatoduodenal branches,
which arise from the superior mesenteric artery. Venous blood is
returned through the splenic and superior mesenteric veins into
the hepatic portal vein.
Pancreatic cancer has the worst prognosis of all types of cancer. This is probably because of the spongy, vascular nature
of this organ and its vital exocrine and endocrine functions. Pancreatic surgery is a problem because the soft, spongy tissue is difficult
to suture.
(a)
(b)
(c)
(d)
Knowledge Check
23. Describe the liver. Where is it located? List the lobes of the
liver and the supporting ligaments.
24. List the principal functions of the liver.
25. Describe the structure of liver lobules and trace the flow of
blood and bile in the lobules.
26. Explain how the liver receives a double blood supply.
27. Explain how the gallbladder fills with bile secretions and
how bile and pancreatic secretions enter the duodenum.
28. Briefly state the exocrine and endocrine functions of the
pancreas. What are the various cellular secretory units of
the pancreas?
of the Digestive System
are beneficial, but some bacteria and protozoa can cause diseases.
Only a few examples of the pathogenic microorganisms will be
discussed here.
Dysentery (dis'en'ter''e) is an inflammation of the intestinal
mucosa, characterized by frequent loose stools containing mucus,
pus, and blood. The most common dysentery is amoebic dysentery, which is caused by the protozoan Entamoeba histolytica. Cysts
from this organism are ingested in contaminated food, and after
the protective coat is removed by HCl in the stomach, the vegetative form invades the mucosal walls of the ileum and colon.
Food poisoning is caused by consuming the toxins produced by pathogenic bacteria. Salmonella is a bacterium that
commonly infects food. Botulism, the most serious type of food
poisoning, is caused by ingesting food contaminated with the
toxin produced by the bacterium Clostridium botulinum. This organism is widely distributed in nature, and the spores it produces
are frequently found on food being processed by canning. For this
reason, food must be heated to 120 C (248 F) before it is
canned. It is the toxins produced by the bacterium growing in
the food that are pathogenic, rather than the organisms themselves. The poison is a neurotoxin that is readily absorbed into
the blood, at which point it can affect the nervous system.
Clinical Problems of the Teeth

and Salivary Glands
Pathogens and Poisons
Dental caries, or tooth decay, is the gradual decalcification of

tooth enamel (fig. 18.37) and underlying dentin, caused by the
acid products of bacteria. These bacteria thrive between teeth,
The GI tract presents a hospitable environment for an array of

parasitic helminths (worms) and microorganisms. Many of these
dysentery: Gk. dys, bad; entera, intestine
CHAPTER 18
Most of the congenital disorders of the digestive system develop

during the fourth or fifth week of embryonic life. A cleft palate,
as described in chapter 17, is a congenital opening between the
oral and nasal cavities; therefore, it involves both the digestive
and respiratory systems (see fig. 17.28). Esophageal atresia (atre'ze-d), or failure to develop the normal structure of the
esophageal-stomach area, is another disorder of the upper GI
tract that requires surgery to correct. Pyloric stenosis is a common abnormality in which the pyloric sphincter muscle is hypertrophied, reducing the size of the lumen. This condition affects
approximately 1 in 200 newborn males and 1 in 1,000 newborn
females. Stenoses, atresias, and malrotations of various portions
of the GI tract may occur as the gut develops. Umbilical problems involving the GI tract are fairly common, as is some form of
imperforate anus, which occurs in about 1 in 5,000 births.
FIGURE 18.37 Clinical problems of the teeth. (a) Trench mouth

and dental caries, (b) severe alveolar bone destruction from periodontitis, (c) pyogenic granuloma and dental caries, and (d) malposition of teeth.

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where food particles accumulate, and form part of the thin layer
of bacteria, proteins, and other debris called plaque that covers
teeth. The development of dental caries can be reduced by
brushing at least once a day and by flossing between teeth at regular intervals.
People over the age of 35 are particularly susceptible to
periodontal disease, which involves inflammation and deterioration of the gingiva, dental alveoli, periodontal membranes, and
the cementum that covers the roots of teeth. Some of the symptoms are loosening of the teeth, bad breath, bleeding gums when
brushing, and some edema. Periodontal disease may result from
impacted plaque, cigarette smoking, crooked teeth, or poor diet.
It accounts for 80% to 90% of tooth loss in adults.
Mumps is a viral disease of the parotid glands, and in advanced stages it may involve the pancreas and testes. In children,
mumps is generally not serious, but in adults it may cause deafness and destroy the pancreatic islet tissue or testicular cells.
Disorders of the Liver
CHAPTER 18
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The liver is a remarkable organ that has the ability to regenerate

even if up to 80% has been removed. The most serious diseases
of the liver (hepatitis, cirrhosis, and hepatomas) affect the
liver throughout, so that it cannot repair itself. Hepatitis is
inflammation of the liver. Certain chemicals may cause hepatitis,
but generally it is caused by infectious viral agents. Hepatitis A
(infectious hepatitis) is a viral disease transmitted through contaminated foods and liquids. Hepatitis B (serum hepatitis) is also
caused by a virus and is transmitted in blood plasma during transfusions or by improperly sterilized needles and syringes. Other
types of viral hepatitis are designated as hepatitis C, D, E, and G.
In cirrhosis (s-ro'sis) the liver becomes infused with fibrous tissue. This causes the liver tissue to break down and become filled with fat. Eventually, all functions of the liver are
compromised. Cirrhosis is most often the result of long-term alcohol abuse, but it can also result from malnutrition, hepatitis, or
other infections.
Jaundice is a yellow staining of the tissues produced by
high blood concentrations of either free or conjugated bilirubin.
Because free bilirubin is derived from hemoglobin, abnormally
high concentrations of this pigment may result from an unusually high rate of red blood cell destruction. This can occur, for
example, as a result of Rh disease (erythroblastosis fetalis) in an
Rh positive baby born to a sensitized Rh negative mother. Jaun-
cirrhosis: Gk. kirrhos, yellow orange

jaundice: L. galbus, yellow
dice may also occur in healthy infants, because excess red blood
cells are normally destroyed at about the time of birth. This condition is called physiological jaundice of the newborn and is not indicative of disease. Premature infants may also develop jaundice
due to inadequate amounts of hepatic enzymes necessary to conjugate bilirubin and excrete it in the bile. In adults, jaundice is
commonly exhibited when the excretion of bile is blocked by
gallstones.
Hepatomas (hep''a-to-'maz) are tumors (usually malignant)
that originate in or secondarily invade the liver. Those that originate in the liver (primary hepatomas) are relatively rare, but
those that metastasize to the liver from other organs (secondary
hepatomas) are common. Carcinoma of the liver is usually fatal.
Disorders of the GI Tract

Peptic ulcers are erosions of the mucous membranes of the stomach (fig. 18.38) or duodenum produced by the action of HCl.
Prolonged exposure to agents that weaken the mucosal lining of
the stomach, such as alcohol and aspirin, and abnormally high
secretions of HCl increase the likelihood of developing peptic ulcers. Chronic stress can impair mucosal defense mechanisms,
thereby increasing mucosal susceptibility to the damaging effects
of HCl. A relatively recent finding is that most people who have
peptic ulcers are infected with a bacterium known as Helicobacter
pylori, which resides in the GI tract. Clinical trials have demonstrated that antibiotics that eliminate this bacterium appear to
help in the treatment of the peptic ulcers. It is now thought that
H. pylori does not itself cause the ulcer, but rather contributes to
the weakening of the mucosal barriers to gastric acid damage.
Enteritis, an inflammation of the mucosa of the small intestine, is frequently referred to as intestinal flu. Causes of enteritis include bacterial or viral infections, irritating foods or fluids
(including alcohol), and emotional stress. The symptoms include
abdominal pain, nausea, and diarrhea. Diarrhea is the passage of
watery, unformed stools. This condition is symptomatic of inflammation, stress, and many other body dysfunctions.
A hernia is a protrusion of a portion of a visceral organ,
usually the small intestine, through a weakened portion of the
abdominal wall. Inguinal, femoral, umbilical, and hiatal hernias
are the most common types. With a hiatal hernia, a portion of
the stomach pushes superiorly through the esophageal hiatus in
the diaphragm and protrudes into the thorax. The potential dangers of a hernia are strangulation of the blood supply followed by
gangrene, blockage of chyme, or ruptureeach of which can
threaten life.
Diverticulosis (di''ver-tik''yu-lo'sis) is a condition in which
the intestinal wall weakens and an outpouching (diverticulum)
occurs. Studies suggest that suppressing the passage of flatus (in-


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Digestive System
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FIGURE 18.38 Common sites of upper GI disorders.

The likely source of bleeding in such a case is from a rupture of the internal portion of the liver involving significant blood vessels, either of
the hepatic arterial or venous circulation, but also possibly of the portal
venous circulation. Trauma to the hepatic plates (see fig. 18.33) may
allow blood to enter bile canaliculi. From there, the course of the blood
is as follows: large hepatic ducts common hepatic duct common
bile duct through duodenal papilla duodenum of the small intestine.
In some cases, this type of bleeding can be stopped by introducing radiographic-guided catheters into the arterial system to block the
blood vessels (angiographic embolization). However, surgery is sometimes required.
CHAPTER 18
testinal gas) may contribute to diverticulosis, especially in the

sigmoid colon. Diverticulitis, or inflammation of a diverticulum,
can develop if fecal material becomes impacted in these pockets.
Peritonitis is inflammation of the peritoneum lining the
abdominal cavity and covering the viscera. The causes of peritonitis include bacterial contamination of the abdominal cavity
through accidental or surgical wounds in the abdominal wall or
perforation of the intestinal wall (as with an ulcer or a ruptured
appendix).

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A 69year-old gentleman comes to your office because of abdominal pain in the left
lower quadrant for 2 days. The pain is constant and deep. He has also developed mild
diarrhea since the pain started.
In your office the patient does not have
a fever. You can hear normal bowel sounds
with your stethoscope. His abdominal muscles are relaxed, but the left lower quadrant is
indeed tender, and you feel a fullness on deep
palpation. A stool sample and lab work are
normal. You decide to send him home on a
liquid diet and oral antibiotics. Hes soon

better, and in 2 weeks you order a barium
enema (i.e., an abdominal X ray taken after
barium is introduced into the colon).
QUESTIONS
1. What are the many blebs seen on the
adjacent X ray of the large colon?
2. Explain this mans earlier abdominal
pain.
3. What other complication can
accompany this condition?
CHAPTER 18
A 47-year-old male presents to the emergency room with worsening abdominal pain
and anorexia. He reports the pain began
three days ago and was generalized, but has
now localized to the lower right quadrant.
He describes the pain as a sharp, stabbing
pain. He also reports fevers but no diarrhea.
Upon physical exam, the patient is febrile
(feverish) and has abdominal tenderness that
localizes to the lower right quadrant, specifically McBurneys point. He also has rebound
tenderness, a sign of peritoneal irritation. Lab
results indicate an increase in the while blood
count. You order a CT scan of the abdomen.
QUESTIONS
1. What is McBurneys point?
2. After looking at the CT scan, what is
your diagnosis? (The cecum is indicated
with the letter C.)
3. What is the white density indicated by
the arrow? How does it contribute to
the disease process?
4. What is the appropriate treatment for
this patient?
Important Clinical Terminology

chilitis (ki-li'tis) Inflammation of the lips.
colitis (ko-li'tis) Inflammation of the colon.
colostomy (ko-los'to-me) The formation of an
abdominal exit from the GI tract by bringing
a loop of the colon through the abdominal
wall to its outside surface. If the rectum is
removed because of cancer, the colostomy
provides a permanent outlet for the feces.
cystic fibrosis An inherited disease of the
exocrine glands, particularly the pancreas.
Pancreatic secretions are too thick to drain
easily, causing the ducts to become inflamed
and promoting connective tissue formation
that occludes drainage from the ducts still
further.
gingivitis (jin-j-vi'tis) Inflammation of the
gums. It may result from improper hygiene,
poorly fitted dentures, improper diet, or
certain infections.
halitosis (hal--to'sis) Offensive breath odor. It

may result from dental caries, certain
diseases, eating particular foods, or smoking.
heartburn A burning sensation of the
esophagus and stomach. It may result from
the regurgitation of gastric juice into the
lower portion of the esophagus.
hemorrhoids (hem'o-roidz) Varicose veins of
the rectum and anus.
nausea Gastric discomfort and sensations of
illness with a tendency to vomit. This feeling
is symptomatic of motion sickness and other
diseases, and may occur during pregnancy.
pyorrhea (pi''o-re'a) The discharge of pus at
the base of the teeth at the gum line.
regurgitation (vomiting) The forceful
expulsion of gastric contents into the mouth.
Nausea and vomiting are common symptoms
of almost any dysfunction of the digestive
system.
trench mouth A contagious bacterial

infection that causes inflammation,
ulceration, and painful swelling of the floor
of the mouth. It is generally contracted
through direct contact by kissing an infected
person. Trench mouth can be treated with
penicillin and other medications.
vagotomy (va-got'o-me) The surgical removal
of a section of the vagus nerve where it
enters the stomach in order to eliminate
nerve impulses that stimulate gastric
secretion. This procedure may be used to
treat chronic ulcers.


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Chapter 18
Digestive System
673
Chapter Summary
Introduction to the Digestive System
(pp. 635636)
1. The digestive system mechanically and
chemically breaks down food to forms
that can be absorbed through the
intestinal wall and transported by the
blood and lymph for use at the cellular
level.
2. The digestive system consists of a
gastrointestinal (GI) tract and accessory
digestive organs.
Serous Membranes and Tunics of the

Gastrointestinal Tract (pp. 636640)
Mouth, Pharynx, and Associated

Structures (pp. 640648)
1. The oral cavity is formed by the cheeks,
lips, and hard palate and soft palate. The
tongue and teeth are contained in the
oral cavity.
(a) Lingual tonsils and papillae with
taste buds are located on the tongue.
(b) Structures of the palate include
palatal folds, a cone-shaped
projection called the palatine uvula,
and palatine tonsils.
Esophagus and Stomach (pp. 648652)

1. Swallowing (deglutition) occurs in three
phases and involves structures of the oral
cavity, pharynx, and esophagus.
2. Peristaltic waves of contraction push food
through the lower esophageal sphincter
into the stomach.
3. The stomach consists of a cardia, fundus,
body, and pylorus. It displays greater and
lesser curvatures, and contains a pyloric
sphincter at its junction with the
duodenum.
(a) The mucosa of the stomach is thrown
into distensible gastric folds; gastric
pits and gastric glands are present in
the mucosa.
(b) The parietal cells of the gastric glands
secrete HCl, and the principal cells
secrete pepsinogen.
Small Intestine (pp. 652656)

1. Regions of the small intestine include the
duodenum, jejunum, and ileum; the
common bile duct and pancreatic duct
empty into the duodenum.
2. Fingerlike extensions of mucosa, called
intestinal villi, project into the lumen,
and at the bases of the intestinal villi the
mucosa forms intestinal glands.
(a) New epithelial cells are formed in the
intestinal crypts.
(b) The membrane of intestinal

epithelial cells is folded to form
microvilli; this brush border of the
mucosa increases the absorptive
surface area.
3. Movements of the small intestine include
rhythmic segmentation, pendular
movement, and peristalsis.
Large Intestine (pp. 656660)

1. The large intestine absorbs water and
electrolytes from the chyme and passes
fecal material out of the body through the
rectum and anal canal.
2. The large intestine is divided into the
cecum, colon, rectum, and anal canal.
(a) The appendix is attached to the
inferior medial margin of the cecum.
(b) The colon consists of ascending,
transverse, descending, and sigmoid
portions.
(c) Haustra are bulges in the walls of the
large intestine.
3. Movements of the large intestine include
peristalsis, haustral churning, and mass
movement.
Liver, Gallbladder, and Pancreas

(pp. 660669)
1. The liver is divided into right, left,
quadrate, and caudate lobes. Each lobe
contains liver lobules, the functional units
of the liver.
(a) Liver lobules consist of plates of
hepatic cells separated by modified
capillaries called sinusoids.
(b) Blood flows from the periphery of
each lobule, where branches of the
hepatic artery and hepatic portal vein
empty, through the sinusoids and out
the central vein.
(c) Bile flows within the hepatic plates,
in bile canaliculi, to the biliary
ductules at the periphery of each
lobule.
2. The gallbladder stores and concentrates
the bile; it releases the bile through the
cystic duct and common bile duct into
the duodenum.
3. The pancreas is both an exocrine and an
endocrine gland.
(a) The endocrine portion, consisting of
the pancreatic islets, secretes the
hormones insulin and glucagon.
(b) The exocrine acini of the pancreas
produce pancreatic juice, which
contains various digestive enzymes.
CHAPTER 18
1. Peritoneal membranes line the abdominal

wall and cover the visceral organs. The
GI tract is supported by a double layer of
peritoneum called the mesentery.
(a) The lesser omentum and greater
omentum are folds of peritoneum that
extend from the stomach.
(b) Retroperitoneal organs are positioned
2. The layers (tunics) of the abdominal GI
tract are, from the inside outward, the
mucosa, submucosa, tunica muscularis,
and serosa.
(a) The mucosa consists of a simple
columnar epithelium, a thin layer of
connective tissue called the lamina
propria, and thin layers of smooth
muscle called the muscularis
mucosae.
(b) The submucosa is composed of
connective tissue; the tunica
muscularis consists of layers of
smooth muscle; and the serosa is
composed of connective tissue
covered with the visceral peritoneum.
(c) The submucosa contains the
submucosal plexus, and the tunica
muscularis contains the myenteric
plexus of autonomic nerves.
2. The incisors and canines have one root

each; the bicuspids and molars have two
or three roots.
(a) Humans are diphyodont; they have
deciduous and permanent sets of teeth.
(b) The roots of teeth fit into sockets
called dental alveoli that are lined
with a periodontal membrane. Fibers
in the periodontal membrane insert
into the cementum covering the
roots, firmly anchoring the teeth in
the sockets.
(c) Enamel forms the outer layer of the
tooth crown; beneath the enamel is
dentin.
(d) The interior of a tooth contains a
pulp cavity, which is continuous
through the apical foramen of the
root with the connective tissue
around the tooth.
3. The major salivary glands are the parotid
glands, the submandibular glands, and the
sublingual glands.
4. The muscular pharynx provides a
passageway connecting the oral and nasal
cavities to the esophagus and larynx.

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Review Activities
CHAPTER 18
Objective Questions
1. Viscera are the only body organs that are
(a) concerned with digestion.
(b) located in the abdominal cavity.
(c) covered with peritoneal membranes.
(d) located within the thoracic and
abdominal cavities.
2. Which of the following types of teeth are
found in the permanent but not in the
deciduous dentition?
(a) incisors
(c) premolars
(b) canines
(d) molars
3. The double layer of peritoneum that
supports the GI tract is called
(a) the visceral peritoneum.
(b) the mesentery.
(c) the greater omentum.
(d) the lesser omentum.
4. Which of the following tissue layers in
the small intestine contains the lacteals?
(a) the submucosa
(b) the muscularis mucosae
(c) the lamina propria
(d) the tunica muscularis
5. Which of the following organs is not
considered a part of the digestive system?
(a) the pancreas (c) the tongue
(b) the spleen
(d) the gallbladder
6. The numerous small elevations on the
surface of the tongue that support taste
buds and aid in handling food are called
(a) cilia.
(c) intestinal villi.
(b) rugae.
(d) papillae.
7. Most digestion occurs in
(a) the mouth.
(b) the stomach.
(c) the small intestine.
(d) the large intestine.
8. Stenosis (constriction) of the sphincter of
ampulla (of Oddi) would interfere with
(a) transport of bile and pancreatic juice.
(b) secretion of mucus.
(c) passage of chyme into the small
intestine.
(d) peristalsis.
9. The first organ to receive the blood-borne
products of digestion is
(a) the liver.
(c) the heart.
(b) the pancreas. (d) the brain.

hepatic portal blood is true?
(a) It contains absorbed fat.
(b) It contains ingested proteins.
(c) It is mixed with bile in the liver.
(d) It is mixed with blood from the
hepatic artery in the liver.
Essay Questions
1. Define digestion. Differentiate between
the mechanical and chemical aspects of
digestion.
2. Distinguish between the gastrointestinal
tract, viscera, accessory digestive organs,
and gut.
3. List the specific portions or structures of
the digestive system formed by each of the
three embryonic germ layers.
4. Define serous membrane. How are the
serous membranes of the abdominal
cavity classified and what are their
functions?
5. Describe the structures of the four tunics
in the wall of the GI tract.
6. Why are there two autonomic
innervations to the GI tract? Identify the
specific sites of autonomic stimulation in
the tunic layers.
7. Define the terms dental formula,
diphyodont, deciduous teeth, permanent
teeth, and wisdom teeth.
8. Outline the stages of deglutition. What
biomechanical roles do the tongue, hard
palate and soft palate, pharynx, and hyoid
bone perform in deglutition?
9. How does the stomach protect itself from
the damaging effects of HCl?
10. Describe the kinds of movements in the
small intestine and explain what they
accomplish.
11. Diagram an intestinal villus and explain
why intestinal villi are considered the
functional units of the digestive system.
12. What are the regions of the large
intestine? In what portion of the
abdominal cavity and pelvic cavity is each
region located?

13. Describe the location and gross structure

of the liver. Draw a labeled diagram of a
liver lobule.
14. Describe how the gallbladder is filled with
and emptied of bile fluid. What is the
function of bile?
15. List the functions of the large intestine.
What are the biomechanical movements
of the large intestine that make these
functions possible?
16. Define cirrhosis and explain why this
condition is so devastating to the liver.
What are some of the causes of cirrhosis?
1. Technically, ingested food is not in the
body. Neither are feces excreted from
within the body (except bile residue).
Explain these statements. Why would this
information be important to a drug
company interested in preparing a new
oral medication?
2. The deciduous (milk) teeth dont matter
because they fall out anyway. Do you
agree or disagree with this statement?
Explain.
3. Which surgery do you think would have
the most profound effect on digestion:
(a) removal of the stomach (gastrectomy),
(b) removal of the pancreas
(pancreatectomy), or (c) removal of the
gallbladder (cholecystectomy)? Explain
your reasoning.
4. Describe the adaptations of the GI tract
that make it more efficient by either
increasing the surface area for absorption
or increasing the time of contact between
food particles and digestive enzymes.
5. During surgery to determine the cause of
an intestinal obstruction, why might the
surgeon elect to remove a healthy
appendix?
6. Explain why a ruptured appendix may
result in peritonitis, while an inflamed
kidney (nephritis) generally does not
result in peritonitis.


Body
19. Urinary System
Urinary System
The McGrawHill
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19
Introduction to the Urinary
System 676
Kidneys 676
Ureters, Urinary Bladder,
and Urethra 684
The Urinary System 689
Chapter Summary 695
Clinical Case Study

A 17-year-old male was involved in a knife fight in which he sustained two stab wounds to the
anterior abdomen. He was brought to the emergency room where he complained of mild abdominal pain and an urgent need to urinate. Although neither of the stab wounds was bleeding
externally, examination by the surgeon revealed signs of moderate hemorrhagic shock. One
wound was 3 cm below the right costal margin at the midclavicular line; the other was just medial to the right anterior superior iliac spine. The surgeon immediately ordered preparations for
an emergency exploratory laparotomy. She noted that the patients urinary bladder was quite
full and chided the intern for not having placed a urinary catheter. Placement of the catheter
yielded a brisk flow of bright red blood.
How would you explain the phenomenon of hematuria (blood in the urine) in this case?
Which of the two stab wounds is most likely associated with the hematuria? Regarding the
blood draining into the catheter, trace and explain its course of flow. Begin at a point in the abdominal aorta, and end the course with drainage into the catheter. Assuming that the surgeon
would be prompted to remove the kidney in order to quickly control hemorrhage, what possible
anatomical variant should she keep in mind?
Hints: Study the positions of the kidneys within the abdominal cavity and note the location of
the supportive and protective serous membrane. Carefully examine the specific location of
urine production and the route of urine passage through the urinary system.
FIGURE: The placement of a urinary

catheter is a common procedure for patients
who have abdominal trauma or abdominal
surgery. A laboratory assessment (urinalysis)
of the collected urine may help reveal the
extent of the trauma or a patients progress in
healing.

676
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19. Urinary System
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INTRODUCTION
TO THE URINARY SYSTEM
The urinary system maintains the composition and properties of
the body fluid that establishes the internal environment of the
body cells. The end product of the urinary system is urine, which
is voided from the body during micturition.
Objective 1
Inferior
vena cava
Kidney
List the functions of the urinary system.
Objective 2
CHAPTER 19
Identify the arteries that transport blood to the

urinary system for filtration.
The urinary system, along with the respiratory, digestive, and integumentary systems, excretes substances from the body. For this
reason, these systems are occasionally referred to as excretory systems. In the process of cellular metabolism, nutrients taken in by
the digestive system and oxygen from inhaled air are used to synthesize a variety of substances while providing energy needed for
body maintenance. Metabolic processes, however, produce cellular wastes that must be eliminated if homeostasis is to be maintained. Just as the essential nutrients are transported to the cells
by the blood, the cellular wastes are removed through the circulatory system to the appropriate excretory system. Carbon dioxide is eliminated through the respiratory system; excessive water,
salts, nitrogenous wastes, and even excessive metabolic heat are
removed through the integumentary system; and various digestive wastes are eliminated through the digestive system.
The urinary system is the principal system responsible for
water and electrolyte balance. Electrolytes are compounds that
separate into ions when dissolved in water. Electrolyte balance is
achieved when the number of electrolytes entering the body
equals the number leaving. Hydrogen ions, for example, are
maintained in precise concentration so that an acid-base, or pH,
balance exists in the body.
A second major function of the urinary system is the excretion of toxic nitrogenous compoundsspecifically, urea and creatinine. Other functions of the urinary system include the
elimination of toxic wastes that may result from bacterial action
and the removal of various drugs that have been taken into the
body. All of these functions are accomplished through the formation of urine by the kidneys.
The urinary system consists of two kidneys, two ureters, the
urinary bladder, and the urethra (fig. 19.1). Tubules in the kidneys
are intertwined with vascular networks of the circulatory system
to enable the production of urine. After the urine is formed, it is
moved through the ureters to the urinary bladder for storage.
Micturition, or voiding of urine from the urinary bladder, occurs
through the urethra.
Blood to be processed by a kidney enters through the large
renal artery. After the filtration process (see chapter 3), it exits
through the renal vein. The importance of filtration of the blood
is demonstrated by the fact that during normal resting conditions
the kidneys receive approximately 20% to 25% of the entire cardiac output. Every minute, the kidneys process approximately
1,200 ml of blood.
Abdominal
aorta
Ureter
Urinary
bladder
Urethra
FIGURE 19.1 The organs of the urinary system are the two kidneys, two ureters, urinary bladder, and urethra.
Knowledge Check
1. Drawing on your knowledge of the functions of the urinary
system, list the basic substances that compose normal urine.
2. Explain the role of the renal vessels in maintaining homeostasis. Approximately how much blood is processed in the
kidneys each minute?
KIDNEYS
The kidney consists of an outer renal cortex and an inner renal
medulla that contains the renal pyramids. Urine is formed as a filtrate from the blood at the nephrons and collects in the calyces
and renal pelvis before flowing from the kidney via the ureter.
Objective 3
Describe the gross structure of the kidney.
Objective 4
Describe structure of a nephron and explain

how its components are oriented within the kidney.
Objective 5
Describe the position of cortical and

juxtamedullary nephrons with respect to the gross structure
of the kidney.


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Chapter 19
Twelfth
thoracic
vertebra
Twelfth
rib
Minor
calyx
Urinary System
677
Although the kidneys are firmly supported by the renal adipose capsule, renal fascia, and even the renal vessels, under
certain conditions these structures may give in to the force of gravity
and the kidneys may drop a bit in position. This condition is called
renal ptosis (to'sis) and generally occurs in extremely thin elderly
people, who have insufficient amounts of supportive fat in the adipose capsular layer. It also may affect victims of anorexia nervosa,
who suffer from extreme weight loss. The potential danger of renal
ptosis is that the ureter may kink, blocking the flow of urine from the
affected kidney.
Renal
pelvis
Kidney
Ureter
Urinary
bladder
FIGURE 19.2 A color-enhanced radiograph of the calyces

and renal pelvises of the kidneys, the ureters, and the urinary bladder. (Note the position of the kidneys relative to the vertebral column
and ribs.)
Position and Appearance of the Kidneys
hilum: L. hilum, a trifle
A coronal section of the kidney shows two distinct regions and a

major cavity (figs. 19.4b and 19.5). The outer renal cortex, in
contact with the renal capsule, is reddish brown and granular in
appearance because of its many capillaries. The deeper renal
medulla is darker, and the presence of microscopic tubules and
blood vessels gives it a striped appearance. The renal medulla is
composed of 8 to 15 conical renal pyramids. Portions of the
renal cortex extend between the renal pyramids to form the
renal columns. The apexes of the renal pyramids are known as
the renal papillae (pa-pil'e). These nipplelike projections are directed toward the inner region of the kidney.
The cavity of the kidney collects and transports urine from
the kidney to the ureter. It is divided into several portions. The
papilla of a renal pyramid projects into a small depression called
the minor calyx (ka'liksin the plural, calyces). Several minor
calyces unite to form a major calyx. In turn, the major calyces
join to form the funnel-shaped renal pelvis. The renal pelvis collects urine from the calyces and transports it to the ureter.
Microscopic Structure of the Kidney

The nephron (nef'ron) is the functional unit of the kidney that is
responsible for the formation of urine. Each kidney contains
more than a million nephrons surrounded by associated small
blood vessels. Fluid formed by capillary filtration enters the
nephron and is subsequently modified by transport processes.
The resulting fluid that leaves the nephron is urine.
Renal Blood Vessels

The kidneys have an extensive circulatory network to allow for
the continuous cleansing and modification of large volumes of
blood (fig. 19.6). Arterial blood enters the kidney at the hilum
through the renal artery, which divides into interlobar (in''terlo'bar) arteries that pass between the renal pyramids through the
renal columns. Arcuate (ar'kyoo-at) arteries branch from the interlobar arteries at the boundary of the renal cortex and renal
medulla. Small interlobular arteries radiate from the arcuate arteries and project into the renal cortex. Microscopic afferent
glomerular arterioles arise from branches of the interlobular
ptosis: Gk. ptosis, a falling
arcuate: L. arcuare, to bend
CHAPTER 19
The reddish brown kidneys are positioned against the posterior

wall of the abdominal cavity between the levels of the twelfth
thoracic and the third lumbar vertebrae (fig. 19.2). The right
kidney is usually 1.5 to 2.0 cm lower than the left because of the
large area occupied by the liver on the right side.
The kidneys are retroperitoneal, which means that they are
located behind the parietal peritoneum (fig. 19.3). Each adult
kidney is a lima-bean-shaped organ about 11.25 cm (4 in.) long,
5.5 to 7.7 cm (23 in.) wide, and 2.5 cm (1 in.) thick. The lateral
border of each kidney is convex, whereas the medial border is
strongly concave (figs. 19.2 and 19.3). The hilum (h'lum) of the
kidney is the depression along the medial border through which
the renal artery enters and the renal vein and ureter (yoo're'ter)
exit. The hilum is also the site for drainage of lymph vessels and
innervation of the kidney. The superior border of each kidney is
capped by the adrenal gland (see figs. 14.21 and 19.4a).
Each kidney is embedded in a fatty fibrous pouch consisting
of three layers. The renal capsule (fibrous capsule), the innermost
layer, is a strong, transparent fibrous attachment to the surface of
the kidney. The renal capsule protects the kidney from trauma and
the spread of infections. Surrounding the renal capsule is a firm
protective mass of fatty tissue called the renal adipose capsule
(fig. 19.3). The outermost layer, the renal fascia, is composed of
dense irregular connective tissue. It is a supportive layer that anchors the kidney to the peritoneum and the abdominal wall.
Gross Structure of the Kidney

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Vertebra
Kidney
Renal
adipose capsule
Renal fascia
Parietal
peritoneum
Spleen
Renal
capsule
Abdominal
aorta
Inferior
vena cava
Stomach
Pancreas
FIGURE 19.3 The position of the kidneys as seen in cross section through the upper abdomen. The kidneys are embedded in adipose tissue
Renal
capsule
CHAPTER 19
Renal
column
(a)
(b)
FIGURE 19.4 The kidney. (a) The anterior surface and (b) a coronal section.
arteries. The afferent glomerular arterioles transport the blood into
ball-shaped capillary networks, the glomeruli (glo-mer'yu-li), which
produce a blood filtrate that enters the urinary tubules. The blood
remaining in the glomerulis leaves through efferent glomerular
arterioles. This blood vessel arrangement is unique because blood
usually flows out of a capillary bed into venules rather than into
glomerulus: L. diminutive of glomus, ball
other arterioles. From the efferent glomerular arterioles, the blood

enters either the peritubular capillaries surrounding the convoluted tubules or the vasa recta surrounding the ascending and descending tubules (fig. 19.7). From these capillary networks, the
blood is drained into veins that parallel the course of the arteries
in the kidney. These are the interlobular veins, arcuate veins,
and interlobar veins. The interlobar veins descend between the
renal pyramids, converge, and then leave the kidney as a single
renal vein that empties into the inferior vena cava.


Body
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Chapter 19
Urinary System
679
Renal
cortex
Renal cortex
Nephron
Renal medulla
Minor calyx
Major calyx
Renal pyramid
Renal
medulla
Renal capsule
Renal
artery
Renal column
Renal
vein
Opening of
renal calyx
into renal pelvis
Renal pelvis
Ureter
Renal
papilla
Renal papilla
(b)
(a)
Minor calyx
Glomerular
capsule
Distal convoluted
tubule
Papillary duct
Proximal convoluted
tubule
Creek
(c)
FIGURE 19.5 The internal structures of a kidney. (a) A coronal section showing the structure of the renal cortex, renal medulla, and renal
pelvis. (b) A diagrammatic magnification of a renal pyramid to depict the renal tubules. (c) A diagram of a single nephron and a papillary duct.
In summary, vessels are at a capillary level between the arterioles in each glomerulus of the kidney. The blood pressure at
the glomerulus is strong enough to force water and dissolved
wastes from the blood into the urinary tubular portion of the
nephron. Thus, this capillary network produces the filtrate. A
secondary capillary network of peritubular capillaries and vasa
recta surrounds various tubular portions of the nephron. This
capillary bed, however, is adapted for absorption rather than fil-
trate formation. It reabsorbs water and other substances that

should not be excreted with the urine, reclaiming most of the filtrate produced in the glomerulus.
Although the kidneys are generally well protected in that
they are encapsulated retroperitoneally, they may be injured by a
hard blow to the lumbar region. Such an injury can produce
blood in the urine, because the highly vascular kidneys are particularly susceptible to hemorrhage.
CHAPTER 19
Nephron loop

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Body
19. Urinary System
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Nephron
The tubular nephron consists of a glomerular capsule, proximal
convoluted tubule, descending limb of the nephron loop (loop of
Henle), ascending limb of the nephron loop, and distal convoluted
tubule (fig. 19.7).
The glomerular (Bowman's) capsule surrounds the
glomerulus. The glomerular capsule and its associated glomerulus
are located in the renal cortex of the kidney and together constitute the renal corpuscle (fig. 19.8). The glomerular capsule contains an inner visceral layer of epithelium, in contact with the
glomerular capillaries and an outer parietal layer. The space between these two layers, called the capsular space, is where the
glomerular filtrate collects.
The glomerular epithelium contains tiny pores called fenestrae (fe-nes'tre) that permit the filtrate to pass from the blood
into the glomerular capsular space (fig. 19.8). Although the fenestrae are large, they are still small enough to prevent the passage
of blood cells, platelets, and most plasma proteins into the
FIGURE 19.6 The principal arteries and veins of a kidney.
Bowmans capsule: from Sir William Bowman, English anatomist, 181692
Glomerular capsule
CHAPTER 19
Efferent glomerular
arteriole
Afferent glomerular
arteriole
FIGURE 19.7 A simplified illustration of blood flow from a glomerulus to an efferent glomerular arteriole, to the peritubular capillaries, and to
the venous drainage of the kidney.


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Chapter 19
Afferent glomerular arteriole
Urinary System
681
Efferent glomerular arteriole
Blood flow
Primary process
Blood flow
Capillary
endothelium
Pedicel
Glomerulus
Fenestrae
External wall of
glomerular capsule
Filtration
Podocyte
Proximal convoluted tubule
Internal wall of
glomerular capsule
(a)
Slit pore
Pedicel
Slit pores
Loechel
Primary process
of podocyte
Glomerulus
CHAPTER 19
(b)
FIGURE 19.8 The structure of a glomerulus. (a) A renal corpuscle is composed of a glomerulus and a glomerular (Bowmans) capsule. Note
that the diameter of the efferent glomerular arteriole carrying blood away from the glomerulus is smaller than that of the afferent glomerular arteriole transporting blood into the glomerulus. This is one of the contributing factors in the maintenance of high blood pressure within the glomerulus.
The first step of urine formation is the filtration through the glomerular membrane (arrows) into the glomerular space of the glomerular capsule.
(b) A scanning electron micrograph of a glomerulus (8,000).

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Body
19. Urinary System
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Pedicels
Slit
pores
Basement membrane
Capillary lumen
Fenestra
(a)
(b)
CHAPTER 19
FIGURE 19.9 Glomerular filtration. (a) A transmission electron micrograph of a glomerular capillary and the glomerular membranes, and (b)
accompanying diagram. Substances in the blood are filtered through capillary fenestrae. The filtrate then passes across the basement membrane and through slit pores between the pedicels and enters the capsular space. From here, the filtrate is transported to the lumen of the proximal convoluted tubule (see fig. 19.8).
filtrate. The inner layer of the glomerular capsule is composed of

unique cells called podocytes (pod'o-sts) with numerous cytoplasmic extensions known as pedicels (ped'-selz). Pedicels interdigitate, like the fingers of clasped hands, as they wrap around
the glomerular capillaries (figs. 19.8 and 19.9). The narrow slit
pores between adjacent pedicels provide the passageways
through which filtered molecules must pass to enter the interior
of the glomerular capsule.
Filtrate in the glomerular capsule passes into the lumen of
the proximal convoluted tubule. The wall of the proximal convoluted tubule consists of a single layer of cuboidal cells containing millions of microvilli; these serve to increase the surface area
for reabsorption. In the process of reabsorption, salt, water, and
other molecules needed by the body are transported from the
lumen through the tubular cells and into the surrounding peritubular capillaries.
The glomerulus, glomerular capsule, and proximal convoluted tubule are located in the renal cortex. Fluid passes from the
proximal convoluted tubule to the nephron loop (loop of
Henle). This fluid is carried into the renal medulla in the descending limb of the nephron loop and returns to the renal cortex in the ascending limb of the loop. Back in the renal cortex,
the tubule becomes coiled again, and is called the distal convoluted tubule. The distal convoluted tubule is shorter than the
loop of Henle: from Friedrich Gustav Jacob Henle, German physician, anatomist,
and pathologist, 180985
proximal convoluted tubule and has fewer microvilli. It is the

last segment of the nephron, and terminates as it empties into a
papillary duct (collecting duct). Passing through the renal
papilla, the papillary ducts empty fluid into the minor calyx,
which in turn passes into the major calyx. Once within the calyces, this fluid is known as urine. From the major calyces, urine
collects in the renal pelvis before it passes from the kidney into
the ureter.
Two principal types of nephrons are classified according to
their positions in the kidney and the lengths of their nephron
loops. Nephrons that have their glomulari in the inner one-third
of the cortexcalled juxtamedullary (juk''sta-med'yu-ler-e)
nephronshave longer loops than the cortical nephrons that
have their glomeruli in the outer two-thirds of the renal cortex
(fig. 19.10).
The kidneys have an autonomic nerve supply derived from
the renal plexus of the tenth, eleventh, and twelfth thoracic
nerves. Sympathetic stimulation of the renal plexus produces a
vasomotor vascular network response in the kidney. This response determines the circulation of the blood by regulating the
diameters of arterioles.
The functions of the nephron are summarized in table 19.1.
Urine from a healthy individual is virtually bacteria-free but easily becomes contaminated after voiding because its organic
components serve as a nutrient source for contaminating microbes.
The breakdown of urine by bacterial action produces ammonia.
A urinalysis is a common procedure in any routine physical examination. The appearance and pH of the urine are noted, as well as
the presence of such abnormal constituents as albumin, blood, glucose, and acetone. Abnormal urine is symptomatic of a variety of diseases or problems in the urinary system.


Body
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Chapter 19
Urinary System
683
FIGURE 19.10 Cortical nephrons are located almost exclusively within the renal cortex. Juxtamedullary nephrons are located, for the most
part, within the outer portion of the renal medulla. (Note the flow of blood through the glomerulus indicated with arrows.)
Structure
Function
Renal corpuscle
Glomerulus
Filtration of water and dissolved substances from blood plasma
Glomerular capsule
Nephron
Proximal convoluted
tubule
Descending limb of
the nephron loop
Ascending limb of
the nephron loop
Distal convoluted
tubule
Papillary duct
Receives glomerular filtrate

Reabsorption of water by osmosis
Reabsorption of glucose, amino acids, creatine, lactic acid, citric acid, uric acid, ascorbic acid, phosphate ions, sulfate ions,
calcium ions, potassium ions, and sodium ions by active transport
Reabsorption of proteins by pinocytosis
Reabsorption of chloride ions and other negatively charged ions by electrochemical attraction
Active secretion of such substances as penicillin, histamine, and hydrogen ions
Reabsorption of chloride ions by active transport and passive reabsorption of sodium ions
Reabsorption of sodium ions by active transport
Active secretion of hydrogen ions
Passive secretion of potassium ions by electrochemical attraction
Passive reabsorption of water by osmosis; drains urine from several nephrons
CHAPTER 19
TABLE 19.1 Functions of the Nephron in Urine Formation

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Knowledge Check
3. Describe the general appearance of the renal cortex and
renal medulla.
4. Trace the course of blood through the kidney from the
renal artery to the renal vein.
5. Trace the course of tubular fluid from the glomerular capsules to the ureter.
6. Draw a diagram of a nephron and label the renal cortex and
renal medulla. Also label the structures within each region.
URETERS, URINARY BLADDER,

AND URETHRA
Lumen
Transitional
epithelium
Mucosa
Muscularis
Adventitia
FIGURE 19.11 A photomicrograph of a ureter in cross section.

(Note the transitional epithelium lining the lumen.)
Urine is channeled from the kidneys to the urinary bladder by the

ureters and expelled from the body through the urethra. The mucosa of the urinary bladder permits distension, and the muscles of
the urinary bladder and urethra function in the control of micturition.
Objective 6
Describe the location, structure, and function of
the ureters.
Objective 7
Describe the gross and histological structure

and the innervation of the urinary bladder.
Objective 8
Describe the micturition reflex.
Objective 9
Compare and contrast the structure of the male

urethra with that of the female.
CHAPTER 19
Ureters
The ureters (yoo-re'terz), like the kidneys, are retroperitoneal.
These tubular organs, each about 25 cm (10 in.) long, begin at
the renal pelvis and course inferiorly to enter the urinary bladder
at the posterolateral angles of its base. The thickest portion of a
ureter, near where it enters the urinary bladder, is approximately
1.0 cm (0.4 in.) in diameter.
The wall of the ureter consists of three layers, or tunics.
The inner mucosa is continuous with the linings of the renal
tubules and the urinary bladder. The mucosa consists of transitional epithelium (fig. 19.11). The cells of this layer secrete a
mucus that coats the walls of the ureter with a protective film.
The middle layer of the ureter is called the muscularis.
It consists of inner longitudinal and outer circular layers of
smooth muscle fibers. In addition, the proximal one-third of
the ureter contains another longitudinal layer to the outside
of the circular layer. Muscular peristaltic waves move the
urine through the ureter. The peristaltic waves are initiated
by the presence of urine in the renal pelvis, and their frequency is determined by the volume of urine. The waves,
which occur from once every few seconds to once every few
minutes, force urine through the ureter and cause it to spurt
into the urinary bladder.
FIGURE 19.12 A renal stone (kidney stone) placed next to a dime

for size comparison. Factors contributing to renal stone formation
may include the ingestion of excessive mineral salts, a decrease in
water intake, and overactivity of the parathyroid glands. A renal stone
generally consists of calcium oxalate, calcium phosphate, and uric
acid crystals.
The outer layer of the ureter is called the adventitia
(ad''ven-tish'a) The adventitia is composed of loose connective
tissue that covers and protects the underlying layers. In addition,
extensions of the connective tissue anchor the ureter in place.
The arterial supply of the ureter comes from several sources.
Branches from the renal artery serve the superior portion. The
testicular (or ovarian) artery (also called gonadal artery) supplies
the middle portion, and the superior vesicular artery serves the
pelvic region. The venous return is through corresponding veins.
A urinary stone (calculus) may develop in any organ of the urinary system. A renal stone (kidney stone) is one that forms in
a kidney (fig. 19.12). A renal stone may obstruct the ureter and
greatly increase the frequency of peristaltic waves in an attempt to
pass through. The pain from a lodged urinary stone is extreme and
extends throughout the pelvic area. A lodged urinary stone also
causes a sympathetic ureterorenal reflex that results in constriction of
renal arterioles, thus reducing the production of urine in the kidney
on the affected side.
calculus: L. calculus, small stone


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Chapter 19
Urinary System
685
Ureter
Uterus
Symphysis
pubis
Urinary bladder
Symphysis pubis
Rectum
Rectum
Vagina
Urethra
(a)
(b)
FIGURE 19.13 A sagittal section of the male and female pelves. In the male (a), the urinary bladder is located between the symphysis pubis
and the rectum. In the female (b), the urinary bladder is located between the symphysis pubis and the uterus and upper portion of the vagina. In
a female, the volume capacity of the urinary bladder is diminished during the last trimester of pregnancy, when the greatly enlarged uterus exerts
constant pressure on the urinary bladder.
Urinary Bladder
trigone: L. trigonum, triangle
CHAPTER 19
The urinary bladder is a saccular organ for storage of urine. It is

located just posterior to the symphysis pubis, anterior to the rectum (fig. 19.13). In females, the urinary bladder is in contact
with the uterus and vagina. In males, the prostate is positioned
below the urinary bladder.
The shape of the urinary bladder is determined by the volume of urine it contains. An empty urinary bladder is pyramidal;
as it fills, it becomes ovoid and bulges upward into the abdominal
cavity. The median umbilical ligament, a fibrous remnant
of the embryonic urachus (see Developmental Exposition,
pp. 689690), extends from the anterior and superior border of
the urinary bladder toward the umbilicus. The base of the urinary
bladder receives the ureters, and the urethra exits at the inferior
angle, or apex. The region surrounding the urethral opening is
known as the neck of the urinary bladder.
The wall of the urinary bladder consists of four layers. The
mucosa (fig. 19.14), the innermost layer, is composed of transitional epithelium that becomes thinner as the urinary bladder
distends and the cells are stretched. Further distension is permitted by folds of the mucosa, called rugae (roo'je), which can be
seen when the urinary bladder is empty. Fleshy flaps of mucosa,
located where the ureters pierce the urinary bladder, act as valves
to prevent a reverse flow of urine toward the kidneys as the urinary bladder fills. A triangular area known as the trigone (tri'gon)
FIGURE 19.14 The histology of the urinary bladder (50).

is formed on the mucosa between the two uretal openings and
the single urethral opening (fig. 19.15). The internal trigone
lacks rugae; it is therefore smooth in appearance and remains relatively fixed in position as the urinary bladder changes shape
during distension and contraction.

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Urinary
bladder
Urinary
bladder
Urethra
Prostatic part
of urethra
Membranous
part of urethra
Bulbourethral
gland
(b)
Corpus
cavernosum
penis
CHAPTER 19
Spongy part
of urethra
(a)
FIGURE 19.15 A frontal section of the male and female urinary bladder and urethra. (a) The urethra of a male transports both urine and seminal fluid. It consists of a prostatic part that passes through the prostate, a membranous part that passes through the urogenital diaphragm, and a
spongy part that passes through the penis. (b) The considerably shorter urethra of a female transports only urine.


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Chapter 19
Urinary System
687
The second layer of the urinary bladder, the submucosa,

functions to support the mucosa. The muscularis consists of
three interlaced smooth muscle layers collectively called the detrusor (de-troo'sor) muscle. At the neck of the urinary bladder,
the detrusor muscle is modified to form the superior (called the
internal urethral sphincter) of two muscular sphincters surrounding the urethra. The outer covering of the urinary bladder is the
adventitia. It appears only on the superior surface of the urinary
bladder and is actually a continuation of the parietal peritoneum.
The arterial supply to the urinary bladder comes from the superior and inferior vesicular arteries, which arise from the internal
iliac arteries. Blood draining the urinary bladder enters a vesicular
venous plexus and then empties into the internal iliac veins.
The autonomic nerves serving the urinary bladder are derived from pelvic plexuses. Sympathetic innervation arises from
the last thoracic and first and second lumbar spinal nerves to
serve the trigone, urethral openings, and blood vessels of the urinary bladder. Parasympathetic innervation arises from the second, third, and fourth sacral nerves to serve the detrusor muscle.
The sensory receptors of the urinary bladder respond to distension and relay impulses to the central nervous system via the
pelvic spinal nerves.
The urethra of the male serves both the urinary and reproductive systems. It is about 20 cm (8 in.) long, and is S-shaped
because of the shape of the penis (see fig. 19.13). Three portions
can be identified in the male urethra: the prostatic part, the
membranous part, and the spongy part (fig. 19.15).
The prostatic part of the urethra is the proximal portion,
about 2.5 cm long, that passes through the prostate located near
the neck of the urinary bladder. The portion of the urethra receives drainage from small ducts of the prostate and two ejaculatory ducts of the reproductive system.
The membranous part of the urethra is the short portion
(0.5 cm) that passes through the urogenital diaphragm and proximal portion of the penis. The external urethral sphincter muscle
is located in this portion.
The spongy part of the urethra is the longest portion (15
cm), extending from the outer edge of the urogenital diaphragm
to the external urethral orifice on the glans penis. This portion is
surrounded by erectile tissue as it passes through the corpus spongiosum of the penis. The ducts of the bulbourethral glands (Cowpers glands) of the reproductive system attach to the spongy part
of the urethra near the urogenital diaphragm.
The urinary bladder becomes infected easily, and because a

womans urethra is so much shorter than a mans, women are
particularly susceptible to urinary bladder infections. A urinary bladder infection, called cystitis, may easily ascend from the urinary bladder to the ureters, because the mucous linings are continuous. An
infection that involves the renal pelvis is called pyelitis; if it continues
into the nephrons, it is known as pyelonephritis. To reduce the risk of
these infections, a young girl should be taught to wipe her anal region in a posterior direction, away from the urethral orifice, after a
bowel movement.
Micturition
Urethra
detrusor: L. detrudere, thrust or forced down
Cowpers glands: from William Cowper, English surgeon, 16661709
CHAPTER 19
The tubular urethra (yoo-re'thra) (fig. 19.15) conveys urine from

the urinary bladder to the outside of the body. The urethral wall
has an inside lining of mucous membrane surrounded by a relatively thick layer of smooth muscle, the fibers of which are directed longitudinally. Specialized urethral glands, embedded in
the urethral wall, secrete protective mucus into the urethral canal.
Two muscular sphincters surround the urethra. The involuntary smooth muscle sphincter, the superior of the two, is the
internal urethral sphincter, which is formed from the detrusor
muscle of the urinary bladder. The lower sphincter, composed of
voluntary skeletal muscle fibers, is called the external urethral
sphincter (fig. 19.15).
The urethra of the female is a straight tubular organ, about
4 cm (1.5 in.) long, that empties urine through the urethral orifice into the vestibule between the labia minora. The urethral
orifice is positioned between the clitoris and vaginal orifice (see
fig. 21.15). The female urethra has a single function: to transport
urine to the exterior.
Micturition (mik''tu-rish'un) commonly called urination, is a reflex action that expels urine from the urinary bladder. It is a
complex function that requires a stimulus from the urinary bladder and a combination of involuntary and voluntary nerve impulses to the appropriate muscular structures of the urinary
bladder and urethra.
In young children, micturition is a simple reflex action that
occurs when the urinary bladder becomes sufficiently distended.
Voluntary control of micturition is normally developed by the
time a child is 2 or 3 years old. Voluntary control requires the development of an inhibitory ability by the cerebral cortex and
maturation of various portions of the spinal cord.
The volume of urine produced by an adult averages about
1,200 ml a day, but it can range between 600 and 2,500 ml. The
average capacity of the urinary bladder is 700 to 800 ml. A volume of 200 to 300 ml will distend the urinary bladder enough to
stimulate stretch receptors and trigger the micturition reflex, creating a desire to urinate.
The micturition reflex center is located in the second,
third, and fourth sacral segments of the spinal cord. Following
stimulation of this center by impulses arising from stretch receptors in the urinary bladder, parasympathetic nerves that stimulate
the detrusor muscle and the internal urethral sphincter are activated. Stimulation of these muscles causes a rhythmic contraction of the urinary bladder wall and a relaxation of the internal
urethral sphincter. At this point, a sensation of urgency is perceived in the brain, but there is still voluntary control over
the external urethral sphincter. At the appropriate time, the

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19. Urinary System
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FIGURE 19.16 Innervation of the ureter, urinary bladder, and urethra.
CHAPTER 19
TABLE 19.2 Events of Micturition

1. The urinary bladder becomes distended as it fills with urine.
2. Stretch receptors in the urinary bladder wall are stimulated, and
impulses are sent to the micturition center in the spinal cord.
3. Parasympathetic nerve impulses travel to the detrusor muscle and the
internal urethral sphincter.
4. The detrusor muscle contracts rhythmically, and the internal urethral
sphincter relaxes.
5. The need to urinate is sensed as urgent.
6. Urination is prevented by voluntary contraction of the external
urethral sphincter and by inhibition of the micturition reflex by
impulses from the midbrain and cerebral cortex.
7. Following the decision to urinate, the external urethral sphincter is
relaxed, and the micturition reflex is facilitated by impulses from the
pons and the hypothalamus.
8. The detrusor muscle contracts, and urine is expelled through the
urethra.
9. Neurons of the micturition reflex center are inactivated, the detrusor
muscle relaxes, and the urinary bladder begins to fill with urine.
conscious activity of the brain activates the motor nerve fibers

(S4) to the external urethral sphincter via the pudendal nerve
(S2, S3, and S4), causing the sphincter to relax and urination
to occur.
The innervation of the ureter, urinary bladder, and urethra
is shown in figure 19.16, and the events of micturition are summarized in table 19.2.
Urinary retention, or the inability to void, may occur postoperatively, especially following surgery of the rectum, colon, or internal reproductive organs. The difficulty may be due to nervous
tension, the effects of anesthetics, or pain and edema at the site of
the operation. If urine is retained beyond 6 to 8 hours, catheterization

may become necessary. In this procedure, a catheter (tube) is
passed through the urethra into the urinary bladder so that urine can
flow freely.
Knowledge Check
7. Describe the location and the structure of the ureters and
indicate the function of the muscularis layer.
8. Describe the structure of the urinary bladder. What structural modifications permit the organ to be distended?
9. Compare the male urinary system with that of the female.
10. Explain the mechanisms by which micturition is controlled, with reference to the structures involved.
Urology (yoo-rol'a-je) is the medical specialty concerned with
dysfunctions of the urinary system. Urinary dysfunctions can
be congenital or acquired; they may result from physical
trauma or from conditions that secondarily involve the urinary
organs.
of the Urinary Organs
Abnormalities of the organs of the urinary system occur in about
12% of newborn babies. These deviations range from insignificant anomalies to those that are incompatible with life.


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The Urinary System
formed. The separation in the male is not totally complete, however, because the urethra serves to transport both urine and
semen. The development of both systems is initiated during the
embryonic stage, but the development of the urinary system starts
and ends sooner than that of the reproductive system.
Three successive types of kidneys develop in the human
embryo: the pronephros, mesonephros, and metanephros (exhibit I).
The third type, or metanephric kidney, persists as the permanent
kidney.
EXPLANATION
The urinary and reproductive systems originate from a specialized
elevation of mesodermal tissue called the urogenital ridge. The
two systems share common structures for part of the developmental period, but by the time of birth two separate systems have
(a)
(a1)
(b1)
(b)
(b2)
EXHIBIT I The embryonic development of the kidney. (a) An anterolateral view of an embryo at 5 weeks showing the position of a transverse cut depicted in (a1). During the sixth week (b and b1), the kidney is forming in the pelvic region and the mesonephric duct and gonadal ridge are prominent. The kidney begins migrating during the seventh week (b2), and the urinary bladder and gonad are formed.
689


Body
19. Urinary System
The pronephros (pro-nef'ros) develops during the fourth

week after conception and persists only through the sixth week.
Of the three developmental kidneys, it is the most superior in position on the urogenital ridge and is connected to the embryonic
cloaca by the pronephric duct. Although the pronephros is nonfunctional and degenerates in humans, most of its duct is used by
the mesonephric kidney (exhibit I), and a portion of it is important in the formation of the metanephros.
The mesonephros (mez''o-nef'ros) develops toward the
end of the fourth week as the pronephros becomes vestigial.
The mesonephros forms from an intermediate portion of the urogenital ridge and functions throughout the embryonic period of
development.
Although the metanephros (met''a-nef'ros) begins its formation during the fifth week, it does not become functional until
immediately before the start of the fetal stage of development, at
the end of the eighth week. The paired metanephric kidneys continue to form urine throughout fetal development. The urine is
expelled through the urinary system into the amniotic fluid.
The metanephros develops from two mesodermal sources
(exhibit II). The glomerular part of the kidney forms from a specialized caudal portion of the urogenital ridge called the
metanephrogenic mass. The tubular drainage portion of the kidneys forms as a diverticulum that emerges from the wall of the
mesonephric duct near the cloaca. This diverticulum, known as
the ureteric (yoo''re-ter'ik) bud, expands into the metanephrogenic mass to form the drainage pathway for urine. The stalk of
the ureteric bud develops into the ureter, whereas the expanded
terminal portion forms the renal pelvis, calyces, and papillary
ducts. A combination of the ureteric bud and metanephrogenic
mass forms the other tubular channels within the kidney.
Once the metanephric kidneys are formed, they begin to
migrate from the pelvis to the upper posterior portion of the abdomen. The renal blood supply develops as the kidneys become
positioned in the posterior body wall.
The McGrawHill
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(a)
(b)
(c)
The development of the kidneys illustrates the concept

that ontogeny (embryonic development) recapitulates
phylogeny (evolution). This means that the development of the
three kidney types follows an evolutionary pattern. The larvae of a
few of the more primitive vertebrates have functional pronephric
kidneys. Adult fishes and amphibians have mesonephric kidneys,
whereas adult reptiles, birds, and mammals have metanephric
kidneys.
The urinary bladder develops from the urogenital sinus,

which is connected to the embryonic umbilical cord by the fetal
membrane called the allantois (exhibit I). By the twelfth week,
the two ureters are emptying into the urinary bladder, the urethra
is draining, and the connection of the urinary bladder to the allantois has been reduced to a supporting structure called the urachus (yoo'ra-kus).
Occasionally a patent urachus is present in a newborn and
is discovered when urine is passed through the umbilicus,
especially if there is a urethral obstruction. Usually, however, it remains undiscovered until an enlarged prostate in an elderly male
obstructs the urethra and forces urine through the patent urachus
and out the umbilicus (navel).
690
(d1)
(d)
EXHIBIT II The development of the matanephric kidney (a) at

4 weeks, (b) 5 weeks, (c) 7 weeks, and (d) at birth. (d1) A magnified view of the arrangement of papillary ducts within a papilla.


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Chapter 19
Urinary System
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Unilateral double kidney

Pelvic kidney
Creek
Epispadias
Hypospadias
FIGURE 19.18 Epispadias and hypospadias of the male urethra.
Rosette kidney
Horseshoe kidney
Ureters
Aberrant renal artery

causing urinary
obstruction
Aberrant renal
arteries
Creek
Urinary Bladder
Protrusion of the posterior wall of the urinary bladder, called exstrophy of the urinary bladder, occurs when the wall of the perineum
fails to close. Associated with this condition, which occurs in about
1 in 40,000 births, are defective urethral sphincter muscles.
FIGURE 19.17 Congenital anomalies involving the kidneys.
Urethra
Kidneys
Common malformations of the kidneys are illustrated in
figure 19.17. One common deformity is renal agenesis (a-jen'esis), the unilateral absence of a kidney as a result of failure of a
uretic bud to develop. Renal ectopia is the condition in which
one or both kidneys are in an abnormal position. Generally, this
condition occurs when a kidney remains in the pelvic area.
Horseshoe kidney refers to a fusion of the kidneys across the
midline. The incidence of this asymptomatic condition is about
1 in 600.
The most common anomaly of the urethra is hypospadias

(hi''pospad'e-as), a condition in which the urethra of the male
opens on the underside of the penis instead of at the tip of the
glans penis (fig. 19.18). There is a similar defect in the female in
which the urethra opens into the vagina. Epispadias is a failure
of closure on the dorsum of the penis. Hypospadias and epispadias can be corrected surgically.
Symptoms and Diagnosis

of Urinary Disorders
Normal micturition is painless. Dysuria (dis-yur'e-a), or painful
urination, is a sign of a urinary tract infection or obstruction of
the urethraas in an enlarged prostate in a male. Hematuria
CHAPTER 19
Polycystic kidney
Duplication of the ureters and the associated renal pelvis is a

frequent anomaly of the urinary tract. Unilateral duplication occurs in 1 in 200 births, whereas bilateral duplication occurs in 1
in 1,200 births. Occasionally there is partial duplication, which
increases the propensity for urinary infections. A completely duplicated ureter frequently opens into areas other than the urinary
bladder requiring surgical correction.

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Penis
Urinary
bladder
ence of microscopic bacteria, crystals, and casts. Casts are accumulations of proteins that leaked through the glomeruli and were
pushed through the tubules, like toothpaste through a tube.
Infections of the Urinary Organs

Testis
CHAPTER 19
FIGURE 19.19 Cystoscopic examination of a male.
means blood in the urine and is usually associated with trauma.

Bacteriuria means bacteria in the urine, and pyuria is the term
for pus in the urine, which may result from a prolonged infection. Oliguria is a scanty output of urine, whereas polyuria is an
excessive output. Low blood pressure and kidney failure are two
causes of oliguria. Uremia (yoo-re'me-a) is a condition in which
substances ordinarily excreted in the urine accumulate in the
blood. Enuresis (en''yu-re'sis), or incontinence, is the inability to
control micturition. It may be caused by psychological factors or
by structural impairment.
The palpation and inspection of urinary organs is an important aspect of physical assessment. The right kidney is palpable in
the supine position; the left kidney usually is not. The distended
urinary bladder is palpable along the superior pelvic rim.
The urinary system may be examined using radiographic
techniques. An intravenous pyelogram (pi'e-logram) (IVP) permits radiographic examination of the kidneys following the injection of radiopaque dye. In this procedure, the dye that has
been injected intravenously is excreted by the kidneys so that
the renal pelvises and the outlines of the ureters and urinary
bladder can be observed in a radiograph.
Cystoscopy (s-stos'ko-pe) is the inspection of the inside of
the urinary bladder using an instrument called a cystoscope
(fig. 19.19). By means of this technique, tissue samples can be
obtained, as well as urine samples from each kidney prior to mixing in the urinary bladder. Once the cystoscope is in the urinary
bladder, the ureters and pelvis can be viewed through urethral
catheterization and inspected for obstructions.
A renal biopsy is a diagnostic test for evaluating certain
types and stages of kidney diseases. The biopsy is performed either through a skin puncture (closed biopsy) or through a surgical incision (open biopsy).
Urinalysis is a simple but important laboratory aspect of a
physical examination. The voided urine specimen is tested for
color, specific gravity, chemical composition, and for the pres-
Urinary tract infections (UTIs) are a significant cause of illness

and are also a major factor in the development of chronic renal
failure. Females are more susceptible to urinary tract infections
than are males because the urethra is shorter in females and the
urethral and anal openings are closer together. The incidence of
infection increases directly with sexual activity and aging in
both sexes.
Infections of the urinary tract are named according to the
infected organ. An infection of the urethra is called urethritis
(yoo''re-thri'tis) and involvement of the urinary bladder is cystitis
(sis-ti'tis). Cystitis is frequently a secondary infection from some
other part of the urinary tract. Nephritis is inflammation of the
kidney tissue. Glomerulonephritis (glo-mer''yu-lo-nefri'tis) is inflammation of the glomeruli. Glomerulonephritis may occur following an upper respiratory tract infection because antibodies
produced against streptococci bacteria can produce an autoimmune inflammation in the glomeruli. This inflammation may
permanently change the glomeruli and figure significantly in the
development of chronic renal disease and renal failure.
Any interference with the normal flow of urine, such as
from a renal stone or an enlarged prostate in a male, causes stagnation of urine in the renal pelvis and may lead to pyelitis.
Pyelitis is an inflammation of the renal pelvis and its calyces.
Pyelonephritis is inflammation involving the renal pelvis, the
calyces, and the tubules of the nephron within one or both kidneys. Bacterial invasion from the blood or from the lower urinary
tract is another cause of both pyelitis and pyelonephritis.
Trauma to the Urinary Organs

and Functional Impairments
Trauma
A sharp blow to a lumbar region of the back may cause a contusion or rupture of a kidney. Symptoms of kidney trauma include
hematuria and pain in the upper abdominal quadrant and flank
on the injured side.
Pelvic fractures from accidents may result in perforation of
the urinary bladder and urethral tearing. On a long automobile
trip, it is advisable to stop to urinate at regular intervals because
an attached seat belt over the region of a full urinary bladder can
cause rupture of the urinary bladder in even a relatively minor
accident. Urethral injuries are more common in men than in
women because of the position of the urethra in the penis. In a
straddle injury, for example, a man walking along a raised beam
may slip and compress his urethra and penis between the hard
surface and his pubic arch, rupturing the urethra.


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Chapter 19
Blood in
Urinary System
693
Blood out
Semipermeable
membrane
Wash
solution
in
Wash
solution
out
Waste
products
Blood elements
FIGURE 19.20 The hemodialysis process.
Obstruction
Renal Failure and Hemodialysis

Renal output of 50 to 60 cc of urine per hour is considered
normal. If the output drops to less than 30 cc per hour, it may
indicate renal failurethe loss of the kidneys ability to
maintain fluid and electrolyte balance and to excrete waste
products. Renal failure can be either acute or chronic. Acute
renal failure is the sudden loss of kidney function caused by
shock and hemorrhage, thrombosis, or other physical trauma
CHAPTER 19
The urinary system can become obstructed anywhere along the

tract. Calculi (stones) are the most common cause, but blockage
can also come from trauma, strictures, tumors or cysts, spasms or
kinks of the ureters, or congenital anomalies. If not corrected, an
obstruction causes urine to collect behind the blockage and generate pressure that may cause permanent functional and
anatomic damage to one or both kidneys. As a result of pressure
buildup in a ureter, a distended ureter, or hydroureter, develops.
Dilation in the renal pelvis is called hydronephrosis.
Urinary stones (calculi) are generally the result of infections or metabolic disorders that cause the excretion of large
amounts of organic and inorganic substances (see fig. 19.12). As
the urine becomes concentrated, these substances may crystalize
and form granules in the renal calyces. The granules then serve
as cores for further precipitation and development of larger calculi. This becomes dangerous when a calculus grows large
enough to cause an obstruction. The calculus also causes intense
pain when it passes through the urinary tract.
to the kidneys. The kidneys may sustain a 90% loss of their

nephrons through tissue death and still continue to function
without apparent difficulty. If a patient suffering acute renal
failure is stabilized, the nephrons have an excellent capacity
to regenerate.
A person with chronic renal failure cannot sustain life
independently. Chronic renal failure is the end result of kidney disease in which the kidney tissue is progressively destroyed. As renal tissue continues to deteriorate, the options
for sustaining life are hemodialysis (he''mo-di-al'-sis) or kidney
transplantation.
Hemodialysis equipment is designed to filter the wastes
from the blood of a patient who has chronic renal failure. The
patients blood is pumped through a tube from the radial artery
and passes through a machine, where it is cleansed and then returned to the body through a vein (fig. 19.20). The cleaning
process involves pumping the blood past a semipermeable cellophane membrane that separates the blood from an isotonic solution containing molecules needed by the body (such as glucose).
Waste products diffuse out of the blood through the membrane,
while glucose and other molecules needed by the body remain in
the blood.
More recent hemodialysis techniques include the use of the
patients own peritoneal membranes for filtering. Dialysis fluid is
introduced into the peritoneal cavity, and then, after a period of
time, discarded after wastes have accumulated. This procedure,
called continuous ambulatory peritoneal dialysis (CAPD), can be
performed several times a day by the patients themselves on an
outpatient basis.

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Urinary Incontinence
The inability to voluntarily retain urine in the urinary bladder is
known as urinary incontinence. It has a number of causes and may
be temporary or permanent. Emotional stress is a cause of temporary incontinence in adults. Permanent incontinence may result
from neurological trauma, various urinary diseases, tissue damage
within the urinary bladder or urethra, or weakness of the pelvic
floor muscles. Remarkable advances have been made in treating
permanent urinary incontinence through the implantation of an
artificial urethral sphincter.

The hematuria experienced by the patient is probably the result of laceration caused by the upper abdominal knife wound. The lower right
quadrant stab most likely did not violate the urinary tract. The course of
blood seen in the catheter begins and proceeds as follows: abdominal
aorta right renal artery smaller parenchymal artery through the
lacerated vessel(s) into the lacerated urinary collecting system, either
at the calyx or the renal pelvis or proximal right ureter urinary bladder into catheter. During the operation, the surgeon should keep in
mind that in 2% to 4% of the population only one kidney is present. If,
therefore, she is prompted to remove the damaged kidney, she should
first confirm the presence of a second functioning kidney. If a second
kidney is not present, every effort should be made to correct the problem without performing a nephrectomy, which would consign the patient to chronic hemodialysis or kidney transplant.
CHAPTER 19

A 41-year-old male presents in your office
because his urine has a pink tinge. He also
reports an increase in his frequency of urination. The patient denies fever, chills, pain,
or foul odor from his urine.
The physical exam is normal. A routine
urinalysis reveals red blood cells too numerous
to count, no white blood cells, and no bacteria. You order an intravenous urogram (IVU)
(a), which is an injection of intravenous contrast material for an exam of the urinary system, and CT scan (b). (B = bladder.)
QUESTIONS
1. What is the large dark mass (filling
defect) seen in the urinary bladder
(indicated with the letter B) on the
IVU? Is it the source of the bleeding?
2. On the CT, white contrast fills the
lumen of the urinary bladder (indicated
with an arrow). Why does the patient
have increased frequency of urination?
3. Transitional carcinoma has a 510%
incidence of synchronous tumors.
Synchronous tumors are tumors of the
same cell type occuring at the same
time in separate locations. What other
structures must be evaluated for the
presence of tumors?
(b)
(a)


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19. Urinary System
The McGrawHill
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Chapter 19
Urinary System
695

A 41-year-old female presents at the emergency room with left lumbar pain that at
times radiates to the left groin. She describes
the pain as intermittent and crampy, but denies any fever or blood in her urine.
Physical exam shows a nontender abdomen and no evidence of costovertebral
angle tenderness. Routine urinalysis shows
minimal red blood cells, no white blood
cells, and no bacteria. You order an intravenous urogram.
QUESTIONS
1. On the precontrast image, what is the
density indicated by the arrow?
2. What effect does this have on the
ureter as demonstrated by the
postcontrast image?
3. Why is the patient having intermittent
pain?
Precontrast image
Postcontrast image
Chapter Summary
Introduction to the Urinary System
(p. 676)
Kidneys (pp. 676684)

1. The kidneys are retroperitoneal, embedded
in a renal adipose capsule.
2. Each kidney is contained by a renal
capsule and divided into an outer renal
cortex and an inner renal medulla.
(a) The renal medulla is composed of
renal pyramids separated by renal
columns.
(b) The renal papillae empty urine into
the minor calyces and then into the
major calyces, which drain into the
renal pelvis. From there, urine flows
through the ureter.
Ureters, Urinary Bladder, and Urethra

(pp. 684688)
1. Urine is channeled from the kidneys to
the urinary bladder by the ureters and
expelled from the urinary bladder through
the urethra. The detrusor muscle of the

urinary bladder and the sphincter muscles
of the urethra are used in the control of
micturition.
(a) Each ureter contains three layers: the
mucosa, muscularis, and adventitia.
(b) The lumen of the urinary bladder is
lined by transitional epithelium,
which is folded into rugae. These
structures enhance the ability of the
urinary bladder to distend.
(c) The urethra has an internal sphincter
of smooth muscle and an external
sphincter of skeletal muscle.
(d) The male urethra conducts urine
during urination and seminal fluid
during ejaculation. The female
urethra is much shorter than that of a
male and conducts only urine.
(e) The male urethra is composed of
prostatic, membranous, and spongy
portions.
2. Micturition is controlled by reflex centers
in the second, third, and fourth sacral
segments of the spinal cord.
CHAPTER 19
1. The urinary system consists of two

kidneys, two ureters, the urinary bladder,
and the urethra.
2. The urinary system maintains the
composition and properties of body fluid,
which establishes the extracellular
environment. The end product of the
urinary system is urine, which is voided
from the body during micturition.
3. Each kidney contains more than a million

microscopic functional units called
nephrons.
(a) Filtration occurs in the glomeruli,
which receive blood from afferent
glomerular arterioles.
(b) Glomerular blood is drained by
efferent glomerular arterioles that
deliver blood to peritubular capillaries
surrounding the nephron tubules.
(c) The glomerular capsules and distal
convoluted tubules are located in the
renal cortex.
(d) The nephron loops are located in the
renal medulla.
(e) Filtrate from the distal convoluted
tubules are drained into papillary
ducts that extend through the renal
medulla to empty urine into the
calyces.

696
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19. Urinary System
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Review Activities
CHAPTER 19
Objective Questions
the renal pyramids is false?
(a) They are located in the renal
medulla.
(b) They contain glomeruli.
(c) They contain papillary ducts.
(d) They are supported by renal columns.
2. Renal vessels and the ureter attach at the
concave medial border of the kidney
called
(a) the renal
(c) the calyx.
pelvis.
(d) the hilum.
(b) the urachus.
3. The renal medulla of the kidney contains
(a) glomerular capsules.
(b) glomeruli.
(c) renal pyramids.
(d) adipose capsules.
4. Urine flowing from the papillary ducts
enters directly into
(a) the renal calyces.
(b) the ureter.
(c) the renal pelvis.
(d) the distal convoluted tubules.
5. Which of the following statements
concerning the kidneys is false?
(a) They are retroperitoneal.
(b) They each contain 8 to 15 renal
pyramids.
(c) They each have two distinct
regionsthe renal cortex and renal
medulla.
(d) They are positioned between the
third and fifth lumbar vertebrae.
6. A renal stone (calculus), would most
likely cause stagnation of urine in which
portion of the urinary system?
(a) the urinary bladder
(b) the renal column
(c) the ureter
(d) the renal pelvis
(e) the urethra
7. Distention of the urinary bladder is
possible because of the presence of
(a) rugae.
(b) the trigone.
(c) the adventitia.
(d) the transitional epithelium.
(e) both a and d.
8. The detrusor muscle is located in
(a) the kidneys.
(b) the ureters.
(c) the urinary bladder.

(d) the urethra.
9. The internal urethral sphincter is
innervated by
(a) sympathetic neurons.
(b) parasympathetic neurons.
(c) somatic motor neurons.
(d) all of the above.
metanephric kidneys is true?
(a) They become functional at the end of
the eighth week.
(b) They are active throughout fetal
development.
(c) They are the third pair of kidneys to
develop.
(d) All of the above are true.
Essay Questions
1. Describe the location of the kidneys in
relation to the abdominal cavity and the
peritoneal membranes.
2. Diagram the kidney structures that can be
identified in a coronal section.
3. Describe how the kidney is supported
against the posterior abdominal wall. How
is this support related to the condition
called renal ptosis?
4. Trace a drop of blood from an interlobular
artery through a glomerulus and into an
interlobular vein. List in order all the
vessels through which the blood passes.
How do structural differences in afferent
and efferent glomerular arterioles ensure
the high blood pressure needed for filtrate
formation?
5. In a male, trace the path of urine from the
site of filtration at the renal corpuscle to
the outside of the body. List in order all
the structures through which the urine
passes.
6. What is a nephron? Describe the two
types of nephrons found in a kidney. Why
are nephrons considered the functional
units of the urinary system?
7. Describe the mechanism involved in the
passage of urine from the renal pelvis to
the urinary bladder.
8. Describe the urinary bladder with regard
to position, histological structure, blood
supply, and innervation.
9. Compare and contrast the urethra in the
male and female.

10. What is the micturition reflex? Discuss

the physiological and functional events of
a voluntary micturition response.
11. What is a metanephrogenic mass? A
ureteric bud? Discuss the sequential
development of these embryonic
structures to form the urinary system.
How can a greater knowledge of this
development process lead to a better
understanding of congenital
abnormalities?
12. Briefly describe the purpose of cytoscopy
and urinalysis.
13. List four common congenital
malformations of the urinary system.
Which of these require surgical
correction?
14. Define dysuria, hematuria, bacteriuria,
pyuria, oliguria, polyuria, uremia, and
enuresis.
1. Why is it more accurate to refer to the
kidneys and associated structures as the
urinary system rather than the excretory
system?
2. Treatment with sulfa medications such as
Gantrisin (sulfisoxazole) and broadspectrum antibiotics such as tetracycline
or ampicillin usually clear up the
symptoms of cystitis very quickly. What is
the danger of discontinuing the prescribed
medication as soon as the symptoms
are gone?
3. The neighborhood day-care center wont
accept children who are still in diapers.
Youve tried to toilet train your
15-month-old boy, but you havent made
any progress at all. Should you persist in
your efforts, or would it be better to wait?
Explain.
4. What functions of a real kidney does an
artificial kidney (dialysis machine) fail to
duplicate?
5. Your friends baby is due next month, and
she is constantly running to the bathroom
to urinate. Can you explain why?
6. Explain why a male should be particularly
concerned if he has difficulty voiding
urine.


Development
20. Male Reproductive

System
Male Reproductive System
The McGrawHill
Companies, 2001
20
Introduction to the Male
Reproductive System 698
Perineum and Scrotum 700
Testes 702
Spermatic Ducts, Accessory
Reproductive Glands,
and the Urethra 707
Penis 710
Mechanisms of Erection, Emission,
and Ejaculation 712
The Reproductive System 716
Chapter Summary 723
Clinical Case Study

During a routine physical exam, a 27-year-old man mentioned to his family doctor that he and
his wife had been unable to conceive a child after nearly 2 years of trying. He added that his
wife had taken the initiative of having a thorough gynecological evaluation in an attempt to
find out what was causing the problem. Her test findings revealed no physical conditions that
could be linked to infertility. Upon palpating the patients testes, the doctor found nothing unusual. When he examined the scrotal sac above the testes, however, the doctor appeared perplexed. He informed his patient that two tubular structures, one for each testis, appeared to be
absent, and that they probably had been missing since birth. During a follow-up visit, the doctor told the patient that examination of his ejaculate revealed azoospermia (no viable sperm).
Explain how the result of the semenalysis relates to the patients physical exam findings.
What are the missing structures? Does it seem peculiar that the patient is capable of producing
an ejaculate? Why or why not?
FIGURE: Fertility specialists have made

remarkable advancements during the past
decade in treating couples with fertility
problems. A semenalysis reveals the relative
number of sperm (sperm count), sperm vitality,
and chemical nature of the fluid medium in a
sample ejaculate.

698
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Development

System
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INTRODUCTION TO THE MALE

REPRODUCTIVE SYSTEM
The organs of the male and female reproductive systems are
adapted to produce and allow the union of gametes that contain
specific genes. A random combination of the genes during sexual
reproduction results in the propagation of individuals with genetic
differences.
(the blastocyst), pregnancy, and delivery of a baby. The more

complex reproductive system of the female also provides a means
for nourishing the baby through the secretion of milk from the
mammary glands. In addition, like in the male, another function
is to produce and secrete sex hormones, which maintain the female sex organs and contribute to the female libido.
In this chapter we will consider the anatomy of the male
reproductive system; the female reproductive system is the focus
of chapter 21.
Objective 1
Explain why sexual reproduction is biologically

advantageous.
Objective 2
List the functions of the male reproductive

system and compare them with those of the female.
Objective 3
Distinguish between primary and secondary
CHAPTER 20
sex organs.
Unlike other body systems, the reproductive system is not essential for the survival of the individual; it is, however, required for
the survival of the species. It is through reproduction that additional individuals of a species population are produced and the
genetic code passed from one generation to the next. This can be
accomplished by either asexual or sexual reproduction. But sexual reproduction, in which genes from two individuals are combined in random ways with each new generation, offers the
overwhelming advantage of introducing great variability into a
population. This variability of genetic constitution helps ensure
that some members of a population will survive changes in the
environment over evolutionary time.
The reproductive system is unique in two other respects.
First, the fact that it does not become functional until it is
turned on at puberty by the actions of sex hormones sets the reproductive system apart. By contrast, all of the other body systems
are functional at birth, or shortly thereafter. Second, whereas the
other organ systems of the body exhibit slight sexual differences,
no other system approaches the level of dissimilarity of the reproductive system. Because sexual reproduction requires the production of two types of gametes (gam'ets), or sex cells, the species has
a male and female form, each with its own unique reproductive
system. The male and female reproductive systems complement
each other in their common purpose of producing offspring.
The functions of the male reproductive system are to produce the male gametes, spermatozoa (sper-mat''o-zo'a), and to
transfer them to the female through the process of coitus (sexual
intercourse) or copulation. Another function of the male reproductive system is to produce and secrete sex hormones, which
maintain the male sex organs and contribute to the male libido
(sex drive). The female not only produces her own gametes
(called oocytes [o'o-sts], or ova) and receives the sperm from the
male, but her reproductive organs are specialized to provide sites
for fertilization, implantation of the developing embryonic mass
gamete: Gk. gameta, husband or wife

spermatozoa: Gk. sperma, seed; zoon, animal
Categories of Reproductive Structures

The structures of the male reproductive system can be categorized on a functional basis as follows:
Primary sex organs. The primary sex organs are called gonads; specifically, the testes in the male. Gonads produce
the gametes, or spermatozoa, and produce and secrete sex
hormones. The secretion of male sex hormones, called androgens, at the appropriate times and in sufficient quantities causes the development of secondary sex organs and
the expression of secondary sex characteristics.
Secondary sex organs. Secondary sex organs are those
structures that are essential in caring for and transporting
spermatozoa. The three categories of secondary sex organs
are the sperm-transporting ducts, the accessory glands, and
the copulatory organ. The ducts that transport sperm include the epididymides, ductus deferentia, ejaculatory ducts,
and urethra. The male accessory reproductive glands are
the seminal vesicles, the prostate, and the bulbourethral
glands. The penis, which contains erectile tissue, is the copulatory organ. The scrotum is a pouch of skin that encloses
and protects the testes.
Secondary sex characteristics. Secondary sex characteristics are features that are not essential for the reproductive
process but are generally considered sexual attractants. In
the male, they include body physique, body hair, and
voice pitch.
The organs of the male reproductive system are shown in
figure 20.1 and their functions are summarized in table 20.1.
Knowledge Check
1. What is the primary importance of sexual reproduction?
2. What are the functions of the male reproductive system?
3. List the organs of the male reproductive system and indicate whether they are primary or secondary sex organs.
4. With respect to the other body systems, discuss the latent
(delayed) development of the reproductive system.
androgen: Gk. andros, male producing


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Chapter 20
699
Ureter
Ampulla of
ductus deferens
Ampulla of
ductus deferens
Ejaculatory duct of
ductus deferens
FIGURE 20.1 Organs of the male reproductive system. (a) a sagittal view and (b) a posterior view.
CHAPTER 20
Testis (sectioned
to show testicular
lobules)

Unit 7
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Development

System
TABLE 20.1 Reproductive Organs

of the Male
Organ(s)
Function(s)
Testes
Seminiferous
tubules
Produce spermatozoa
Interstitial cells
Secrete male sex hormones
Epididymides
Site of sperm maturation; store and convey

spermatozoa to the ductus deferentia
Ductus deferentia
Store spermatozoa; convey spermatozoa to the

ejaculatory ducts
Ejaculatory ducts
Receive spermatozoa and additives to produce

seminal fluid
Seminal vesicles
Secrete alkaline fluid containing nutrients and

prostaglandins; helps neutralize the acidic
environment of the vagina
Prostate
Secretes acidic fluid that enhances motility of

spermatozoa
Bulbourethral glands
Secrete fluid that lubricates the urethra and end

of the penis
Scrotum
Encloses and protects the testes; helps maintain

constant temperature for spermatogenesis
Penis
Conveys urine and seminal fluid to outside of

body; copulatory organ
PERINEUM AND SCROTUM

The perineum is the specific portion of the pelvic region that contains the external genitalia and the anal opening. The scrotum, a
pouch that supports the testes, is divided into two internal compartments by a connective tissue septum.
Objective 4
Describe the location of the perineum and

distinguish between the urogenital and anal triangles.
Objective 5
Discuss the structure and function of the scrotum.
Perineum
CHAPTER 20
The McGrawHill
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The perineum (per''-ne'um) is a diamond-shaped region between the symphysis pubis and the coccyx (fig. 20.2). It is the
muscular region of the outlet of the pelvis (see fig. 9.24). The
perineum is divided into a urogenital triangle in front and an anal
triangle in back. In the male perineum, the penis and scrotum are
attached at the anterior portion of the urogenital triangle and the
anus is located within the posterior portion of the anal triangle.
Scrotum
The saclike scrotum (skro'tum) is suspended immediately behind
the base of the penis (see figs. 20.1 and 20.2). The functions of
the scrotum are to support and protect the testes and to regulate
their position relative to the pelvic region of the body. The soft,
textured skin of the scrotum is covered with sparse hair in mature males and is darker in color than most of the other skin of
the body. It also contains numerous sebaceous glands.
The external appearance of the scrotum varies at different
times in the same individual as a result of the contraction and relaxation of the scrotal muscles. The dartos (dar'tos) is a layer of
smooth muscle fibers in the subcutaneous tissue of the scrotum.
The cremaster (kre-mas'ter) is a small band of skeletal muscle extending through the spermatic cord, a tube of fascia that also encloses the ductus deferens and testicular vessels and nerves (fig.
20.3). Both the dartos and the cremaster involuntarily contract
in response to low temperatures to move the testes closer to the
heat of the body in the pelvic region. The cremaster muscle is a
continuation of the internal abdominal oblique muscle of the abdominal wall, which is derived as the testes descend into the
scrotum. Because it is a skeletal muscle, it can be contracted voluntarily as well. When the scrotal muscles are contracted, the
scrotum appears tightly wrinkled as the testes are pulled closer to
the warmth of the body wall. High temperatures cause the dartos
and cremaster muscles to relax and the testes to be suspended
lower in the relaxed scrotum. The temperature of the testes is
maintained at about 35 C (95 F), or about 3.6 F below normal
body temperature, by the contraction or relaxation of the scrotal
muscles. This temperature is optimal for the production and storage of spermatozoa.
Although uncommon, male infertility may result from an excessively high temperature of the testes over an extended period
of time. Frequent hot baths or saunas may destroy sperm cells to the
extent that the sperm count will be too low to enable fertilization.
The scrotum is subdivided into two longitudinal compartments by a fibrous scrotal septum (fig. 20.3b). The purpose of
the scrotal septum is to compartmentalize each testis so that infection of one will generally not affect the other. In addition, the
left testis is generally suspended lower in the scrotum than the
right so that the two are not as likely to be compressed forcefully
together. The site of the scrotal septum is apparent on the surface of the scrotum along a median longitudinal ridge called the
perineal raphe (ra'fe). The perineal raphe extends forward to the
undersurface of the penis and backward to the anal opening (see
fig. 20.2).
The blood supply and innervation of the scrotum are extensive. The arteries that serve the scrotum are the internal pudendal branch of the internal iliac artery, the external pudendal
branch of the femoral artery, and the cremasteric branch of the
inferior epigastric artery. The venous drainage follows a pattern
similar to that of the arteries. The scrotal nerves are primarily
sensory; they include the pudendal nerves, ilioinguinal nerves,
and posterior cutaneous nerves of the thigh.
dartos: Gk. dartos, skinned or flayed
cremaster: Gk. cremaster, a suspender, to hang
septum: L. septum, a partition
pudendal: L. pudeo, to feel ashamed
raphe: Gk. raphe, a seam


Development

System
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Chapter 20
701
FIGURE 20.2 A superficial view of (a) the male perineum and (b) the female perineum (see chapter 21). The perineum is the region between
the symphysis pubis and coccyx. It is divided into urogenital and anal triangles, indicated in red and purple respectively.
(b)
FIGURE 20.3 Structural features of a testis within the scrotum. (a) a longitudinal view and (b) a transverse view.
CHAPTER 20
(a)

702
Unit 7

Development

System
The McGrawHill
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(a)
FIGURE 20.4 A diagram of the seminiferous tubules. (a) A sagittal section of a testis and (b) a transverse section of a seminiferous tubule.
Knowledge Check
5. Describe the location of the perineum.
6. Explain how the temperature of the testes is maintained.
Why is it important to maintain a particular testicular
temperature?
7. Describe the innervation and the blood supply to the scrotum.
TESTES
CHAPTER 20
Located within the scrotum, the testes produce spermatozoa and

androgens. Androgens regulate spermatogenesis and the development and functioning of the secondary sex organs.
Objective 6

of the testes.
Objective 7
Describe the effects of hormones on the growth

and development of the male reproductive organs and
explain how hormones are related to the development of
male secondary sex characteristics.
Objective 8
List the events of spermatogenesis and

distinguish between spermatogenesis and spermiogenesis.
Objective 9
Diagram the structure of a sperm cell and

explain the function of each of its parts.
Structure of the Testes

The testes (tes'tezsingular, testis) are paired, whitish, ovoid organs, each about 4 cm (1.5 in.) long and 2.5 cm (1 in.) in diameter. Each testis weighs between 10 and 14 g. Two tissue layers, or
tunics, cover the testes. The outer tunica vaginalis is a thin
serous sac derived from the peritoneum during the descent of the
testes. The tunica albuginea (al''byoo-jin'e-a) is a tough fibrous
membrane that directly encapsulates each testis (fig. 20.3). Fibrous inward extensions of the tunica albuginea partition the
testis into 250 to 300 wedge-shaped testicular lobules.
Each lobule of the testis contains tightly convoluted seminiferous tubules (fig. 20.4) that may exceed 70 cm (28 in.) in
length if uncoiled. The seminiferous tubules are the functional
units of the testis because it is here that spermatogenesis, the production of spermatozoa, occurs. Spermatozoa are produced at the
rate of thousands per secondmore than 100 million per day
throughout the life of a healthy, sexually mature male.
Various stages of meiosis can be observed in a section of a
seminiferous tubule (see fig. 20.7b). The process begins as specialized germinal cells, called spermatogonia (sper-mat''o-go'ne-a), undergo meiosis to produce, in order of advancing maturity, the
tunica: L. tunica, a coat

vaginalis: L. vagina, a sheath
albuginca: L. albus, white
spermatogonia: Gk, sperma, seed; gone, generation


Development

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Chapter 20
primary spermatocytes, secondary spermatocytes, and spermatids (see
fig. 20.6). Forming the walls of the seminiferous tubules are sustentacular (Sertoli) cells (also called nurse cells) that produce
and secrete nutrients for the developing spermatozoa embedded
between them (see fig. 20.7a). The spermatozoa are formed, but
not fully mature, by the time they reach the lumen of a seminiferous tubule.
Between the seminiferous tubules are specialized endocrine
cells called interstitial cells (cells of Leydig). The function of
these cells is to produce and secrete the male sex hormones. The
testes are thus considered mixed exocrine and endocrine glands
because they produce both spermatozoa and androgens.
Once the spermatozoa are produced, they move through
the seminiferous tubules and enter a tubular network called the
rete (re'te) testis for further maturation (fig. 20.4). Cilia are located on some of the cells of the rete testis. The spermatozoa are
transported out of the testis and into the epididymis through a
series of efferent ductules.
In all, it takes from 8 to 10 weeks for a spermatogonium to
become a mature spermatozoon. In the spermatic ducts, spermatozoa can remain fertile for several months, in a state of suspended animation. If they are not ejaculated, they eventually
degenerate and are absorbed into the blood.
The testes receive blood through the testicular arteries,
which in turn arise from the abdominal aorta immediately below
the origin of the renal arteries. The testicular veins drain the
testes. The testicular vein of the right side enters directly into
the inferior vena cava, whereas the testicular vein of the left side
empties into the left renal vein (see fig. 16.34).
Testicular nerves innervate the testes with both motor and
sensory neurons arising from the tenth thoracic segment of the
spinal cord. Motor innervation is primarily through sympathetic
neurons, but there is limited parasympathetic stimulation as well.
703
TABLE 20.2 Effect of Androgens

in the Male
Category
Effect
Sex differentiation
Development and growth of the embryonic

mesonephric ducts into the epididymides,
ductus deferentia, seminal vesicles, and
ejaculatory ducts
Development of the urogenital sinus and
tubercle into the prostate
Development of the male external genitalia
Spermatogenesis
At puberty: Meiotic division and early

maturation of spermatids
After puberty: Maintenance of
spermatogenesis
Secondary sex
characteristics
Growth and maintenance of the accessory

sex organs
Growth of the penis
Growth of facial and axillary hair
Body growth
Anabolic effects
Protein synthesis and muscle growth

Growth of bones
Growth of other organs (including the
larynx)
Erythropoiesis (red blood cell formation)
Brain and behavioral effects
Increased mass of sexual nuclei; male sex

drive and other aspects of male behavior
Androgens stimulate growth of the larynx (resulting in a

deepening of the voice), hemoglobin synthesis (males have
higher hemoglobin levels than females), and bone growth. The
effect of androgens on bone growth is self-limiting, however, because androgens are ultimately responsible for the conversion of
cartilage to bone in the epiphyseal plates, thus sealing the plates
and preventing further lengthening of the bones.
Endocrine Functions of the Testes
Spermatogenesis
Testosterone (tes-tos'te-ron) is by far the major androgen secreted

by the adult testes. Androgens are sometimes called anabolic
steroids because they stimulate the growth of muscles and other
structures (table 20.2). Increased testosterone secretion during
puberty is also required for the growth of the accessory sex organs, primarily the seminal vesicles and prostate. The removal of
androgens by castration results in atrophy of these organs.
Spermatogenesis (sper-mat''o-jen'e-sis) is the sequence of events

in the seminiferous tubules of the testes that leads to the production of spermatozoa. The germ cells that migrate from the yolk
sac to the testes during early embryonic development become
stem cells, or spermatogonia, within the outer region of the seminiferous tubules. Spermatogonia are diploid (2n) cells (with 46
chromosomes) that give rise to mature haploid (1n) gametes by a
process of reductive cell division called meiosis (mi-o'sis).
Meiosis occurs within the testes of males who have gone
through puberty and involves two nuclear divisions, as summarized in table 20.3. During the first part of this process, the DNA
duplicates (prophase I) and homologous chromosomes are separated (during anaphase I), producing two daughter cells (at
Sertoli cells: from Enrico Sertoli, Italian histologist, 18421910

cells of Leydig: from Franz von Leydig, German anatomist, 18211908
efferent ductules: L. efferre, to bring out; ducere, to lead
CHAPTER 20
The most common cause of male infertility is a condition called

varicocele (var'i-ko-sel). Varicocele occurs when one or both
of the testicular veins draining from the testes becomes swollen, resulting in poor circulation in the testes. A varicocele generally occurs
on the left side because the left testicular vein drains into the renal
vein. The blood pressure is higher here than it is in the inferior vena
cava, into which the right testicular vein empties.

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TABLE 20.3 Stages of Meiosis

Stage
Events
First Meiotic Division
Events
Second Meiotic Division

Chromosomes appear double-stranded.
The two strands, called chromatids,
contain identical DNA and are joined
together by a structure known as a
centromere.
Prophase II
Chromosomes appear, each containing

two chromatids.
Metaphase II
Chromosomes line up single file along

equator as spindle formation is
completed.
Homologous chromosomes pair up side

by side.
Anaphase II
Centromeres split and chromatids move

to opposite poles.
Metaphase I
Homologous chromosome pairs line up at

equator; spindle apparatus is completed.
Telophase II
Anaphase I
Homologous chromosomes separate; the

two members of a homologous pair
move to opposite poles.
Cytoplasm divides to produce two haploid

cells from each of the haploid cells
formed at telophase I.
Telophase I
Cytoplasm divides to produce two

haploid cells.
Prophase I
CHAPTER 20
Stage
telophase I). Because each daughter cell contains only one of

each homologous pair of chromosomes, the cells formed at the
end of this first meiotic division contain 23 chromosomes each and
are haploid (fig. 20.5). Each of the 23 chromosomes at this stage,
however, consists of two strands (called chromatids) of identical
DNA. During the second meiotic division, these duplicate chromatids are separated (at anaphase II), producing daughter cells
(at telophase II). Meiosis of one diploid spermatogonia cell
therefore produces four haploid daughter cells.
Actually, only about 1,500 stem cells migrate from the yolk
sac to the embryonic testes. In order to produce many millions of
spermatozoa throughout adult life, these spermatogonia duplicate
themselves by mitotic division, and only one of the two cells
now called a primary spermatocyteundergoes meiotic division
(Fig. 20.6). In this way, spermatogenesis can occur continuously
without exhausting the number of spermatogonia.
When a diploid primary spermatocyte completes the first
meiotic division (at telophase I), the two haploid daughter cells
thus produced are called secondary spermatocytes. At the end of
the second meiotic division, each of the two secondary spermatocytes produces two haploid spermatids (sper'ma-tidz). One primary spermatocyte therefore produces four spermatids.
The sequence of events in spermatogenesis is reflected in
the wall of the seminiferous tubule. The epithelial wall of the
tubule, called the germinal epithelium, is indeed composed of
germ cells in different stages of spermatogenesis. The spermatogonia and primary spermatocytes are located toward the outer
side of the tubule, whereas the spermatids and mature spermatozoa are located on the side facing the lumen (fig. 20.7).
At the end of the second meiotic division, the four spermatids produced by the meiosis of two secondary spermatocytes
are interconnectedtheir cytoplasm does not completely pinch
off at the end of each division. The development of these interconnected spermatids into separate mature spermatozoa (a
process called spermiogenesis) requires the participation of another type of cell in the tubulesthe sustentacular (nurse) cells,
or Sertoli cells.
Sustentacular (sus''ten-tak'yu-lar) cells are the only
nongerminal cell type in the tubules. Connected by tight junctions, they form a continuous layer around the circumference of
each tubule. In this way, the sustentacular cells form a blood-testis
barrier. The molecules from the blood must pass through the cytoplasm of sustentacular cells before entering germinal cells.
In the process of spermiogenesis (the conversion of spermatids to spermatozoa), most of the spermatid cytoplasm is eliminated. This occurs through phagocytosis by the sustentacular
cells of the residual bodies of cytoplasm from the spermatids. It
is believed that phagocytosis of the residual bodies may transmit
informational molecules from the germ cells to the sustentacular
cells. The sustentacular cells, in turn, may provide molecules
needed by the germ cells. It is known, for example, that the X
chromosome of the germ cells is inactive during meiosis. Because
this chromosome contains the genes needed to produce many essential molecules, it is believed that these molecules are provided
by the sustentacular cells while meiosis is taking place.
Structure of Spermatozoa
A mature sperm cell, or spermatozoon (sper-mat''o-zo'on) is a
microscopic, tadpole-shaped structure approximately 0.06 mm
long (fig. 20.7c and fig. 20.8). It consists of an oval head and an
elongated flagellum. Although the head of a spermatozoon lacks
mitochondria, it does contain a nucleus with 23 chromosomes.
The tip of the head, called the acrosome (ak'ro-som), contains


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Chapter 20
705
FIGURE 20.5 Meiosis, or reduction division. In the first meiotic division, the homologous chromosomes of a diploid parent cell are separated
into two haploid daughter cells. Each of these chromosomes contains duplicate strands, or chromatids. In the second meiotic division, these
chromatids are distributed to two new haploid daughter cells.
Recent findings indicate that ejaculated spermatozoa (plural of

spermatozoon) can survive up to 5 days at body temperature
longer than previously thought. In the average fertile male, up to 20%
of these sperm cells are defective, however, and are of no value. It is
not uncommon for them to have enlarged heads, dwarfed and misshapen heads, two flagella, or a flagellum that is bent. Spermatozoa
such as these are unable to propel themselves adequately.
Knowledge Check
8. Describe the location and structure of the testes.
9. What are the functions of the seminiferous tubules, germinal epithelial cells, and interstitial cells?
10. Discuss the effect of testosterone on the production of sperm
cells and the development of secondary sex characteristics.
11. Diagram a spermatozoon and its adjacent sustentacular cell
and explain the functions of sustentacular cells in the seminiferous tubules.
CHAPTER 20
enzymes that help the spermatozoon penetrate the ovum. The body
of the flagellum contains numerous mitochondria spiraled around a
filamentous core. The mitochondria provide the energy necessary
for locomotion. The flagellum propels the spermatozoon with a lashing movement. The maximum unassisted rate of spermatozoa movement is about 3 mm per hour.

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Spermatogonia
(46 chromosomes)
Wall of
seminiferous
tubule
Primary
spermatocytes
(46 chromosomes)
Secondary
spermatocytes
(23 chromosomes)
Spermatids
(23 chromosomes)
Sustentacular cell
Lumen of
seminiferous
tubule
Spermatozoa
(23 chromosomes)
(a)
(b)
Acrosome of head
Head
Body of flagellum
CHAPTER 20
Flagellum
FIGURE 20.6 The process of spermatogenesis. Spermatogonia

undergo mitotic division to replace themselves and produce a
daughter cell that will undergo meiotic division. This cell is called a
primary spermatocyte. Upon completion of the first meiotic division,
the daughter cells are called secondary spermatocytes. Each
of them complete a second meiotic division to form spermatids. Notice that the four spermatids produced by the meiosis of a primary
spermatocyte are interconnected. Each spermatid forms a mature
spermatozoon.
(c)
FIGURE 20.7 Seminiferous tubules and a spermatozoon. (a) The

stages of spermatogenesis within the germinal epithelium of a seminiferous tubule, in which the relationship between sustentacular cells
and developing spermatozoa is shown. (b) A photomicrograph of a
seminiferous tubule and surrounding interstitial tissue, and (c) a diagram of a spermatozoon.


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Chapter 20
707
Pseudostratified
ciliated columnar
epithelium
Smooth muscle
Sperm in lumen
of duct
Connective
tissue
FIGURE 20.9 The histology of the epididymis showing sperm in

the lumina (50).
FIGURE 20.8 A scanning electron micrograph of a spermatozoon

in contact with an egg.
SPERMATIC DUCTS, ACCESSORY

REPRODUCTIVE GLANDS,
AND THE URETHRA
The spermatic ducts store spermatozoa and transport them from
the testes to the outside of the body by way of the urethra. The
accessory reproductive glands provide additives to the spermatozoa to form semen, which is discharged from the erect penis during ejaculation.
Objective 10
Describe the location and structure of each

segment of the male duct system.
Objective 11
Describe the structure and contents of the

spermatic cord.
Objective 12

of the ejaculatory ducts, seminal vesicles, prostate, and
bulbourethral glands.
The spermatic ducts store spermatozoa and transport them from

the testes to the urethra. The accessory reproductive glands provide additives to the spermatozoa in the formation of semen.
Ductus Deferens
The ductus deferens (duk'tus def'er-enzplural, ductus
deferentia) is a fibromuscular tube about 45 cm (18 in.) long and
2.5 mm thick (see fig. 20.1) that conveys spermatozoa from the
epididymis to the ejaculatory duct. Also called the vas deferens
(plural, vasa deferentia), it exits the scrotum as it ascends along
the posterior border of the testis. From here, it penetrates the inguinal canal, enters the pelvic cavity, and passes to the side of
the urinary bladder on the medial side of the ureter. The ampulla
(am-pool'a) of the ductus deferens is the terminal portion that
joins the ejaculatory duct.
The histological structure of the ductus deferens includes a
layer of pseudostratified ciliated columnar epithelium in contact
with the tubular lumen and surrounded by three layers of tightly
packed smooth muscle (fig. 20.10). Sympathetic nerves from the
pelvic plexus serve the ductus deferens. Stimulation through
these nerves causes peristaltic contractions of the muscular layer,
which propel the stored spermatozoa toward the ejaculatory duct.
Much of the ductus deferens is located within a structure
known as the spermatic cord (see figs. 20.3 and 20.14). The
spermatic cord extends from the testis to the inguinal canal and
consists of the ductus deferens, the testicular artery and venous
Epididymis
The epididymis (ep''-did'-musplural, epididymides) is an elongated organ attached to the posterior surface of the testis (see
figs. 20.1 and 20.3). If it were uncoiled, it would measure 5.5 m
deferens: L. deferens, conducting away

ampulla: L. ampulla, a two-handled bottle
CHAPTER 20
Spermatic Ducts
(about 17 ft). The highly coiled, tubular tail portion contains

spermatozoa in their final stages of maturation (fig. 20.9). The
upper expanded portion is the head, and the tapering middle section is the body. The tail is continuous with the beginning portion of the ductus deferens; both store spermatozoa to be
discharged during ejaculation. The time required to produce mature spermatozoafrom meiosis in the seminiferous tubules to
storage in the ductus deferensis approximately 2 months.

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Seminal crypt
Smooth muscle
FIGURE 20.10 The histology of the ductus deferens (250).

plexus, nerves, the cremaster muscle, lymph vessels, and connective tissue. The portion of the spermatic cord that passes anterior
to the pubic bone can be easily palpated. The inguinal (ing'gwnal) canal is a passageway for the spermatic cord through the abdominal wall; it is a potentially weak area and a common site for
development of a hernia.
Ejaculatory Duct
The ejaculatory (e-jak'yoo-la-tor-e) duct is about 2 cm (1 in.) long
and is formed by the union of the ampulla of the ductus deferens
and the duct of the seminal vesicle. The ejaculatory duct then
pierces the capsule of the prostate on its posterior surface and
continues through this gland (see fig. 20.1). Both ejaculatory
ducts receive secretions from the seminal vesicles and then eject
the spermatozoa with its additives into the prostatic urethra to be
mixed with secretions from the prostate. The urethra, discussed
shortly, serves as a passageway for both semen and urine. It is the
terminal duct of the male duct system.
CHAPTER 20
Accessory Reproductive Glands

The accessory reproductive glands of the male include the seminal vesicles, the prostate, and the bulbourethral glands (see
fig. 20.1). The contents of the seminal vesicles and prostate are
mixed with the spermatozoa during ejaculation to form semen
(seminal fluid). The fluid from the bulbourethral glands is released in response to sexual stimulation prior to ejaculation.
Seminal Vesicles
The paired seminal vesicles, each about 5 cm (2 in.) long, are
convoluted club-shaped glands lying at the base of the urinary
bladder, in front of the rectum. They secrete a sticky, slightly alkaline, yellowish substance that contributes to the motility and
viability of spermatozoa. The secretion from the seminal vesicles
Glandular
epithelium
FIGURE 20.11 The histology of the seminal vesicle (10).

contains a variety of nutrients, including fructose, that provide
an energy source for the spermatozoa. It also contains citric acid,
coagulation proteins, and prostaglandins. The discharge from the
seminal vesicles makes up about 60% of the volume of semen.
Histologically, the seminal vesicle has an extensively
coiled mucosal layer (fig. 20.11). This layer partitions the lumen
into numerous intercommunicating spaces that are lined by pseudostratified columnar and cuboidal secretory epithelia (referred
to as glandular epithelium).
Blood is supplied to the seminal vesicles by branches from
the middle rectal arteries. The seminal vesicles are innervated by
both sympathetic and parasympathetic neurons. Sympathetic
stimulation causes the contents of the seminal vesicles to empty
into the ejaculatory ducts on their respective sides.
Prostate
The firm prostate (pros'tat) is the size and shape of a chestnut. It
is about 4 cm (1.6 in.) across and 3 cm (1.2 in.) thick and lies
immediately below the urinary bladder, where it surrounds the
beginning portion of the urethra (see fig. 20.1). The prostate is
enclosed by a fibrous capsule and divided into lobules formed by
the urethra and the ejaculatory ducts that extend through the
gland. The ducts from the lobules open into the urethra. Extensive bands of smooth muscle course throughout the prostate to
form a meshwork that supports the glandular tissue (fig. 20.12).
Contraction of the smooth muscle expels the contents from the
gland and provides part of the propulsive force needed to ejaculate the semen. The thin, milky-colored prostatic secretion assists sperm cell motility as a liquefying agent, and its alkalinity


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Chapter 20
709
Urethral glands
Tunica muscularis
Glandular alveoli
Lumen of urethra
Smooth muscle
Transitional
epithelium
Corpus
spongiosum
penis
Urethra
Ejaculatory duct
FIGURE 20.13 The histology of the urethra (10).

FIGURE 20.12 The histology of the prostate (50).
protects the sperm in their passage through the acidic environment of the female vagina. The prostate also secretes the enzyme
acid phosphatase, which is often measured clinically to assess
prostate function. The discharge from the prostate makes up
about 40% of the volume of the semen.
Blood is supplied to the prostate from branches of the middle rectal and inferior visceral arteries. The venous return forms
the prostatic venous plexus, along with blood draining from the
penis. The prostatic venous plexus drains into the internal iliac
veins. The prostate has both sympathetic and parasympathetic
innervation arising from the pelvic plexuses.
A routine physical examination of the male includes rectal palpation of the prostate. Enlargement or overgrowth of the glandular substance of the prostate, called benign prostatic hypertrophy
(BPH), is relatively common in older men. It constricts the urethra,
causing difficult urination. This condition may require surgery. In a
transurethral prostatic resection, excess prostatic tissue is removed
by use of a device inserted through the urethra.
Bulbourethral Glands
Urethra
The urethra of the male serves as a common tube for both the
urinary and reproductive systems. However, urine and semen
cannot pass through simultaneously because the nervous reflex
during ejaculation automatically inhibits micturition. The male
The wall of the urethra has an inside lining of mucous membrane, composed of transitional epithelium (fig. 20.13) and surrounded by a relatively thick layer of smooth muscle tissue called
the tunica muscularis. Specialized urethral glands, embedded in
the urethral wall, secrete mucus into the lumen of the urethra.
Knowledge Check
12. List in order the structures through which spermatozoa pass
as they travel from the testes to the urethra.
13. Differentiate between the ductus deferens and the spermatic cord.
14. Describe the structure and location of each of the accessory
reproductive glands.
15. Why is the position of the prostate of clinical importance?
16. Describe the three parts of the male urethra.
CHAPTER 20
The paired, pea-sized, bulbourethral (Cowpers) glands are located below the prostate (see fig. 20.1). Each bulbourethral gland
is about 1 cm (0.4 in.) in diameter and drains by a 2.5-cm (1-in.)
duct into the urethra. Upon sexual arousal and prior to ejaculation, the bulbourethral glands are stimulated to secrete a mucoid
substance that coats the lining of the urethra to neutralize the
pH of the urine residue. It also lubricates the tip of the penis in
preparation for coitus.
urethra is about 20 cm (8 in.) long, and S-shaped because of the

shape of the penis. As described in chapter 19 (see fig. 19.15), it
is divided into three regions, which are briefly reviewed here.
1. The prostatic part of the urethra is the 2.5-cm (1-in.)
proximal portion that passes through the prostate. The
prostatic urethra receives drainage from the small ducts of
the prostate and the two ejaculatory ducts.
2. The membranous part of the urethra is the 0.5-cm (0.2-in.)
portion that passes through the urogenital diaphragm. The
external urethral sphincter muscle is located in this region.
3. The spongy part of the urethra is the longest portion, extending from the outer edge of the urogenital diaphragm
to the external urethral orifice on the glans penis. About
15 cm (6 in.) long, this portion is surrounded by erectile
tissue as it passes through the corpus spongiosum of the
penis. The paired ducts from the bulbourethral glands attach to the spongy part of the urethra near the urogenital
diaphragm.

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Ureter
Ductus deferens
Urinary bladder
Ampulla of
ductus deferens
Seminal vesicle
Suspensory ligament
of penis
Prostate
Body of penis
Root of
the penis
Crus of penis
Bulb of penis
Dorsal vein of penis

Dorsal artery of penis
Spermatic cord
Dorsal nerve of penis

External spermatic
fascia
Corona glandis
Glans penis
Cremasteric muscle
Epididymis
Prepuce
Scrotum
Dorsal vein of penis
Deep dorsal vein of penis
Dorsal nerve of penis
Dorsal artery of penis
Deep artery of penis
Corpora cavernosa penis
Skin
Tunica albuginea
Areolar tissue
Septum penis
Tunica albuginea
Deep fascia
Skin
Urethra
Loose
connective tissue
Corpus spongiosum penis
Deep fascia
Frenulum of penis
Waldrop
FIGURE 20.14 The structure of the penis showing the attachment, blood and nerve supply, and the arrangement of the erectile tissue.
CHAPTER 20
PENIS
The penis, containing the spongy urethra and covered with
loose-fitting skin, is specialized with three columns of erectile tissue to become engorged with blood for insertion into the vagina
during coitus.
Objective 13
Describe the structure and function of the penis.
The penis (pe'nis) when distended, serves as the copulatory

organ of the male reproductive system. The penis and scrotum,
which are suspended from the perineum, constitute the external
genitalia of the male. Under the influence of sexual stimulation,
the penis becomes engorged with blood. This engorgement results from the filling of intricate blood sinuses, or spaces, in the
erectile tissue of the penis.
The penis is divided into a proximal attached root, which is
attached to the pubic arch; an elongated tubular body, or shaft; and
a distal cone-shaped glans penis (fig. 20.14). The root of the penis
expands posteriorly to form the bulb of the penis and the crus
(krus) of the penis. The bulb is positioned in the urogenital triangle of the perineum, where it is attached to the undersurface of
the urogenital diaphragm and enveloped by the bulbocavernosus
crus: L. crus, leg, resembling a leg

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FIGURE 21.19 The cycle of ovulation and menstruation.
CHAPTER 21
TABLE 21.2 Principal Events Surrounding Ovulation and Menstruation

1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
The hypothalamus releases GnRH, which stimulates the anterior pituitary.

The anterior pituitary releases small amounts of FSH and LH.
FSH stimulates the maturation of a follicle.
Follicular cells produce and secrete estrogen.
(a) Estrogen maintains secondary sexual traits.
(b) Estrogen causes the uterine lining to thicken.
Toward the middle of the cycle, a surge in the release of LH from the anterior pituitary causes ovulation.
Follicular cells become corpus luteum cells, which secrete estrogen and progesterone.
(a) Estrogen continues to stimulate the development of the endometrium.
(b) Progesterone stimulates the endometrium to become more glandular and vascular.
(c) Estrogen and progesterone inhibit the secretion of FSH and LH from the anterior pituitary.
If the oocyte is not fertilized, the corpus luteum degenerates.
As concentrations of estrogen and progesterone decline, blood vessels in the stratum basale of the endometrium constrict.
The stratum functionale of the endometrium disintegrates and sloughs away as menstrual flow.
The anterior pituitary, which is no longer inhibited, again secretes FSH and LH.
The cycle is repeated.
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The Female
Reproductive System
EXPLANATION
Although genetic sex is determined at fertilization, both sexes develop similarly through the indifferent stage of the eighth week.
The gonads of both sexes develop from the gonadal ridges medial
to the mesonephros. Primary sex cords form within the gonadal
ridges during the sixth week. The genital tubercle also develops
during the sixth week as an external swelling cephalic to the cloacal membranes.
The ovaries develop more slowly than do the testes. Ovarian
development begins at about the tenth week when primordial follicles begin to form within the ovarian medulla. Each of the primordial follicles consists of an oogonium (o''o-go'ne-um)
surrounded by a layer of follicular cells. Mitosis of the oogonia occurs during fetal development, so that thousands of germ cells are
formed. Unlike the male reproductive system, in which spermatogonia are formed by mitosis throughout life, all oogonia are formed
prenatally and their number continuously decreases after birth.
Ovary (before descent)

Ovarian ligament
Uterine tube
Urinary bladder
Ureter
Uterus
Tract of
mesonephric duct
Round ligament
of uterus
Urethra
Vaginal plate
The genital tract includes the uterus and the uterine tubes.
These organs develop from a pair of embryonic tubes called the
paramesonephric (mllerian) ducts. The paramesonephric ducts
form to the sides of the mesonephric ducts that give rise to the
kidneys. As the mesonephric ducts regress, the paramesonephric
ducts develop into the female genital tract (exhibit I). The lower
portions of the ducts fuse to form the uterus. The upper portions
remain unfused and give rise to the uterine tubes.
The epithelial lining of the vagina develops from the endoderm of the urogenital sinus. The formation of a thin membrane called the hymen (see page 718) separates the lumen of
the vagina from the urethral sinus. The hymen is usually perforated during later fetal development.
The external genitalia of both sexes appear the same during the indifferent stage of the eighth week (see page 718). A
prominent phallus (fal'us) forms from the genital tubercle, and a
urethral groove forms on the ventral side of the phallus. Paired
urethral folds surround the urethral groove on the lateral sides.
In the male embryo, these indifferent structures become masculinized by testosterone secreted by the testes. In the female embryo, in the absence of testosterone, feminization occurs.
In the process of feminization, growth of the phallus is inhibited, and the relatively small clitoris is formed. The urethral
folds remain unfused to form an inner labia minora, and the
labioscrotal swellings remain unfused to form the prominent labia
majora (table 21.3). The external genitalia of a female are completely formed by the end of the twelfth week.
The structure of the hymen is characterized by tremendous individual variation. The hymen of a baby girl may
be absent, or it may partially (or occasionally completely) cover
the vaginal orifice, in which case it is called an imperforate
hymen. An imperforate hymen is usually not detected until the first
menstruation (menarche), when the discharge cannot be expelled. If the hymen is present, it may be ruptured during childhood in the course of normal exercise. On the other hand, a
hymen may be so elastic that it persists even after coitus. Therefore, the presence of a hymen is not a reliable sign of virginity.
Vestibular
gland
(a)
Ovary (after descent)
Ovarian ligament
TABLE 21.3 Summary of Homologous

Structures
Uterine tube
Round ligament
of uterus
Inguinal canal
Vagina
Hymen
(b)
Creek
Labia majora
EXHIBIT I The development of the female genital tract. (a) A
Indifferent Stage
Male
Female
Gonads
Testes
Ovaries
Urethral groove
Membranous urethra
Vaginal vestibule
Genital tubercle
Glans penis
Clitoris
Urethral folds
Spongy urethra
Labia minora
Labioscrotal swellings
Scrotum
Labia majora
Bulbourethral glands
Vestibular glands
lateral view and (b) an anterior view.
mllerian ducts: from Johannes P. Mller, German physician, 180158

primordial: L. prima, first; ordior, to begin
follicle: L. diminutive of follis, bag
hymen: Gk. (mythology) Hymen was god of marriage; hymen, thin skin or
membrane
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Lumen of uterine tube
"Spill" of radiopaque medium

into peritoneal cavity
Cavity of uterus
Application tube for
hysterosalpingogram
FIGURE 21.20 A contrast radiograph of the uterine cavity and lumina of the uterine tubes (hysterosalpingogram).
Females are more prone to dysfunctions and diseases of the reproductive organs than are males because of cyclic changes in reproductive events, problems associated with pregnancy, and the
susceptibility of the female breasts to infections and neoplasms.
The termination of reproductive capabilities at menopause can
also cause complications as a result of hormonal changes. Gynecology (gi''ne-kol'o-je) is the specialty of medicine concerned with
dysfunction and diseases of the female reproductive system. Obstetrics is the specialty dealing with pregnancy and childbirth.
Frequently a physician will specialize in both obstetrics and gynecology (OBGYN).
A comprehensive discussion of the numerous clinical aspects of the female reproductive system is beyond the scope of
this text; thus, only the most important conditions will be addressed in the following sections. Coverage includes diagnostic
procedures, developmental abnormalities, problems involving
the ovaries and uterine tubes, problems involving the uterus, diseases of the vagina and vulva, and diseases of the breasts and
mammary glands. In addition, the more popular methods of birth
control are described.
CHAPTER 21
Diagnostic Procedures
A gynecological, or pelvic, examination is generally included in
a thorough physical examination of an adult female, especially
prior to marriage, during pregnancy, or if problems involving the
reproductive organs are suspected. In a gynecological examination, the physician inspects the vulva for irritations, lesions, or
abnormal vaginal discharge and palpates the vulva and internal
organs. Most of the internal organs can be palpated through the
vagina, especially if they are enlarged or tender. Inserting a lubricated speculum into the vagina allows visual examination of the
cervix and vaginal walls. A speculum is an instrument for opening or distending a body opening to permit visual inspection.
FIGURE 21.21 A tubal ligation involves the removal of a portion of

each uterine tube. In actual practice, cautery, clips, or rings are used
for tubal closure more often than ligation with suture thread.
In special cases, it may be necessary to examine the cavities

of the uterus and uterine tubes by hysterosalpingography (his''ter-osal''ping-gog'ra-fe) (fig. 21.20). This technique involves injecting
a radiopaque dye into the reproductive tract. The patency of the
uterine tubes, irregular pregnancies, and various types of tumors
may be detected using this technique. A laparoscopy (lap''a-ros'kope) permits visualization of the internal reproductive organs. The
entrance for the laparoscope may be through the umbilicus,
through a small incision in the lower abdominal wall, or through
the posterior fornix of the vagina into the rectouterine pouch.
Although a laparoscope is used primarily in diagnosis, it can be
used when performing a tubal ligation (fig. 21.21), a method of
sterilizing a female by tying off the uterine tubes.


Development

System
The McGrawHill
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Chapter 21
745
FIGURE 21.22 The technique of conducting a breast self-examination (BSE) involves (a) visual inspection and (b) palpation. The sites and incidence of occurrence of breast cancer are shown in (c).
left and right breast will not match exactlyfew womens

breasts do. Finally, squeeze the nipple of each breast gently
between the thumb and index finger to check for discharge. Any discharge from the nipple should be reported
to a physician.
2. Palpation during bathing. Examine the breasts during a
bath or shower when the hands will glide easily over wet
skin. With the fingers flat, move gently over every part of
each breast. Use the right hand to examine the left breast,
and the left hand for the right breast. Palpate for any lump,
hard knot, or thickening. If the breasts are normally fibrous
or lumpy (fibrocystic tissue), the locations of these lumps
should be noted and checked each month for changes in
size and location.
3. Palpation while lying down. To examine the right breast,
put a pillow or folded towel under the right shoulder. Place
the right hand behind the headthis distributes the breast
tissue more evenly on the rib cage. With the fingers of the
left hand held flat, press gently in small circular motions
around an imaginary clock face. Begin at outermost top of
CHAPTER 21
One diagnostic procedure that should be routinely performed by a woman is a breast self-examination (BSE). The importance of a BSE is not to prevent diseases of the breast but to
detect any problems before they become life-threatening. One in
nine women will develop breast cancer during her lifetime. Early
detection of breast cancer and follow-up medical treatment minimizes the necessary surgical treatment and improves the patients
prognosis. Breast cancer is curable if it is caught early.
A woman should examine her breasts monthly. If she has
not yet reached menopause, the ideal time for a BSE is 1 week
after her period ends because the breasts are less likely to be
swollen and tender at that time. A woman no longer menstruating should just pick any day of the month and do a BSE on that
same day on a monthly basis. Visual inspection and palpation
(Fig. 21.22) are equally important in doing a BSE. The steps involved in this procedure are the following:
1. Observation in front of a mirror. Inspect the breasts with
the arms at the sides. Next, raise the arms high overhead.
Look for any changes in the contour of each breasta
swelling, dimpling of skin or changes in the nipple. The

746
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(a)
(b)
(c)
(d)
(e)
(f)
FIGURE 21.23 Congenital uterine abnormalities. (a) A double uterus and double vagina, (b) a double uterus with a single vagina, (c) a bicornuate uterus, (d) a bicornuate uterus with a rudimentary left horn, (e) a septate uterus, and (f ) a unicornuate uterus.
the right breast for 12 oclock, then move to 1 oclock, and

so on around the circle back to 12 oclock. A ridge of firm
tissue in the lower curve of each breast is normal. Then
move in an inch, toward the nipple, and keep circling to
examine every part of the breast, including the nipple.
Also examine the armpit carefully for enlarged lymph
nodes. Repeat the procedure on the left breast.
malities and others are diagrammed in figure 21.23. In about 1 in

4,000 females, the vagina is absent. This is usually accompanied
by an absence of the uterus as well.
Another important diagnostic procedure is a Papanicolaou

(Pap) smear. The Pap smear permits a microscopic examination
of cells covering the tip of the cervix. Samples of cells are obtained by gently scraping the surface of the cervix with a specially designed wooden spatula. Women should have periodic
Pap smears for the early detection of cervical cancer.
Nonmalignant ovarian cysts are lined by cuboidal epithelium

and filled with a serous albuminous fluid. These abnormal
growths often can be palpated during a gynecological examination and may require surgical removal if they exceed about 4 cm
in diameter. They are generally removed as a precaution because
it is impossible to determine by palpation whether the mass is
malignant or benign.
Ovarian tumors, which occur most often in women over
the age of 60, can grow to be massive. Ovarian tumors as heavy
as 5 kg (14 lb) are not uncommon, and some weighing as much
as 110 kg (300 lb) have been reported. Some ovarian tumors
produce estrogen and thus cause feminization in elderly women,
including the resumption of menstrual periods. The prognosis for
women with ovarian tumors varies depending on the type of
tumor, whether or not it is malignant, and if it is, the stage of
the cancer.
Two frequent problems involving the uterine tubes are
salpingitis and ectopic pregnancies. Salpingitis (sal-pin-gi-tis) is
an inflammation of one or both uterine tubes. Infection of the
uterine tubes is generally caused by a sexually transmitted disease, although secondary bacterial infections from the vagina
may also cause salpingitis. Salpingitis may cause sterility if the
uterine tubes become occluded.
Ectopic pregnancy results from implantation of the blastocyst in a location other than the body or fundus of the uterus.
The most frequent ectopic site is in the uterine tube, where an
CHAPTER 21
of the Female Reproductive System
Many of the developmental problems of the female reproductive
system also occur in the reproductive system of the male and
were discussed in the previous chapter. Hermaphroditism and irregularities of the sex chromosomes, for example, are developmental conditions that cause a person to develop both male and
female characteristics.
Other developmental abnormalities of the female reproductive system may occur during the formation of the uterus and
vagina. Failure of the paramesonephric ducts to fuse normally
can result in a double uterus, a bicornuate (bi-kor'nyoo-at)
uterus, or a unicornuate uterus. These types of uterine abnor-
Pap smear: from George N. Papanicolaou, American anatomist and physician,

18831962
bicornuate: L. bi, two; cornu, horn
Problems Involving the Ovaries

and Uterine Tubes


Development

System
The McGrawHill
Companies, 2001
Chapter 21
implanted blastocyst causes what is commonly called a tubal
pregnancy. One danger of a tubal pregnancy is the enlargement,
rupture, and subsequent hemorrhage of the uterine tube where
implantation has occurred. A tubal pregnancy is frequently
treated by removing the affected tube.
Infertility, or the inability to conceive, is a clinical problem that may involve the male or female reproductive system.
On the basis of number of people who seek help for this problem,
it is estimated that 10% to 15% of couples have impaired fertility. Generally, when a male is infertile, it is because of inadequate sperm counts. Female infertility is frequently caused by an
obstruction of the uterine tubes or abnormal ovulation.
Nonmalignant
747
Malignant
Ectopic tubal
pregnancy
Tubal
carcinoma
Submucous
leiomyoma (fibroid)
Endometrial
carcinoma
Endometrial polyp
Endometrial hyperplasia
Atrophic endometrium
Cervical
carcinoma
Cervicitis
Atrophic vaginitis
Vaginal
carcinoma
Vulval
carcinoma
FIGURE 21.24 Sites of various conditions and diseases of the female reproductive tract, all of which could cause an abnormal discharge of blood.
Uterine neoplasms are an extremely common problem of

the female reproductive tract. They include cysts, polyps, and
smooth muscle tumors (leiomyomas), and most of them are benign. Any of these conditions may provoke irregular menstruations and may cause infertility if the neoplasms are massive.
Cancer of the uterus is the most common malignancy of
the female reproductive tract. The most common site of uterine
cancer is the cervix (fig. 21.24). Cervical cancer, which is second only to cancer of the breast in frequency of occurrence, is a
disease of relatively young women (ages 30 through 50), especially those who have had frequent sexual intercourse with multiple partners during their teens and onward. If detected early
through regular Pap smears, the disease can be cured before it
metastasizes. The treatment of cervical cancer depends on the
stage of the malignancy and the age and health of the woman. In
the case of women for whom fertility is not an issue, a hysterectomy (his''te-rek'to-me) (surgical removal of the uterus) is usually
performed.
neoplasm: Gk. neos, new; plasma, something formed
CHAPTER 21
Abnormal menstruations are among the most common disorders

of the female reproductive system. Abnormal menstruations
may be directly related to problems of the reproductive organs
and pituitary gland or associated with emotional and psychological stress.
Amenorrhea (a-men''o-re-a) is the absence of menstruation
and may be categorized as normal, primary, or secondary. Normal
amenorrhea follows menopause, occurs during pregnancy, and in
some women may occur during lactation. Primary amenorrhea is
the failure to have menstruated by the age when menstruation
normally begins. Primary amenorrhea is generally accompanied
by lack of development of the secondary sex characteristics. Endocrine disorders may cause primary amenorrhea and abnormal
development of the ovaries or uterus.
Secondary amenorrhea is the cessation of menstruation in
women who previously have had normal menstrual periods
and who are not pregnant and have not gone through
menopause. Various endocrine disturbances and psychological
factors may cause secondary amenorrhea. It is not uncommon,
for example, for young women who are in the process of making major changes or adjustments in their lives to miss menstrual periods. Secondary amenorrhea is also frequent in
women athletes during periods of intense training. A low percentage of body fat may be a contributing factor. Sickness, fatigue, poor nutrition, or emotional stress also may cause
secondary amenorrhea.
Dysmenorrhea is painful or difficult menstruation accompanied by severe menstrual cramps. The causes of dysmenorrhea
are not totally understood but may include endocrine disturbances (inadequate progesterone levels), a faulty position of the
uterus, emotional stress, or some type of obstruction that prohibits menstrual discharge.
Abnormal uterine bleeding includes menorrhagia (men''ora'je-a), or excessive bleeding during the menstrual period, and
metrorrhagia, or spotting between menstrual periods. Other
types of abnormal uterine bleeding are menstruations of excessive
duration, too-frequent menstruations, and postmenopausal bleeding. These abnormalities may be caused by hormonal irregularities, emotional factors, or various diseases and physical
conditions.
Cervical polyp
Cre
ek
Problems Involving the Uterus

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Endometriosis (en''do-me''tre-o'sis) is a condition characterized by the presence of endometrial tissues at sites other than
the inner lining of the uterus. Frequent sites of ectopic endometrial cells are on the ovaries, on the outer layer of the uterus, on
the abdominal wall, and on the urinary bladder. Although it is
not certain how endometrial cells become established outside the
uterus, it is speculated that some discharged endometrial tissue
might be flushed backward from the uterus and through the uterine tubes during menstruation. Women with endometriosis will
bleed internally with each menstrual period because the ectopic
endometrial cells are stimulated along with the normal endometrium by ovarian hormones. The most common symptoms
of endometriosis are extreme dysmenorrhea and a feeling of fullness during each menstrual period. Endometriosis can cause infertility. It is treated by suppressing the endometrial tissues with
oral contraceptive pills or by surgery. An oophorectomy, or removal of the ovaries, may be necessary in extreme cases.
Uterine displacements are relatively common in elderly
women. When uterine displacements occur in younger women,
they are important because they may cause dysmenorrhea, infertility, or problems with childbirth. Retroversion, or retroflexion,
is a displacement backward; anteversion, or anteflexion, is displacement forward. Prolapse of the uterus is a marked downward displacement of the uterus into the vagina.
An abortion is defined as the termination of a pregnancy
before the twenty-eighth week of gestation. A spontaneous
abortion, or miscarriage, occurs without mechanical aid or medicinal intervention and may occur in as many as 10% of all pregnancies. Spontaneous abortions usually occur when there is
abnormal development of the fetus or disease of the maternal reproductive system. An induced abortion is removal of the fetus
from the uterus by mechanical means or drugs. Induced abortions
are a major issue of controversy because of questions regarding
individual rights (those of the mother and those of the fetus), the
definition of life, and morality.
Diseases of the Vagina and Vulva
CHAPTER 21
The McGrawHill
Companies, 2001
Pelvic inflammatory disease (PID) is a general term for inflammation of the female reproductive organs within the pelvis. The infection may be confined to a single organ, or it may involve all of the
internal reproductive organs. The pathogens generally enter
through the vagina during coitus, induced abortion, or childbirth.
Inflammation of the ovaries is called oophoritis (o''of-o-ri'tis) and
inflammation of the uterine tube is salpingitis (sal''pin-ji'tis).
The vagina and vulva are generally resistant to infection because of the acidity of the vaginal secretions. Occasionally, however, localized infections and inflammations do occur. These are
termed vaginitis, if confined to the vagina, or vulvovaginitis, if
both the vagina and external genitalia are affected. The symptoms
of vaginitis are a discharge of pus (leukorrhea) and itching (pruritus). The two most common organisms that cause vaginitis are the
protozoan Trichomonas vaginalis and the fungus Candida albicans.
Diseases of the Breasts

and Mammary Glands
The breasts and mammary glands of females are highly susceptible to infections, cysts, and tumors. Infections involving the
mammary glands usually follow the development of a dry and
cracked nipple during lactation. Bacteria enter the wound and
establish an infection within the lobules of the gland. During an
infection of the mammary gland, a blocked duct frequently
causes a lobe to become engorged with milk. This localized
swelling is usually accompanied by redness, pain, and an elevation of temperature. Administering specific antibiotics and applying heat are the usual treatments.
Nonmalignant cysts are the most frequent diseases of
the breast. These masses are generally of two types, neither of
which is life-threatening. Dysplasia (fibrocystic disease) is a
broad condition involving several nonmalignant diseases of
the breast. All dysplasias are benign neoplasms of various sizes
that may become painful during or prior to menstruation.
Most of the masses are small and remain undetected. Dysplasia affects nearly 50% of women over the age of 30 prior to
menopause.
A fibroadenoma (fi''bro-ad''e-no'ma) is a benign tumor of
the breast that frequently occurs in women under the age of 35.
Fibroadenomas are nontender rubbery masses that are easily
moved about in the mammary tissue. A fibroadenoma can be excised in a physicians office under local anesthetic.
Carcinoma of the breast is the most common malignancy in women. One in nine women will develop breast cancer and one-third of these will die from the disease. Breast
cancer is the leading cause of death in women between 40 and
50 years of age. Men are also susceptible to breast cancer, but
it is 100 times more frequent in women. Breast cancer in men
is usually fatal.
The causes of breast cancer are not known, but women
who are most susceptible are those who are over age 35, who
have a family history of breast cancer, and who are nulliparous
(never having given birth). The early detection of breast cancer is important because the prognosis worsens as the disease
progresses.
Confirming suspected breast cancer generally requires mammography (fig. 21.25). If the mammogram suggests breast cancer, a
biopsy (tissue sample) is obtained and the tumor assessed. If the
tumor is found to be malignant, surgery is performed, the extent
of which depends on the size of the tumor and whether metastasis
has occurred. The surgical treatment for breast cancer is generally
some degree of mastectomy (mas-tekto-me). A simple mastectomy is
removal of the entire breast but not the underlying lymph nodes.
A modified radical mastectomy is the complete removal of the
breast, the lymphatic drainage, and perhaps the pectoralis major
muscle. A radical mastectomy is similar to a modified except that
the pectoralis major muscle is always removed, as well as the pectoral lymph nodes and adjacent connective tissue.


Development

System
The McGrawHill
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Chapter 21
(a)
749
(b)
FIGURE 21.25 (a) In mammography, the breast is placed alternately on a metal plate and radiographed from above and from the side. (b) A
mammogram of a patient with carcinoma of the breast. (Note the presence of a neoplasm indicated with an arrow.)
Methods of Contraception
CHAPTER 21
The rhythm method of birth control continues to be used by many

people, but the popularity of this technique has declined as more
successful methods of contraception have been introduced
(fig. 21.26). In the rhythm method, an attempt is made to predict the day of the womans ovulation and restrict coitus to safe
times of the cycle that allow no chance for fertilization to occur.
The day of ovulation can be determined by a rise in basal body
temperature or by a change in mucous discharge from the vagina.
This technique has one of the highest failure rates of any of the
widely used birth control methods, largely because womens cycles are often irregular.
More popular methods of birth control include sterilization,
oral contraceptives, intrauterine devices (IUDs), and barrier
methodsincluding condoms for the male and female and diaphragms for the female. All of these techniques are effective, but
they vary with respect to safety, side effects, and degree of efficacy.
Sterilization techniques include vasectomy for the male and
tubal ligation for the female. In the latter technique (which accounts for over 60% of sterilization procedures performed in the
United States), the uterine tubes are cut and tied. This is analogous to the procedure performed on the ductus deferentia in a
vasectomy. It prevents fertilization of the ovulated ovum.
About 10 million women in the United States and 60 million women in the world are currently using oral steroid contraceptives (the Pill). These contraceptives usually consist of a
synthetic estrogen combined with a synthetic progesterone in
the form of pills that are taken once each day for 3 weeks after
the last day of a menstrual period. This procedure causes an immediate increase in blood levels of ovarian steroids (from the
pill), which is maintained for the normal duration of a monthly
cycle. As a result of negative feedback inhibition of gonadotrophin
secretion, ovulation never occurs. The entire cycle is like a false
luteal phase, with high levels of progesterone and estrogen and
low levels of gonadotrophins.
Because the contraceptive pills contain ovarian steroid
hormones, the endometrium proliferates and becomes secretory,
just as it does during a normal cycle. In order to prevent an abnormal growth of the endometrium, women stop taking the pill
after 3 weeks. This causes estrogen and progesterone levels to
fall, and permits menstruation to occur. The contraceptive pill is
an extremely effective method of birth control, but it does have
potentially serious side effectsincluding an increased incidence
of thromboembolism and cardiovascular disorders. It has been
pointed out, however, that the mortality risk associated with
contraceptive pills is still much lower than the risk of death from
complications of pregnancyor from automobile accidents.

CHAPTER 21
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Development

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Companies, 2001
(a)
(b)
(d)
(e)
(f)
(g)
(c)
(h)
FIGURE 21.26 Various types of birth control devices. (a) IUD, (b) contraceptive sponge, (c) diaphragm, (d ) birth control pills, (e) vaginal
spermicides, (f ) condom, (g) female condom, and (h) Norplant.


Development

System
The McGrawHill
Companies, 2001
Chapter 21
Another way in which to deliver hormonal contraceptives
to a womans body is by means of a subdermal implant. Implants
are 2-in. rods filled with a hormonal contraceptive drug. They
are implanted just under the skin, usually on the upper arm,
through a tiny incision. The contraceptive hormone gradually
leaches out through the walls of the rods and enters the bloodstream, preventing pregnancy for at least 5 years.
Intrauterine devices (IUDs) do not prevent ovulation,
but instead prevent implantation of the blastocyst in the uterine
wall in the event that fertilization occurs. The mechanisms by
which their contraceptive effects are produced are not well understood but appear to involve their ability to cause inflammatory reactions in the uterus. Uterine perforations are the
foremost complication associated with the use of IUDs. Because
of the potential problems with IUDs, their use has diminished.
Barrier contraceptivescondoms, diaphragms, and cervical capsare only slightly less effective than hormonal contraceptives or IUDs, but they do not have serious side effects.
Barrier contraceptives are most effective when they are used in
conjunction with spermicidal (sperm-killing) foams and gels.
Many couples avoid them, however, because they detract from
the spontaneity of sexual intercourse. Latex condoms offer an additional benefit; they provide some protection against sexually
transmitted diseases, including AIDS.
Several new chemical contraceptives are being developed
that should be soon available. These include hormonal patches,
hormonal IUDs, and injectible contraceptives. A hormonal patch
is intended to be placed on the skin directly over an ovary. De-
751
signed to resist moisture, the patch releases a weeks worth of estrogen and progestin. A hormonal IUD is a T-shaped structure
that fits into the uterine cavity where it slowly releases progestin
up to a five-year period. Improved monthly injectible contraceptives have fewer side effects than Depro-Provera, which has been
available since 1992.

An ectopic pregnancy is any pregnancy that implants outside the uterine
cavity. This is most likely to occur in the uterine tube (see fig. 21.24).
The events leading up to our patients problem are as follows: An oocyte
is extruded from the ovary during ovulation and received into the uterine tube. Soon after it enters the tube, the oocyte is fertilized, creating a
zygote. Up to this point, the events are no different from those that
occur in a normal pregnancy. In the case of tubal pregnancy, however,
the transporting ability of the uterine tube fails, causing the conceptus
to be retained in the tube. Implantation then occurs within tissues that
are not well suited for that purpose. For example, the uterine tube does
not expand well to accommodate a growing embryo, nor does it possess
the necessary epithelium and glandular structures as does the endometrium of the uterus. The result is overexpansion and erosion of the
tubal wall, leading to rupture and hemorrhage. Because the uterine tube
is basically exposed to the peritoneal cavity, blood from the site of rupture can flow into and collect in the rectouterine pouch (pouch of Douglas or Douglas cul-de-sac), which lies posterior to the vaginal fornix.
There, it is easily aspirated by a needle placed through the vaginal wall.
QUESTIONS
1. These adnexal masses are consistent
with endometriomas, foci of
endometrial tissue outside the uterus on
the ovaries. How would these masses

cause the patients symptoms?
2. How does endometrial tissue get to the
ovaries?
3. Endometrial implants can be found
throughout the peritoneal space from
direct spread and have even been
reported in the pleural space.
Remembering that these implants
cause irritation of, as well as, scarring
around the organs on which they
are implanted, what symptoms would
the patient have if implants were found
on the following organs: (a) rectum,
(b) urinary bladder, (c) stomach, or
(d) diaphragm?
A
U
CHAPTER 21
A 32-year-old female comes to you because

she and her husband have been unsuccessfully attempting to become pregnant for
more than a year. In questioning her, you
find that she has been suffering from vague
pelvic pain during her periods for many
years. She also reports occasional pain during intercourse over that same time. On
pelvic exam, you find bilateral tender adnexal masses, flanking a normal-sized uterus.
An MRI of the pelvis confirms these findings. [uterus (U), adnexal masses (A)]

752
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System
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Companies, 2001
Chapter Summary
Introduction to the Female Reproductive
System (pp. 726728)
1. The reproductive period of a female is the
period between puberty (about age 12)
and menopause (about age 50). In the
course of this age span, cyclic ovulation
and menstruation patterns occur in
nonpregnant females.
2. The functions of the female reproductive
system are to produce ova; secrete sex
hormones; receive sperm from the male;
provide sites for fertilization,
implantation, and development of the
embryo and fetus; facilitate parturition;
and secrete milk from the mammary
glands.
3. The female reproductive system consists
of (a) primary sex organsthe ovaries;
(b) secondary sex organsthose that are
essential for sexual reproduction,
characterized by latent development; and
(c) secondary sex characteristics
features that are sexual attractants,
expressed after puberty.
Structure and Function of the Ovaries

(pp. 728732)
CHAPTER 21
1. The ovaries are supported by the

mesovarium, which extends from the
broad ligament, and by the ovarian and
suspensory ligaments.
2. The ovarian follicles within the ovarian
cortex undergo cyclic changes.
(a) Primary oocytes, arrested at prophase
I of meiosis, are contained within
primordial follicles.
(b) Upon stimulation by gonadotropic
hormones, some of the primordial
follicles enlarge to become primary
follicles.
(c) When a follicle develops a fluid-filled
antrum, it is called a secondary
follicle.
(d) Generally only one follicle continues
to grow to become a vesicular ovarian
follicle.
(e) The vesicular ovarian follicle

contains a secondary oocyte, arrested
at metaphase II of meiosis.
(f) In the process of ovulation, the
vesicular ovarian follicle ruptures and
releases its secondary oocyte, which
becomes a zygote upon fertilization.
(g) After ovulation, the empty follicle
becomes a corpus luteum.
Secondary Sex Organs (pp. 732738)

1. The uterine tube, which conveys ova
from the ovary to the uterus, provides a
site for fertilization.
(a) The open-ended portion of each
uterine tube is expanded; its margin
bears fimbriae that extend over the
lateral surface of the ovary.
(b) Movement of an ovum is aided by
ciliated cells that line the lumen and
by peristaltic contractions in the wall
of the uterine tube.
2. The uterus is supported by the broad
ligaments, uterosacral ligaments, cardinal
ligaments, and round ligaments. The
regions of uterus are the fundus, body, and
cervix. The cervical canal opens into the
vagina at the uterine ostium.
(a) The endometrium consists of a
stratum basale and a stratum
functionale; the superficial stratum
functionale is shed during
menstruation.
(b) The myometrium produces the
muscular contractions needed for
labor and parturition.
3. The vagina serves to receive the erect
penis during coitus, to convey menses to
the outside during menstruation, and to
transport the fetus during parturition.
(a) The vaginal wall is composed of an
inner mucosa, a middle muscularis,
and an outer fibrous layer.
(b) The vaginal orifice may be partially
covered by a thin membranous
hymen.
4. The external genitalia, or vulva, include

the mons pubis, labia majora and minora,
clitoris, vaginal vestibule, vestibular
bulbs, and vestibular glands.
5. Impulses through parasympathetic nerves
stimulate erectile tissues in the clitoris
and vestibular bulbs; in orgasm, muscular
contraction occurs in the perineum,
uterus, and uterine tubes.
Mammary Glands (pp. 738740)

1. Mammary glands, located within the
breasts, are modified sweat glands.
(a) Each mammary gland is composed of
15 to 20 lobes; the lobes are
subdivided into lobules that contain
mammary alveoli.
(b) During lactation, the mammary
alveoli secrete milk. The milk passes
through mammary ducts, lactiferous
ducts, and lactiferous sinuses and is
discharged through the nipple.
2. The nipple is a cylindrical projection near
the center of the breast, surrounded by
the circular pigmented areola.
Ovulation and Menstruation

(pp. 740742)
1. Ovulation and menstruation are
reproductive cyclic events that are
regulated by hormones secreted by the
hypothalamus, the anterior pituitary, and
the ovaries.
2. The menstrual cycle is divided into
menstrual, proliferative, and secretory
phases.
3. The principal hormones that regulate
ovulation and menstruation are estrogen,
progesterone, follicle-stimulating
hormone (FSH), and luteinizing
hormone (LH).
Review Activities
Objective Questions
1. The cervix is a portion of
(a) the vulva.
(b) the vagina.
(c) the uterus.
2. Fertilization normally occurs in

(a) the ovary.
(b) the uterine tube.
(c) the uterus.
(d) the vagina.
3. The secretory phase of the endometrium

corresponds to which of the following
ovarian phases?
(a) the follicular phase
(b) ovulation


Development

System
The McGrawHill
Companies, 2001
Chapter 21
4.
5.
6.
7.
8.
9.
(c) the luteal phase

(d) the menstrual phase
Which of the following statements about
oogenesis is true?
(a) Oogonia, like spermatogonia, form
continuously during postnatal life.
(b) Primary oocytes are haploid.
(c) Meiosis is completed prior to
ovulation.
(d) During ovulation, a secondary oocyte
is released from a vesicular ovarian
follicle.
The function of the mesovarium is
(a) movement of the sperm to the ova.
(b) nourishment of the ovarian walls.
(c) muscular contraction of the uterus.
(d) suspension of the ovary.
Which of the following layers is shed as
menses?
(a) the perimetrial layer
(b) the fibrous layer
(c) the functionalis layer
(d) the menstrual layer
The transverse folds in the mucosal layer
of the vagina are called
(a) perineal folds. (c) fornices.
(b) vaginal rugae. (d) labia gyri.
Which of the following is not a part of the
vulva?
(a) the mons pubis
(b) the clitoris
(c) the vaginal vestibule
(d) the vagina
(e) the labia minora
The paramesonephric (mllerian) ducts
give rise to
(a) the uterine tubes.
(b) the uterus.
(c) the pudendum.
(d) both a and b.
(e) both b and c.
10. In a female, the homologue of the male

scrotum is/are
(a) the labia majora.
(b) the labia minora.
(c) the clitoris.
(d) the vestibule.
Essay Questions
1. Define puberty, ovulation, menstruation,
and menopause.
2. Describe the follicular changes within the
ovarian cortex during the events that
precede and follow ovulation.
3. Define oogenesis. When is the process
initiated and when is it completed?
4. Describe the gross and histologic structure
of the uterus and comment on the
significance of the two endometrial layers.
5. Identify the secondary sex organs and
explain their functions.
6. Summarize the events of a menstrual
cycle and explain the roles of estrogen
and progesterone.
7. List the functions of the vagina and
describe its structure.
8. Distinguish between the labia majora and
labia minora, between the pudendal cleft
and vaginal vestibule, and between the
vestibular bulbs and vestibular glands.
9. List the events that cause an erection of
the female genitalia, and explain the
process of orgasm.
10. Describe the structure and position of
the mammary glands in the breasts.
11. List the homologous reproductive organs
of the male and female reproductive
systems and note the undifferentiated
structures from which they develop.
12. Define gynecology, obstetrics, speculum,
laparoscopy, breast self-examination, and
Pap smear.
753
13. Distinguish between normal, primary, and

secondary amenorrhea. What causes
each?
14. List the various kinds of uterine
neoplasms. How is cancer of the uterus
generally detected?
15. Distinguish between dysplasia,
fibroadenoma, and carcinoma of the
breast.
1. Why is the hormonal regulation of the
reproductive system much more complex
in females than it is in males?
2. Your 14-year-old daughter is serious about
her gymnastics lessons and exercises
rigorously for at least 2 hours every day.
She has not yet had her first menstrual
period and is beginning to get worried.
How would you reassure her?
3. A wide variety of commercial douche
preparations and scented aerosols can be
purchased in drug stores and supermarkets
as feminine hygiene products. Why
should women avoid using them?
4. Both sterility and impotence are sexual
dysfunctions in men. Only one of these is
associated with women, however.
Explain.
5. Contraceptive pills trick the brain into
thinking youre pregnant. Explain what
is meant by this statement.
6. With respect to homologous structures,
explain why men have nipples? Why do
mammary glands normally function only in
females?
7. If a woman is on estrogen replacement
therapy following menopause, why is it
important to monitor her uterine lining?

CHAPTER 21


Development
22
22. Developmental
Anatomy, Postnatal
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Developmental Anatomy,
Postnatal Growth,
and Inheritance
Fertilization 755
Preembryonic Period 757
Embryonic Period 762
Fetal Period 772
Labor and Parturition 775
Periods of Postnatal Growth 775
Inheritance 782

Genetic Disorders
of Clinical Importance 790
Chapter Summary 791
Clinical Case Study

A 27-year-old woman gave birth to twin boys, followed by an apparently single placenta. After
examining the two infants, the pediatrician informed the mother that one of them had a cleft
palate but that the other was normal. She added that cleft palate could be hereditary and asked
if any family members had the problem. The mother said that she knew of none. Further examination of the placenta revealed two amnions and only one chorion.
Does the presence of two amnions and one chorion indicate monozygotic or dizygotic
twins? How would you account for the fact that one baby has a cleft palate, whereas the other
does not?
Hints: Read the section on multiple pregnancy at the end of the chapter and carefully study
figure 22.37. Can any conclusions be drawn regarding genetic similarities between the two infants? Note that identical twins are always the same gender.
FIGURE: A cleft lip and an accompanying

cleft palate are congenital disorders in which
one or two sides of the upper lip and hard
palate fail to fuse. Cleft lips and cleft palates
can be treated very effectively with surgery.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
FERTILIZATION
Upon fertilization of an ovum by a spermatozoon in the uterine tube,
meiotic development is completed and a diploid zygote is formed.
Objective 1
Define fertilization, capacitation, and

morphogenesis.
Objective 2
Describe the changes that occur in the

spermatozoon and ovum prior to, during, and immediately
following fertilization.
Fertilization refers to the penetration of an ovum (egg) by a spermatozoon (fig. 22.1), with the subsequent union of their genetic
material. It is this event that determines a persons gender (see
p. 716) and biological inheritance. Fertilization cannot occur,
however, unless certain conditions are met. First of all, an ovum
must be present in the uterine tubeit can be there for at most
24 hours before it becomes incapable of undergoing fertilization.
Second, large numbers of spermatozoa must be ejaculated to ensure fertilization. Although a recent study has shown that sperm
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cells can remain viable 5 days after ejaculation, this still leaves a
window of fertility of only 6 days each monththe day of ovulation and the 5 days leading up to it.
As described in chapter 21, a woman usually ovulates one
ovum a month, totaling about 400 during her reproductive years.
Each ovulation releases an ovum that is actually a secondary
oocyte arrested at metaphase of the second meiotic division. An
ovum is surrounded by a thin layer of protein and polysaccharides, called the zona pellucida, and a layer of granulosa cells,
called the corona radiata (fig. 22.1). These layers provide a protective shield around the ovum as it enters the uterine tube.
During coitus, a male ejaculates between 100 million and
500 million spermatozoa into the females vagina. This tremendous number is needed because of the high fatality rateonly
about 100 sperm cells survive to encounter the ovum in the uterine tube. In addition to deformed sperm cellsup to 20% in the
zona pellucida: L. zone, a girdle; pellis, skin

corona radiata: Gk. korone, crown; radiata, radiate
Corona radiata
First polar body
Zona pellucida
Second meiotic spindle
Oocyte membrane
Cytoplasm
(a)
(c)
Nucleus
containing
chromosome
Acrosome
containing
enzymes
Perforations
in acrosome wall
CHAPTER 22
1
3
(b)
Creek
FIGURE 22.1 The process of fertilization. (a,b) As the head of the sperm encounters the corona radiata of the oocyte (2), digestive enzymes
are released from the acrosome (3, 4), clearing a path to the oocyte membrane. When membrane of the sperm contacts the oocyte membrane
(5), the membranes become continuous, and the nucleus and other contents of the sperm move into the egg cytoplasm of the oocyte. (c) A
scanning electron micrograph of a sperm cell bound to the surface of an oocyte.

756
Unit 7

Development
22. Developmental
Anatomy, Postnatal
Cell
membrane
Acrosomal
cap
Nucleus
Basal
plate
FIGURE 22.2 A transmission electron micrograph showing the

head of a human spermatozoon with its nucleus and acrosome.
average fertile maleanother 25% will perish as soon as they contact

the vaginas acidic environment. Still others are destroyed by the
womans immune cells, which recognize them as foreign cells. Even if
they manage to reach the uterine ostium (see fig. 21.12), many sperm
cells will get stuck there and never make it through the uterus.
If it finally encounters an ovum, the spermatozoon must
penetrate the protective corona radiata and zona pellucida for
fertilization to occur. To do this, the head of each sperm cell is
capped by an organelle called an acrosome (ak'ro-som) (figs. 22.1
and 22.2). The acrosome contains a trypsinlike proteindigesting enzyme and hyaluronidase (hi''a-loo-ron'-das), which
digests hyaluronic acid, an important constituent of connective
tissue. When a spermatozoon meets an ovum in the uterine
tube, an acrosomal reaction allows the spermatozoon to penetrate the corona radiata and the zona pellucida. A spermatozoon that comes along relatively lateafter many others
have undergone acrosomal reactions to expose the ovum
membraneis more likely to be the one that finally achieves
penetration of the egg.
CHAPTER 22
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Experiments confirm that freshly ejaculated spermatozoa are

infertile and must be in the female reproductive tract for at
least 7 hours before they can fertilize a secondary oocyte. Their
membranes must become fragile enough to permit the release of the
acrosomal enzymesa process called capacitation. During in vitro
fertilization, capacitation is induced artificially by treating the ejaculate with a solution of gamma globulin, free serum, follicular fluid,
dextran, serum dialysate, and adrenal gland extract to chemically
mimic the conditions of the female reproductive tract.
As soon as a spermatozoon penetrates the zona pellucida, a rapid chemical change in this layer prevents other
spermatozoa from attaching to it. Therefore, only one spermatozoon is permitted to fertilize an oocyte. With the penetration of a single spermatozoon through its cell membrane, the
oocyte is stimulated to complete its second meiotic division
(fig. 22.3). Like the first meiotic division, the second produces one cell that contains most of the cytoplasm and one
polar body. The cell containing the cytoplasm is the mature
ovum, and the second polar body, like the first, ultimately
fragments and disintegrates.
At fertilization, the head of the sperm cell enters the cytoplasm of the much larger ovum. Within 12 hours, the nuclear
membranes in both the sperm cell and ovum disappear, and the
haploid number of chromosomes (23) in the ovum is joined by
the haploid number of chromosomes from the spermatozoon. A
fertilized egg, or zygote (zi'got), containing the diploid number of
chromosomes (46) is thus formed.
Within hours after conception, the structure of the body
begins to form from this single fertilized egg, culminating some
38 weeks later with the birth of a baby. The transformation involved in the growth and differentiation of cells and tissues is
known as morphogenesis (mor''fo-jen'-sis), and it is through
this awesome process that the organs and systems of the body
are established in a functional relationship. Moreover, there are
sensitive periods of morphogenesis for each organ and system,
during which genetic or environmental factors may affect normal development.
Prenatal development can be divided into a preembryonic
period, which is initiated by the fertilization of an ovum; an embryonic period, during which the bodys organ systems are
formed; and a fetal period, which culminates in parturition
(childbirth).
Knowledge Check
1. Explain why capacitation and the acrosomal reaction of
spermatozoa are necessary to accomplish fertilization of a
secondary oocyte.
2. Discuss the changes that occur in a spermatozoon from the
time of ejaculation to the time of fertilization. What
changes occur in a secondary oocyte following ovulation to
the time of fertilization?
3. Define morphogenesis. When does this process begin? What
does it accomplish?
haploid: Gk. haplous, single; L. ploideus, multiple in form

acrosome: Gk. akron, extremity; soma, body
capacitation: L. capacitas, capable of
zygote: Gk. zygotos, yolked, joined

diploid: Gk. diplous, double; L. ploideus, multiple in form
morphogenesis: Gk. morphe, form; genesis, beginning


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
Primary oocyte
Secondary oocyte
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Secondary oocyte
at metaphase II
First
polar
body
First
polar
body
First
meiotic
division
Spindle
apparatus
Ovulation
First polar
body
757
Second
polar body
Fertilization
Nucleus
of ovum
Sperm cell
inside ovum
Chromosomes
Zygote
Nuclear membrane
disappearing
Sperm cell
nucleus
FIGURE 22.3 A secondary oocyte, arrested at metaphase II of meiosis, is released at ovulation. If this cell is fertilized, it will become a mature
ovum, complete its second meiotic division, and produce a second polar body. The fertilized ovum is known as a zygote.
PREEMBRYONIC PERIOD
The events of the 2-week preembryonic period include fertilization, transportation of the zygote through the uterine tube, mitotic divisions, implantation, and the formation of primordial
embryonic tissue.
Objective 3
Describe the events of preembryonic

development that result in the formation of the blastocyst.
Objective 4
Discuss the role of the trophoblast in the

implantation and development of the placenta.
Objective 5
Explain how the primary germ layers develop

and list the structures produced by each layer.
cells. The morula floats freely in the uterine cavity for about
3 days. During this time, the center of the morula fills with fluid
passing in from the uterine cavity. As the fluid-filled space develops within the morula, two distinct groups of cells form, and the
structure becomes known as a blastocyst (blas'to-sist) (fig. 22.4).
The hollow, fluid-filled center of the blastocyst is called the blastocyst cavity. The blastocyst is composed of an outer layer of
cells, known as the trophoblast, and an inner aggregation of cells,
called the embryoblast (internal cell mass) (see fig. 22.6). With
further development, the trophoblast differentiates into a structure called the chorion (kor'e-on), which will later become a portion of the placenta. The embryoblast will become the embryo. A
diagrammatic summary of the ovarian cycle, fertilization, and the
morphogenic events of the first week is presented in figure 22.5.
Objective 6
Define gestation and explain how the

parturition date is determined.
Cleavage and Formation of the Blastocyst
morula: L. morus, mulberry
The process of implantation begins between the fifth and seventh day following fertilization. This is the process by which the
blastocyst embeds itself into the endometrium of the uterine wall
(fig. 22.6a). Implantation is made possible by the secretion of
proteolytic enzymes by the trophoblast, which digest a portion of
the endometrium. The blastula sinks into the depression, and endometrial cells move back to cover the defect in the wall. At the
same time, the part of the uterine wall below the implanting
blastocyst: Gk. blastos, germ; kystis, bladder

chorion: Gk. chorion, membrane
implantation: L. im, in; planto, to plant
CHAPTER 22
Within 30 hours following fertilization, the zygote undergoes a

mitotic division called cleavage. This first division results in the
formation of two identical daughter cells called blastomeres
(fig. 22.4). Additional cleavages occur as the structure passes
down the uterine tube and enters the uterus on about the third
day. It is now composed of a ball of 16 or more cells called a
morula (mor'yu-la). Although the morula has undergone several
mitotic divisions, it is not much larger than the zygote because no
additional nutrients necessary for growth have been entering the
Implantation

758
Unit 7

Development
22. Developmental
Anatomy, Postnatal
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Polar body
Zona pellucida
Blastomere
2-cell stage
4-cell stage
8-cell stage
Embryoblast
Degenerating
zona pellucida
Blastocyst cavity
Trophoblast
Morula
Early blastocyst
Late blastocyst
FIGURE 22.4 Sequential illustrations from the first cleavage of the zygote to the formation of the blastocyst. (Note the deterioration of the zona
pellucida in the early blastocyst.)
Cleavage
(30 hours)
Zygote
Egg nucleus
Morula
(72 hours)
8-cell
stage
4-cell
stage
2-cell
stage
Sperm nucleus
Sperm
cells
Ovary
CHAPTER 22
Fertilization
Blastocyst
(4 days)
Corpus
luteum
Maturing
follicle
Ovulation
Secondary
oocyte
Implanted
blastocyst
(6 days)
FIGURE 22.5 A diagram of the ovarian cycle, fertilization, and the events of the first week. Implantation of the blastocyst begins between the
fifth and seventh day, and is generally completed by the tenth day.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
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Blastocoel
Uterine cavity
Trophoblast
Embryonic endoderm
Embryoblast
Cytotrophoblasts
Blastocyst
cavity
Embryoblast
Endometrium
Trophoblast
Endometrial
epithelium
Endometrial
capillary
Endometrial
gland
Glandular
secretion
Endometrial
storma
(a)
Creek
(b)
FIGURE 22.6 The blastocyst adheres to the endometrium on about the sixth day as seen in (a), a photomicrograph. By the seventh day (b),
specialized syncytiotrophoblasts from the trophoblast have begun to invade the endometrium. The syncytiotrophoblasts secrete human chorionic
gonadotropin (hCG) to sustain pregnancy and will eventually participate in the formation of the placenta for embryonic and fetal sustenance.
All pregnancy tests assay for the presence of hCG in the

blood or urine because this hormone is secreted only by the
blastocyst. Because there is no other source of hCG, the presence of
this hormone confirms a pregnancy. Modern pregnancy tests detect
the presence of hCG by use of antibodies against hCG or by the use
of cellular receptor proteins for hCG.
Human chorionic
gonadotropin
Estrogen
Progesterone
FIGURE 22.7 Human chorionic gonadotropin (hCG) is secreted

by syncytiotrophoblasts during the first trimester of pregnancy. This
pituitary-like hormone maintains the mothers corpus luteum for the
first 512 weeks of pregnancy. The placenta then assumes the role of
estrogen and progesterone production, and the corpus luteum degenerates.
CHAPTER 22
blastocyst thickens, and specialized cells of the trophoblast

produce fingerlike projections, called syncytiotrophoblasts
(sin-sit''e-o-trof'o-blasts), into the thickened area. The syncytiotrophoblasts arise from a specific portion of the trophoblast called the cytotrophoblast (fig. 22.6b), located next to
the embryoblast.
The blastocyst saves itself from being aborted by secreting
a hormone that indirectly prevents menstruation. Even before
the sixth day when implantation begins, the syncytiotrophoblasts
secrete human chorionic gonadotropin (kor''e-on-ik go-nad-otro'pin) (hCG). This hormone is identical to LH in its effects,
and therefore is able to maintain the corpus luteum past the time
when it would otherwise regress. The secretion of estrogen and
progesterone is maintained, and menstruation is normally prevented (fig. 22.7).
The secretion of hCG declines by the tenth week of pregnancy. Actually, this hormone is required only for the first 5 to
6 weeks of pregnancy because the placenta itself becomes an active steroid-secreting gland by this time. At the fifth to sixth
week, the mothers corpus luteum begins to regress (even in the
presence of hCG), but the placenta secretes more than sufficient
amounts of steroids to maintain the endometrium and prevent
menstruation.

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Unit 7

Development
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Anatomy, Postnatal
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Chorion
Ectoderm
Germ
layers
Amnion
Mesoderm
Amniotic cavity
Endoderm
Body stalk
(umbilical cord)
Yolk sac
of embryo
Chorionic villi
FIGURE 22.8 The completion of implantation occurs as the primary germ layers develop at the end of the second week.
TABLE 22.1 Summary of Preembryonic Development

Morphogenic Stage
Time Period
Principal Events
Zygote
24 to 30 hours following
ovulation
Egg is fertilized; zygote has 23 pairs of chromosomes (diploid) from haploid sperm and haploid
egg and is genetically unique
Cleavage
30 hours to third day
Mitotic divisions produce increased number of cells
Morula
Third to fourth day
Solid ball-like structure forms, composed of 16 or more cells
Blastocyst
Fifth day to end of second week
Hollow ball-like structure forms, a single layer thick; embryoblast and trophoblast form;
implantation occurs; embryonic disc forms, followed by primary germ layers
CHAPTER 22
Formation of the Germ Layers

week of development, the embryoblast undergoes marked differentiation. A slitlike space called the amniotic (am''ne-ot'ik)
cavity forms between the embryoblast and the trophoblast
(fig. 22.8). The embryoblast flattens into the embryonic disc (see
fig. 22.10), which consists of two layers: an upper ectoderm, which
is closer to the amniotic cavity, and a lower endoderm, which borders the blastocyst cavity. A short time later, a third layer called
the mesoderm forms between the endoderm and ectoderm. These
three layers constitute the primary germ layers (fig. 22.8). Once
they are formed, at the end of the second week, the preembryonic
period is completed and the embryonic period begins.
The primary germ layers are especially significant because all of

the cells and tissues of the body are derived from them. Ectodermal
cells form the nervous system; the outer layer of skin (epidermis), including hair, nails, and skin glands; and portions of the sensory organs. Mesodermal cells form the skeleton, muscles, blood,
reproductive organs, dermis of the skin, and connective tissue. Endodermal cells produce the lining of the GI tract, the digestive organs,
the respiratory tract and lungs, and the urinary bladder and urethra.
The events of the preembryonic period are summarized in
table 22.1. Refer to figure 22.9 for an illustration and a listing of the organs and body systems that derive from each of the primary germ layers.
The period of prenatal development is referred to as gestation.
Normal gestation for humans is 9 months. Knowing this and the
pattern of menstruation make it possible to determine the babys delivery date. In a typical reproductive cycle, a woman ovulates 14 days
prior to the onset of the next menstruation and is fertile for approximately 20 to 24 hours following ovulation. Adding 9 months, or
38 weeks, to the time of ovulation gives one the estimated delivery date.

gestation: L. gestatus, to bear


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
Amnion
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Digestive tract
Amniotic fluid
Skin
Heart
Spinal cord
Chorion
Body stalk with

umbilical vessels
Brain
Nels
on
Yolk sac
Endoderm
Mesoderm
Ectoderm
FIGURE 22.9 The body systems and the primary germ layers from which they develop.
CHAPTER 22
Epidermis of skin and epidermal

derivatives: hair, nails, glands
of the skin; linings of oral, nasal,
anal, and vaginal cavities
Pituitary gland
Adrenal medulla
Muscle: smooth, cardiac,

and skeletal
Connective tissue: embryonic,
connective tissue proper,
cartilage, bone, blood
Epithelium of blood vessels,
lymphatic vessels, body
cavities, joint cavities
Kidneys and ureters
Adrenal cortex
Epithelium of pharynx, external
acoustic canal, tonsils, thyroid,
parathyroid, thymus, larynx,
trachea, lungs, GI tract, urinary
bladder and urethra, and vagina
Liver and pancreas

762
Unit 7

Development
22. Developmental
Anatomy, Postnatal
Knowledge Check
4. List the structural characteristics of a zygote, morula, and
blastocyst. Approximately when do each of these stages of
the preembryonic period of development occur?
5. Discuss the process of implantation and describe the physiological events that ensure pregnancy.
6. Describe the development of the placenta.
7. List the major structures that derive from each germ layer.
8. Define gestation. What is the average gestation time for humans? How is the parturition date determined?
EMBRYONIC PERIOD
The events of the 6-week embryonic period include the differentiation of the germ layers into specific body organs and the formation
of the placenta, the umbilical cord, and the extraembryonic membranes. Through these morphogenic events, the needs of the embryo are met.
Objective 7
Define embryo and describe the major events

of the embryonic period of development.
Objective 8
List the embryonic needs that must be met to

avoid a spontaneous abortion.
Objective 9
Describe the structure and function of each of

the extraembryonic membranes.
Objective 10
Describe the development and functions of

the placenta and the umbilical cord.
During the embryonic periodfrom the beginning of the third

week to the end of the eighth weekthe developing organism is
correctly called an embryo. It is at this period that all of the
body tissues and organs form, as well as the placenta, umbilical
cord, and extraembryonic membranes. The term conceptus refers
to the embryo, or to the fetus later on, and all of the extraembryonic structuresthe products of conception.
CHAPTER 22
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Embryology is the study of the sequential changes in an organism as the various tissues, organs, and systems develop.
Chick embryos are frequently studied because of the easy access
through the shell and their rapid development. Mice and pig embryos
are also extensively studied as mammalian models. Genetic manipulation, induction of drugs, exposure to disease, radioactive tagging or
dyeing of developing tissues, and X-ray treatments are some of the
commonly conducted experiments that provide information that can
be applied to human development and birth defects.
During the preembryonic period of cell division and differentiation, the developing structure is self-sustaining. The embryo, however, must derive sustenance from the mother. For
morphogenesis to continue, certain immediate needs must be
met. These needs include (1) formation of a vascular association
between the uterus of the mother and the embryo so that nutrients and oxygen can be provided and metabolic wastes and car-
bon dioxide can be removed; (2) establishment of a constant,

protective environment around the embryo that is conducive to
development; (3) establishment of a structural foundation for
embryonic morphogenesis along a longitudinal axis; (4) provision for structural support for the embryo, both internally and
externally; and (5) coordination of the morphogenic events
through genetic expression. If these needs are not met, a spontaneous abortion will generally occur.
The first and second of these needs are provided for by extraembryonic structures; the last three are provided for intraembryonically. The extraembryonic membranes, the placenta, and
the umbilical cord will be considered separately, prior to a discussion of the development of the embryo.
Serious developmental defects usually cause the embryo to
be naturally aborted. About 25% of early aborted embryos
have chromosomal abnormalities. Other abortions may be caused by
environmental factors, such as infectious agents or teratogenic
drugs (drugs that cause birth defects). In addition, an implanted embryo is regarded as foreign tissue by the immune system of the
mother, and is rejected and aborted unless maternal immune responses are suppressed.
Extraembryonic Membranes
At the same time that the internal organs of the embryo are
being formed, a complex system of extraembryonic membranes
is also developing (fig. 22.10). The extraembryonic membranes
are the amnion, yolk sac, allantois, and chorion. These membranes
are responsible for the protection, respiration, excretion, and
nutrition of the embryo and subsequent fetus. At parturition,
the placenta, umbilical cord, and extraembryonic membranes
separate from the fetus and are expelled from the uterus as the
afterbirth.
Amnion
The amnion (am'ne-on) is a thin membrane, derived from ectoderm and mesoderm. It loosely envelops the embryo, forming an
amniotic sac that is filled with amniotic fluid (fig. 22.11b). In
later fetal development, the amnion expands to come in contact
with the chorion. The development of the amnion is initiated
early in the embryonic period, at which time its margin is attached to the free edge of the embryonic disc (see fig. 22.10). As
the amniotic sac enlarges during the late embryonic period (at
about 8 weeks), the amnion gradually sheaths the developing
umbilical cord with an epithelial covering (fig. 22.12).
The buoyant amniotic fluid performs several functions for
the embryo and subsequent fetus.
It ensures symmetrical structural development and growth.
It cushions and protects by absorbing jolts that the mother
may receive.
It helps maintain consistent pressure and temperature.
It helps eliminate metabolic wastes.
It permits freedom of fetal movement, which is important
for skeletomuscular development and blood flow.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
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Amniotic
fluid
Embryonic
disc
Amnion
Allantois
Implanted
embryo
Yolk sac
Body of
uterus
Chorion
(a)
Amnion
(b) Connecting
stalk
(a)
Umbilical cord
Placenta
Yolk sac
Allantois
Chorion
frondosum
(c)
Villi of chorion
frondosum
Creek
FIGURE 22.10 The formation of the extraembryonic membranes
Chorion
Amnion
during a single week of rapid embryonic development. (a) At

3 weeks, (b) at 312 weeks, and (c) at 4 weeks.
Amniotic sac
containing
amniotic fluid
Amniotic fluid is formed initially as an isotonic fluid absorbed from the maternal blood in the endometrium surrounding
the developing embryo. Later, the volume is increased and the
concentration changed by urine excreted from the fetus into the
amniotic sac. Amniotic fluid also contains cells that are sloughed
off from the fetus, placenta, and amniotic sac. Because all of these
cells are derived from the same fertilized egg, all have the same
genetic composition. Many genetic abnormalities can be detected
by aspirating a sample of this fluid and examining the cells obtained, in a procedure called amniocentesis (am''ne-o-sen-te'sis).
Amniotic fluid is normally swallowed by the fetus and absorbed in the GI tract. The fluid enters the fetal blood, and the
waste products it contains enter the maternal blood in the plaDown syndrome: from John L. H. Down, English physician, 182896
(b)
Umbilical blood vessels
FIGURE 22.11 An implanted embryo at approximately 412 weeks.

(a) The interior of a uterus showing the implantation site. (b) The developing embryo, extraembryonic membranes, and placenta.
centa. Prior to delivery, the amnion is naturally or surgically ruptured, and the amniotic fluid (bag of waters) is released.
As the fetus grows, the amount of amniotic fluid increases.
It is also continually absorbed and renewed. For the near-term
baby, almost 8 liters of fluid are completely replaced each day.
Yolk Sac
The yolk sac is established during the end of the second week as
cells from the trophoblast form a thin exocoelomic (ek''so-selo'mik) membrane. Unlike the yolk sac of many vertebrates, the
CHAPTER 22
Amniocentesis (fig. 22.13) is usually performed at the fourteenth or fifteenth week of pregnancy, when the amniotic sac
contains 175225 ml of fluid. Genetic diseases, such as Down
syndrome, or trisomy 21 (in which there are three instead of two
number21 chromosomes), can be detected by examining chromosomes. Diseases such as Tay-Sachs disease, in which there is a defective enzyme involved in formation of myelin sheaths, can be
detected by biochemical techniques from these fetal cells.
Yolk sac

764
Unit 7

Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
Companies, 2001
Decidua
basalis
Chorion
frondosum
Maternal
vein
Umbilical
cord
Chorion
Maternal
artery
Amnion
Amniotic sac containing

amniotic fluid
Placenta
FIGURE 22.12 The embryo, extraembryonic membranes, and placenta at approximately 7 weeks of development. Blood from the embryo is
CHAPTER 22
carried to and from the chorion frondosum by the umbilical arteries and vein. The maternal tissue between the chorionic villi is known as the decidua basalis; this tissue, together with the villi, form the functioning placenta.
human yolk sac contains no nutritive yolk, but it is still an essential structure during early embryonic development. Attached to
the underside of the embryonic disc (see figs. 22.10 and 22.11), it
produces blood for the embryo until the liver forms during the
sixth week. A portion of the yolk sac is also involved in the formation of the primitive gut. In addition, primordial germ cells
form in the wall of the yolk sac. During the fourth week, they
migrate to the developing gonads, where they become the primitive germ cells (spermatogonia or oogonia).
The stalk of the yolk sac usually detaches from the gut by
the sixth week. Following this, the yolk sac gradually shrinks as
pregnancy advances. Eventually, it becomes very small and
serves no additional function.
blood cells and gives rise to the fetal umbilical arteries and vein.
It also contributes to the development of the urinary bladder.
The extraembryonic portion of the allantois degenerates
during the second month. The intraembryonic portion involutes
to form a thick urinary tube called the urachus (yoo'ra-kus) (see
p. 690). After birth, the urachus becomes a fibrous cord called
the median umbilical ligament that attaches to the urinary bladder.
Chorion
The allantois forms during the third week as a small outpouching, or diverticulum, near the base of the yolk sac (see
fig. 22.10). It remains small but is involved in the formation of
The chorion (kor'e-on) is the outermost extraembryonic membrane. It contributes to the formation of the placenta as small fingerlike extensions, called chorionic villi, penetrate deeply into the
uterine tissue (see fig. 22.10). Initially, the entire surface of the
chorion is covered with villi. But those chorionic villi on the surface toward the uterine cavity gradually degenerate, producing a
smooth, bare area known as the smooth chorion. As this occurs,
the chorionic villi associated with the uterine wall rapidly increase
in number and branch out. This portion of the chorion is known
allantois: Gk. allanto, sausage; iodos, resemblance
chorion: Gk. chorion, external fetal membrane
Allantois


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
The McGrawHill
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Two
chorions
787
Two
amnions
Implantation
of blastocysts
Two
zygotes
Two-cell
stage
Blastocysts
Separate
chorionic sacs
Separate placentas
(a)
Two
chorions
fused
(b)
Two
amnions
Fused placentas
Cre
ek
Fused chorionic sacs
(c)
FIGURE 22.36 The formation of dizygotic twins. Twins of this type are fraternal rather than identical and may have (a) separate or (b) fused
placentas. (c) A photograph of fraternal twins at 11 weeks.
Fetoscopy (fig. 22.39) goes beyond amniocentesis by allowing direct examination of the fetus. Using fetoscopy, physicians
scan the uterus with pulsed sound waves to locate fetal structures,
the umbilical cord, and the placenta. Skin samples are taken from
the head of the fetus and blood samples extracted from the placenta. The principal advantage of fetoscopy is that external features of the fetus (such as fingers, eyes, ears, mouth, and genitals)
can be carefully observed. Fetoscopy is also used to determine
several diseases, including hemophilia, thalassemia, and sicklecell anemia cases, 40% of which are missed by amniocentesis.
Most hospitals are now equipped with instruments that
monitor fetal heart rate and uterine contractions during labor.
These instruments can detect any complication that may arise
during the delivery. The procedure is called Electronic Monitoring
amniocentesis: Gk. amnion, lamb (fetal membrane); kentesis, puncture
fetoscopy: L. fetus, offspring; skopein, to view
CHAPTER 22
sonography employs a mechanical vibration of high frequency to

produce a safe, high-resolution (sharp) image of fetal structure
(fig. 22.38). Ultrasonic imaging is a reliable way to determine
pregnancy as early as 6 weeks after ovulation. It can also be used
to determine fetal weight, length, and position, as well as to diagnose multiple fetuses.
Amniocentesis is a technique used to obtain a small sample of amniotic fluid so that it can be assessed genetically and
biochemically (see fig. 22.13). A wide-bore needle is inserted
into the amniotic sac with guidance by ultrasound, and 510 ml
of amniotic fluid is withdrawn with a syringe. Amniocentesis is
most often performed to determine fetal maturity, but it can also
help predict serious disorders such as Down syndrome or
Gauchers disease (a metabolic disorder).

788
Unit 7

Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
Companies, 2001
One placenta
One chorionic sac

Two amniotic sacs
Implantation
of blastocyst
Zygote
Two-cell
stage
Blastocyst with two

embryoblasts
(a)
Single
placenta
(b)
C re
ek
Anastomosis of
placental vessels
FIGURE 22.37 The formation of monozygotic twins. Twins of this type develop from a single zygote and are identical. Such twins have two
amnions but only one chorion, and they share a common placenta.
Amniotic fluid
Placenta
Left cerebral
hemisphere
Syringe
withdraws
blood
Endoscope (guided
to exact location by
pulsed sound waves)
Orbit of eye
Left hand
Uterine wall
CHAPTER 22
Thorax
FIGURE 22.38 A color-enhanced ultrasonogram of a fetus during

the third trimester. The left hand is raised, as if waving to the viewer.
Placenta
Fiber optics
Needle punctures fetal
vein on placenta
FIGURE 22.39 Fetoscopy.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
The McGrawHill
Companies, 2001
789
Fetal monitor
Amplifier
Cardiotachometer
Amplifier
Fetal
heart rate
Uterine
activity
Pressure transducer
FIGURE 22.40 Monitoring the fetal heart rate and uterine contractions using an FHRUC device.
of Fetal Heart Rate and Uterine Contractions (FHR-UC Monitoring). The extent of stress to the fetus from uterine contractions
can be determined through FHR-UC monitoring (fig. 22.40).
Long, arduous deliveries are taxing to both the mother and fetus.
If the babys health and vitality are presumed to be in danger because of a difficult delivery, the physician may decide to perform
a cesarean section.
Congenital Defects
Major developmental problems called congenital malformations
occur in approximately 2% of all newborn infants. They may be
caused by genetic inheritance, mutation (genetic change), or environmental factors. About 15% of neonatal deaths are attributed to congenital malformations. The branch of developmental
biology concerned with abnormal development and congenital

Two amnions and one chorion in all but very unusual cases prove the
twins to be monozygotic. The two infants, therefore, are genetically
identical. Thus, when considering genetic disorders such as cleft palate,
one would expect a high degree of concordance (both twins of a
monozygotic pair exhibit a particular anomaly). Many such defects however can be present in only one twina consequence of nongenetic factors, such as intrauterine environment. An example would be
inadequate blood supply to only one twin, resulting in a defect in that
twin but not in the other.
teratology: Gk. teras, monster; logos, study of
CHAPTER 22
congenital: L. congenitus, born with
malformations is called teratology (tar''a-tol'o-je). Many congenital problems have been discussed in previous chapters, in connection with the body system in which they occur.

790
Unit 7

Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
Companies, 2001

A 31-year-old female comes to your office
for her annual gynecologic exam. She reports no menstrual abnormalities. On
physical exam, you note a mass in her left
adnexa (accessory component of the main
organ). Because of this finding, you order a
CT scan to further evaluate this area. (B =
urinary bladder.)
QUESTIONS
1. What is the heterogeneous mass
(arrow) seen to the left of the urinary
bladder (B)?
2. What organ and cell line does this mass
arise from?
3. What is the significance of this mass?
Genetic Disorders of Clinical Importance
CHAPTER 22
cystic fibrosis An autosomal recessive disorder

characterized by the formation of thick mucus
in the lungs and pancreas that interferes with
normal breathing and digestion.
familial cretinism An autosomal recessive
disorder characterized by a lack of thyroid
secretion because of a defect in the iodine
transport mechanism. Untreated children are
dwarfed, sterile, and may be mentally retarded.
galactosemia (ga-lak''te-se'me-a) An autosomal
recessive disorder characterized by an
inability to metabolize galactose, a
component of milk sugar. Patients with this
disorder have cataracts, damaged livers, and
mental retardation.
gout An autosomal dominant disorder
characterized by an accumulation of uric acid
in the blood and tissue resulting from an
abnormal metabolism of purines.
hepatic porphyria (por-fer'e-a) An autosomal
dominant disorder characterized by painful GI
disorders and neurologic disturbances resulting
from an abnormal metabolism of porphyrins.
Huntingtons chorea: from George Huntington,

American physician, 18501916
Marfans syndrome: from Antoine Bernard-Jean
Marfan, French physician 18581942
hereditary hemochromatosis (he''mo-kro''mato'sis) A sex-influenced autosomal dominant

disorder characterized by an accumulation of
iron in the pancreas, liver, and heart,
resulting in diabetes, cirrhosis, and heart
failure.
hereditary leukomelanopathy (loo''ko-mel''anop'a-the) An autosomal recessive disorder
characterized by decreased pigmentation in
the skin, hair, and eyes and abnormal white
blood cells. People with this condition are
generally susceptible to infections and early
deaths.
Huntingtons chorea An autosomal dominant
disorder characterized by uncontrolled
twitching of skeletal muscles and the
deterioration of mental capacities. A latent
expression of this disorder allows the mutant
gene to be passed to children before
symptoms develop.
Marfans syndrome An autosomal dominant
disorder characterized by tremendous growth
of the extremities, extreme looseness of the
joints, dislocation of the lenses of the eyes,

and congenital cardiovascular defects.
phenylketonuria (fen''il-ket''n-oor'e-a)
(PKU) An autosomal recessive disorder
characterized by an inability to metabolize
the amino acid phenylalanine. It is
accompanied by brain and nerve damage and
mental retardation. (Newborns are routinely
tested for PKU; those that are affected are
placed on a diet low in phenylalanine.)
pseudohypertrophic muscular dystrophy A
sex-linked recessive disorder characterized by
progressive muscle atrophy. It usually begins
during childhood and causes death in
adolescence.
retinitis pigmentosa A sex-linked recessive
disorder characterized by progressive atrophy
of the retina and eventual blindness.
Tay-Sachs disease An autosomal recessive
disorder characterized by a deterioration of
physical and mental abilities, early blindness,
and early death. It has a disproportionately high
incidence in Jews of Eastern European origin.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
The McGrawHill
Companies, 2001
791
Chapter Summary
Fertilization (pp. 755757)
1. Upon fertilization of a secondary oocyte
by a spermatozoon in the uterine tube,
meiotic development is completed and a
diploid zygote is formed.
2. Morphogenesis is the sequential
formation of body structures during the
prenatal period of human life. The
prenatal period lasts 38 weeks and is
divided into a preembryonic, an
embryonic, and a fetal period.
3. A capacitated spermatozoon digests its
way through the zona pellucida and
corona radiata of the secondary oocyte to
complete the fertilization process and
formation of a zygote.
2.
Preembryonic Period (pp. 757762)
Embryonic Period (pp. 762771)

1. The events of the 6-week embryonic
period include the differentiation of the
germ layers into specific body organs and
3.
4.
5.
Fetal Period (pp. 772774)

1. A small amount of tissue differentiation and
organ development occurs during the fetal
period, but for the most part fetal development
is primarily limited to body growth.
2. Between weeks 9 and 12, ossification

centers appear, the genitalia are formed,
and the digestive, urinary, respiratory, and
muscle systems show functional activity.
3. Between weeks 13 and 16, facial features
are formed and the fetal heartbeat can be
detected with a stethoscope.
quickening can be felt by the mother, and
vernix caseosa and lanugo cover the skin
of the fetus.
substantial weight gain occurs and the
fetal skin becomes wrinkled and pinkish.
6. Toward the end of the 26-to-29-week
period, the eyes have opened, the gonads
have descended in a male, and the fetus is
developed to the extent that it might
survive if born prematurely.
7. At 38 weeks, the fetus is full-term; the
normal gestation is 266 days.
Labor and Parturition (p. 775)

1. Labor and parturition are the culmination
of gestation and require the action of
oxytocin, secreted by the posterior
pituitary, and prostaglandins, produced in
the uterus.
2. Labor is divided into dilation, expulsion,
and placental stages.
Periods of Postnatal Growth

(pp. 775782)
1. The course of human life after birth is
seen in terms of physical and
physiological changes and the attainment
of maturity in the neonatal period,
infancy, childhood, adolescence, and
adulthood.
2. The neonatal period, extending from birth to
the end of the fourth week, is characterized
by major physiological changes.
(a) The most critical need of the newborn
is to establish adequate respiratory and
heart rates. A normal respiratory rate
is 30 to 40 respirations per minute,
and a normal heart rate ranges from
120 to 160 beats per minute.
(b) The four reflexes in the newborn
critical to survival are the suckling
reflex, the rooting reflex, the crying
reflex, and the breathing reflex.
3. Infancy, extending from 4 weeks through
the second year, is characterized by
tremendous growth, increased
coordination, and mental development.
(a) By 2 years, most infants weigh about
4 times their birth weight and average
between 32 and 36 inches in length.
CHAPTER 22
1. Cleavage of the zygote is initiated within

30 hours and continues until a morula
forms; the morula enters the uterine
cavity on about the third day.
2. A hollow, fluid-filled space forms within
the morula, at which point it is called a
blastocyst.
3. Implantation begins between the fifth and
seventh day and is enabled by the
secretion of enzymes that digest a portion
of the endometrium.
(a) During implantation, the trophoblast
cells secrete human chorionic
gonadotropin (hCG), which prevents
the breakdown of the endometrium
and menstruation.
(b) The secretion of hCG declines by the
tenth week as the developed placenta
secretes steroids that maintain the
endometrium.
4. The embryoblast of the implanted
blastocyst flattens into the embryonic
disc, from which the primary germ layers
of the embryo develop.
(a) Ectoderm gives rise to the nervous
system, the epidermis of the skin and
epidermal derivatives, and to portions
of the sensory organs.
(b) Mesoderm gives rise to bones,
muscles, blood, reproductive organs,
the dermis of the skin, and
connective tissue.
(c) Endoderm gives rise to linings of the
GI tract, digestive organs, the
respiratory tract and lungs, and the
urinary bladder and urethra.
the formation of the placenta, the

umbilical cord, and the extraembryonic
membranes. These events make it possible
for morphogenesis to continue.
The extraembryonic membranes include
the amnion, yolk sac, allantois, and
chorion.
(a) The amnion is a thin membrane
surrounding the embryo. It contains
amniotic fluid that cushions and
protects the embryo.
(b) The yolk sac produces blood for the
embryo.
(c) The allantois also produces blood for
the embryo and gives rise to the
umbilical arteries and vein.
(d) The chorion participates in the
The placenta, formed from both maternal
and embryonic tissue, has a transport role
in providing for the metabolic needs of
the fetus and in removing its waste.
(a) The placenta produces steroid and
polypeptide hormones.
(b) Nicotine, drugs, alcohol, and viruses
can cross the placenta to the fetus.
The umbilical cord, containing two
umbilical arteries and one umbilical vein,
is formed as the amnion envelops the
tissues on the underside of the embryo.
From the third to the eighth week, the
structure of all the body organs, except
the genitalia, becomes apparent.
(a) During the third week, the primitive
node forms from the primitive line,
which later gives rise to the
notochord and intraembryonic
mesoderm.
(b) By the end of the fourth week, the
heart is beating; the primordial tissues
of the eyes, brain, spinal cord, lungs,
and digestive organs are properly
positioned; and the superior and
inferior limb buds are recognizable.
(c) At the end of the fifth week, the
sense organs are formed in the
enlarged head and the appendages
have developed with digital primordia
evident.
(d) During the seventh and eighth weeks,
the body organs are formed, except
for the genitalia, and the embryo
appears distinctly human.

792
Unit 7

Development
22. Developmental
Anatomy, Postnatal
The McGrawHill
Companies, 2001
(b) Growth is a differential process

resulting in gradual changes from
infant to adult body proportions.
4. Childhood, extending from the end of
infancy to adolescence, is characterized by
steady growth until preadolescence, at
which time there is a marked growth spurt.
(a) During childhood, the average child
becomes thinner and stronger each
year as he or she grows taller.
(b) The fact that disease and death are
relatively rare during childhood may
be due to the fact that lymphoid
tissue is at its peak of development at
this time; it is also present in greater
amounts in children than in adults.
5. Adolescence is the period of growth and
development between childhood and
adulthood.
(a) Puberty is the stage of early
adolescence when the secondary sex
characteristics are expressed and the
sex organs become functional.
(b) The end result of puberty is the
structural expression of gender, or
sexual dimorphism.
(c) Menarche generally occurs in
adolescent girls at the age of 13, but it
may range from 9 to 17 years. At this
time, vaginal secretions change from
alkaline to acid.
(d) The first physical indications of
puberty are the appearance of breast
buds in females and the growth of the

testes and the appearance of sparse
pubic hair in males.
(e) Although semen may be ejaculated at
age 13, sufficient mature spermatozoa
for fertility are not produced until 14
to 16 years of age.
6. Adulthood, the final period of human
physical change, is characterized by
gradual senescence as a person ages.
(a) Although skeletal maturity is reached
in early adulthood, anatomical and
physiological changes continue
throughout adulthood and are part of
the aging process.
(b) Sexual dimorphism in human adults
is evident anatomically,
physiologically, metabolically, and
behaviorally.
(c) Male and female differences in the
stature, proportions, and composition
of the body may become more
apparent with age.
Inheritance (pp. 782785)

1. Inheritance is the passage of hereditary
traits carried on the genes of
chromosomes from one generation to
another.
2. Each zygote contains 22 pairs of
autosomal chromosomes and 1 pair of sex
chromosomesXX in a female and XY in
a male.
3. A gene is the portion of a DNA molecule

that contains information for the
production of one kind of protein
molecule. Alleles are different forms of
genes that occupy corresponding positions
on homologous chromosomes.
4. The combination of genes in an
individuals cells constitutes his or her
genotype; the observable expression of the
genotype is the persons phenotype.
(a) Dominant alleles are symbolized by
uppercase letters and recessive alleles
are symbolized by lowercase letters.
(b) The three possible genotypes are
homozygous dominant, heterozygous,
and homozygous recessive.
5. A Punnett square is a convenient means
for expressing probability.
(a) The probability of a particular
genotype is 1 in 4 (.25) for
homozygous dominant and
homozygous recessive, and 1 in 2
(.50) for heterozygous.
(b) A single trait is studied in a
monohybrid cross; two traits are
studied in a dihybrid cross.
6. Sex-linked traits such as color blindness
and hemophilia are carried on the sexdetermining chromosome.
Review Activities
CHAPTER 22
Objective Questions
1. The preembryonic period is completed
when
(a) the blastocyst implants.
(b) the placenta forms.
(c) the blastocyst reaches the uterus.
(d) the primary germ layers form.
2. The yolk sac produces blood for the
embryo until
(a) the heart is functional.
(b) the kidneys are functional.
(c) the liver is functional.
(d) the baby is delivered.
3. Which of the following is a function of
the placenta?
(a) production of steroids and hormones
(b) diffusion of nutrients and oxygen
(c) production of enzymes
(d) all of the above apply
4. The decidua basalis is
(a) a component of the umbilical cord.
(b) the embryonic portion of the villous
chorion.
5.
6.
7.
8.
(c) the maternal portion of the placenta.

(d) a vascular membrane derived from
the trophoblast.
Which of the following could diffuse
across the placenta?
(a) nicotine
(c) heroin
(b) alcohol
(d) all of the above
During which week following conception
does the embryonic heart begin pumping
blood?
(a) the fourth week
(b) the fifth week
(c) the sixth week
(d) the eighth week
Which of the following is the period of growth
from birth to the end of the fourth week?
(a) the neonatal period
(b) the fetal period
(c) infancy
(d) the suckling period
The normal newborn heart rate is
(a) 7080 beats/min.
(b) 120160 beats/min.
(c) 100120 beats/min.

(d) 180200 beats/min.
9. The continuum of physical change in
adolescence that regulates the growth of
body hair is known as
(a) puberty.
(b) pubal progression.
(c) pubescence.
(d) dimorphism.
10. Which condition is not characteristic of
the female as compared to the male?
(a) lower blood pressure
(b) higher basal metabolic rate
(c) lower red blood cell count
(d) faster heart rate
11. The first physical indication of puberty in
females is generally
(a) alkaline vaginal secretions.
(b) a widening pelvis.
(c) breast buds.
(d) axillary hair.


Development
22. Developmental
Anatomy, Postnatal
Chapter 22
12. Twins that develop from two zygotes
resulting from the fertilization of two ova
by two spermatozoa in the same ovulatory
cycle are referred to as
(a) monozygotic. (c) dizygotic.
(b) conjoined.
(d) identical.
13. Match the genotype descriptions in the
left-hand column with the correct
symbols in the right-hand column.
homozygous recessive
Bb
heterozygous
bb
homozygous dominant
BB
14. An allele that is not expressed in a
heterozygous genotype is called
(a) recessive.
(c) genotypic.
(b) dominant.
(d) phenotypic.
15. If the genotypes of both parents are Aa
and Aa, the offspring probably will be
(a) 12 AA and 12 aa.
(b) all Aa.
(c) 14 AA, 12 Aa, 14 aa.
(d) 34 AA and 14 aa.
Essay Questions
1. Describe the implantation of the
blastocyst into the uterine wall and the
involvement of the trophoblast in the
2. Explain how the primary germ layers
form. What major structures does each
germ layer give rise to?
3. Explain why development is so critical
during the embryonic period and list the
embryonic needs that must be met for
morphogenesis to continue.
4. State the approximate time period (in
weeks) for the following occurrences:
(a) appearance of the arm and leg buds.
(b) differentiation of the external
genitalia.
5.
6.
7.
8.
9.
10.
11.
12.
The McGrawHill
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(c) perception of quickening by the

mother.
(d) functioning of the embryonic heart.
(e) initiation of bone ossification.
(f) appearance of lanugo and vernix caseosa.
(g) survival of fetus if born prematurely.
(h) formation of all major body organs
completed.
Define the term infancy and discuss the
growth and developmental events
characteristic of this period of life.
Define the term childhood and discuss the
growth and developmental events
characteristic of this period of life.
Define the term adolescence and discuss
the role of puberty in this period of life for
both males and females.
Distinguish between puberty and pubescence.
Define the term adulthood and describe
the characteristics of this period of life.
List some of the structural features of
adults that exemplify sexual dimorphism.
Define anthropometry and give examples
of anthropometric characteristics.
State the features of a genetic disorder
that would lead one to believe that it was
a form of sex-linked inheritance.
1. Write a short paragraph about pregnancy
that includes the terms ovum, blastocyst,
implantation, embryo, fetus, gestation, and
parturition.
793
2. If a pregnant woman had a pack-a-day

cigarette habit, what predictions could
you make about the health of her baby?
Justify your responses.
3. The sedative thalidomide was used by
thousands of pregnant women in the
1960s to alleviate their morning sickness.
This drug inhibited normal limb
development and resulted in tragically
deformed infants with flipperlike arms
and legs. At what period of prenatal
development did such abnormalities
originate? What lessons were learned from
the thalidomide tragedy?
4. Your friend just learned that shes
pregnant and she wants to name the baby
either Louis or Louise. How soon can she
know for sure whether she is carrying a
boy or a girl? Describe the technique by
which the sex can be determined in vitro.
5. Cesarean sections must be performed
when a baby cannot be delivered breech.
Can you think of some other reasons for a
C-section?
6. Hemophilia used to be called a royal disease
because it plagued many male members of
the royal families of Europe and Russia.
Why are female hemophiliacs rare, as their
absence from royal pedigrees shows?
7. In many states, laws prohibit
consanguineous marriagesthose
between blood relatives such as siblings or
first cousins. Why is it likely that a
geneticist would endorse such restrictions?

CHAPTER 22

Back Matter
Appendix A: Answers to
Objective Questions with
Explanations
The McGrawHill
Companies, 2001
Appendix A
Answers to Objective Questions with Explanations
APPENDIX A
Chapter 1
1. (a) The word anatomy is derived
from the Greek prefix ana,
meaning up and the suffix tome,
a cutting. In ancient times, the
Greek word anatomize was more
commonly used than the word
dissect.
2. (a) In William Harveys time,
there were many misconceptions
regarding the flow of blood. Using
the scientific method, Harvey
demonstrated the flow of blood
and described the function of
valves within veins.
3. (c) The concept of body humors
was widely accepted by physicians
in Greek and Roman times as an
explanation for a persons
disposition and general health. It
was not until the Renaissance that
this concept was gradually
discarded in the light of scientific
research.
4. (d) The Greeks held the body in
great esteem and were eager to
learn about its structure and
function.
5. (a) Hippocrates is regarded as the
father of medicine because of the
sound principles of medical
practice that he established.
Although the Hippocratic oath
cannot be directly credited to him,
it undoubtedly represents his ideals
and principles.
6. (c) Phlegm was thought to be the
body humor associated with the
lungs. It was believed that too
much phlegm, or not enough,
could result in respiratory
disorders.
7. (c) Andreas Vesalius completed
De Humani Corporis Fabrica in A.D.
1543 when he was 28 years old.
This beautifully illustrated book
corrected many anatomical errors
that had been handed down as fact
and provided a visual guide for
studying anatomy.
8. (d) The development of movable
type allowed for widespread access
to printed material, and was
therefore important in ushering in
the Renaissance.
794
9. (b) Aristotle thought that the

heart was the seat of intelligence
and that the warm blood from the
heart was cooled by the fluids
surrounding the brain.
10. (a) Wilhelm Konrad Roentgen
discovered X rays in 1895. His
achievement was recognized in
1901, when he was awarded the
Nobel Prize.
Chapter 2
1. (b) Although chordates have a
distinct head, thorax, and
abdomen, so do many invertebrate
organisms.
2. (a) Because we have prehensile
hands, digits modified for grasping,
and large, well-developed brains,
we are classified as primates along
with monkeys and great apes.
3. (b) The tissue level of body
organization is intermediate
between the cell and organ levels.
4. (a) The urinary system maintains
blood homeostasis. Basically, the
kidneys are filtering organs of the
blood.
5. (b) The cubital fossa is located on
the anterior surface of the upper
extremity, at the junction of the
brachium and antebrachium. It is
the anterior surface of the elbow.
6. (c) An organ is a body structure
composed of two or more tissue
types.
7. (c) By definition, both the coronal
and sagittal planes extend
vertically through the body, but
there is no vertical plane
designation.
8. (b) The external genitalia are
located within the perineum,
which is the area between the
coccyx and the symphysis pubis.
9. (b) Within the mediastinum are
the principal (primary) bronchi,
esophagus, and heart, along with
major vessels that transport blood
toward and away from these
organs.
10. (b) The epigastric region of the
abdomen is centrally located,
medial to the right and left
hypochondriac regions.
11. (c) The mesenteries are modified

serous membranes that support
portions of the small and large
intestines within the abdominal
cavity.
12. (c) A coronal section divides the
body into front (anterior) and back
(posterior) portions.
13. (c) The anatomical position
provides a consistent frame of
reference when describing
structures of the body.
14. (d) Some listening can be
accomplished without the aid of an
instrument, but the technique of
choice is to use a stethoscope.
Chapter 3
1. (c) Electrolytes are formed from
the breakdown of inorganic
compounds in water. All three
kinds of electrolytesacids, bases,
and saltsare important for
normal cellular function.
2. (d) The elements carbon, oxygen,
nitrogen, and hydrogen compose
over 95% of the body and are often
linked to form molecules and
compounds.
3. (a) The strong hydrolytic enzymes
in lysosomes digest foreign
molecules and worn and damaged
cells.
4. (e) Ciliated cells promote
movement of substances through
the tubes or ductules of sections
of the respiratory and reproductive
systems. For example, mucus
containing trapped particles is
moved in the bronchioles and
trachea, sperm are moved in the
ductus deferentia and ova are
moved in the uterine tubes.
5. (b) Osmosis is movement of
water and solvent molecules
through a semipermeable
membrane as a result of
concentration differences.
6. (c) Metaphase is the stage of
mitosis immediately preceding the
separation of the chromatids.
7. (d) Anaphase is the stage of
mitosis immediately following
metaphase; it precedes the
formation of two identical cells.
8. (a) The Golgi complex synthesizes

carbohydrate molecules in the
production of glycoproteins.
9. (b) Compensatory hypertrophy is
an increase in cell mass in response
to greater demands, such as the
enlargement of a muscle fiber as a
result of exercise.
10. (b) Hyperplasia is a protective
mechanism that ensures the
availability of an adequate number
of cells to perform a particular task.
11. (d) cytosine always pairs with
guanine.
12. (b) It contains the sugar ribose.
Chapter 4
1. (b) The four principal types of
tissues are epithelial tissues,
connective tissues, muscular
tissues, and nervous tissues. The
integument, or skin, is an organ.
2. (c) Blood is a fluid tissue that flows
through blood vessels.
3. (a) Many body structures derive
from mesoderm, including
cartilage, bone, and other
connective tissue; smooth, cardiac,
and skeletal muscle; and the
dermis of the skin.
4. (b) Elastic and collagenous fibers
are characteristic of certain kinds
of connective tissues.
5. (d) The entire abdominal portion
of the gastrointestinal (GI) tract
contains simple columnar
epithelium lining the lumen.
6. (a) Based on structural classification,
the mammary glands are classified as
compound acinar. Based on secretory
classification, the mammary glands
are classified as apocrine.
7. (c) Dense regular connective tissue
is the principal component of
tendons, which accounts for their
great strength.
8. (a) Reticular tissue, which contains
white blood cells, is phagocytic in
cleansing body fluids.
9. (b) Cartilage tissue is slow to heal
because it is avascular (without blood
vessels). Cartilage generally derives
its nutrients from surrounding fluids
rather than from permeating blood
vessels and capillaries.

Back Matter
Explanations
Appendix A
10. (d) Like skeletal muscle, cardiac
muscle is striated. As compared to
skeletal muscle or smooth muscle
tissues, however, cardiac muscle
tissue is unique in containing
intercalated discs and experiencing
rhythmic involuntary contractions.
Chapter 5
1. (a) Ectoderm is the outermost of
the three embryonic germ layers,
and it is from this germ layer that
hair, nails, integumentary glands,
and the epidermis of the skin is
formed.
2. (b) The appearance and general
health of nails is an indicator of
general body health and of certain
dietary deficiencies, such as iron
deficiency.
3. (b) The stratum lucidum is an
epidermal layer that occurs only in
the skin of the palms of the hands
and the soles of the feet.
4. (a) The dermal papillae may
contribute to surface features of
the skin, such as print patterns, but
do not contribute to skin
coloration.
5. (c) Mitosis occurs in the stratum
basale, and to a limited extent, in
the stratum spinosum.
6. (a) The sebum from sebaceous
glands is emptied into a hair
follicle, where it is dissipated along
the shaft of the hair to the surface
of the skin.
7. (c) Lanugo, or fetal hair, is
thought to be important in the
development of the hair follicles.
8. (b) Melanoma is an aggressive
malignant skin cancer that is life
threatening if untreated.
9. (b) A second-degree burn is
serious, but it generally does not
require skin grafts.
10. (d) A comedo is a blackhead or
whitehead resulting from a small,
localized skin infection.
Chapter 6
Chapter 7
1. (a) When in anatomical position,
a person is erect and facing
anteriorly. The subscapular fossa is
found on the anterior side of the
scapula. The supraspinatus and
infraspinatus fossae are located on
the posterior side.
2. (c) The sternal extremity of the
clavicle articulates with the
manubrium of the sternum. The
acromial extremity of the clavicle
articulates with the acromion of
the scapula.
3. (d) The clavicle has a conoid
tubercle near its acromial
extremity.
4. (b) The bony prominence of the
elbow is the olecranon of the ulna.
5. (b) Sesamoid bones are formed in
tendons, and none of the carpal
bones are sesamoid bones.
Sesamoid bones are fairly common,
however, at specific joints of the
digits.
6. (d) Pelvimetry measures the
dimension of the lesser pelvis in a
pregnant woman to determine
whether a cesarean delivery might
be necessary.
7. (e) The linea aspera is a vertical
ridge on the posterior surface of
the body of the femur, where the
posterior gluteal muscles of the hip
attach.
8. (b) The intertrochanteric line is

located on the anterior side of the
femur between the greater
trochanter and the lesser
trochanter.
9. (d) Sex-related structural
differences in the pelvis reflect
modifications for childbirth. For
example, the female pelvis as
compared to that of the male has a
wider pelvic outlet, a shallower
and shorter symphysis pubis, and a
wider pubic arch.
10. (a) Talipes is a congenital
malformation in which the sole of
the foot is twisted medially.
Chapter 8
1. (b) Some joints are inmovable,
some are slightly movable, and only
synovial joints are freely movable.
2. (c) Synchondroses are
cartilaginous joints that have
hyaline cartilage between the bone
segments.
3. (d) Syndesmoses are fibrous joints
found in the antebrachium
(forearm) and leg, where adjacent
bones are held together by
interosseous ligaments.
4. (d) Syndesmoses do not occur
within the skull; rather, they are
located only in the upper and
lower extremities.
5. (c) The only saddle joint in the
body is at the base of the thumb,
where the trapezium articulates
with the first metacarpal bone.
6. (d) Only the knee joints contain
menisci.
7. (c) A pivotal joint is a synovial
joint that permits rotational
movement.
8. (b) The coxal (hip) joint has a
wide range of movement,
including hyperextension as the
lower extremity is moved
posteriorly beyond the vertical
position of the body in anatomical
position.
9. (b) The shoulder joint, with its
relatively shallow socket and weak
ligamentous support, is vulnerable
to dislocation. In addition, we
often place our arms in vulnerable
positions as we engage in various
activities.
10. (d) Rheumatoid arthritis is a
chronic disease that frequently
leaves the patient crippled. It
occurs most commonly between
the ages of 30 and 35.
795
Chapter 9
1. (b) The neuromuscular cleft is a
slight gap within the
neuromuscular junction where a
motor nerve fiber and a skeletal
muscle fiber meet.
2. (c) When there are many motor
units present within a muscle, a
person can be more selective at
which ones are recruited, and thus
have greater dexterity than when
few motor units are present.
3. (e) Muscles are named on the basis
of structural features, location,
attachment, relative position, or
function (action).
4. (d) When a nerve impulse reaches
an axon terminal, a
neurotransmitter chemical is
released into the neuromuscular
cleft at the neuromuscular
junction.
5. (c) With respect to the muscles of
the thigh, a single motor neuron
serves a large number of muscle
fibers and, therefore, has a low
innervation ratio.
6. (a) The corrugator supercilli muscle
is located beneath the medial
portion of the eyebrow. When the
muscle contracts, the eyebrow is
drawn toward the midline.
7. (a) A description of a muscles
contraction is always made in
reference to a person in anatomical
position. In anatomical position,
the shoulder joint is positioned
vertically at 180. When the
pectoralis muscle is contracted
while in this position, the angle of
the shoulder joint is decreased;
therefore, the joint is flexed.
8. (b) Although it is positioned on
the anterior surface of the
humerus, the biceps brachii muscle
arises from the coracoid process of
the scapula and from the tuberosity
above the glenoid cavity of the
scapula. Both heads insert on the
radial tuberosity.
9. (c) Spanning both the hip and
knee joints, the rectus femoris
muscle can flex the hip joint and
extend the knee joint.
10. (b) The tibialis posterior muscle is
deep to the soleus muscle. Its
tendinous insertion passes across the
arches of the foot and inserts on the
plantar surfaces of a number of foot
bones. In this position, the tibialis
posterior muscle supports the arches
of the foot and plantar flexes and
inverts the foot as it contracts.
APPENDIX A
1. (c) A bone, such as the femur, is

considered to be an organ because
it is composed of more than two
tissues integrated to perform a
particular function.
2. (a) Skin, rather than bone, is
involved in the synthesis of
vitamin D.
3. Matching: 1(c), 2(d), 3(e),
4(a), 5(b)
4. (c) The mandible articulates with
the mandibular fossae of the
temporal bones.
5. (a) The sella turcica of the

sphenoid bone is located
immediately superior to the
sphenoidal sinus, where it supports
the pituitary gland.
6. (d) Osteoclasts are important in
releasing stored minerals within
bone tissue and in the continuous
remodeling of bone.
7. (a) Located in the squamous part
of the temporal bone, the
mandibular fossa is the depression
for articulation of the condyloid
process of the mandible.
8. (b) The crista galli is the vertical
extension of the ethmoid bone,
which provides an attachment site
for the dura mater.
9. (d) Cervical vertebrae contain
transverse foramina that permit
the passage of the vertebral vessels
to and from the brain.
10. (c) A person may gain some
protection against osteoporosis by
maintaining a healthy diet and a
regular exercise program.
The McGrawHill
Companies, 2001

796
Appendix A
Explanations
The McGrawHill
Companies, 2001
Chapter 10
1. (e) The scalp is attached anteriorly
to the supraorbital ridges. The
eyebrows are attached to the scalp,
just above the supraorbital ridges.
2. (b) The ala is a portion of the
nose, lateral to the apex.
3. (d) The concha is a bony
projection into the nasal cavity.
4. (d) The conjunctiva is a thin
mucous membrane that covers the
anterior surface of the eyeball and
lines the undersurface of the
eyelids.
5. (d) The cervix of the neck is the
anterior portion. The cervical
vertebrae can be palpated in the
posterior portion, called the
nucha.
6. (a) The carotid triangle is located
on the lateral side of the neck and
is bordered by the
sternocleidomastoid, posterior
digastric, and omohyoid muscles.
Located within the carotid triangle
are the common carotid artery,
internal jugular vein, and vagus
nerve.
7. (a) The ulnar nerve passes through
the ulnar sulcus of the elbow. A
fracture of the olecranon of the
ulna often damages the ulnar
nerve.
8. (b) The scapular muscles are
usually obscure on an obese
person.
9. (c) The axilla is the depression
commonly known as the armpit.
The anterior axillary fold is formed
by the pectoralis major muscle and
the posterior axillary fold consists
of the latissimus dorsi muscle.
10. (a) The great saphenous vein and
small saphenous vein are
superficial veins of the leg that are
frequently varicosed in elderly
people.
Chapter 11
APPENDIX A
Back Matter
1. (d) The cerebellum is a structure

within the metencephalon of the
brain.
2. (a) The cerebral cortex is the outer
portion of the cerebrum, and the
cerebrum is a structure within the
telencephalon of the brain.
3. (e) The medulla oblongata is a
structure within the
myelencephalon of the brain.
4. (c) As the nervous system matures
in an infant, the neurolemmocytes
wrap around the axons and some
of the dendrites of neurons within
the peripheral nervous system.
Myelination is the process of
forming myelin layers that protect
neurons and aid conduction.
5. (d) Ganglia are collections of

nerve cell bodies outside the CNS.
Collections of nerve cell bodies
within the CNS are called nuclei.
6. (a) A pseudounipolar neuron has a
single process that divides into
two, and its cell body is located in
posterior root ganglia of the spinal
and cranial nerves.
7. (a) Depolarization of an axon is
produced by the movement of Na+
into the axon and K+ out of the
axon. Once the action potential
(nerve impulse) has completed its
course to the axon terminal,
repolarization of the axon is
produced by the movement of
K+ into the axon and the
movement of Na+ out of the axon.
8. (a) The corpus collosum is
composed of commissural fibers
that connect the two cerebral
hemispheres.
9. (d) The thalamus autonomically
responds to pain by activating the
sympathetic nervous system. It also
relays pain sensations to the
parietal lobes of the cerebrum for
perception.
10. (b) The basal nuclei consist of cell
bodies of motor neurons that regulate
contraction of skeletal muscles. Basal
metabolic rate is regulated, for the
most part, in the hypothalamus and
medulla oblongata.
11. (c) Located within the
mesencephalon, the corpora
quadrigemina is concerned with
visual and hearing reflexes, the red
nucleus is concerned with motor
coordination and posture
maintenance, and the substantia
nigra is thought to inhibit forced
involuntary movements.
12. (d) The fourth ventricle is located
inferior to the cerebellum within
the metencephalon and contains
cerebrospinal fluid.
Chapter 12
1. (a) Anatomically speaking, the
PNS consists of all of the
structures of the nervous system
outside of the CNS. That means,
then, that all nerves, sensory
receptors, neurons, ganglia, and
plexuses are part of the PNS.
2. (b) The oculomotor nerve
innervates the medial rectus eye
muscles that, when simultaneously
contracted, cause the eyes to be
directed medially.
3. (b) The oculomotor nerve
innervates the levator palpebrae
superioris muscle with motor
fibers. This muscle elevates the
upper eyelid when contracted.
4. (c) The vestibulocochlear nerve

serves the vestibular organs of the
inner ear with sensory fibers.
These organs are associated with
equilibrium and balance.
5. (a) Passing through the
stylomastoid foramen, the facial
nerve innervates the muscles of
facial expression with motor fibers
and the taste buds on the anterior
two-thirds of the tongue with
sensory fibers.
6. (b) The accessory nerve innervates
several muscles that move the
head and neck with motor fibers.
7. (c) The four spinal nerve plexuses
are the cervical, brachial, lumbar,
and sacral.
8. (d) Only the brachial plexus
consists of roots, trunks, divisions,
and cords. The nerves of the upper
extremity arise from the cords.
9. (a) The median nerve arises from
the brachial plexus.
10. (c) The knee-jerk reflex is an
ipsilateral reflex because the
receptor and effector organs are on
the same side of the spinal cord.
Chapter 13
1. (d) Postganglionic neurons from
the superior mesenteric ganglion
innervate the small intestine and
colon.
2. (d) Parasympathetic neurons
within the oculomotor, facial, and
glossopharyngeal nerves synapse in
ganglia located in the head.
3. (c) Parasympathetic ganglia, also
called terminal ganglia, synapse
with the effector cells near or
within the organs being served.
4. (c) Secreted from synaptic vesicles,
the neurotransmitter chemical
acetylcholine facilitates
transmission across a synapse.
5. (d) Because the preganglionic
neurons of the sympathetic
division of the autonomic nervous
system exit the vertebral column
from the first thoracic to the
second lumbar levels, the
sympathetic division is also called
the thoracolumbar division.
6. (c) Nerve impulses through the
postganglionic sympathetic
neurons in the heart release
norepinephrine, which stimulates
the heart to contract.
7. (b) The cooperative effect of
sympathetic and parasympathetic
stimulation is evident in the
erection of the penis
(parasympathetic response) and
the ejaculation of semen
(sympathetic response).
8. (d) Most blood vessels dilate in

response to sympathetic
stimulation. A few blood vessels
(e.g., those serving the external
genitalia) constrict in response to
parasympathetic stimulation.
Visceral blood vessels do not
constrict in response to
parasympathetic stimulation.
9. (e) Parasympathetic stimulation
increases digestive activity,
constricts pupils, and decreases the
heart rate. Because atropine blocks
parasympathetic stimulation, it
results in decreased mucus
secretion and GI tract movement
and causes dilation of the pupils. It
also results in an increased heart
rate.
10. (c) The medulla oblongata is the
structure of the brain that most
directly controls the activity of the
ANS. The medulla oblongata
contains control centers for the
circulatory, respiratory, urinary,
reproductive, and digestive
systems.
Chapter 14
1. (e) The adenohypophysis derives
from the hypophyseal pouch in the
roof of the oral cavity.
2. (b) The neurohypophysis derives
from the neurohypophyseal bud of
neuroectoderm within the
developing brain.
3. (d) The adrenal medulla derives
from neural crest ectoderm within
the primitive coelomic cavity.
4. (c) The pancreas derives from an
outpouching of the endoderm
along the developing foregut.
5. (a) The thyroid gland derives from
the thyroid diverticulum of
endoderm within the developing
pharynx.
6. (c) The sella turcica is a depression
within the sphenoid bone that
supports the pituitary gland.
7. (d) Secreted from the thyroid
gland, thyroxine determines the
basic metabolic rate of most organs
and promotes the maturation of
the brain.
8. (c) Activation of the adrenal
medulla causes the secretion of
epinephrine and norepinephrine.
Both of these hormones prepare
the body for greater physical
performancethe fight-or-flight
response.
9. (e) Insulin is secreted from beta
cells within the pancreatic islets in
response to a rise in blood glucose.
Insulin stimulates the production
of glycogen and fat.

Back Matter
Explanations
Appendix A
10. (d) The adrenal medulla releases
epinephrine upon receiving
sympathetic nerve impulse
stimulation.
11. (a) The thyroid gland releases
thyroxine upon receiving TSH
stimulation from the
adenohypophysis of the pituitary
gland.
12. (b) The adrenal cortex releases
corticosteroids upon receiving
ACTH stimulation from the
adenohypophysis of the pituitary
gland in response to stress.
13. (e) The adenohypophysis of the
pituitary gland releases ACTH in
response to receiving CRF
stimulation from the
hypothalamus.
14. (d) Both the adrenal cortex and
the gonads secrete steroid
hormones.
15. (a) In the case of an endemic
goiter, growth of the thyroid is due
to excessive TSH secretion, which
results from low levels of thyroxine
secretion.
Chapter 15
Chapter 16
1. (c) Fibrinogen is a protein in blood
plasma that aids in clotting.
2. (a) An excessive leukocyte count,
called leukocytosis, is generally
diagnostic of infections or diseases
within the body.
3. (b) The pulmonary arteries
transport deoxygenated blood from
the right ventricle of the heart to
the lungs.
4. (c) The right atrium receives
venous blood from the superior
and inferior vena cavae and the
coronary sinus. The coronary sinus
collects venous blood from
coronary circulation prior to
delivery into an opening of the
right atrium.
5. (f) Closure of the atrioventricular
valves causes the lub sound of
the heart.
6. (c) The QRS complex represents
the depolarization of the
ventricles. During this interval,
the ventricles of the heart are in
systole and blood is being ejected
from the heart.
7. (b) Arising from ascending aorta,
the coronary arteries feed directly
into the myocardium of the heart
to ensure a rich blood supply to the
cardiac muscles.
8. (a) The external carotid arteries
supply blood to the entire head,
excluding the brain. The paired
internal carotid arteries and
vertebral arteries (that unite to
form the basilar artery) supply
blood to the brain.
9. (a) Branching from the external
carotid artery at the level of the
mandibular condyle, the maxillary
artery supplies blood to the teeth
and gums of the upper jaw and the
superficial temporal artery supplies
blood to the parotid gland and to
the superficial temporal region.
10. (a) The hepatic portal vein drains

nutrient-rich blood into the liver,
where the blood is processed by a
venous portal system. An arteriole
portal system provides the pituitary
gland with blood.
11. (b) Although the heart begins
pumping blood at 25 days following
conception, its development is not
complete until the end of the fifth
week (at about 35 days).
12. (c) The umbilical vein receives
oxygenated blood from the placenta
and transports it to the fetal heart.
Chapter 17
1. (b) Inhaled air is cleansed,
moistened, and warmed prior to its
arrival at the pulmonary alveoli.
2. (a) The paired palatine bones
support the nasal septum but are
not part of its structure.
3. (b) Adenoid is the common name
of the pharyngeal tonsil. An
adenoidectomy is the removal of
both pharyngeal tonsils.
4. (a) There are four paranasal sinuses,
each named according to the bone
in which it is located. Hence, we
have ethmoidal, sphenoidal,
frontal, and maxillary sinuses.
5. (e) Unlike the right lung that has
three lobes, the left lung has only a
superior lobe and an inferior lobe.
6. (a) The parietal pleura lines the
wall of the thoracic cavity and the
visceral pleura covers the lung.
The space between the parietal
pleura and the visceral pleura is
referred to as the pleural cavity.
7. (d) When contracted, the muscles
of inspiration increase the
dimensions of the thoracic cavity,
causing a decrease in the air
pressure surrounding the lungs. Air
flows through the respiratory tract,
inflating the lungs.
8. (d) The vocal folds (cords) are
attached between the arytenoid
and thyroid cartilages on either
lateral side of the glottis.
9. (c) The vital capacity is the
greatest amount of air that can be
exhaled following a maximal
inhalationan approximate
volume of 4,500 cc of air.
10. (a) The nuclei for normal
breathing are located within the
medulla oblongata of the brain.
Chapter 18
1. (d) Viscera are the organs located
within the trunk of the body. In
the thoracic cavity, the viscera
include the heart, lungs, and
esophagus. In the abdominopelvic
cavity, the viscera include the
2.
3.
4.
5.
6.
7.
8.
9.
10.
797
stomach, small and large intestines,

liver, gallbladder, spleen, pancreas,
kidneys, and adrenal glands, along
with major vessels.
(c) The deciduous dentition
includes 8 incisors, 4 canines, and
8 molars. The permanent dentition
includes 8 incisors, 4 canines, 8
premolars, and 12 molars. The
third molars are called wisdom
teeth.
(b) It is through the supporting
mesentery that vessels and nerves
supply the abdominal viscera.
(c) Lacteals are lymph ductules
found within the lamina propria of
the intestinal villi.
(b) Composed of lymphoid tissue,
the spleen stores red blood cells
and is an organ of the circulatory
system.
(d) The papillae on the surface of
the tongue provide a roughened
surface for physically handling
food. The papillae also support
taste buds for responding to the
chemical stimuli of various foods.
(c) Following the formation of
chyme within the stomach, the
food entering into the small
intestine is ready for additional
digestion and absorption. Bile and
pancreatic juice enter the lumen of
the duodenum to continue the
chemical breakdown of food.
Intestinal movements aid in its
mechanical breakdown. The
nutrients from digested food enter
the bloodstream as absorption
occurs within the small intestine.
(a) As the sphincter of ampulla
opens, bile and pancreatic juice
enter the duodenum. Stenosis of
the sphincter of ampulla would
prohibit the entry of these
products.
(a) The hepatic portal vein
transports absorbed nutrients
within the bloodstream to the
liver, where they are processed.
(d) Fats are absorbed into the
lymphatic system. Proteins are
broken down into amino acids
before they are absorbed. Bile is
secreted into the duodenum. The
liver is served with blood from the
hepatic portal vein and the hepatic
artery; thus, it has a double blood
supply, which becomes mixed at
the capillary level.
Chapter 19
1. (b) The glomeruli are enclosed
within the glomerular capsules in
the renal cortex.
APPENDIX A
1. (d) In order for perception to

occur, there must be a stimulus at a
receptor site that causes a nerve
impulse to be conducted to the
cerebrum of the brain.
2. (b) Located within the dermis of
the skin, in certain visceral organs,
and near synovial joints,
lamellated corpuscles respond to
heavy pressures.
3. (d) Located within connective
tissue capsules in synovial joints,
joint kinesthetic receptors are
stimulated by changes in position
caused by movements at the joints.
4. (c) Because sensations of referred
pain are consistent from one
person to another, an
understanding of this phenomenon
is of immense clinical importance
in diagnosing organ dysfunction.
5. (d) When an object is viewed at a
distance of at least 20 feet by
someone with normal vision, the
suspensory ligaments of the eyes are
taut and the lenses are flat because
the ciliary muscles are relaxed.
6. (a) Composed of tightly bound
elastic and collagenous fibers, the
toughened sclera is avascular but
does contain sensory receptors for
pain.
7. (b) In dim light, sympathetic
motor impulses cause the radially
arranged smooth muscle fibers
within the iris to contract,
permitting the pupil to become
larger.
8. (a) The semicircular ducts contain

endolymph and hair cells that
respond to movements of the head
and convey sensations to the brain
that are important for maintaining
equilibrium and balance.
9. (d) The vestibular window is at
the footplate of the stapes and the
cochlear window borders between
the scala tympani and the
tympanic cavity. Both windows
separate the middle ear from the
cochlea within the inner ear.
10. (b) A concave lens corrects
myopia (nearsightedness) by
causing the light waves to focus
deeper within the posterior cavity
upon the fovea centralis.
The McGrawHill
Companies, 2001

798
Appendix A
Explanations
The McGrawHill
Companies, 2001
2. (d) The hilum of the kidney is the

concave medial surface where the
renal vessels enter and exit and
where the ureter is located.
3. (c) Renal pyramids, containing the
papillary ducts, are located within
the renal medulla.
4. (a) The minor calyx receives urine
directly from the papillary ducts.
The urine then passes through the
major calyx and into the renal
pelvis.
5. (d) The kidneys are located high
in the abdominal cavity between
the levels of the twelfth thoracic
and the third lumbar vertebrae.
6. (d) A renal calculus is most likely
to cause a blockage of a ureter and
backup of urine into the renal
pelvis.
7. (e) The rugae of the empty urinary
bladder permits distension as does
the transitional epithelium that
makes up the mucosa.
8. (c) Collectively, the three layers of
smooth muscle within the wall of
the urinary bladder are referred to
as the detrusor muscle. The
detrusor muscle plays an active
role in forcing urine from the
urinary bladder during micturition.
9. (b) The internal urethral sphincter
is actually a modified portion of
the detrusor muscle and is
innervated by parasympathetic
neurons.
10. (d) Metanephros are the last type
of developmental kidneys. They
are continuously functioning
throughout the fetal period, with
the urine produced being expelled
into the amniotic fluid.
Chapter 20
APPENDIX A
Back Matter
1. (c) The perineum is of functional

and clinical importance because
the external genitalia are located
there.
2. (a) The dartos muscle is embedded
within the wall of the scrotum.
Along with the cremasteric
muscle, it regulates the position of
the testes within the scrotum
through involuntary contraction in
response to cold temperatures.
3. (b) Produced by interstitial cells,
testosterone maintains male
sexuality, including production of
spermatozoa, activity of accessory
sex organs, expression of secondary
sex characteristics, and
determination of sex drives.
4. (e) As components of the root of
the penis, the bulb attaches to the
undersurface of the urogenital
5.
6.
7.
8.
9.
10.
diaphragm and the crus is the

expanded proximal portion of the
corpora cavernosa penis.
(b) The spermatic cord contains
the ductus deferens, which is a
spermatic duct. The spermatic
cord, however, also contains
spermatic vessels, a nerve, and the
cremasteric muscle.
(a) The epididymides are long,
flattened organs that store sperm
in their last stage of maturation.
Sperm are also stored in the ductus
deferentia.
(a) Mucus secreted by the urethral
glands lubricates the urethra and
retards the entry of pathogens into
the urinary bladder.
(c) Emission is movement of stored
sperm from the epididymides and
ductus deferentia to the
ejaculatory ducts. Ejaculation is
the forceful discharge of semen
from the erect penis. Emission and
ejaculation occur only if
stimulation is sufficient, as in
masturbation or coitus.
(a) The ovum from a female may
be fertilized by a sperm cell
containing either an X or a Y
chromosome. If the sperm cell
contains an X chromosome, it will
pair with the X chromosome of the
ovum and a female child will
develop. A sperm cell carrying a Y
chromosome results in an XY
combination, and a male child will
develop. The Y chromosome,
therefore, is responsible for the
subsequent production of
androgens, which cause
masculinization.
(d) The epididymides, ductus
deferentia, and seminal vesicles
derive from the mesonephric duct.
The prostate arises from an
endodermal outgrowth of the
urogenital sinus.
Chapter 21
1. (c) The cervix of the uterus is the
inferior constricted portion that
opens into the vagina.
2. (b) The usual site of fertilization is
within a uterine tube. From there,
the fertilized egg continues to
develop and enters into the cavity
of the uterus about 3 days after
conception.
3. (c) The secretory phase of the
endometrium is characterized by
an increase in glandular secretion
and blood flow into the
endometrium in preparation for
implantation. During this time the
4.
5.
6.
7.
8.
9.
10.
ovary is in its luteal phase,

characterized by a regressive corpus
luteum.
(d) A secondary oocyte is
discharged from an ovary during
ovulation. If a spermatozoon passes
through the corona radiata and
zona pellucida and enters the
cytoplasm of the secondary oocyte,
the second meiotic division is
completed and a mature ovum is
formed.
(d) In addition to the mesovarium,
each ovary is supported by an
ovarian ligament and a suspensory
ligament.
(c) The endometrium consists of a
superficial stratum functionale
layer that is shed as menses during
menstruation and a deeper stratum
basale layer that replenishes the
stratum functionale layer after
each menstruation.
(b) Vaginal rugae permit
distension of the vagina during
coitus and also during parturition.
(d) The vulva is the collective
term for the female external
genitalia. The vagina is an internal
reproductive organ.
(d) The paramesonephric ducts
give rise to the genital tract of the
female reproductive system, which
includes the uterus and the uterine
tubes.
(a) Both the labia majora of the
female and the scrotum of the
male derive from the embryonic
labioscrotal swellings.
Chapter 22
1. (d) The preembryonic period of
developments lasts 14 days and is
completed when the three primary
germ layers have been formed and
are in place to begin migration and
differentiation.
2. (c) The yolk sac only produces
blood for about a 2-week period
(the third to the fifth week) prior
to the formation of the liver. The
liver then produces blood until the
red bone marrow is sufficiently
developed to carry out the task.
3. (d) The placenta is an organ that
serves many physiological
functions, including the exchange
of gases and other molecules
between the maternal and fetal
blood and the production of
enzymes. In addition, it serves as a
barrier against many harmful
substances and as an endocrine
gland in secreting steroid and
glycoprotein hormones.
4. (c) The embryonic portion of the

placenta is the chorion frondosum,
and the maternal portion is the
decidua basalis.
5. (d) Although the placenta is an
effective barrier against diseases of
bacterial origin, viruses and certain
blood-borne diseases can diffuse
through the vascular tissues.
Furthermore, most drugs ingested
by a pregnant woman can readily
pass through the placenta,
including nicotine, alcohol, and
heroin.
6. (a) The embryonic heart begins
pumping blood on about day 25, or
during the fourth week of
development.
7. (a) The neonatal period from birth
to the end of the fourth week is
characterized by major
physiological changes, including
the establishment of a stable heart
rate and respiratory rate, and a
consistent body temperature.
8. (b) A rapid heart rate of 120160
beats/min ensures an adequate
oxygen supply to all of the cells
and helps to maintain a constant
body temperature.
9. (c) Pubescence accompanies
puberty and refers to the
continuum of physical changes
that occur during this period of
maturation, especially with regard
to body hair.
10. (b) As compared to males, females
have a lower basal metabolic rate.
This may account, in part, for the
longer life span of females.
11. (c) The average age at which
breast buds appear in healthy girls
in the United States is about 11
years, but the range is from 9 to 13
years.
12. (c) Dizygotic, or fraternal, twins
may be of the same sex or of
different sexes and are not any
more alike than brothers or sisters
born at different times.
13. homozygous recessive (bb)
heterozygous (Bb)
homozygous dominant (BB)
14. (a) A recessive allele is not
expressed in a heterozygous
genotype. This means that the
particular recessive trait is not
physically apparent.
15. (c) In a monohybrid cross, the
probability of an offspring having a
particular genotype is one in four
for homozygous dominant and
homozygous recessive and one in
two for heterozygous.

Back Matter
Appendix B: Answers to
Clinical Practicum
Questions
The McGrawHill
Companies, 2001
Appendix B
Answers to Clinical Practicum Questions
Clinical Practicum 5.1
1. This rash has the classic appearance and history for poison oak exposure.
The rash is the result of an inflammatory reaction to pathogens deposited by
contact with the poison oak plant. These pathogens are deposited as the
plant is brushed over the skin, therefore it is in a linear pattern.
Inflammatory reaction to this pathogen results in a type of rash known as a
contact dermatitis.
2. Exposure to poison oak deposits allergic pathogens on the skin which incite
an inflammatory response. The cells of the immune system already within
the skin release locally acting cytokines which result in recruitment of other
inflammatory cells to the area as well as increased permeability of the blood
vessels in the area. The release of cytokines and histamines by the
inflammatory cells results in the local pruritis (itching). The increased
permeability of the blood vessels causes local edema. This edema seeps into
intercellular spaces particularly within the stratum spinosum. Increased
intracellular fluid in this area may cause blistering. Because a full-fledged
reaction to the allergic pathogen requires recruitment of additional
inflammatory cells, appearance of symptoms is delayed approximately 24
hours after exposure.
3. Topical steroids are the basis of treatment for poison oak to inhibit the
inflammatory response. Often antihistamine medications are given to
decrease pruritis in order to diminish risk of secondary infection because of
scratching of the rash by the patient. The antihistamines may also relieve
patient discomfort.

1. In the subdural space on the left there is a fluid collection causing mass
effect on the brain. This pressure on the brain can explain the significant
changes in the patients mental status. With changes in mental status and
evidence of mass effect on the brain, concern for herniation of brain tissue is
real. The brain can herniate laterally under the falx cerebri or downward
past the tentorium or through the foramen magnum. Any of these changes is
life threatening.
2. A defect is seen in the posterior wall of the frontal sinus. This defect is
caused by erosion of bone secondary to the patients sinus infection. This
communication of the frontal sinus with the subdural space allows infected
fluid to move from one space to the other. Immune reaction also contributes
to the fluid collection in the subdural space.
3. The diagnosis is a subdural empyema secondary to the patients sinus
infection.
4. The fluid in the subdural space must be removed for two reasons. First, it
must be removed to relieve the pressure on the brain and lessen the risk of
herniation. Second, it must be removed in order to help clear the infection.
Antibiotic treatment must also be administered.

1. This patient has suffered a fracture of the L2 vertebral body.
2. Retropulsion of fracture fragments into the spinal canal causes compression
of the spinal cord. At the level of this injury the portion of the cord injured
is most likely the cauda equina. The injury may be due to compression and
swelling secondary to the fracture or due to damage to the nerves
themselves. The nature of this injury could be further delineated by MRI.
The patients neurologic deficits are manifest in his legs because the nerves
innervating the legs originate at the lumbar and sacral levels of the spine.
3. The seat belt fixes this portion of the body (L1L3) and causes it to become
a fulcrum on which the rest of the body bends during the extreme
deceleration of an automobile accident. The extreme force of bending
focused at this spot leads to compression and fracture of the vertebral body.

1. This is a compound comminuted fracture of the distal radius and ulna with
dorsal displacement and angulation. It is a comminuted fracture because
there are multiple fragments in each fracture. It is an open fracture because
the fractured bones are exposed to the outside of the body through a soft
tissue defect. The fracture fragments are displaced dorsally because of the
mechanism of injury. This type of fracture of the distal radius is classically
known as a Colles fracture.
2. In an open fracture the ends of the fracture fragments are exposed to the
environment through an open wound. This exposes the bone marrow cavity
to infectious environmental pathogens. The bone marrow is a good growth
medium for bacteria. These fractures are at greater risk of developing
infection within the bone which may interfere with healing.
3. Considering the amount of displacement seen on the radiographs, vascular
and nervous injuries associated with the bony trauma would be quite possible.
Vascular injury may be manifest as a pulseless cold hand. Neurologic injury
could have symptoms of either weakness or decreased sensation. These are
emergencies which would require immediate surgical attention. Muscular
injury is also very likely. Continuing vascular and neurologic exams must be
performed on the patient over the ensuing 24 to 48 hours to watch for
possible compartment syndrome. Compartment syndrome occurs when
swelling or hemorrhage secondary to an injury causes increased pressure
within one of the compartments defined by the fascial planes between the
muscle groups of an extremity. This increased pressure within the
compartment causes vascular compromise leading to neurologic injury.

1. There is erosion of the bone of the acetabulum. This is demonstrated as an
area of decreased density just superior to the acetabulum on the plain film
and is clearly shown on the CT scan. This may represent several different
entities, but considering the patients history of thyroid cancer, this is most
likely an osteolytic bony metastasis. The proximity of this lesion to the
acetabulum explains why this patient has increased pain with walking or
weight bearing. These activities put increased pressure on the abnormal bone
causing pain.
2. Metastatic bone lesions are prone to pathologic fracture. By eroding the
bone they weaken its structure. Often bony metastasis and other bony
lesions will initially present as fractures only to have the lesion found on
radiographic evaluation. The risk of pathologic fracture in this patient is
great considering its location near the acetabulum and the weightbearing/weight-distributing function of the pelvis.
799
APPENDIX B
1. Examination of the radiograph shows narrowing of the joint spaces of several

of the distal and proximal interphalangeal joints. This joint space narrowing
is due to erosion of the articular cartilage. Chronic rubbing of bone against
bone has resulted in bony remodeling as evidenced by the presence of
subchondral sclerosis and osteophyte formation. Osteophytes are bony
excressences which form at the margins of joints affected by degenerative
change. Soft tissue swelling is also seen around several of the affected joints.

800
Appendix B
Back Matter
Clinical Practicum
Questions
2. Osteoarthritis. The distribution of joint involvement, symptoms, and physical

findings are classical for idiopathic osteoarthritis. This is a common
degenerative disease of the joints that affects many people as they age. They
will report pain with use of the affected joints as well as morning stiffness in
the joints which resolves quickly with movement. In this disease the articular
cartilage on the ends of bones degenerates leading to joint space narrowing
and bony remodeling secondary to bone-on-bone articulation. Osteoarthritis
of the hands classically involves the distal and proximal interphalangeal
joints and the wrist, but sparing the metacarpophalangeal joints. This differs
from rheumatoid arthritis, which usually involves the metacarpophalangeal
joints first. Swelling around the distal interphalangeal joints is a classic
physical finding of osteoarthritis known as Heberdens nodes.
3. Osteoarthritis is a common degenerative disease that affects many joints of
the body as people age. The most common joints affected include distal and
proximal interphalangeal joints, wrist, acromioclavicular, hips, knees, and
facet joints of the cervical and lumbar spine.

1. The patient has dislocated his shoulder. The head of the humerus has been
forced out of the glenoid cavity by the force of his fall.
2. The dislocation must be reduced by moving the head of the humerus back
into alignment with the glenoid cavity. This can be very painful and is often
performed under anesthesia, because it requires a great amount of traction on
the arm. A radiograph is taken after reduction to check that the head of the
humerus is correctly positioned. Then the shoulder is immobilized to allow
healing of the muscles of the rotator cuff.
3. Because of the structures that pass through the axilla, a dislocation
can damage nerves, arteries, or muscles around and within the shoulder.
Nerve damage may result in weakness and numbness in the shoulder. Arterial
damage may result in pain and impairment of blood flow to the arm. Muscle
damage may result in recurrent dislocations of the shoulder, even after minor
injuries, as well as limited range of motion if rotator cuff injuries occur.

1. A low-density area with peripheral enhancement is seen adjacent to the
psoas major muscle. Low-density collections with peripheral enhancement
such as this are suggestive of an abscess. This abscess is closely related to the
psoas major muscle and does not seem to be related to any of the
surrounding organs. These abscesses generally form from hematologic spread
of bacteria secondary to an infection in another part of the body.
2. The iliopsoas test is a physical examination maneuver performed when there
is suspicion of an intra-abdominal inflammation. The patient lies on the
right (unaffected) side and tries to extend her left leg at the hip against the
examiners hand. Abdominal pain upon extension of the left leg occurs
because of the stretching of the inflammed psoas major muscle. This is
known as a positive psoas sign.
3. The abscess must be drained. This is done by inserting a tube into the
abscess and allowing it to drain over several days. Antibiotics must also be
administered to deal with the infection.
APPENDIX B
The McGrawHill
Companies, 2001
1. This patient has suffered a tibial tuberosity avulsion fracture.

2. The muscles that extend the knee joint are the rectus femoris, vastus
medialis, vastus intermedius, and vastus lateralis. These muscles comprise
the quadriceps femoris group and have a common tendon, the quadriceps
tendon, which extends to the patella. Between the patella and the tibia is
the patellar ligament that attaches at the tibial tuberosity. This patient is
unable to extend his leg at the knee because the tibial tuberosity has avulsed
from the tibia. Therefore the quadriceps femoris muscle cannot exert force
on the tibia.
3. Two major muscle groups, the quadriceps femoris and the hamstrings span
the knee. The quadriceps group extends the knee joint, whereas the
hamstring group flex the knee joint. In the relaxed individual these muscle
groups are in direct opposition to one another and hold the knee in a
neutral, or straight, position through their resting muscle tone. Since the
quadriceps femoris can no longer exhibit force on the tibia because of the
injury, the hamstrings are acting unopposed on the knee. The resting muscle
tone of the hamstrings holds the knee in a slightly flexed position.

A.
B.
C.
D.
E.
F.
liver
psoas major muscle
kidney
gallbladder containing gallstones
spleen
parietal peritoneum

1. This collection of blood is known as an epidural hematoma, and it is usually
caused by a skull fracture which severs the middle meningeal artery. The arterial
blood collects between the dura mater and the skull. These two boundaries give
the hematoma its smooth borders. Because the dura mater is adherent to the
cranium, it resists being stripped from the bone by the accumulating blood. This
is what gives the hematoma its pointed tips. The shape is said to be lenticular
(lens-shaped) and is characteristic of this type of intracranial hematoma.
2. The initial trauma to the brain in these types of injuries is usually minimial,
as evidenced by her brief loss of consciousness. The danger occurs later as
the growing hematoma presses on the brain, raising the intracranial pressure
until coma is induced.
3. The expanding hematoma forces the brain to shift toward the opposite side,
as seen in this CT scan. As the brain shifts, the medial portion of the
temporal lobe often compresses the oculomotor nerve against the tentorium,
causing the pupil to dilate.

1. The patient herniated an intervertebral disc of the lumbar spine. Each disc has
tough circular fibers which surround and contain a soft jellylike center known as
the nucleus pulposus. In a herniated disc, the soft nucleus pulposus has pushed
(herniated) through a weakened area of the fibers to press on a nerve root.
Initially, this causes pain in the distribution of the affected nerve. If continued, it
can also result in numbness/pain and motor deficits specific to the nerve root.
2. By knowing neuroanatomy you can surmise which nerve root is irritated and
therefore which disc is herniated:
Disc
Nerve Root Numbness/Pain Motor Deficit
Lost Reflex
L3L4
L4
Anterior leg
Tibialis anterior
muscle
Patellar
L4L5
L5
Superior foot
L5S1
S1
Lateral foot
Extensor hallucis None

longus muscle
Gastrocnemius
Tendo calcaneous
muscle
3. The fibers that surround the nucleus pulposus are naturally weakest in the
right and left posterior aspects. This means that herniations usually bulge to
the right or to the left, thereby affecting only one leg. If the nucleus pulposus
herniates straight back, however, it presses on the cauda equina instead of
just a single nerve root. This may cause numbness of the perineum and
dysfunction of bowel or urinary bladder control. This is known as cauda
equina syndrome and it requires immediate referral to a specialist.

1. Although several conditions of the lung can look like this on the radiograph,
this mans history favors one diagnosis. He is a lifetime smoker, he has
developed a chronic cough, and he has been losing weight. He most
assuredly has lung cancer.
2. The apex of the lung is in close proximity to the brachial plexus. This apical
tumor is either pressing on the trunk or chords of the brachial plexus, or it
has actually eroded through the pleura and is invading the plexus, causing
the arm symptoms.
3. The apex of the lung is also very close to the sympathetic chain in the upper
thorax and the neck. Tumor invasion in this location afflicts sympathetic
innervation to one-half the head. The result is loss of muscle tone in the eyelid,
loss of pupil dilation, and inability of the skin to sweat on that half of the face.

Back Matter
Clinical Practicum
Questions
The McGrawHill
Companies, 2001
Appendix B
1. The patients recent change in facial appearance and the change to his
hands and feet are due to excessive growth hormone. Growth hormone is
released by the pituitary gland. This man has developed a pituitary tumor
(seen on the MR image) that is secreting excess growth hormone, and which
will be confirmed by measuring the hormone in the lab.
Excess growth hormone during childhood greatly accelerates growth,
resulting in gigantism. However, if this occurs after a person has reached full
stature (i.e., after the epiphyseal plates of the long bones have already ossified),
the result is increased growth of other bones such as facial bones and increased
soft tissue growth of the hands and feet. This condition is termed acromegaly.
2. A MRI of the head is necessary to determine the size of the pituitary tumor.
3. The optic chiasma sits superior and just anterior to the pituitary gland. If a
pituitary tumor gets large enough it compresses the optic chiasma, interfering
with neuronal transmission through the optic fibers that cross in this
structure.

1. The fracture involves the floor of the orbit, which is mainly composed of the
maxillary bone. Small portions of the palatine and zygomatic bones make up
the remainder of the orbital floor. In this case, the fracture involves the
maxillary bone.
2. There are four paranasal sinuses: frontal, ethmoidal, sphenoidal, and
maxillary. The maxillary sinus lies just inferior to the orbit inside the
maxillary bone. The inferior wall of the orbit makes up the superior wall of
the sinus. Therefore, a fracture of the inferior wall of the orbit must also
involve the maxillary sinus.
3. As in this case, when fracture fragments of an inferior orbital wall fracture
are displaced into the maxillary sinus, herniation of the orbital contents into
the sinus is a common occurrence. In most cases, this involves only orbital
fat. However, in this case as well as others, the inferior rectus muscle may
also herniate into the sinus and become entrapped. Entrapment of the
inferior rectus muscle prevents movement of the eye.

1. A dissection flap. In a dissection, there is a defect in the tunica intima of the
vessel. This allows blood to enter into the potential space between the
tunica intima and tunica media. In a high-pressure vessel such as the aorta
this blood can dissect between these layers for some distance creating a true
and false lumen to the vessel. The dissection flap therefore represents the
tunica intima separating the true from the false lumina.
2. The descending aorta. Because this dissection involves only the descending
aorta the treatment would involve controlling the patients hypertension.
No surgical treatment is necessary. However, dissections of the ascending
aorta are surgical emergencies because they may extend into the coronary
arteries affecting the blood supply to the heart.
3. The dissection has extended into the left subclavian artery as it comes off
the aortic arch. The raising of the dissection flap narrows the lumen of the
artery. This decreases blood flow and in turn blood pressure in the left arm.
801

1. Although the patient has an underlying pulmonary infection causing some
shortness of breath, the acute worsening of his symptoms is due to a tension
pneumothorax. The tension pneumothorax occurred secondary to a
bronchopleural fistula resulting from the patients underlying infection. This
makes ventilation of the left lung very difficult. A tension pneumothorax is
a type of pneumothorax in which gas collects in the pleural space resulting
in increased intrathoracic pressure which affects both cardiac and pulmonary
function in a life-threatening manner.
2. As the left pleural space fills with air, it shifts the mediastinum to the
opposite side putting pressure on both the heart and the lung. This makes it
more difficult for the right lung to expand during inspiration and for the
heart to fill with blood. With each breath the patient takes, more air is
drawn into the left pleural space. The more the patient struggles for breath,
the more the pleural space fills with air.
3. Treatment of a tension pneumothorax is a medical emergency requiring
placement of a chest tube to relieve the increased intrathoracic pressure and
the reexpansion of the collapsed lung. If the patients clinical condition is
critical, decreasing the intrathoracic pressure becomes the priority and may
be accomplished through the placement of a needle through the chest wall.
A chest tube would then be placed when the patient stabilized.

1. No normal lung is identified in the left hemithorax. Multiple air-filled
tubular structures are seen in the left hemithorax consistent with loops of
bowel. These loops of bowel also exhibit some mass effect on the
mediastinum shifting it to the right.
2. A congenital diaphragmatic hernia allowed multiple loops of both large and
small bowel to herniate into the left hemithorax. This results in compression
of the lung causing the patients respiratory distress.
3. No, her breathing will improve, but it will take a significant amount of time
before it will become normal. The left lung has not developed normally
because it was not allowed to expand in utero. Normal development of the
lung requires expansion of the lung through inspiration of amniotic fluid by
the fetus. In the case of a large diaphragmatic hernia, bowel occupies the
space into which the lung would normally expand. By preventing this
expansion, the normal development of the lung has been retarded. Once the
hernia has been repaired and the bowel returned to the abdomen, the lung
may then expand and develop over time, with the infants normal
inspiration.

1. The blebs are called diverticulae and this condition is called diverticulosis.
Each colonic diverticulum is a herniation of mucosa through the muscularis
layer of the colon. Diverticulae tend to occur at the weakest parts of the
muscularis where nutrient vessels penetrate the muscle. They are more
numerous with advancing age and are thought to result from the high
pressures the colon generates when moving hard feces.
2. His abdominal pain was likely the result of diverticular inflammation
(diverticulitis). It is believed that bacteria and undigested food can clog a
diverticular sac, leading to infection. Symptoms can range from mild to
severe depending on the extent of infection.
3. Erosion of a diverticular vessel can produce massive, painless, rectal
bleeding.

1. McBurneys point is a point on a line connecting the right anterior superior
iliac spine and the umbilicus. The point lies approximately one-third of the
way from the anterior superior iliac spine to the umbilicus. This point on the
skin overlies the expected location of the appendix. In acute appendicitis
pain is said to eventually localize to McBurneys point. This is also a
landmark for surgeons where the incision for an appendectomy is made.
APPENDIX B
1. The noise is called a bruit (pronounced broo-ee). It is caused by turbulent

blood flow through a narrowed artery, and it is analogous to the noise of
water flowing through a kinked garden hose. The arteriogram shows stenosis
of the left renal artery, which is likely the source of the abdominal bruit.
2. The stenosis of the renal artery is, in effect, starving the left kidney of blood.
The kidney interprets this decrease in blood supply as a sign that the body is
low on blood (such as after massive hemorrhage). It subsequently activates a
hormone system called the renin-angiotensin-aldosterone system whose
actions increase blood pressure by direct vasoconstriction and by retention of
salt and water. This mechanism is life-saving if the bodys blood volume is
truly low, but it is detrimental if triggered by renal artery stenosis.
3. Renal retention of salt and water is accomplished at the expense of
potassium loss in the urine.

802
Appendix B
Back Matter
Clinical Practicum
Questions
2. This CT is consistent with the diagnosis of acute appendicitis. The appendix

is dilated with a thickened wall. The periappendiceal fat is infiltrated and
edematous. These findings are consistent with inflammation of the
appendix. An appendicolith is also seen near the orifice of the appendix.
3. This white object is an appendicolith. It is a calcified fragment of fecal
matter lodged within the lumen of the appendix. By lodging with and
obstructing the appendix, the appendicolith leads to the initial
inflammation and infection causing acute appendicitis.
4. Appropriate treatment is surgery to remove the appendix.

1. There is a large filling defect in the urinary bladder seen on the intravenous
urogram. Normally the mucosa of a distended urinary bladder has a mucosa
of smooth appearance. In this case there is a large irregularity demonstrated
in the urinary bladder. This most likely represents a transitional cell
carcinoma of the bladder, a malignant neoplasm of the urothelium.
Considering the patients history of hematuria, this is the most likely source
of his bleeding.
2. The CT scan shows a mass filling a significant portion of the lumen of the
urinary bladder. This greatly decreases the functional volume of the bladder.
Therefore a smaller amount of urine will fill the urinary bladder. This leads
to stimulation of the stretch receptors of the bladder, leading to the urge to
urinate.
3. Transitional epithelium lines the renal calyces, renal pelvis, ureters, urinary
bladder, and urethra. Any of these structures may be involved by
synchronous tumor. Continued surveillance of all these organs must also be
performed to evaluate for recurrence of cancer.

1. The identified object has the same density as bone, meaning that it contains
a significant amount of calcium. This density lies within the expected course
of the ureter. These findings are most consistent with a stone within the left
ureter. Either an intravenous urogram or a spiral CT may confirm this
diagnosis.
2. The intravenous urogram demonstrates that there is a stone within the upper
third of the left ureter. This stone is large enough to cause partial obstruction
of the ureter, leading to dilation of the collecting system. Dilation of the
collecting system causes increased pressures within the kidney. This in turn
decreases renal function. If these changes are long-standing, permanent
damage to the kidney occurs.
3. The ureter has a structure similar to many other hollow organs in the body.
Deep to the mucosa there is a layer of smooth muscle which is surrounded by
a serosa. This smooth muscle layer undergoes periodic peristaltic
contractions to propel the urine from the kidney to the urinary bladder. In
the setting of a stone within the ureter these contractions become painful as
the smooth muscle exerts pressure on the stone. Because these peristaltic
waves are only intermittent the pain is intermittent.

1. The testicular self-exam for males is, in a way, equivalent to the breast selfexam for females. It is a screening test for testicular cancer, the most
common malignancy among young adult males. It involves examining both
testes on a monthly basis to check for changes in contour, nodules, or areas
of tenderness. If performed properly and at timely intervals this exam may
result in earlier detection of testicular neoplasms. By detecting these cancers
at earlier stages it is hoped that they may be more effectively treated.
APPENDIX B
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2. The venous drainage of the left testis is via the left renal vein. Abnormal
lymph nodes could be found along the course of this vein in the
retroperitoneal space. Around the left renal vein and in the periaortic region
are also common areas to find abnormal lymph nodes. It would be unusual to
find abnormal nodes isolated to the inguinal area because the inguinal nodes
are not within the direct lymphatic drainage of the scrotum.
3. Ultrasound is an imaging modality which uses the reflection of sound waves
from tissue interfaces to create representative images of the anatomy. No
radiation is used during an ultrasound examination. This is of particular
importance not only in imaging of the testis but also in imaging of the ovary.
Because of the high meiotic rate in these organs, the DNA in these tissues is
particularly susceptible to radiation-induced damage which can potentially
lead to genetic mutation. Therefore ultrasound, a modality that uses no
radiation is safer than radiographic evaluation.

1. These foci of endometrial tissue are susceptible to hormonal influences just
like normal endometrial tissue. During the time of menstruation, these
tissues bleed. Recurrent bleeding in the pelvis leads to scarring which can
distort the pelvic anatomy causing pain and infertility.
2. Several theories exist as to how endometrial tissue is found outside the
uterus in the disease endometriosis. Some aspect of each of these theories is
likely true. Direct spread of endometrial tissue from the uterine cavity back
out through the uterine tubes is the most likely way that ectopic endometrial
tissue forms in the pelvis. Spread of endometrial tissue through vascular and
lymphatic structures can also occur.
3. a. Pain with defecation
b. Pain with urination
c. Postprandial pain
d. Pain with breathing

1. Considering the physical exam findings and the location of this mass on the
CT, it most likely originates from the ovary. Its heterogeneous appearance
on CT tells us that many different types of tissue are found within the mass.
This mass therefore, most likely represents a dermoid cyst.
2. This benign mass arises from the germ cells of the ovary. Because this mass
arises from germ cells it has the potential to form all types of tissue. It is very
common to find teeth, hair, and fat within a dermoid cyst. In this case two
teeth and a significant amount of fat can be identified.
3. The vast majority of dermoid cysts are benign, however a very small
percentage will undergo malignant degeneration. The greater concern with
such a benign mass is that with increasing size there is increased risk of
ovarian torsion. In this condition the ovary twists, causing occlusion of the
ovarian vessels, which may lead to ischemia. When a dermoid cyst becomes
larger than 4 cm the risk of ovarian torsion becomes great enough that
surgical removal is warranted.

Back Matter
Glossary
The McGrawHill
Companies, 2001
Glossary
KEY TO PRONUNCIATION
Most of the words in this glossary are followed by a phonetic spelling that serves as a guide to
pronunciation. The phonetic spellings reflect standard scientific usage and can easily be interpreted following a few basic rules.
1. Any unmarked vowel that ends a syllable or that stands alone as a syllable has the long
sound. For example, ba, ma, and na rhyme with fay; be, de, and we rhyme with fee; bi, di, and
pi rhyme with sigh; bo, do, and mo rhyme with go. Any unmarked vowel that is followed by a
consonant has the short sound (for example, the vowel sounds in hat, met, pit, not, and but).
2. If a long vowel appears in the middle of a syllable (followed by a consonant), it is
marked with a macron () Similarly, if a vowel stands alone or ends a syllable but should
have short sound, it is marked with a breve ().
3. Syllables that are emphasized are indicated by stress marks. A single stress mark (') indicates the primary emphasis; a secondary emphasis is indicated by a double stress mark ('').
A
abdomen (ab'do-men or ab-do'men) The
region of the trunk between the diaphragm
and pelvis.
abduction (ab-duk'shun) Movement of a
body part away from the axis or midline of
the body; movement of a digit away from the
axis of the limb; the opposite of adduction.
accessory organs Organs that assist the
functioning of other organs within a system.
acetabulum (as''e-tab'yu-lum) The cupshaped depression on the lateral surface of
the hipbone (os coxae) with which the head
of the femur articulates.
Achilles (ekil'ez) tendon See tendo
calcaneus.
actin (ak'tin) A protein in muscle fibers that,
together with myosin, is responsible for
contraction.
adduction (a-duk'shun) Movement of a body
part toward the axis or midline of the body;
movement of a digit toward the axis of the
limb; the opposite of abduction.
adipocyte (ad'-po-st) A fat cell found within
adipose tissue.
adrenal cortex (a-dre'nal kor'teks) The outer
part of the adrenal gland that secretes steroid
hormones.
adrenal glands Two small endocrine glands,
one located above each kidney; also called
the suprarenal glands.
adrenal medulla (me-dul'a) The inner part of
the adrenal gland that secretes the hormones
epinephrine and norepinephrine.
adventitia (ad'ven-tish'a) The outermost

epithelial layer of a visceral organ; also called
serosa.
afferent (af'er-ent) Conveying or
transmitting to.
afferent glomerular arteriole (ar-te're-ol) A
vessel within the kidney that supplies blood
to the glomerulus.
afferent neuron (noor'on) A sensory nerve
cell that transmits an impulse toward the
central nervous system.
agonist (ag'o-nist) The prime mover muscle,
which is directly engaged in the contraction
that produces a desired movement.
alimentary (al'-men'tre) canal The tubular
portion of the digestive tract; also called the
gastrointestinal (GI) tract.
allantois (a-lan'to-is) An extraembryonic
membranous sac involved in the formation of
blood cells. It gives rise to the fetal umbilical
arteries and vein, and also contributes to the
formation of the urinary bladder.
all-or-none principle The statement that
muscle fibers of a motor unit contract to their
maximum extent when exposed to a stimulus
of threshold strength.
alveolar (al-ve'o-lar) sacs A cluster of alveoli
that share a common chamber or central
atrium.
alveolus (al-ve'o-lus) 1. A pulmonary alveolus
is an individual air capsule within the lung.
The pulmonary alveoli are the basic
functional units of respiration. 2. A dental
alveolus is a socket that secures a tooth (a
tooth socket).
amniocentesis (am''ne-o-sen-te'sis) A
procedure in which a sample of amniotic
fluid is aspirated to examine cells for various
genetic diseases.
amnion (am'ne-on) The innermost fetal
membranea thin sac that holds the fetus
suspended in amniotic fluid; commonly
known as the bag of waters.
ampulla (am-pool'a) A saclike enlargement of
a duct or tube.
ampulla of Vater (fa'ter) See
hepatopancreatic ampulla.
anal (a'nal) canal The terminal tubular
portion of the large intestine that opens
through the anus of the GI tract.
anal glands Enlarged and modified sweat
glands that empty into the anal opening.
anastomosis (a-nas''to-mo'sis) An anatomical
convergence of blood vessels or nerves that
forms a network.
anatomical position An erect body stance with
the eyes directed forward, the arms at the
sides, the palms of the hands facing forward,
and the fingers pointing straight down.
anatomy The branch of science concerned
with the structure of the body and the
relationship of its parts.
antagonist (an-tag'o-nist) 1. A muscle that
acts in opposition to another muscle. 2. Any
agent, such as a hormone or drug, that
opposes the action of another.
antebrachium (an''te-bra'ke-um) The
forearm.
anterior (ventral) Toward the front; the
opposite of posterior (dorsal).
anterior root The anterior projection of the
spinal cord composed of axons of motor, or
efferent, fibers.
anus (a'nus) The terminal opening of the GI
tract.
aorta (a-or'ta) The major systemic vessel of
the arterial system of the body, emerging
from the left ventricle.
aortic arch The superior left bend of the aorta
between the ascending and descending
portions; also called the arch of aorta.
apex (a'peks) The tip or pointed end of a
conical structure.
apocrine (ap'o-krin) gland A type of sweat
gland that functions in evaporative cooling.
It may respond during periods of emotional
stress.
803

804
Back Matter
Glossary
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Glossary
aponeurosis (ap''o-noo-ro'sis) A fibrous or

membranous sheetlike tendon.
appendix (a-pen'diks) A short pouch that
attaches to the cecum; also called the
vermiform appendix.
aqueous (a'kwe-us) humor The watery fluid
that fills the anterior and posterior chambers
of the eye.
arachnoid (a-rak'noid) The weblike middle
covering (meninx) of the central nervous
system.
arbor vitae (ar'bor vi'te) The branching
arrangement of white matter within the
cerebellum.
arm (brachium) That portion of the upper
extremity extending from the shoulder to the
elbow.
arrector pili (a-rek'tor pi'li); plural,
arrectores pilorum A bundle of smooth
muscle cells attached to a hair follicle that,
upon contraction, pulls the hair into a more
vertical position, resulting in goose bumps.
arteriole (ar-te're-o l) A minute arterial
branch.
artery (ar'te-re) A blood vessel that carries
blood away from the heart.
arthrology (ar-throl'o-je) The scientific study
of the structure and function of joints.
articular cartilage (ar-tik'yu-lar kar't-lij) A
hyaline cartilaginous covering over the
articulating surface of bones of synovial
joints.
articulation (ar-tik'yu-la'shun) A joint.
arytenoid (ar''e-te'noid) cartilages A pair of
small cartilages on the superior aspect of the
larynx.
ascending colon (ko'lon) That portion of the
large intestine between the cecum and the
hepatic (right colic) flexure.
association neuron (noor'on) A nerve cell
located completely within the central
nervous system. It conveys impulses in an arc
from sensory to motor neurons; also called an
internuncial neuron or interneuron.
atom The smallest particle of matter that
characterizes an element.
atrioventricular (a''tre-o-ven-trik'yu-lar)
bundle A group of specialized cardiac fibers
that conducts impulses from the
atrioventricular node to the ventricular
muscles of the heart; also called the bundle of
His or AV bundle.
atrioventricular node A microscopic
aggregation of specialized cardiac fibers
located in the interatrial septum of the heart;
a component of the conduction system of the
heart; also called the AV node.
atrioventricular valve A cardiac valve located
between an atrium and a ventricle of the
heart; also called an AV valve.
atrium (a-tre'um) Either of the two superior

chambers of the heart that receive venous
blood.
atrophy (at'ro-fe) A gradual wasting away or
decrease in the size of a tissue or organ.
auditory Pertaining to the structures of the ear
associated with hearing.
auditory tube A narrow canal that connects
the middle-ear chamber to the pharynx; also
called the eustachian canal.
auricle (or'-kul) 1. The fleshy pinna of the
ear. 2. An ear-shaped appendage of each
atrium of the heart.
autonomic nervous system (ANS) The
sympathetic and parasympathetic portions of
the nervous system that innervate cardiac
muscle, smooth (visceral) muscles, and
glands. The ANS functions largely without
conscious control.
axilla (ak-sil'a) The depressed hollow
commonly called the armpit.
axon (ak'son) The elongated process of a
nerve cell that transmits an impulse away
from the cell body of a neuron.
B
ball-and-socket joint The most freely movable
of all the synovial joints (e.g., the shoulder
and hip joints).
baroreceptor (bar''o-re-sep'tor) A cluster of
neuroreceptors stimulated by pressure
changes. Baroreceptors monitor blood
pressure.
basal ganglion (gang'gle-on) A mass of nerve
cell bodies located deep within a cerebral
hemisphere of the brain.
basement membrane A thin sheet of
extracellular substance to which the basal
surfaces of membranous epithelial cells are
attached; also called the basal lamina.
basophil (ba'so-fil) A granular leukocyte that
readily stains with basophilic dye.
belly The bulging, central part of a skeletal
muscle; also called gaster.
benign (be-nn) Not malignant.
bicuspid (bi-kus'pid) valve The heart valve
located between the left atrium and the left
ventricle; the left AV valve; also called the
mitral valve.
bifurcation (bi''fur-ka'shun) Forked; divided
into two branches.
bile (bl) A liver secretion that is stored and
concentrated in the gallbladder and released
through the common bile duct into the
duodenum. It is essential for the absorption
of fats.
bipennate (bi-pen'at) Denoting muscles

whose fibers are arranged on each side of a
tendon like barbs on a feather shaft.
bipolar neuron (noor'on) A nerve cell with
two processes, one at each end of the cell
body.
blastocyst (blas'to-sist) An early stage of
embryonic development consisting of a
hollow ball of cells with an embryoblast
(inner cell mass) and an outer layer called
the trophoblast.
blood The fluid connective tissue that
circulates through the cardiovascular system
to transport substances throughout the body.
blood-brain barrier A specialized mechanism
that inhibits the passage of certain materials
from the blood into brain tissue and
bolus (bo'lus) A moistened mass of food that
is swallowed from the oral cavity into the
pharynx.
bone 1. The hard, calcified connective tissue
forming the major portion of the skeleton.
2. Any of the more than 200 anatomically
distinct structures making up the skeleton.
bony labyrinth (lab'-rinth) A series of
chambers within the petrous part of the
temporal bone associated with the vestibular
organs and the cochlea.
Bowman's (bo'manz) capsule See glomerular
capsule.
brachial plexus (bra'ke-al plek'sus) A
network of nerve fibers arising from spinal
nerves C5C8 and T1. Nerves arising from
the brachial plexus supply the upper
extremity.
brain The enlarged superior portion of the
central nervous system located in the cranial
cavity of the skull.
brain stem That portion of the brain
consisting of the midbrain, pons, and medulla
oblongata.
bronchial (brong'ke-al) tree The bronchi and
their branching bronchioles.
bronchiole (brong'ke-ol) A small division of a
bronchus within the lung.
bronchus (brong'kus) A branch of the
trachea that leads to a lung.
bulbourethral (bul''bo-yoo-re'thral) glands A
pair of glands that secrete a viscous fluid into
the male urethra during sexual excitement;
also called Cowpers glands.
bundle of His (hiss) See atrioventricular
bundle.
bursa (bur'sa) A saclike cavity filled with
synovial fluid. Bursae are located between
tendons and bones or at points of friction
between moving structures.

Back Matter
Glossary
The McGrawHill
Companies, 2001
Glossary
buttock (but'ok) The fleshy mass on the
posterior aspect of the lower trunk, formed
primarily by the gluteal muscles; the rump.
C
calyx (ka'liks) The cup-shaped portion of the
renal pelvis that encircles a renal papilla.
canaliculus (kan''a-lik'yu-lus) A microscopic
channel in bone tissue that connects
lacunae.
canal of Schlemm (shlem) See scleral venous
sinus.
cancellous (kan'se-lus) bone Spongy bone;
bone tissue with a latticelike structure.
capillary (kap'-lar''e) A microscopic blood
vessel that connects an arteriole and a
venule; the functional unit of the circulatory
system.
carbohydrate Any of the group of organic
molecules composed of carbon, hydrogen,
and oxygen, including sugars and starches. A
carbohydrate usually has the formula CH2O.
carotid (ka-rot'id) sinus An expanded
portion of the internal carotid artery located
immediately above the point of branching
from the external carotid artery. The carotid
sinus contains baroreceptors that monitor
blood pressure.
carpus (kar'pus) The proximal portion of the
hand that contains the eight carpal bones.
cartilage (kar't-lij) A type of connective
tissue with a solid elastic matrix.
cartilaginous (kar''t-laj'-nus) A joint that
lacks a joint cavity, permitting little
movement between the bones held together
by cartilage.
cauda equina (kaw'da e-kwi'na) The
extension from the terminal portion of the
spinal cord, where the roots of spinal nerves
resemble a horses tail.
caudal (kaw'dal) Referring to a position more
toward the tail.
cecum (se'kum) The pouchlike portion of the
large intestine to which the ileum of the
small intestine is attached.
cell The structural and functional unit of an
organism; the smallest structure capable of
performing all of the functions necessary for
life.
cementum (se-men'tum) Bonelike material
that binds the root of a tooth to the
periodontal membrane of the tooth socket
(dental alveolus).
central canal The elongated longitudinal
channel at the center of an osteon in bone
tissue that contains branches of the nutrient
vessels and a nerve; also called a haversian
canal.
central nervous system (CNS) The brain and

the spinal cord.
centrosome (sen'tro-som) A specialized zone
of cytoplasm near the nucleus of a cell that
contains a pair of centrioles.
cerebellar peduncle (ser''e-bel'ar pedung'kul) An aggregation of nerve fibers
connecting the cerebellum with the brain
stem.
cerebellum (ser'' e-bel'um) That portion of
the brain concerned with the coordination of
skeletal muscle contraction. Part of the
metencephalon, it consists of two
hemispheres and a central vermis.
cerebral aqueduct (ser'e-bral ak'we-dukt) See
mesencephalic aqueduct.
cerebral arterial (ar-te're-al) circle An
arterial vessel on the inferior surface of the
brain that encircles the pituitary gland. It
provides alternate routes for blood to reach
the brain should a carotid or vertebral artery
become occluded; also called the circle of
Willis.
cerebral peduncles (pe-dung'kulz) A paired
bundle of nerve fibers along the inferior
surface of the midbrain that conduct impulses
between the pons and the cerebral
hemispheres.
cerebrospinal (ser''e-bro-spi'nal) fluid
(CSF) A fluid produced by the choroid
plexus of the ventricles of the brain. It fills the
ventricles and surrounds the central nervous
system in association with the meninges.
cerebrum (ser'e-brum or se-re'brum) The
largest portion of the brain, composed of
right and left cerebral hemispheres.
ceruminous (se-roo'm-nus) gland A
specialized integumentary gland that secretes
cerumen, or earwax, into the external
acoustic canal.
cervical (ser'v-kal) Pertaining to the neck or
a necklike portion of an organ.
cervical ganglion (gang'gle-on) A cluster of
postganglionic sympathetic nerve cell bodies
located in the neck, near the cervical
vertebrae.
cervical plexus (plek'sus) A network of
spinal nerves formed by the anterior branches
of the first four cervical nerves.
cervix (ser'viks) 1. The narrow necklike
portion of an organ. 2. The inferior end of
the uterus that adjoins the vagina (cervix of
the uterus).
chemoreceptor (ke''mo-re-sep'tor) A
neuroreceptor that is stimulated by the
presence of a chemical in solution.
chiasma (ki-as'ma) A crossing of nerve tracts
from one side of the CNS to the other; also
called a chiasm.
805
choanae (ko-a'ne) The two posterior

openings from the nasal cavity into the
nasopharynx; also called the internal nares.
chondrocranium (kon''dro-kra'ne-um) That
portion of the skull that supports the brain. It
is derived from endochondral bone.
chondrocyte (kon'dro-st) A cartilage cell.
chordae tendineae (kor'de ten-din'e-e)
Tendinous bands that connect papillary
muscles to the cusps of the atrioventricular
valves within the ventricles of the heart.
chorion (ko're-on) An extraembryonic
membrane that contributes to the formation
of the placenta.
choroid (kor'oid) The vascular, pigmented
middle layer of the wall of the eye.
choroid plexus A mass of vascular capillaries
from which cerebrospinal fluid is secreted
into the ventricles of the brain.
chromatophilic (kro''ma-to-fil'ik)
substances Clumps of rough endoplasmic
reticulum in the cell body of a neuron; also
called Nissl bodies.
chromosomes (kro'mo-somz) Structures in
the nucleus that contain the genes for
genetic expression.
chyme (km) The mass of partially digested
food that passes from the pylorus of the
stomach into the duodenum of the small
intestine.
cilia (sil'e-a) Microscopic hairlike processes
that move in a wavelike manner on the
exposed surfaces of certain epithelial cells.
ciliary (sil'e-er''e) body A portion of the
choroid layer of the eye that secretes aqueous
humor. It contains the ciliary muscle that,
when contracted, changes the shape of the
lens.
circle of Willis See cerebral arterial circle.
circumduction (ser''kum-duk'shun) Circular
movement of a body part in which a coneshaped airspace is traced.
cleavage Rapid cell divisions of the zygote that
occur approximately 30 hours after
fertilization of the secondary oocyte.
clitoris (klit'or-is or kli'tor-is) A small
erectile structure in the vulva of the female,
homologous to the glans penis in the male.
coccygeal (kok-sij'e-al) Pertaining to the
region of the coccyx; the caudal termination
of the vertebral column.
cochlea (kok'le-a) The spiral portion of the
inner ear that contains the spiral organ
(organ of Corti).
cochlear window A round, membranecovered opening between the middle and
inner ear, directly below the vestibular
window; also called the round window.

806
Back Matter
Glossary
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Glossary
coelom (se'lom) The abdominal cavity.

collateral A small branch of a blood vessel or
nerve fiber.
colon (ko'lon) The first portion of the large
intestine.
common bile duct A tube formed by the
union of the hepatic duct and cystic duct
that transports bile to the duodenum.
compact (dense) bone Tightly packed bone,
superficial to spongy bone and covered by the
periosteum.
conceptus (kon-sep'tus) The product of
conception at any point between fertilization
and birth. It includes the embryo or fetus, as
well as the extraembryonic structures.
conduction myofibers Specialized cardiac
muscle fibers that conduct electrical impulses
from the AV bundle into the ventricular
walls; also called Purkinje fibers.
condyle (kon'dl) A rounded prominence at
the end of a long bone, most often for
articulation with another bone.
cone A type of photoreceptor cell in the
retina of the eye that provides for color
vision.
congenital (kon-jen'-tal) Present at the time
of birth.
conjunctiva (con''jungk-ti'va) The thin
membrane covering the anterior surface of
the eyeball and lining the eyelids.
connective tissue One of the four basic tissue
types within the body. It is a binding and
supporting tissue with abundant matrix.
conus medullaris (ko'nus med''yu-lar'is) The
caudal tapering portion of the spinal cord.
convolution (con-vo-loo'shun) 1. An
elevation on the surface of a structure and an
infolding of the tissue upon itself. 2. One of
the convex folds on the surface of the brain.
cornea (kor'ne-a) The transparent convex
anterior portion of the outer layer of the
eyeball.
cornification (kor''n-f-ka'shun) The drying
and flattening of the outer keratinized cells of
the epidermis.
coronal (kor'o-nal or ko-ro'nal) plane A
plane that divides the body into anterior and
posterior portions; also called a frontal plane.
coronary (kor'o-nar''e) circulation The flow
of blood in the arteries and veins in the wall
of the heart; the functional blood supply of
the heart.
coronary sinus A large venous channel on
the posterior surface of the heart into which
the cardiac veins drain.
corpora quadrigemina (kor'por-a kwad''rjem'-na) Four superior lobes of the
midbrain concerned with visual and auditory
functions.
corpus callosum (kor'pus ka-lo'sum) A large

tract of white matter within the brain that
connects the right and left cerebral
hemispheres.
cortex (kor'teks) 1. The outer layer of an
internal organ or body structure, as of the
kidney or adrenal gland. 2. The convoluted
layer of gray matter that covers the surface of
each cerebral hemisphere.
costal (kos'tal) cartilage The cartilage that
connects the ribs to the sternum.
cranial (kra'ne-al) Pertaining to the cranium.
cranial nerve One of 12 pairs of nerves that
arise from the inferior surface of the brain.
cranium (kra'ne-um) The endochondral
bones of the skull that enclose or support the
brain and the organs of sight, hearing, and
balance.
crest A thickened ridge of bone for the
attachment of muscle.
cricoid (kri'koid) cartilage A ring-shaped
cartilage that forms the inferior part of the
larynx.
crista (kris'ta) A crest, such as the crista galli
that extends superiorly from the cribriform
plate.
cubital (kyoo'b-tal) Pertaining to the elbow.
The cubital fossa is the anterior aspect of the
elbow.
cystic (sis'tik) duct The tube that transports
bile from the gallbladder to the common bile
duct.
cytology (si-tol'o-je) The science dealing with
the study of cells.
cytoplasm (si'to-plaz'em) In a cell, the
material between the nucleus and the cell
(plasma) membrane.
D
deciduous (de-sij'oo-us) Pertaining to
something cast off or shed in a particular
sequence. Deciduous teeth are shed and
replaced by permanent teeth in a predictable
sequence.
decussation (de''kus-a'shun) A crossing of
nerve fibers from one side of the CNS to the
other.
defecation (def''e-ka'shun) The elimination
of feces from the rectum through the anal
canal and out the anus.
deglutition (de''gloo-tish'un) The act of
swallowing.
dendrite (den'drt) A nerve cell process that
transmits impulses toward the cell body of a
neuron.
dentin (den'tin) The main substance of a

tooth, covered by enamel over the crown of
the tooth and by cementum on the root.
dentition (den-tish-un) The type, number,
and arrangement of a set of teeth.
dermal papilla (pa-pil'a) A projection of the
dermis into the epidermis.
dermis (der'mis) The second, or deep, layer of
skin beneath the epidermis.
descending colon That segment of the large
intestine that descends on the left side from
the level of the spleen to the level of the left
iliac crest.
diaphragm (di'a-fram) A sheetlike dome of
muscle and connective tissue that separates
the thoracic and abdominal cavities.
diaphysis (di-af'-sis) The body (shaft) of a
long bone.
diastole (di-as'to-le) The sequence of the
cardiac cycle in which a heart chamber wall
relaxes and the chamber fills with blood;
especially ventricular relaxation.
diencephalon (di''en-sef'a-lon) That part of
the brain lying between the telencephalon
and the mesencephalon. It includes the third
ventricle, pineal gland, epithalamus,
thalamus, hypothalamus, and pituitary gland.
digestion The process by which food is broken
down mechanically and chemically into
simple molecules that can be absorbed and
used by body cells.
diploe (dip'lo-e) The spongy layer of bone
positioned between the inner and outer
layers of compact bone.
distal (dis'tal) Away from the midline or
origin; the opposite of proximal.
dorsal (dor'sal) Pertaining to the back, or
posterior, portion of a body part; the opposite
of ventral.
dorsiflexion (dor''s-flek'shun) Movement at
the ankle or wrist as the dorsum of the foot or
hand is elevated.
ductus arteriosus (duk'tus ar-te''re-o'sus)
The blood vessel that connects the
pulmonary trunk and the aorta in a fetus.
ductus deferens (def''er-enz); plural, ductus
deferentia A tube that carries spermatozoa
from the epididymis to the ejaculatory duct;
also called the vas deferens or seminal duct.
ductus venosus (ven-o'sus) A fetal blood
vessel that connects the umbilical vein and
the inferior vena cava.
duodenum (doo''o-de'num or doo-od'e-num)
The first portion of the small intestine that
leads from the pylorus of the stomach to the
jejunum.
dura mater (door'a ma'ter) The outermost
meninx.

Back Matter
Glossary
The McGrawHill
Companies, 2001
Glossary
E
eccrine (ek'rin) gland Any of the numerous
sweat glands distributed over the body that
function in thermoregulation.
ECG Electrocardiogram; EKG.
ectoderm (ek'to-derm) The outermost of the
three primary germ layers of an embryo.
edema (e-de'ma) Abnormal accumulation of
interstitial (tissue) fluid, causing the tissue to
swell.
effector (e-fek'tor) An organ, such as a gland
or muscle, capable of responding to a
stimulus.
efferent (ef'er-ent) Conveying away from the
center of an organ or structure.
efferent ductules (duk'toolz) A series of
coiled tubules that convey spermatozoa from
the rete testis to the epididymis.
efferent glomerular arteriole (ar-te're-ol) A
vessel within the kidney that conducts blood
away from the glomerulus to the peritubular
capillaries.
efferent neuron (noor'on) A motor nerve cell
that conducts impulses from the central
nervous system to effector organs, such as
muscles or glands.
ejaculation (e-jak''yoo-la'shun) The
discharge of semen from the male urethra
during climax.
ejaculatory (e-jak'yoo-la-tor-e) duct A tube
that transports spermatozoa from the ductus
deferens to the prostatic part of the urethra.
elastic fibers Protein strands found in certain
connective tissue that have contractile
properties.
elbow The synovial joint between the
brachium and the antebrachium.
electrocardiogram (e-lek''tro-kar'de-ogram) A recording of the electrical activity
that accompanies the cardiac cycle; ECG or
EKG.
electroencephalogram (e-lek''tro-en-sef'a-logram) A recording of the brain-wave
patterns or electrical impulses of the brain;
EEG.
electromyogram (e-lek''tro-mi'o-gram) A
recording of the electrical impulses or
activity of a muscle; EMG.
embryology (em''bre-ol'o-je) The study of
prenatal development from conception
through the eighth week in utero.
enamel (e-nam'el) The outer dense substance
covering the crown of a tooth.
endocardium (en''do-kar'de-um) The
endothelial lining of the heart chambers and
valves.
endochondral (en''do-kon'dral) bone Bones
that develop as hyaline cartilage models first,
and then are ossified.
endocrine (en'do-krin) gland A ductless,

hormone-producing gland of the endocrine
system.
endoderm (en'do-derm) The innermost of the
three primary germ layers of a developing
embryo.
endolymph (en'do-limf) A fluid within the
cochlear duct and membranous labyrinth of
the inner ear that aids in the conduction of
vibrations involved in hearing and
maintaining equilibrium.
endometrium (en''do-me'tre-um) The inner
lining of the uterus.
endomysium (en-do-mis'e-um) The
connective tissue sheath surrounding
individual skeletal muscle fibers, separating
them from one another.
endoneurium (en''do-nyoo're-um) The
connective tissue sheath surrounding
individual nerve fibers, separating them from
one another within a peripheral nerve.
endoplasmic reticulum (en''do-plaz'mik retik'yoo-lum) (ER) A system of
interconnected tubules or channels running
through the cytoplasm of a cell. Rough ER
bears ribosomes on its membrane, while
smooth ER does not.
endothelium (en''do-the'le-um) The layer of
epithelial tissue that forms the thin inner
lining of blood vessels and heart chambers.
eosinophil (e''o-sin'o-fil) A type of white
blood cell characterized by the presence of
cytoplasmic granules that become stained by
acidic eosin dye. Eosinophils normally
constitute from 2% to 4% of the white blood
cells.
epicardium (ep''-kar'de-um) The thin outer
layer of the heart; also called the visceral
pericardium.
epicondyle (ep''-kon'dl) A projection of
bone above a condyle.
epidermis (ep'' -der'mis) The outermost layer
of the skin, composed of stratified squamous
epithelium.
epididymis (ep''-did'i-mis); plural,
epididymides (ep''-d-dm'dez) A highly
coiled tube located along the posterior border
of the testis. It stores spermatozoa and
transports them from the seminiferous
tubules of the testis to the ductus deferens.
epidural (ep''-door'al) space A space
between the spinal dura mater and the bone
of the vertebral canal.
epiglottis (ep''-glot'is) A leaflike structure
positioned on top of the larynx. It covers the
glottis during swallowing.
epimysium (ep''-mis'e-um) The fibrous outer
sheath of connective tissue surrounding a
skeletal muscle.
807
epinephrine (ep''-nef'rin) A hormone

secreted from the adrenal medulla whose
actions are similar to those initiated by
sympathetic nervous system stimulation; also
called adrenaline.
epineurium (ep''-nyoo're-um) The fibrous
outer sheath of connective tissue surrounding
a nerve.
epiphyseal (ep''-fiz'e-al) plate A layer of
hyaline cartilage between the epiphysis and
diaphysis of a long bone. It is responsible for
the lengthwise growth of long bones.
epiphysis (e-pif'-sis) The end segment of a
long bone, separated from the diaphysis early
in life by an epiphyseal plate, but later
becoming part of the larger bone.
episiotomy (e-pe''ze-ot'o-me) An incision of
the perineum at the end of the second stage
of labor to facilitate delivery and avoid
tearing of the perineum.
epithelial (ep''-the'le-al) tissue One of the
four basic tissue types; the type of tissue that
covers or lines all exposed body surfaces; also
called epithelium.
eponychium (ep''o-nik'e-um) The thin layer
of stratum corneum of the epidermis of the
skin that overlaps and protects the lunula of
the nail.
erythrocyte (e-rith'ro-st) A red blood cell.
esophagus (e-sof'a-gus) The tubular organ of
the GI tract that leads from the pharynx to
the stomach as it passes through the thoracic
cavity.
estrogen (es'trojen) Any of several female sex
hormones secreted from the ovarian
(graafian) follicle.
etiology (e''te-ol'o-je) The study of cause,
especially of disease, including the origin and
what pathogens, if any, are involved.
eustachian (yoo-sta'shun) canal See auditory
tube.
eversion (e-ver'shun) Movement of the foot
in which the sole is turned outward; the
opposite of inversion.
exocrine (ek'so-krin) gland A gland that
secretes its product to an epithelial surface,
either directly or through ducts.
expiration The process of expelling air from
the lungs through breathing out; also called
exhalation.
extension Movement that increases the angle
between parts of a joint; the opposite of
flexion.
external (superficial) Located on or toward
the surface.
external acoustic meatus (me-a'tus) An
opening through the temporal bone that
connects with the tympanum and the

808
Back Matter
Glossary
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Glossary
middle-ear chamber and through which

sound vibrations pass; also called the external
auditory meatus.
external ear The outer portion of the ear,
consisting of the auricle (pinna) and the
external acoustic canal.
exteroceptor (ek''ster-o-sep'tor) A
specialized sensory neuron located near the
surface of the body that responds to stimuli
from the external environment.
extraocular (ek''stra-ok'yu-lar) muscles The
muscles that insert onto the sclera of the eye
and act to change the position of the eye in
its orbit (as opposed to the intraocular
muscles, such as those of the iris and ciliary
body within the eye); also called extrinsic
ocular muscles.
extrinsic (eks-trin'sik) Pertaining to an
outside or external origin.
F
face 1. The anterior aspect of the head not
supporting or covering the brain. 2. The
exposed surface of a structure.
facet (fas'et) A flattened, shallow articulating
surface on a bone.
falciform (fal's-form) ligament The
extension of parietal peritoneum that
separates the right and left lobes of the liver.
fallopian (fa-lo'pe-an) tube See uterine tube.
false vocal cords See vestibular folds.
falx cerebelli (falks ser''e-bel'i) A fold of
dura mater anchored to the occipital bone. It
projects inward between the cerebellar
hemispheres.
falx cerebri (ser'e-bri) A fold of dura mater
anchored to the crista galli. It extends
between the right and left cerebral
hemispheres.
fascia (fash'e-a) A tough sheet or band of
fibrous connective tissue binding the skin to
underlying muscles (superficial fascia) or
supporting and separating muscles (deep
fascia).
fasciculus (fa-sik'yu-lus) A small bundle of
muscle or nerve fibers ensheathed in
connective tissue.
fauces (faw'sez) The passageway between the
mouth and the oropharynx.
feces (fe'sez) Material expelled from the GI
tract by way of the anal canal during
defecation; composed of food residue,
bacteria, and secretions; also called stool.
fetus (fe'tus) A prenatal human after 8 weeks
of development.
fibroblast (fi'bro-blast) An elongated
connective tissue cell with cytoplasmic
extensions that is capable of forming
collagenous or elastic fibers.
fibrous joint A type of articulation bound by

fibrous connective tissue that allows little or
no movement (e.g., a syndesmosis).
filiform papillae (fil'-form pa-pil'e)
Numerous small projections scattered
irregularly over the entire surface of the
tongue in which taste buds are absent.
filum terminale (fi'lum ter-m-nal'e) A
fibrous, threadlike continuation of the pia
mater, that extends inferiorly from the
terminal end of the spinal cord to the coccyx.
fimbriae (fim'bre-e) Fringelike extensions
from the borders of the open end of the
uterine tube.
fissure (fish'ur) A groove or narrow cleft that
separates two parts, such as the cerebral
hemispheres of the brain.
flagellum (fla-jel'um) A whiplike structure
that provides motility for spermatozoa.
flexion Movement that decreases the angle
between parts of a joint; the opposite of
extension.
fontanel (fon''ta-nel') A soft membranous
gap between the incompletely formed cranial
bones of a fetus or baby; commonly known as
a soft spot.
foot The terminal portion of the lower
extremity, consisting of the tarsus,
metatarsus, and digits.
foramen (fo-ra'men); plural, foramina (foram'-na) An opening in an anatomical
structure, usually in a bone, for the passage of
a blood vessel or nerve.
foramen ovale (o-val'e) An opening through
the interatrial septum of the fetal heart.
forearm That portion of the upper extremity
between the elbow and the wrist; also called
the antebrachium.
formed elements The cellular portion of
blood.
fornix (for'niks) 1. A recess surrounding the
cervix of the uterus where it protrudes into
the vagina. 2. A tract within the brain
connecting the hippocampus with the
mammillary bodies.
fossa (fos'a) A depressed area, usually on a
bone.
fourth ventricle (ven'tr-kul) A cavity within
the brain, between the cerebellum and the
medulla oblongata and pons, containing
fovea centralis (fo've-a sen-tra'lis) A
depression on the macula lutea of the eye
where only cones are located; the area of
keenest vision.
frenulum (fren'yu-lum) A membranous
structure that serves to anchor and limit the
movement of a body part.
frontal 1. Pertaining to the forehead or frontal
bone. 2. Pertaining to the frontal plane.
frontal plane See coronal plane.

fungiform papillae (fun'j-form papil'e) Flattened, mushroom-shaped
projections interspersed among the filiform
type on the surface of the tongue. Fungiform
papillae contain taste buds.
G
gallbladder A pouchlike organ attached to the
underside of the liver; serves as a storage
reservoir for bile secreted by the liver.
gamete (gam'et) A haploid sex cell; either an
egg cell or a sperm cell.
ganglion (gang'gle-on) An aggregation of
nerve cell bodies located outside the central
nervous system.
gastrointestinal tract (GI tract) A continuous
tube through the anterior (ventral) body
cavity that extends from the mouth to the
anus; also called the digestive tract.
gene That portion of the DNA of a
chromosome containing the information
needed to synthesize a particular protein
molecule.
gingiva (jin-ji'va) The fleshy covering over
the mandible and maxilla through which the
teeth protrude within the mouth; also called
the gum.
gland A cell, tissue, or organ that produces a
specific substance or secretion for use in or
for elimination from the body.
glans penis (glanz pe'nis) The cone-shaped
terminal portion of the penis, formed from
the expanded corpus spongiosum.
gliding joint A type of synovial joint in which
the articular surfaces are flat, permitting only
side-to-side and back-and-forth movements.
glomerular (glo-mer'yu-lar) capsule The
double-walled proximal portion of a renal
tubule that encloses the glomerulus of a
nephron; also called Bowmans capsule.
glomerulus (glo-mer'yu-lus) The coiled tuft
of capillaries surrounded by the glomerular
capsule. It filters urine from the blood.
glottis (glot'is) A slitlike opening into the
larynx, positioned between the vocal folds.
goblet cell A mucus-secreting unicellular
gland associated with columnar epithelia;
also called a mucous cell.
Golgi (gol'je) complex A network of stacked,
flattened membranous sacs within the
cytoplasm of a cell. It serves to concentrate
and package proteins for secretion from the
cell.
Golgi tendon organ See neurotendinous
receptor.

Back Matter
Glossary
The McGrawHill
Companies, 2001
Glossary
gomphosis (gom-fo'sis) A fibrous joint
between the root of a tooth and the
periodontal ligament of the tooth socket.
gonad (go'nad) A reproductive organtestis
or ovarythat produces gametes and sex
hormones.
gray matter The region of the central nervous
system composed of nonmyelinated nerve
tissue.
greater omentum (o-men'tum) A doublelayered peritoneal membrane that originates
on the greater curvature of the stomach. It
hangs inferiorly, like an apron, over the
contents of the abdominal cavity.
gross anatomy The branch of anatomy
concerned with structures of the body that
can be studied without a microscope.
gustation (gus-ta'shun) The sense of taste.
gut The GI tract or a portion thereof;
generally used in reference to the embryonic
digestive tube, consisting of the foregut,
midgut, and hindgut.
gyrus (gi'rus) A convoluted elevation or
ridge.
H
hair A threadlike appendage of the epidermis
consisting of keratinized dead cells that have
been pushed up from a dividing basal layer.
hair cells Specialized sensory cells, such as in
the spiral organ, possessing numerous surface
microvilli; receptors for the senses of hearing
and equilibrium.
hair follicle A tubular depression in the
dermis of the skin in which a hair develops.
hand The terminal portion of the upper
extremity, consisting of the carpus,
metacarpus, and digits.
hard palate (pal'it) The bony partition
between the oral and nasal cavities, formed
by the maxillae and palatine bones and lined
by mucous membrane.
haustra (haws'tra) Sacculations, or pouches,
of the colon.
haversian (ha-ver'shun) canal See central
canal.
haversian system See osteon.
head The uppermost portion of a human that
contains the brain and major sense organs;
the proximal articulating part of a long bone.
heart A four-chambered muscular pumping
organ positioned in the thoracic cavity,
slightly to the left of midline.
hemoglobin (he'mo-glo''bin) A substance in
red blood cells consisting of the protein
globin and the iron-containing red pigment
heme. It functions in the transport of oxygen
and carbon dioxide.
hemopoiesis (hem''o-poi-e'sis) The

production of red blood cells.
hepatic (he-pat'ik) duct A duct formed from
the union of several bile ducts that drain bile
from the liver. It merges with the cystic duct
from the gallbladder to form the common
bile duct.
hepatic portal circulation The retum of
venous blood from the digestive organs and
spleen through a capillary network within
the liver before draining into the heart.
hepatopancreatic ampulla (hep''a-to-pan''kreat'ik am-pool'a) A short ductule formed by
the combined pancreatic and common bile
ducts within the duodenal papilla; also called
the ampulla of Vater.
hiatus (hi-a'tus) An opening or fissure; a
foramen.
hilum (hi'lum) A concave or depressed area
where vessels or nerves enter or exit an
organ.
hinge joint A type of synovial joint in which
the convex surface of one bone fits into the
concave surface of another, confining
movement to one plane (e.g., the knee and
interphalangeal joints).
histology (his-tol'o-je) Microscopic study of
the structure and function of tissues.
horizontal (transverse) plane A directional
plane that divides the body, an organ, or an
appendage into superior and inferior or
proximal and distal portions.
hormone (hor'mon) A chemical substance
produced in an endocrine gland and secreted
into the bloodstream that acts on target cells
to produce a specific effect.
hyaline (hi'a-lin) cartilage A cartilage with a
homogeneous matrix. It is the most common
type, occurring at the articular ends of bones,
in the trachea, and within the nose. Most of
the bones in the body are formed from
hyaline cartilage.
hymen (hi'men) A developmental remnant
(vestige) of membranous tissue that partially
covers the vaginal opening.
hyperextension Extension beyond the normal
anatomical position of 180.
hypertension Elevated or excessive blood
pressure.
hypodermis (hi''po-der'mis) Subcutaneous
tissue that binds the dermis to underlying
organs.
hyponychium (hi''po-nik'e-um) A thickened
supportive layer of the stratum corneum at
the distal end of a digit, under the free edge
of the nail.
hypothalamus (hi''po-thal'a-mus) An
important autonomic and neuroendocrine
control center located below the thalamus
within the diencephalon.
809
I
ileocecal (il'e-o-se'kal) valve A modification
of the mucosa at the junction of the small
intestine and large intestine that forms a
one-way passage and prevents the backflow
of food materials.
ileum (il'e-um) The terminal portion of the small
intestine between the jejunum and cecum.
incus (ing'kus) The middle of three auditory
ossicles within the middle-ear chamber;
commonly known as the anvil.
inferior vena cava (ve'na ka'va) A large
systemic vein that collects blood from the
body regions inferior to the level of the heart
and returns it to the right atrium.
infundibulum (in''fun-dib'yoo-lum) The
portion of the pituitary stalk that attaches
the hypothalamus of the brain to the
posterior pituitary.
ingestion The process of taking food or liquid
into the body by way of the oral cavity.
inguinal (ing'gw-nal) Pertaining to the groin
region.
inguinal canal The passage in the abdominal
wall through which a testis descends into the
scrotum.
inner ear The innermost region of the ear,
containing the cochlea and vestibular organs.
insertion The more movable attachment of a
muscle, usually the more distal; opposite the
origin.
inspiration The act of breathing air into the
pulmonary alveoli of the lungs; also called
inhalation.
insula (in'su-la) A cerebral lobe lying deep to
the lateral sulcus. It is covered by portions of
the frontal, parietal, and temporal lobes.
integument (in-teg'yoo-ment) The skin; the
largest organ of the body.
intercalated (in-ter'ka-lat-ed) disc A
thickened portion of the sarcolemma that
extends across a cardiac muscle fiber and
delimits the boundary between cells.
intercellular substance The matrix or
material between cells that largely
determines tissue types.
internal (deep) Toward the center, away from
the surface of the body.
interstitial fluid Fluid between the cells; also
called tissue fluid.
intervertebral (in''ter-ver'te-bral) disc A pad
of fibrocartilage positioned between the
bodies of adjacent vertebrae.
intestinal gland A simple tubular digestive
gland that opens onto the surface of the
intestinal mucosa and secretes digestive
enzymes; also called the crypt of Lieberkhn.
intestinal villus (vil'us) A minute projection
from the free surface of the tunica mucosa of

810
Back Matter
Glossary
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Companies, 2001
Glossary
the small intestine; the basic functional unit

of the digestive system.
intramembranous (in''tra-mem'bra-nus)
ossification A type of bone formation in which
layers of bone are formed from mesenchymal
cells without any cartilage model.
intrinsic (in-trin'zik) Situated in or
pertaining to internal origin.
inversion Movement of the foot in which the
sole is turned inward; the opposite of
eversion.
iris (i'ris) The pigmented portion of the
vascular tunic of the eye that surrounds the
pupil and regulates its diameter.
islets of Langerhans (i'letz of lang'erhanz) See pancreatic islets.
isthmus (is'mus) A narrow neck or portion of
tissue connecting two structures.
J
jejunum (je-joo'num) The middle portion of
the small intestine, located between the
duodenum and the ileum.
joint The point of juncture between two
bones; an articulation.
joint capsule The fibrous tissue that encloses
the joint cavity of a synovial joint.
K
keratin (ker'a-tin) An insoluble protein
present in the epidermis and in epidermal
derivatives, such as hair and nails.
kidney (kid'ne) One of the paired organs of
the urinary system that, among other
functions, filters wastes from the blood in the
formation of urine.
kinesiology (k-ne''se-ol'o-je) The study of
body movement.
knee The region in the lower extremity
between the thigh and the leg that contains a
synovial hinge joint.
L
labial frenulum (la'be-al fren'yu-lum) A
longitudinal fold of mucous membrane that
attaches the lips to the gum along the
midline of both the upper and lower lip.
labia majora (la'be-a ma-jor'a); singular,
labium majus The portion of the female
external genitalia consisting of two
longitudinal folds of skin that extend
downward and backward from the mons
pubis.
labia minora (m-nor'a), singular, labium

minus Two small folds of skin, devoid of hair
and sweat glands, lying between the labia
majora of the female external genitalia.
labyrinth (lab'-rinth) A complex system of
interconnecting tubes within the inner ear
that includes the semicircular ducts and the
cochlear and vestibular labyrinths.
lacrimal canaliculus (lak'r-mal kan''a-lik'yulus) A drainage duct for tears at the medial
corner of the eyelid. It conveys the tears
medially into the nasolacrimal sac.
lacrimal gland A tear-secreting gland on the
superior lateral portion of the eyeball,
underneath the upper eyelid.
lactation (lak-ta'shun) The production and
secretion of milk by the mammary glands.
lacteal (lak'te-al) A small lymphatic duct
associated with an intestinal villus of the
small intestine.
lacuna (la-kyoo'na) A small, hollow chamber
that houses an osteocyte in mature bone
tissue or a chondrocyte in cartilage tissue.
lambdoid suture (lam'doid soo'chur) The
immovable joint in the skull between the
parietal bones and the occipital bone.
lamella (la-mel'a) A concentric ring of matrix
surrounding the central canal in an osteon of
mature bone tissue.
lamellated corpuscle (lam'e-la'ted
kor'pus'l) A sensory receptor for pressure,
found in tendons, around joints, and in
visceral organs; also called a pacinian
corpuscle.
lamina (lam'-na) A thin plate of bone that
extends superiorly from the body of a
vertebra to form both sides of the arch of a
vertebra.
lanugo (la-noo'go) Short, silky fetal hair,
which may be present for a short time on a
premature infant.
large intestine The last major portion of the
GI tract, consisting of the cecum, colon,
rectum, and anal canal.
laryngopharynx (la-ring''go-far'ingks) The
lowermost portion of the pharynx, extending
inferiorly from the level of the hyoid bone to
the larynx.
larynx (lar'ingks) The short tubular organ
between the pharynx and trachea that houses
the vocal folds (cords); commonly known as
the voice box.
lateral Pertaining to the side; farther from the
median plane.
lateral ventricle (ven'tr-kul) A cavity within
the cerebral hemisphere of the brain that is
filled with cerebrospinal fluid.
leg That portion of the lower extremity
extending from the knee to the ankle.
lens 1. A transparent refracting medium,

usually made of glass or plastic. 2. The
transparent biconvex organ of the eye lying
posterior to the pupil and iris. It focuses light
rays entering through the pupil to form an
image on the retina.
lesser omentum (o-men'tum) A peritoneal
fold of tissue extending from the lesser
curvature of the stomach to the liver.
leukocyte A white blood cell; variant spelling,
leucocyte.
ligament (lig'a-ment) A tough cord or fibrous
band of connective tissue that binds bone to
bone to strengthen a joint and provide
flexibility. It also may support viscera.
limbic (lim'bik) system A group of deep brain
structures encircling the brain stem,
including the cingulate gyrus, the
hypothalamus, the hippocampus, the
amygdaloid nucleus, and various fiber tracts.
The limbic system is associated with aspects
of emotion and behavior and autonomic
functions.
linea alba (lin'e-a al'ba) A vertical fibrous
band running down the center of the anterior
abdominal wall.
lingual frenulum (ling'gwal fren'yu-lum) A
longitudinal fold of mucous membrane that
attaches the tongue to the floor of the oral
cavity.
lipid (lip'id) Any of a group of organic
molecules, including fats, phospholipids, and
steroids, that are generally insoluble in water.
liver A large visceral organ lying inferior to
the diaphragm in the right hypochondriac
region. The liver detoxifies the blood and
modifies the concentration of glucose,
triglycerides, ketone bodies, and proteins in
the blood plasma.
loop of Henle See nephron loop.
lower extremity A lower appendage,
including the hip, thigh, knee, leg, and foot.
lumbar (lum'bar) Pertaining to the region of
the loins; the part of the back between the
thorax and pelvis.
lumbar plexus (plek'sus) A network of nerves
formed by the anterior branches of spinal
nerves L1 through L4.
lumen (loo'men) The space within a tubular
structure through which a substance passes.
lung Either of the two major organs of
respiration positioned within the thoracic
cavity on either lateral side of the
mediastinum.
lunula (loo'nyoo-la) The crescent-shaped
whitish area at the proximal portion of a nail.

Back Matter
Glossary
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Glossary
luteinizing (loo'te--ni''zing) hormone
(LH) A hormone secreted by the
adenohypophysis (anterior lobe) of the
pituitary gland that stimulates ovulation and
progesterone secretion by the corpus luteum,
influences milk secretion by the mammary
glands in females, and stimulates testosterone
secretion by the testes in males.
lymph (limf) A clear, plasmalike fluid that
flows through lymphatic vessels.
lymph node A small ovoid mass of reticular
tissue located along the course of lymph
vessels.
lymphocyte (lim'fo-st) A type of white blood
cell characterized by a granular cytoplasm.
Lymphocytes usually constitute about
20%25% of the white cell count.
lymphoid tissue A type of connective tissue
dominated by lymphocytes.
M
macrophage (mak'ro-faj) A wandering
phagocytic cell.
macula lutea (mak'yu-la loo'te-a) A yellowish
depression in the retina of the eye that
contains the fovea centralis, the area of
keenest vision.
malignant Threatening to life; virulent. Of a
tumor, cancerous, tending to metastasize.
malleus (mal'e-us) The largest and outermost
of three auditory ossicles; attached to the
tympanic membrane and articulating with
the incus; commonly known as the
hammer.
mammary (mam'er-e) gland The gland of the
female breast responsible for lactation and
nourishment of the young.
marrow (mar'o) The soft connective tissue
found within the inner cavity of certain
bones that produces red blood cells.
mastication (mas't-ka'shun) The chewing of
food.
matrix (ma'triks) The intercellular substance
of a tissue.
meatus (me-a'tus) A passageway or opening
into a structure.
mechanoreceptor (mek''a-no-re-sep'tor) A
sensory receptor that responds to a
mechanical stimulus.
medial Toward or closer to the midline of the
body.
mediastinum (me''de-a-sti'num) 1. A septum
or cavity between two principal portions of
an organ. 2. The region in the center of the
thorax separating the lungs; contains the
heart and all of the thoracic viscera except
the lungs.
medulla (me-dul'a) The innermost part; the

central portion of such organs as the adrenal
gland and the kidney.
medulla oblongata (ob''long-ga'ta) That
portion of the brain stem between the spinal
cord and the pons.
medullary (med'l-er''e) (marrow) cavity The
hollow core of the diaphysis of a long bone in
which yellow bone marrow is found.
meiosis (mi-o'sis) A specialized type of cell
division by which gametes, or haploid sex
cells, are formed.
Meissners corpuscle (ms'nerz kor'pusl) A
sensory receptor found in the papillary layer
of the dermis of the skin; responsible for fine,
discriminative touch.
melanin (mel'a-nin) A dark pigment found
within the epidermis or epidermal derivatives
of the skin.
melanocyte (mel'a-no-st) A specialized
melanin-producing cell found in the deepest
layer of epidermis.
melanoma (mel''a-no'ma) A dark malignant
tumor of the skin; frequently forms in moles.
membranous (mem'bra-nus) bone Bone that
forms from membranous connective tissue
rather than from cartilage; also called
intramembranous bone.
menarche (me-nar'ke) The first menstrual
discharge; occurs normally between the ages
of 9 and 17.
meninges (me -nin'jez); singular, meninx
(me'ningks) A group of three fibrous
membranes covering the central nervous
system, composed of the dura mater,
arachnoid, and pia mater.
meniscus (me-nis'kus); plural, menisci
(menis'ki or me-nis'i) A wedge-shaped
fibrocartilage in certain synovial joints.
menopause (men''o-pawz) The period marked
by the cessation of menstrual periods in the
human female.
menstrual (men'stroo'al) cycle The rhythmic
female reproductive cycle characterized by
physical changes in the uterine lining.
menstruation (men''stroo-a'shun) The
discharge of blood and tissue from the uterus
at the end of menstrual cycle.
mesencephalic aqueduct (mez''en-se-fal'ik
ak'we-dukt) The channel that connects the
third and fourth ventricles of the brain; also
called the cerebral aqueduct or the aqueduct of
Sylvius.
mesencephalon (mez''en-sef'a-lon) The
midbrain, which contains the corpora
quadrigemina, the cerebral peduncles, and
specialized nuclei that help to control posture
and movement.
811
mesenchyme (mez'en-km) An embryonic

connective tissue that can migrate, and from
which all connective tissues arise.
mesenteric (mes''en-ter'ik) patches Clusters
of lymph nodes on the walls of the small
intestine; also called Peyers patches.
mesentery (mes'en-ter''e) A fold of
peritoneal membrane that attaches an
abdominal organ to the abdominal wall.
mesoderm (mes'o-derm) The middle layer of
the three primary germ layers of the
developing embryo.
mesothelium (mes''o-the'le-um) A simple
squamous epithelial tissue that lines body
cavities and covers visceral organs; also
called serosa.
mesovarium (mes''o-va're-um) The
peritoneal fold that attaches an ovary to the
broad ligament of the uterus.
metabolism (me-tab'o-liz-em) The sum total
of the chemical changes that occur within a
cell.
metacarpus (met''a-kar'pus) That region of
the hand between the wrist and the digits,
including the five bones that support the
palm of the hand.
metarteriole (met''ar-te're-ol) A small blood
vessel that emerges from an arteriole, passes
through a capillary network, and empties
into a venule.
metastasis (me-tas'ta-sis) 1. The spread of
pathogens or cancerous cells from an original
site to one or more sites elsewhere in the
body. 2. A secondary cancerous growth
formed by transmission of cancerous cells
from a primary growth located elsewhere in
the body.
metatarsus (met''a-tar'sus) The region of the
foot between the ankle and the digits that
includes five bones.
metencephalon (met''en-sef'a-lon) The most
superior region of the hindbrain, containing
the cerebellum and the pons.
microglia (mi-krog'le-a) Small phagocytic
cells found in the central nervous system.
microvilli (mi''kro-vil'i) Microscopic hairlike
projections of the cell membranes of certain
epithelial cells.
micturition (mik''tu-rish'un) The process of
voiding urine; also called urination.
midbrain That portion of the brain between
the pons and the forebrain.
middle ear The middle region of the ear,
containing the three auditory ossicles.
midsagittal (mid-saj'-tal) plane A plane that
divides the body or an organ into right and
left halves; also called the median plane.

812
Back Matter
Glossary
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Glossary
mitosis (mi-to'sis) The process of cell division

that results in two identical daughter cells,
containing the same number of
chromosomes.
mitral (mi'tral) valve See bicuspid valve.
mixed nerve A nerve that contains both
motor and sensory nerve fibers.
monocyte (mon'o-st) A phagocytic type of
white blood cell, normally constituting about
3%8% of the white blood cell count.
mons pubis (monz pyoo'bis) A fatty tissue
pad over the symphysis pubis in the female,
covered by coarse pubic hair.
morphogenesis (mor''fo-jen'e-sis) During
prenatal development, the transformation
involved in the growth and differentiation of
cells and tissues.
morula (mor''yu-la) An early stage of
embryonic development characterized by a
solid ball of cells.
motor area A region of the cerebral cortex
from which motor impulses to muscles or
glands originate.
motor nerve A nerve composed of motor
nerve fibers.
motor neuron (noor'on) A nerve cell that
conveys action potentials away from the
central nervous system to effector organs
(muscles and glands). Motor neurons form
the anterior roots of the spinal nerves.
motor unit A single motor neuron and the
muscle fibers it innervates.
mucosa (myoo-ko'sa) A mucous membrane
that lines cavities and tracts opening to the
exterior.
mucous (myoo'kus) cell A specialized
unicellular gland that produces and secretes
mucus; also called a goblet cell.
multipolar neuron (noor'on) A nerve cell
with many processes originating from the cell
body.
muscle A major type of tissue adapted to
contract. The three kinds of muscle are
cardiac, smooth, and skeletal.
muscularis (mus''kyu-lar'is) A muscular layer
or tunic of an organ, composed of smooth
muscle tissue.
myelencephalon (mi''e-len-sef'a-lon) The
posterior region of the hindbrain that
contains the medulla oblongata.
myelin (mi'e-lin) A lipoprotein material that
forms a sheathlike covering around nerve
fibers.
myeloid (mi'e-loid) tissue The red bone
marrow in which blood cells are produced.
myenteric plexus (mi''en-ter'ik plek'sus) A
network of sympathetic and parasympathetic
nerve fibers located in the muscular layer of
the wall of the small intestine; also called the
plexus of Auerbach.
myocardium (mi''o-kar'de-um) The middle

layer of the heart wall, composed of cardiac
muscle.
myofibril (mi''o-fi'bril) A bundle of
contractile fibers within muscle cells.
myofilament The filament that constitutes
myofibrils. It is composed of either actin or
myosin.
myogram (mi'o-gram) A recording of
electrical activity within a muscle.
myology (mi-ol'o-je) The science or study of
muscle structure and function.
myometrium (mi'o-me'tre-um) The layer or
tunic of smooth muscle within the uterine
wall.
myoneural (mi''o-noor'al) junction The site
of contact between the axon of a motor
neuron and a muscle fiber.
myopia (mi-o'pe-a) A visual defect in which
objects can be seen distinctly only when very
close to the eyes; also called nearsightedness.
myosin (mi'o-sin) A thick filament of protein
that, together with actin, causes muscle
contraction.
N
nail A hardened, keratinized plate that
develops from the epidermis and forms a
protective covering over the distal phalanges
of fingers and toes.
nares (na'rez) The openings into the nasal
cavity; also called the nostrils.
nasal cavity A mucosa-lined space above the
oral cavity, divided by a nasal septum. It is
the first chamber of the respiratory system.
nasal concha (kong'ka); plural, conchae
(kong'ke) A scroll-like bone extending
medially from the lateral wall of the nasal
cavity; also called a turbinate.
nasal septum A bony and cartilaginous
partition that separates the nasal cavity into
two portions.
nasopharynx (na''zo-far'ingks) The first or
uppermost portion of the pharynx, positioned
behind the nasal cavity and extending down
to the soft palate.
neck 1. Any constricted portion, such as the
neck of an organ. 2. The cervical region of
the body between the head and thorax.
necrosis (ne-kro'sis) Cell death or tissue
death as a result of disease or trauma.
neonatal (ne''o-na'tal) Concerning the period
from birth to the end of 4 weeks.
nephron (nef'ron) The functional unit of the
kidney, consisting of a glomerulus,
glomerular capsule, convoluted tubules, and
the nephron loop.
nephron loop The U-shaped part of the

nephron, consisting of descending and
ascending limbs; also called the loop of Henle.
nerve A bundle of nerve fibers outside the
neurilemma (noor''-lem'a) A thin,
membranous covering surrounding the
myelin sheath of a nerve fiber.
neurofibril (noor'o-fi'bril) One of many
delicate threadlike structures within the
cytoplasm of a cell body and the axon hillock
of a neuron.
neurofibril node A gap in the myelin sheath
of a nerve fiber; also called the node of
Ranvier.
neuroglia (noo-rog'le-a) Specialized
supportive cells of the central nervous
system; also called glial cells or glia.
neurohypophysis (noor''o-hi-pof'-sis) The
posterior lobe of the pituitary gland.
neurolemmocyte (noor''o-lem'o-st) A
specialized neuroglial cell that surrounds an
axon fiber of a peripheral neuron and forms
the myelin sheath; also called a Schwann cell.
neuron (noor'on) The structural and
functional cell of the nervous system,
composed of a cell body, dendrites, and an
axon; also called a nerve cell.
neurotendinous (noor''o-ten'din-us)
receptor A proprioceptor found near the
junction of tendons and muscles. It senses
muscle tension and acts to prevent overuse of
a muscle; also called a Golgi tendon organ.
neutrophil (noo'tro-fil) A type of phagocytic
white blood cell, normally constituting about
60%70% of the white cell count.
nipple A pigmented, cylindrical projection at
the apex of the breast.
Nissl (nis'l) bodies See chromatophilic
substances.
node of Ranvier (rahn-ve-a' or ran'ver) See
neurofibril node.
notochord (no'to-kord) flexible rod of tissue
that extends the length of the back of an
embryo.
nuclear membrane A double-walled
membrane composed of protein and lipid
molecules that surrounds the nucleus of a
cell.
nucleoplasm (noo'kle-o-plaz''em) The
protoplasmic contents of the nucleus of a
cell.
nucleus (noo'kle-us) The spherical or oval
body within a cell that contains the genetic
code.
nucleus pulposus (pul-po'sis) The soft, pulpy
core of an intervertebral disc; a remnant of
the notochord.

Back Matter
Glossary
The McGrawHill
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Glossary
O
olfaction (ol-fak'shun) The sense of smell.
olfactory bulb A ganglion-like expansion of
the olfactory tract, lying inferior to the
frontal lobe of the cerebrum on either side of
the crista galli of the ethmoid bone; receives
the olfactory nerves from the nasal cavity.
olfactory tract The tract of axons that
conveys impulses from the olfactory bulb to
the olfactory portion of the cerebral cortex.
oligodendrocyte (ol''-go-den'dro-st) A type
of neuroglial cell involved in the formation
of the myelin of nerve fibers within the
oocyte (o''o-sit) A developing egg cell.
oogenesis (o''o-jen'e -sis) The process of
female gamete formation.
optic Pertaining to the eye.
optic chiasma (ki-az'ma) An X-shaped
structure on the inferior aspect of the brain,
anterior to the pituitary gland, where there is
a partial crossing over of fibers in the optic
nerves; also called the optic chiasm.
optic disc A small region of the retina where
the fibers of the ganglion neurons exit from
the eyeball to form the optic nerve; also
called the blind spot.
optic tract A bundle of sensory axons between
the optic chiasma and the thalamus that
conveys visual impulses from the
photoreceptors within the eye.
oral Pertaining to the mouth.
ora serrata (o'ra se-ra'-ta) The jagged
peripheral margin of the retina.
organ A structure consisting of two or more
tissues that performs a specific function.
organ of Corti (kor'te) See spiral organ.
organelle (or''ga-nel') A specialized cellular
structure that performs a specific function for
the cell as a whole.
organism Any individual life form; either
unicellular or multicellular.
orifice (or'-fis) An opening into a body
cavity or tube.
origin The place of muscle attachment
usually the more stationary point or proximal
bone; opposite the insertion.
oropharynx (o''ro-far'ingks) The middle
portion of the pharynx, located posterior to
the oral cavity and extending from the soft
palate to the level of the hyoid bone.
ossicle (os'-kul) One of the three bones of
the middle ear; also called the auditory ossicle.
ossification (os''-f-ka'shun) The process of
bone tissue formation.
osteoblast (os'te-o-blast) A bone-forming
cell.
osteoclast (os'te-o-klast) A cell that erodes or
resorbs bone tissue.
osteocyte (os'te-o-st) A mature bone cell.

osteology (os''te-ol'o-je) The study of the
structure and function of bone and the entire
skeleton.
osteon (os'te-on) A group of osteocytes and
concentric lamellae surrounding a central
canal, constitutes the basic unit of structure
in bone tissue; also called a haversian system.
oval window See vestibular window.
ovarian (o-va're-an) follicle A developing
ovum and its surrounding epithelial cells.
ovarian ligament A cordlike connective tissue
that attaches the ovary to the uterus.
ovary (o'va-re) The female gonad in which
ova and certain sex hormones are produced.
oviduct (o'v-duct) The tube that transports
ova from the ovary to the uterus; also called
the uterine tube or fallopian tube.
ovulation (ov-yu-la'shun) The rupture of an
ovarian follicle with the release of an ovum.
ovum (o'vum) The female reproductive cell
or gamete; an egg cell.
P
pacinian corpuscle (pa-sin'e-an kor'pus'l)
See lamellated corpuscle.
palate (pal'at) The roof of the oral cavity.
palatine (pal'a-tin) Pertaining to the palate.
palmar (pal'mar) Pertaining to the palm of
the hand.
palpebra (pal'pe-bra) An eyelid.
pancreas (pan'kre-us) A mixed organ in the
abdominal cavity that secretes gastric juices
into the GI tract and insulin and glucagon
into the blood.
pancreatic (pan'kre-at'ik) duct A drainage
tube that carries pancreatic juice from the
pancreas into the duodenum of the
hepatopancreatic ampulla.
pancreatic islets A cluster of endocrine gland
cells within the pancreas that secretes insulin
and glucagon; also called islets of Langerhans.
papillae (pa-pil'e) Small, nipplelike
projections.
papillary (pap'-ler-e) muscle Muscular
projections from the ventricular walls of the
heart to which the chordae tendineae are
attached.
paranasal sinus (par''a-na'zal si'nus) An air
chamber lined with a mucous membrane that
communicates with the nasal cavity.
parasympathetic (par''a-sim''pa-thet'ik)
division Pertaining to the division of the
autonomic nervous system concerned with
activities that, in general, inhibit or oppose
the physiological effects of the sympathetic
division of the autonomic nervous system.
813
parathyroid (par''a-thi'roid) gland One of

four small endocrine glands embedded in the
posterior surface of the thyroid gland;
secretes parathyroid hormone.
parietal (pa-ri'etal) Pertaining to the wall of
an organ or cavity.
parietal pleura (ploor'a) The thin serous
membrane attached to the thoracic walls of
the pleural cavity.
parotid (pa-rot'id) gland One of the paired
salivary glands located over the masseter
muscle just anterior to the ear and connected
to the oral cavity by the parotid duct.
parturition (par''tyoo-rish'un) The act of
giving birth to young; childbirth.
pectoral (pek'to-ral) Pertaining to the chest
region.
pectoral girdle The portion of the skeleton
that supports the upper extremities.
pedicle (ped'kul) 1. A constricted portion or
stalk. 2. The bony process that projects
backward from the body of a vertebra,
connecting with the lamina on each side.
pelvic (pel'vik) Pertaining to the pelvis.
pelvic girdle That portion of the skeleton to
which the lower extremities are attached.
pelvis (pel'vis) A basinlike bony structure
formed by the sacrum and ossa coxae.
penis (pe'nis) The male organ of copulation,
used to introduce spermatozoa into the
female vagina and through which urine
passes during urination.
pennate (pen'at) Pertaining to a skeletal
muscle fiber arrangement in which the fibers
are attached to tendinous slips in a
featherlike pattern.
perforating canal A minute duct by means of
which blood vessels and nerves from the
periosteum penetrate into compact bone; also
called Volkmanns canal.
pericardium (per''-kar'de-um) The
protective serous membrane that surrounds
the heart.
perichondrium (per''-kon'dre-um) A sheet
of fibrous connective tissue that surrounds
some kinds of cartilage.
perikaryon (per''i-kar'e-on) The cell body of
a neuron.
perilymph (per''-limf) A fluid of the inner
ear that serves as a conducting medium for
the vibrations involved in hearing and
maintaining equilibrium.
perimysium (per''-mis'e-um) Fascia
(connective tissue) surrounding a bundle
(fascicle) of muscle fibers.
perineum (per''-ne'um) 1. The pelvic floor
and associated structures. 2. The external
region between the scrotum and anus in a
male or between the vulva and anus in a
female.

814
Back Matter
Glossary
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Glossary
perineurium (per''-noor'e-um) Connective

tissue surrounding a bundle (fascicle) of
nerve fibers.
periodontal (per''e-o-don'tal) ligament The
fibrous connective tissue lining the dental
alveoli.
periosteum The fibrous connective tissue
covering the outer surface of bone.
peripheral (pe'rif'er-al) nervous system
(PNS) The nerves and ganglia of the
nervous system that lie outside the brain and
spinal cord.
peristalsis (per''-stal'sis) Rhythmic
contractions of smooth muscle in the walls of
various tubular organs by which the contents
are forced onward.
peritoneum (per''-to-ne'um) The serous
membrane that lines the abdominal cavity
and covers the abdominal visceral organs.
Peyers (pi'ers) patches See mesenteric
patches.
phalanx (fa'langks); plural, phalanges (falan'jez) A bone of a digit (finger or toe).
pharynx (far'ingks) The tubular organ of the
digestive system and respiratory system
located at the back of the oral and nasal
cavities that extends to the larynx anteriorly
and to the esophagus posteriorly; also called
the throat.
phenotype (fe-no-tp) Observable features in
an individual that result from expression of
the genotype.
photoreceptor (fo''to-re-sep'tor) A sensory
nerve ending capable of being stimulated by
light.
physiology (fiz''e-ol'o-je) The science that
deals with the study of body functions.
pia mater (pi'a ma'ter) The innermost
meninx, in direct contact with the brain and
spinal cord.
pineal (pin'e-al) gland A small cone-shaped
gland located in the roof of the third
ventricle.
pinna (pin'a) The fleshy outer portion of the
external ear; also called the auricle.
pituitary (p-too'-tar-e) gland A small, peashaped endocrine gland situated in the sella
turcica of the sphenoid bone and connected
to the hypothalamus by the pituitary stalk;
consists of anterior and posterior lobes; also
called the hypophysis.
pivot joint A synovial joint in which the
rounded head of one bone articulates with
the depressed cup of another, permitting
rotational movement.
placenta (pla-sen'ta) The organ of metabolic
exchange between the mother and the fetus.
It is expelled following birth.
plantar (plan'tar) Pertaining to the sole of the
foot.
plasma (plaz'ma) The clear, yellowish fluid

portion of blood in which cells are
suspended.
platelets (plat'letz) Small fragments of specific
bone marrow cells that function in blood
clotting; also called thrombocytes.
pleural (ploor'al) Pertaining to the serous
membranes associated with the lungs.
pleural cavity The potential space between
the visceral pleura and the parietal pleura
surrounding a lung.
pleural membranes Serous membranes that
surround the lungs and provide protection
and compartmentalization.
plexus (plek'sus) A network of nerve fibers,
blood vessels, or lymphatics.
plexus of Auerbach (ow'er-bak) See
myenteric plexus.
plexus of Meissner (ms'ner) See submucosal
plexus.
plicae circulares (pli'se ser-kyu-lar-e z) Deep
folds in the wall of the small intestine that
increase the absorptive surface area.
pneumotaxic (noo''mo-tak'sik) area A
portion of the respiratory control center in
the pons that has an inhibitory effect on the
inspiratory center in the medulla oblongata.
pons (ponz) The portion of the brain stem
just above the medulla oblongata and
anterior to the cerebellum.
popliteal (pop''l-te'al or pop-lit'e-al)
Pertaining to the concave region on the
posterior aspect of the knee.
posterior Toward the back; also called dorsal.
posterior root An aggregation of sensory
neuron fibers lying between a spinal nerve
and the posterolateral aspect of the spinal
cord; also called the dorsal root or sensory
root.
posterior root ganglion (gang'gle-on) A
cluster of cell bodies of sensory neurons
located along the posterior root of a spinal
nerve; also called a sensory ganglion.
postganglionic (post''gang-gle-on'ik)
neuron The second neuron in an autonomic
motor pathway. Its cell body is outside the
central nervous system, and it terminates at
an effector organ.
postnatal After birth.
preganglionic neuron The first neuron in an
autonomic motor pathway. Its cell body is
within the central nervous system, and it
terminates on a postganglionic neuron.
pregnancy The condition in which a female is
carrying a developing offspring within the
uterus.
prenatal Existing or occurring before birth.
prepuce (pre'pyoos) A fold of loose
retractable skin covering the glans of the
penis or clitoris; also called the foreskin.
prime mover The muscle most directly

responsible for a particular movement.
pronation (pro-na'shun) A rotational
movement of the forearm that turns the palm
of the hand posteriorly; the opposite of
supination.
proprioceptor (pro''pre-o-sep'tor) A sensory
nerve ending that responds to changes in
tension in a muscle or tendon.
prostate (pros'tat) A walnut-shaped gland
surrounding the male urethra just below the
urinary bladder. It secretes an additive to
seminal fluid during ejaculation.
prosthesis (pros-the'sis) An artificial device
to replace a diseased or worn body part.
protein Any of a group of large organic
molecules made up of amino acid subunits
linked by peptide bonds.
protraction (pro-trak'shun) Forward
movement of a body part, such as the
mandible, on a plane parallel with the
ground; the opposite of retraction.
proximal (prok's-mal) Closer to the midline
of the body or origin of an appendage; the
opposite of distal.
pseudounipolar neuron (soo''do-yoo''npo'lar noor'on) A nerve cell in which only
one process extends from the cell body;
results from the fusion of two processes
during embryonic development.
puberty (pyoo'ber-tee) The period of
development in which the reproductive
organs become functional and the secondary
sex characteristics are expressed.
pulmonary (pul'mo-ner''e) Pertaining to the
lungs.
pulmonary circulation The circuit of blood
flow between the heart and the lungs.
Oxygen-poor blood from the right ventricle
is oxygenated in the lungs and then returned
to the left atrium of the heart.
pulp cavity A cavity within the center of a
tooth that contains blood vessels, nerves, and
lymphatics.
pupil The opening through the iris that
permits light to enter the posterior cavity of
the eyeball and be refracted by the lens.
Purkinje (pur-kin'je) fibers See conduction
myofibers.
pyloric sphincter (pi-lor'ik sfingk'ter) A
thick ring of smooth muscle encircling the
opening between the stomach and the
duodenum. It regulates the passage of food
material into the small intestine and prevents
backflow.
pyramid (pir'amid) Any structure of the body
with a pyramidal shape, including the renal
pyramids in the kidney and the medullary
pyramids on the inferior surface of the brain.

Back Matter
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The McGrawHill
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Glossary
R
ramus (ra'mus) A branch of a bone, artery, or
nerve.
raphe (ra'fe) A seamlike line or ridge between
two similar parts of a body organ, as in the
scrotum.
receptor (re-sep'tor) A sense organ or the
specialized distal end of a sensory neuron that
receives stimuli from the environment.
rectouterine (rek''to-yoo'ter-in) pouch A
pocket formed by the deflection of the
parietal peritoneum between the uterus and
the rectum; also called the pouch of Douglas
or Douglas cul-de-sac.
rectum (rec'tum) That portion of the GI tract
between the sigmoid colon and the anal
canal.
red bone marrow (mar'o) A hematopoietic
tissue found within the spongy bone tissue of
certain bones.
red nucleus(noo'kle-us) An aggregation of
gray matter of a reddish color located in the
upper portion of the midbrain. It sends fibers
to certain brain tracts and helps to
coordinate muscular movements.
reflex A rapid involuntary response to a
stimulus.
reflex arc The basic conduction pathway
through the nervous system, consisting of a
sensory neuron, association neuron, and a
motor neuron.
regional anatomy The division of anatomy
concerned with structural arrangement in
specific areas of the body, such as the head,
neck, thorax, or abdomen.
renal (re'nal) Pertaining to the kidney.
renal corpuscle (kor'pusl) A glomerular
capsule and its enclosed glomerulus; also
called the malpighian corpuscle.
renal cortex The outer portion of the kidney,
primarily vascular.
renal medulla (me-dul'a) The inner portion of
the kidney, including the renal pyramids and
renal columns.
renal pelvis The inner cavity of the kidney
formed by the expanded ureter, into which
the major calyces open.
renal pyramid Any of various pyramidal
masses that are seen on longitudinal section
of the kidney and that contain part of the
nephron loops and the collecting tubules.
respiration The exchange of gases between
the external environment and the cells of an
organism.
respiratory center The structure or portion of
the brain stem that regulates the depth and
rate of breathing.
respiratory membrane A thin, moistened

membrane within the lungs, composed of an
alveolar portion and a capillary portion,
through which gaseous exchange occurs.
rete testis (re'te tes'tis) The network of
canals at the termination of the seminiferous
tubules in the testis that is associated with
the production of spermatozoa.
reticular (re-tik'yoo-lar) formation A
network of nervous tissue fibers in the brain
stem that arouses the higher brain centers.
retina (ret'-na) The inner layer of the eyeball
that contains the photoreceptors.
retraction Backward movement of a body
part, such as the mandible, on a plane
parallel with the ground; the opposite of
protraction.
retroperitoneal (ret''ro-per''-tone'al) Positioned behind the parietal
peritoneum.
rhythmicity (rith-mis'-te) area A portion of
the respiratory control center located in the
medulla oblongata. It controls inspiratory
and expiratory phases.
ribosome (ri'bo-som) A cytoplasmic organelle
composed of protein and RNA in which
protein synthesis occurs.
right lymphatic (lim-fat'ik) duct A major
vessel of the lymphatic system that drains
lymph from the upper right portion of the
body into the right subclavian vein.
rod A type of photoreceptor cell in the retina
of the eye that is specialized for colorless,
dim-light vision.
root canal The tubular extension of the pulp
cavity into the root of a tooth. It contains
vessels and nerves.
rotation Movement of a bone around its own
longitudinal axis.
round window See cochlear window.
rugae (roo'je) Folds or ridges of the mucosa of
an organ, such are those of the stomach or
urinary bladder.
S
saccule (sak'yool) The saclike cavity in the
membranous labyrinth within the vestibule
of the inner ear that contains a vestibular
organ for equilibrium.
sacral (sa'kral) Pertaining to the sacrum.
sacral plexus (plek'sus) A network of nerve
fibers arising from spinal nerves L4 through
S3. Nerves arising from the sacral plexus
merge with those from the lumbar plexus to
form the lumbosacral plexus that supplies the
lower extremity.
815
saddle joint A synovial joint in which the

articular surfaces of both bones are concave
in one plane and convex, or saddle shaped, in
the other plane, such as in the distal
carpometacarpal joint of the thumb.
sagittal plane A vertical plane through the
body that divides it into right and left sides.
salivary (sal'-ver-e) gland An accessory
digestive gland that secretes saliva into the
oral cavity.
sarcolemma (sar''ko-lem'a) The cell
membrane of a muscle fiber.
sarcomere (sar'ko-mer) The portion of a
skeletal muscle fiber between a pair of Z
lines; the basic subunit of skeletal muscle
contraction.
sarcoplasm (sar'ko-plaz''em) The cytoplasm
within a muscle fiber.
scala tympani (ska'la tim'pa-ne) The lower
channel of the cochlea that is filled with
perilymph.
scala vestibuli (ve-stib'yu-le) The upper
channel of the cochlea that is filled with
perilymph.
Schwann (shwahn) cell See neurolemmocyte.
sclera (skler'a) The outer white layer of
fibrous connective tissue that serves as a
protective covering for the eyeball and
attachment for the extrinsic ocular muscles.
scleral venous sinus A circular venous
drainage for the aqueous humor from the
anterior chamber of the eye; located at the
junction of the sclera and the cornea; also
called the canal of Schlemm.
scrotum (skro'tum) The musculocutaneous
sac that contains the testes and their
accessory organs.
sebaceous (se-ba'shus) gland An exocrine
gland of the skin that secretes sebum.
sebum (se'bum) An oily waterproofing
secretion of sebaceous glands.
semen (se'men) The thick, whitish secretion
of the reproductive organs of the male,
consisting of spermatozoa and secretions of
the testes, seminal vesicles, prostate, and
bulbourethral glands.
semicircular canals Tubular channels within
the inner ear that contain the receptors for
equilibrium.
semilunar (sem''e-loo'nar) valve Crescentshaped heart valves, positioned at the
entrances to the aorta and the pulmonary
trunk.
seminal vesicles (sem'-nal ves'-kulz) A pair
of male accessory reproductive organs, lying
posterior and inferior to the urinary bladder,
that secrete additives to spermatozoa into the
ejaculatory ducts.

816
Back Matter
Glossary
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Companies, 2001
Glossary
seminiferous (sem''-nif'er-us)
tubules Numerous small ducts in the testes,
where spermatozoa are produced.
senescence (se-nes'ens) The process of
growing old; aging.
sensory area A region of the cerebral cortex
that receives and interprets sensory nerve
impulses.
sensory neuron (noor'un) A nerve cell that
conducts an impulse from a receptor organ to
the central nervous system; also called an
afferent neuron.
septum (sep'tum) A membranous or fleshy
wall dividing two cavities.
serous (se'rus) membrane An epithelial and
connective tissue membrane that lines body
cavities and covers visceral organs within
these cavities; also called serosa.
Sertoli (ser-to'le) cells See sustentacular cells.
serum (se'rum) Blood plasma with the
clotting elements removed.
sesamoid (ses'a-moid) bone A membranous
bone formed in a tendon in response to joint
stress (e.g., the patella).
shoulder The region of the body where the
humerus articulates with the scapula; also
called omo.
sigmoid colon (sig'moid ko'lon) The Sshaped portion of the large intestine between
the descending colon and the rectum.
sinoatrial (si''no-a'tre-al) node A mass of
specialized cardiac tissue in the wall of the
right atrium that initiates the cardiac cycle;
the SA node; also called the pacemaker.
sinus (si'nus) A cavity or hollow space within
a bone or other tissue.
sinusoid A small, blood-filled space in certain
organs, such as the spleen or liver.
skeletal muscle A muscle that is connected at
either or both extremities with a bone. It
consists of elongated, multinucleated, striated
skeletal muscle fibers that contract when
stimulated by motor nerve impulses.
small intestine The portion of the GI tract
between the stomach and the cecum whose
function is the absorption of food nutrients;
consists of the duodenum, jejunum, and
ileum.
smooth muscle A specialized type of
nonstriated muscle tissue, composed of
spindle-shaped fibers with a single nucleus. It
contracts in an involuntary, rhythmic fashion
within the walls of visceral organs.
soft palate (pal'at) The fleshy posterior
portion of the roof of the mouth, extending
from the palatine bones to the uvula.
somatic (so-mat'ik) Pertaining to the
nonvisceral parts of the body.
spermatic (sper-mat'ik) cord A structure of

the male reproductive system that includes
the ductus deferens, spermatic vessels, nerves,
cremaster muscle, and connective tissue. The
spermatic cord extends from a testis to the
inguinal ring.
spermatogenesis (sper-mat''o-jen'-sis) The
production of male gametes, or spermatozoa.
spermatogonia (sper-mat''o-go'ne-a) Sperm
stem cells within the seminiferous tubules of
the testes; the progenitors of spermatocytes.
spermatozoon (sper-mat''o-zo'on); plural,
spermatozoa or, loosely, sperm A mature
male sex cell, or gamete.
sphincter (sfingk'ter) A circular muscle that
normally maintains constriction of a body
opening or the lumen of a tubular structure
and that relaxes as required by normal
physiological functioning.
sphincter of ampulla (am-pool'a) The
muscular constriction at the opening of the
common bile and pancreatic ducts; also
called the sphincter of Oddi.
sphincter of Oddi (o'de) See sphincter of
ampulla.
spinal cord The portion of the central nervous
system that extends downward from the brain
stem through the vertebral canal.
spinal ganglion (gang'gle-on) A cluster of
nerve cell bodies on the posterior root of a
spinal nerve.
spinal nerve One of the 31 pairs of nerves
that arise from the spinal cord.
spinous (spi'nus) process A sharp projection
of bone or a ridge of bone, such as on the
scapula.
spiral organ The functional unit of hearing,
consisting of a basilar membrane that
supports receptor hair cells and a tectorial
membrane located within the cochlea; also
called the organ of Corti.
spleen (sple n) A large, blood-filled, glandular
organ in the upper left quadrant of the
abdomen. It is attached by mesenteries to the
stomach.
spongy bone A type of bone with a latticelike
structure; also called cancellous bone.
squamous (skwa'mus) Flat or scalelike.
stapes (sta'pez) The innermost of the auditory
ossicles of the ear that fits against the
vestibular (oval) window of the inner ear;
commonly known as the stirrup.
statoconia (stat''o-ko'ne-a) Small, hardened
particles of calcium carbonate in the saccule
and utricle of the inner ear that are
associated with the receptors of equilibrium;
also called otoliths.
stomach A pouchlike digestive organ between
the esophagus and the duodenum, lying just
beneath the diaphragm.
stratified (strat'-fd) Arranged in layers, or

strata.
stratum basale (ba-sa'le) The deepest
epidermal layer, where mitotic activity
occurs.
stratum corneum (kor'ne-um) The outer
cornified layer of the epidermis of the skin.
stroma (stro'ma) A connective tissue
framework in an organ, gland, or other tissue.
subarachnoid (sub''a-rak'noid) space The
space within the meninges between the
arachnoid and the pia mater, where
cerebrospinal fluid flows.
subdural (sub-door'al) space The narrow
space between the dura mater and the
arachnoid.
sublingual (sub-ling'gwal) gland One of the
three pairs of salivary glands. It is located
below the tongue and its several sublingual
ducts open into the floor of the mouth.
submandibular (sub''man-dib'yu-lar)
gland One of the three pairs of salivary
glands. It is located below the mandible and
its submandibular duct opens to the side of
the lingual frenulum.
submucosa (sub''myoo-ko'sa) A layer of
supportive connective tissue that underlies a
mucous membrane.
submucosal plexus (plek'sus) A network of
sympathetic and parasympathetic nerve fibers
located in the submucosa of the small
intestine; also called the plexus of Meissner.
sudoriferous (soo'dor-if'er-us) gland An
exocrine gland that excretes perspiration, or
sweat, onto the surface of the skin; also called
a sweat gland.
sulcus (sulkus) A shallow impression or
groove.
superficial Toward or near the surface.
superficial fascia (fash'e-a) A binding layer of
connective tissue between the dermis of the
skin and the underlying muscle.
superior Toward the upper part of a structure
or toward the head; also called cephalic.
superior vena cava (ve'na ka'va) A large
systemic vein that collects blood from regions
of the body superior to the heart and returns
it to the right atrium.
supination (soo''p-na'shun) A rotational
movement of the forearm that turns the palm
of the hand anteriorly; the opposite of
pronation.
surface anatomy The division of anatomy
concerned with the form and markings of the
surface of the body as they relate to the
deeper structures.
surfactant (sur-fak'tant) A substance
secreted by the pulmonary alveolar cells of
the lung that reduces the surface tension and
the tendency for the pulmonary alveoli to
collapse after each expiration.

Back Matter
Glossary
The McGrawHill
Companies, 2001
Glossary
suspensory (su-spen'so-re) ligament 1. A
band of peritoneum that extends laterally
from the surface of the ovary to the wall of
the pelvic cavity. 2. A ligament that supports
an organ or a body part, such as that
supporting the lens of the eye.
sustentacular (sus-ten-tak'yu-lar)
cells Specialized cells within the testes that
supply nutrients to developing spermatozoa;
also called Sertoli cells or nurse cells.
sutural (soo'chur-al) bone A small bone
positioned within a suture of certain cranial
bones; also called a wormian bone.
suture (soo'chur) A type of fibrous joint
found between bones of the skull.
sweat gland See sudoriferous gland.
sympathetic division Pertaining to the
division of the autonomic nervous system
concerned with activities that, in general,
inhibit or oppose the physiological effects of
the parasympathetic nervous system; also
called the thoracolumbar division.
symphysis (sim'f-sis) A type of joint
characterized by a fibrocartilaginous pad
between the articulating bones that provides
slight movement.
symphysis pubis (pyoo'bis) The slightly
movable cartilaginous joint between the two
pubic bones of the pelvic girdle.
synapse (sin'aps) The functional junction
between two neurons or between a neuron
and an effector.
synaptic (s-nap'tik) cleft The minute space
that separates the axon terminal of one
neuron from another neuron or muscle fiber.
Neurotransmitter chemicals diffuse across the
cleft to affect the postsynaptic cell.
synchondrosis (sin''kon-dro'sis) A
cartilaginous joint in which the articulating
bones are separated by hyaline cartilage.
syndesmosis (sin''des-mo'sis) A type of
fibrous joint in which two bones are united
by an interosseous ligament.
synergist (sin'er-jist) A muscle that assists
the action of the prime mover.
synovial (s-no've-al) cavity The space
between the articulating bones of a synovial
joint, filled with synovial fluid; also called a
joint cavity.
synovial joint A freely movable joint in
which a joint (synovial) cavity is present
between the articulating bones; also called a
diarthrotic joint.
synovial membrane The inner membrane of a
joint (synovial) capsule that secretes synovial
fluid into the cavity.
system A group of body organs that function
together.
systemic (sis-tem'ik) Of, relating to, or
affecting the organism as a whole.
systemic anatomy The division of anatomy

concerned with the structure and function of
the various systems of the body.
systemic circulation The circuit of blood flow
from the left ventricle of the heart to the
entire body and back to the heart via the
right atrium; in contrast to the pulmonary
circulation, which involves the lungs.
systole (sis'to-le) The muscular contraction of
a heart chamber during the cardiac cycle;
ventricular contraction, unless otherwise
specified.
systolic (sis-tol'ik) pressure Arterial blood
pressure during the ventricular systolic phase
of the cardiac cycle.
T
tactile (tak'til) Pertaining to the sense of
touch.
taeniae coli (te'ne-e ko'li) The three
longitudinal bands of muscle in the wall of
the large intestine.
target organ A tissue or organ that is affected
by a particular hormone.
tarsal gland An oil-secreting gland that opens
on the exposed edge of each eyelid; also
called the meibomian gland.
tarsus (tar'sus) Pertaining to the ankle; the
proximal portion of the foot that contains
the seven tarsal bones.
taste bud An organ containing the
chemoreceptors associated with the sense of
taste.
tectorial (tek'to''re-al) membrane A
gelatinous membrane positioned over the
hair cells of the spiral organ in the cochlea.
teeth Accessory structures of digestion
adapted to cut, shred, crush, and grind food.
telencephalon (tel''en-sef'a-lon) The anterior
region of the forebrain, constituting the
cerebral hemispheres and related parts.
tendo calcaneus (ten'do kal-ka'ne-us) The
tendon that attaches the calf muscles to the
calcaneus; also called the Achilles tendon.
tendon A band of dense regular connective
tissue that attaches muscle to bone.
tendon sheath A covering of synovial
membrane surrounding certain tendons.
tentorium cerebelli (ten-to're-um ser''e
bel'i) An extension of dura mater separating
the cerebellum from the basal surface of the
occipital and temporal lobes of the cerebral
cortex.
teratogen (te-rat'o-jen) Any agent or factor
that causes a physical defect in a developing
embryo or fetus.
testis (tes'tis) The primary reproductive organ
of a male that produces spermatozoa and
male sex hormones.
817
thalamus (thal'a-mus) An oval mass of gray

matter within the diencephalon that serves
as a sensory relay center.
thigh The proximal portion of the lower
extremity between the hip and the knee in
which the femur is located.
third ventricle (ven'tr-kul) A narrow cavity
between the right and left halves of the
thalamus and between the lateral ventricles
that contains cerebrospinal fluid.
thoracic (thor'a-sik) Pertaining to the chest
region.
thoracic duct A major lymphatic vessel of the
body that drains lymph from the entire body,
except for the upper right quadrant, and
returns it to the left subclavian vein.
thorax (thor'aks) The chest.
thrombocytes (throm'bo-sts) See platelets.
thymus (thi'mus) A bilobed lymphoid organ
positioned in the upper mediastinum,
posterior to the sternum and between the
lungs.
thyroid cartilage The largest cartilage of the
larynx. Its two broad processes join anteriorly
to form the Adams apple.
thyroid gland An endocrine gland located just
below the larynx, in front of the trachea,
consisting of two lobes connected by a
narrow band of tissue called the isthmus.
tissue An aggregation of similar cells and their
binding intercellular substance, joined to
perform a specific function.
tongue A protrusible muscular organ on the
floor of the oral cavity.
tonsil (ton'sil) A mass of lymphoid tissue
embedded in the mucous membrane of the
pharynx.
trabeculae (tra-bek'yu-le) 1. Any of the
supporting strands of connective tissue
projecting into an organ and constituting
part of its framework. 2. Any of the fine
spicules forming a network in spongy bone.
trachea (tra'ke-a) The airway leading from
the larynx to the bronchi, composed of
cartilaginous rings and a ciliated mucosal
lining of the lumen; commonly known as the
windpipe.
tract A bundle of nerve fibers within the
transection A cross-sectional cut.
transverse colon (ko'lon) That portion of the
large intestine extending from right to left
across the abdomen between the hepatic and
splenic flexures.
transverse fissure (fish'ur) The prominent
cleft that horizontally separates the cerebrum
from the cerebellum.
tricuspid (tri-kus'pid) valve The heart valve
located between the right atrium and the
right ventricle; the right AV valve.

818
Back Matter
Glossary
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Glossary
trigone (tri'gon) A triangular area in the

urinary bladder delimited by the openings of
the ureters and the urethra.
trochanter (tro-kan'ter) A broad, prominent
process on the proximolateral portion of the
femur.
trochlea (trok'le-a) A structure having the
shape or function of a pulley, especially the
part of the distal end of the humerus that
articulates with the ulna.
trunk The thorax and abdomen together; also
called torso.
tubercle (too'ber-kul) A small elevated
process on a bone.
tuberosity (too''be-ros'-te) An elevation or
protuberance on a bone.
tunica albuginea (too'n-ka al''byoo-jin'e-a)
A tough connective tissue sheath
surrounding a structure (e.g., the capsule
enclosing a testis).
tympanic (tim-pan'ik) membrane The
membranous eardrum positioned between the
outer and middle ear.
U
umbilical (um-bil'-kal) cord A flexible
cordlike structure that connects the fetus
with the placenta. It contains two umbilical
arteries and one vein that transport
nourishment to the fetus and remove its
waste.
umbilicus (um-bil'-kus) The site where the
umbilical cord was attached to the fetus; also
called the navel.
upper extremity The appendage attached to
the pectoral girdle, consisting of the
shoulder, brachium, elbow, antebrachium,
and hand.
ureter (yoo-re'ter) A tube that transports
urine from the kidney to the urinary bladder.
urethra (yoo-re'thra) A tube that transports
urine from the urinary bladder to the outside
of the body.
urinary (yoo'r-ner''e) bladder A distensible
sac that stores urine; situated in the pelvic
cavity, posterior to the symphysis pubis.
urogenital (yoo''ro-jen'-tal) triangle The
region of the pelvic floor containing the
external genitalia.
uterine (yoo'ter-in) tube The tube through
which the ovum is transported to the uterus;
the site of fertilization; also called the oviduct
or fallopian tube.
uterus (yoo'ter-us) The hollow muscular
organ in which a fertilized egg implants and
develops into a fetus. It is located within the
female pelvis between the urinary bladder
and the rectum; also called the womb.
utricle (yoo'tr-kul) An enlarged portion of

the membranous labyrinth, located within
the vestibule of the inner ear.
uvula (yoo'vyu-la) A fleshy, pendulous
portion of the soft palate that blocks the
nasopharynx during swallowing.
V
vacuole (vak'yoo-ol) A small space or cavity
within the cytoplasm of a cell.
vagina (va-ji'na) A tubular organ leading from
the uterus to the vestibule of the female
reproductive tract that receives the male
penis during coitus.
vallate papillae (val'at pa-pil'e) The largest of
the papillae on the surface of the tongue.
They are arranged in an inverted V-shape at
the back of the tongue. Vallate papillae
contain taste buds.
vasomotor (va-zo-mo'tor) center A cluster of
nerve cell bodies in the medulla oblongata. It
controls the diameter of blood vessels, and
therefore has an important role in regulating
blood pressure.
vein A blood vessel that conveys blood toward
the heart.
vena cava (ve'na ka'va) One of two large
vessels that return deoxygenated blood to the
right atrium of the heart.
ventral (ven'tral) Toward the front or belly
surface; the opposite of dorsal.
ventricle (ven'tr-kul) A cavity within an
organ; especially those cavities in the brain
that contain cerebrospinal fluid and those in
the heart that contain blood to be pumped
from the heart.
venule (ven'yool) A small vessel that carries
venous blood from capillaries to a vein.
vermiform (ver'm-form) appendix See
appendix.
vermis (ver'mis) The narrow, middle lobular
structure that separates the cerebellar
hemispheres.
vertebral (ver'te-bral) canal The tubelike
cavity extending through the vertebral
column that contains the spinal cord; also
called the spinal canal.
vestibular (ve-stib'yu-lar) folds The
supporting folds of tissue for the vocal folds
within the larynx; also called false vocal cords.
vestibular window The oval opening in the
bony wall between the middle and inner ear
on which the footplate of the stapes rests;
also called the oval window.
vestibule (ves't-byool) A space or region at
the beginning of a canal, especially that of
the nose, inner ear, and vagina.
villus (vil'us) A minute projection from the

free surface of a mucous membrane,
especially one of the vascular projections of
the mucosal layer of the small intestine.
viscera (vis'er-a) The organs within the
abdominal or thoracic cavities.
visceral (vis'er-al) Pertaining to the
membranous covering of the viscera.
visceral peritoneum (per''-to-ne'um) A
serous membrane that covers the surfaces of
abdominal viscera.
visceral pleura (ploor'a) A serous membrane
that covers the surfaces of the lungs.
visceroceptor (vis''er-o-sep'tor) A sensory
receptor located within the visceral organs
that responds to internal pain, pressure,
stretch, and chemical changes.
vitreous (vit're-us) humor The transparent
gel that occupies the posterior cavity, located
between the lens and retina of the eyeball.
vocal folds Folds of the mucous membrane in
the larynx that produce sound as they are
pulled taut and vibrated; also called vocal
cords.
Volkmanns (folk'manz) canal See
perforating canal.
vulva (vul'va) The external genitalia of the
female that surround the opening of the
vagina; also called the pudendum.
W
white matter Bundles of myelinated axons
located in the central nervous system.
wormian (wer'me-an) bone See sutural bone.
Y
yellow bone marrow (mar'o) Specialized
tissue within medullary cavities of certain
bones in which lipids are stored.
Z
zygote (zi'got) A fertilized egg cell formed by
the union of a spermatozoon and an ovulated
secondary oocyte (ovum).

Back Matter
Credits
The McGrawHill
Companies, 2001
Credits
Photographs
Chapter 1
Opener: Giraudon/Art Resource;
1.1: Mario Caprio/Visuals
Unlimited; 1.2: A Short History of
Anatomy and Physiology from the Greeks
to Harvey by C. Singer, 1957, Dover
Publications, New York, NY. Reprinted
by permission; 1.3: Published by
permission of the National Museum of
Denmark; 1.5: The Trustees of the
British Museum; 1.6: Egyptian
National Museum; 1.7a: John
Watney/Science Photo Library/Photo
Researchers, Inc.; 1.7b: Szuson J.
Wong/Peter Arnold; 1.8:
Biblioteque Nationale; 1.9, 1.10:
Alinari/Art Resource, NY; 1.11:
Giraudon/Art Resource, NY: 1.12,
1.13: Reproduced by gracious
permission of Her Majesty Queen
Elizabeth, Royal Art Collection, Great
Britain; 1.14: From the Works of
Andrea Vesalius of Brussels by J. Bade,
C. M. Saunders and Charley P. OMalley,
pg. 1096, Dover Publications; 1.15:
Scala/Art Resource, NY; 1.16:
Stock Montage; 1.17a: Debi
Stambaugh; 1.17b: George Mosil/
Visuals Unlimited; 1.17c: CNRI/
SPL/Photo Researchers, Inc.; 1.18a:
Courtesy of Eastman Kodak Co.;
1.18b: Carroll H. Weiss/Camera
M.D. Studios; 1.18c: Courtesy of Utah
Valley Regional Medical Center, Dept.
of Radiation; 1.19a Lester V. Bergman
& Associates, Inc.; 1.19b: From
R.A. Robb, Three-Dimensional
Biomedical Imaging, Vol. L, 1985.
Copyright CRC Press, Inc., Boca Raton,
FL.; 1.19c: Hank Morgan/Science
Source/Photo Researchers, Inc.; 1.19d:
Monte S. Buchsbaum, M.D.
Chapter2
Opener: M. M. Michelle del
Guerico/Photo Researchers, Inc.;
2.2b: Kent M. Van De Graaff;
2.3b: From C. D. Haagensen, Diseases
of the Breast, 2/ed, 1974, W. B. Saunders
Company; 2.4: Dr. Sheril D. Burton;
2.7a: Joan L. Cohen/Photo
Researchers, Inc.; 2.7b John D.
Cunningham/Visuals Unlimited;
2.7c: James L. Shaffer; 2.7d:
L. Lee Rue III/Photo Researchers,
Inc.; 2.7e: Cyril Toker/Photo
Researchers, Inc.; 2.7f: Tom
Hollyman/Photo Researchers, Inc.;
2.11: Dr. Sheril D. Burton;
2.12a, 2.12b, 2.12c: McGraw-Hill
Higher Education, Inc./Rebecca Gray,
photographer/Don Kincaid,
dissections.
Chapter 3
Opener: Maryann Fittipaldi; 3.5a,
3.5b: Reproduced from R. G. Kessel
and R. H. Kardon, Tissues and Organs:
A Text Atlas of Scanning Electron
Microscopy, W. H. Freeman and Co.
1979; 3.6: BioPhoto Associates/
Photo Researchers, Inc.; 3.7a: Keith
R. Porter; 3.8: David M. Phillips/
Visuals Unlimited; 3.9: Keith R.
Porter; 3.10: K. G. Murti/Visuals
Unlimited; 3.11: David M. Phillips/
Visuals Unlimited; 3.13: From Joseph
G. Gall, Microtube fine structure,
Journal of Cell Biology, 31: (1966)
pp. 639643; 3.15: Don Fawcett/
Photo Researchers, Inc.; 3.16a, 3.16b:
Stephen L. Wolfe; 3.17: CNRI/
SPL/Photo Researchers, Inc.;
3.21: Edwin A. Reschke;
3.22: Tony Brain/SPL/Photo
Researchers, Inc.
Chapter 4
Opener: Mauritius GMBH/
Phototake; 4.1a: Edwin A. Reschke;
4.1b: Dr. Kerry L. Openshaw;
4.2b: Ray Simons/Photo Researchers,
Inc.; 4.3b: Edwin A. Reschke/Peter
Arnold; 4.4b, 4.5b, 4.6b, 4.7b,
4.8b, 4.9b: Edwin A. Reschke;
4.10b: CNRI/SPL/Photo Researchers,
Inc.; 4.13b: BioPhoto Associates/
Science Source/Photo Researchers,
Inc.; 4.14b, 4.15b: Edwin A.
Reschke; 4.16b: Biology Media/
Photo Researchers; 4.17b, 4.18b,
4.19b, 4.20b, 4.21b, 4.22b: Edwin
A. Reschke/Peter Arnold; 4.23a, b:
Kent M. Van De Graaff; 4.24b:
Edwin A. Reschke; 4.26a, b, c:
Edwin A. Reschke; 4.27a: John D.
4.27b: 1984 Martin M. Rotker/
Photo Researchers, Inc.
Chapter 5
Opener: Kathy Sloane/Photo
Researchers, Inc.; 5.2: Astrid &
Hanns-Frieder Michler/Science Photo
Library/Photo Researchers, Inc.;
5.3: J. Burgess/SLP/Photo
Researchers, Inc.; 5.4: Victor B.
Eichler; 5.7: Dr. Sheril D. Burton;
5.8: James M. Clayton; 5.10: Lester
V. Bergman & Associates, Inc.;
5.11a: World Health Organization;
5.11b: George P. Bogumill;
5.12a, b: Dr. Sheril D. Burton;
5.13a: Michael Abbey/Photo
Researchers, Inc.; 5.13b: Kerry L.
Openshaw; 5.14b, 5.16: John D.
5.19a: James Stevenson/SPL/Photo
Researchers, Inc.; 5.19b: Zeva
Oelbaum/Peter Arnold; 5.19c:
Dr. P. Marazzi/Photo Researchers,

Inc.; 5.20a: Dorte Groning/Tierbild
Okapia/Photo Researchers, Inc.;
5.20b: James Stevenson/Photo
Researchers, Inc..; 5.20c: John
Radcliff/SPL/Photo Researchers, Inc.;
5.22a, b, c: Courtesy of Drs. David A.
Kappel, E. Phillips Polack, and
Marjorie L. Bush, Plastic Surgery, Inc.;
5.23 Dan McCoy/Rainbow;
5.26: Art Wolfe/Tony Stone Images;
5.27 [child]: From Science Year, The
World Book Science Annual 1973
Field Enterprise Education Corporation.
By permission of World Book, Inc.;
[woman]: Black Star; 5.28: Norman
Lightfoot/Photo Researchers, Inc.;
Table 5.1: 1993 Carroll H. Weiss/
Camera M.D. All rights reserved.
Chapter 6:
Opener: Spencer Grant/PhotoEdit;
6.6a, b: BioPhoto Associates/Photo
Researchers, Inc.; 6.8a: From R. G.
Kessel and K. H. Kardon, Tissues and
Organs: A Text Atlas of Scanning
Electron Microscopy 1979 W. H.
Freeman and Company; 6.8b: Victor
B. Eichler; 6.1b: ScienceVU/Visuals
Unlimited; 6.10: Edwin A. Reschke;
6.11: Courtesy of Utah Valley
Regional Medical Center, Dept. of
Radiology; 6.18: Kent M. Van De
Graaff; 6.22a, b, 6.34, 6.36: Courtesy
of Utah Valley Regional Medical
Center, Dept. of Radiology;
6.41: Blaine L. Hirsche; 6.42: CNRI/
SPL/Photo Researchers, Inc.; Clinical
Practicum 6.1, 6.2: Courtesy of
University of California, San Diego,
Department of Radiology
Chapter 7
Opener: Richard Hutchings/Photo
Researchers, Inc.; 7.3: Kent M. Van
De Graaff; 7.8: Dr. Sheril D. Burton;
7.10, 7.12, 7.17: Courtesy of Utah
of Radiology; 7.19a: Dr. Sheril D.
Burton; 7.19b: Courtesy of Utah Valley
Radiology; 7.21a: Kent M. Van De
Graaff; 7.21: Dr. Blayne L. Hirsche;
7.22: Kent M. Van De Graaff; 7.24e:
Courtesy of Eastman Kodak Company;
Clinical Practicum 7.1, 7.2: Courtesy of
Chapter 8
Opener: Daemmrich/The Image
Works; 8.5: Courtesy of Utah Valley
Radiology; 8.6: Paolo Koch/Photo
Researchers, Inc.; 8.18ah: Dr.
Sheril D. Burton; 8.19a: Kent M.
Van De Graaff; 8.19b, c: Dr. Sheril

D. Burton; 8.19d: Kent M. Van De
Graaff; 8.19e: Dr. Sheril D. Burton;
8.20a, b: Kent M. Van De Graaff;
8.26, 8.28, 8.32: From Clinical
Anatomy Atlas, R. A. Chase, et al.,
Mosby 1996 (Class Project, Stanford
University); 8.34: McGraw-Hill
photographer/Don Kincaid, dissections;
8.35a: Kent M. Van De Graaff;
8.35b: Lester V. Bergman &
Associates, Inc.; 8.36a: James
Stevenson/SPL/Photo Researchers,
Inc.; 8.36: CNRI/SPL/Photo
Researchers, Inc.; 8.37, 8.38ad:
SIU School of Medicine; Clinical
Department of Radiology;
8.40: Kent M. Van De Graaff
Chapter 9
Opener: Michael
Tamborrino/Medichrome; Table 9.1:
Kent M. Van De Graaff; 9.6b:
Edwin M. Reschke; 9.9a, b: Dr.
H. E. Huxley; 9.9c: From R. G. Kessel
and R. H. Kardon. Tissues and Organs:
A Text-Atlas of Scanning Electron
Microscopy 1979 W. H. Freeman and
Company; 9.10: Dr. H. E. Huxley;
9.12b: BioPhoto Associates/Photo
Researchers, Inc.; 9.15: Dr. Sheril D.
Burton; 9.52: John Bird Photography,
Clinical Practicum 9.1, 9.2: Courtesy
of University of California, San Diego,
Chapter 10
Opener: Melanie Brown/PhotoEdit;
10.1: Stock Montage; 10.4a, b,
10.5, 10.6a & b, 10.7, 10.8:
Dr. Sheril D. Burton; 10.9a: Kent
M. Van De Graaff; 10.9b, 10.10a, b:
Dr. Sheril D. Burton; 10.11, 10.12:
From S. D. Vidic and F. R. Suarez,
Photographic Atlas of the Human Body,
1984 (pl29, p. 38) St. Louis, C. V.
Mosby Co.; 10.13: McGraw-Hill
Higher Education/Karl Rubin,
photographer, 10.14: Kent M. Van
De Graaff; 10.16, 10.17: From S.D.
Vidic and F. R. Suarez, Photographic Atlas
of the Human Body, 1984 (pl71 p. 62)
St. Louis, C. V. Mosby Co.; 10.18
through 22: Kent M. Van De
Graaff; 10.23, 10.24: From S.D. Vidic
and F. R. Suarez, Photographic Atlas of
the Human Body, 1984 (pl115 p. 184)
St. Louis, C. V. Mosby Co.; 10.26:
William C. Brown Communications,
Inc./Rubin; 10.27: From S. D. Vidic
St. Louis, C. V. Mosby Co.;
819

820
Back Matter
Credits
The McGrawHill
Companies, 2001
Credits
10.28: From S.D. Vidic and F. R.

Suarez, Photographic Atlas of the Human
Body, 1984 (pl40 p. 60) St. Louis, C. V.
Mosby Co.; 10.29, 10:30: McGrawHill Higher Education, Inc./Karl
Rubin, photographer; 10.33 through
10.41: Kent M. Van De Graaff;
Body, 1984 (pl29 p. 210) St. Louis,
C. V. Mosby Co.; 10.43: From S.D. Vidic
St. Louis, C. V. Mosby Co.; 10:44:
Kent M. Van De Graaff; 10:45,
10:46: Dr. Sheril Burton; 10:47
through 10:50 Kent M. Van De
Graaff; 10.51a, b: Dr. Kenneth A.
Johnson; 10.51c: Lester Bergman &
Associates; 10.52: McGraw-Hill
Body, 1984 (pl206 p. 336) St. Louis,
C. V. Mosby Co.; 10.54, 10.55,
10.56: McGraw-Hill Higher
Education/Karl Rubin, photographer;
10.57: McGraw-Hill Higher
Education, Inc./Karl Rubin,
photographer; Clinical Practicum
10.1: Courtesy of University of
California, San Diego, Department
of Radiology
Chapter 11
Opener: LUNAGRAFIX/Photo
Researchers, Inc.; 11.3: Edwin A.
Reschke; 11.6b: Dr. John Hubbard;
11.8b: Edwin A. Reschke; 11.17a,
b, c, d: Kent M. Van De Graaff;
11.18: Wellcome Dept. of Cognitive
Neurology/SPL/Custom Medical Stock;
11.39b, c: McGraw-Hill Higher
Education, Inc./Rebecca Gray,
11.40: Per Kjeldsen, University of
Michigan, Ann Arbor; Clinical
Practicum 11.1: Courtesy of
Chapter 12
Opener: Courtesy of Utah Valley
Regional Medical Center, Provo, UT;
12.2: From R. G. Kessel and R. H.
Kardon, Tissues and Organs: A TextAtlas of Scanning Electron Microscopy
1979 W. H. Freeman and Company;
12.5b: McGraw-Hill Higher
12.16, 12.26: McGraw-Hill Higher
12.34: Photographed by Dr. Sheril D.
Burton assisted by Dr. Douglas W.
Hacking; Clinical Practicum 12.1:
Courtesy of University of California,
San Diego, Department of Radiology
Chapter 13
Opener: Corbis/Digital Stock/
Business & Agriculture, Vol. 101;
Clinical Practicum 13.1: Courtesy of

Chapter 14
Opener: John Radcliff
Hospital/SPL/Photo Researchers;
14.9: John Cunningham/Visuals
Unlimited; 14.13: SIU/Nawrocki
Stock Photo, Inc.; 14.14: Fred
Hossler/Visuals Unlimited; 14.17:
BioPhoto Associates/Photo
Researchers, Inc.; 14.20: Edwin A.
Reschke; 14.22: R. Calentine/Visuals
Unlimited; 14.26a: Richard Hutchings/
Photo Researchers, Inc.; 14.26b:
Bettina Cirone/Photo Researchers,
Inc.; 14.26c, 14.27, 14.28: Lester
V. Bergman & Associates, Inc.;
Clinical Practicum 14.1: Courtesy
of University of California, San Diego,
Chapter 15
Opener: The Stock
Shop/Medichrome; 15.5b: 1993
Discover Magazine, reprinted with
permission; 15.10, 15.13: Dr. Sheril
D. Burton; 15.14a: Carroll H.
Weiss/Camera M.D. Studios, Inc.;
15.16b: P. N. Farnsworth/University of
Medicine & Dentristy, New Jersey
Medical School; 15.19b: Per
Kjeldsen, University of Michigan, Ann
Arbor; 15.21a: Kent M. Van De
Graaff; 15.21b: A. L. Blum/Visuals
Unlimited; 15.29, 15.30a: Kent M.
Van De Graaff; 15.30b: Southern
Illinois University; 15.35: From R. G.
Kessel and C. Y. Shih, Scanning
Electron Microscopy in Biology, Copyright 1976, Springer-Verlag; 15.39:
Dean E. Hillman; 15.34: Courtesy of Dr.
Stephen Clark; Clinical Practicum 15.1:
Chapter 16
Opener: SuperStock; 16.4: Bill
Longcore/Science Service/Photo
Researchers, Inc.; 16.9: McGrawHill Higher Education/Karl Rubin,
photographer; 16.18: Blair
Seitz/Photo Researchers, Inc.; 16.22b:
From Practische Inteleedkunde from J.
Dankmeyer, H. G. Lambers, and
J. M. F. Landsmearr. Bohn, Scheltma
and Holkema; 16.25b: From Practische
Outleedkunde from J. Dankmeyer and
J. M. F. Landsmearr. Bohn, Scheltma
and Holkema; 16.29, 16.39:
McGraw-Hill Higher Education,
Inc./ Rebecca Gray, photographer/Don
Kincaid, dissections; 16.44: Edwin
A. Reschke; 16.46a: Kent M. Van
De Graaff; 16.46c, 16.47c: Visuals
Unlimited; 16.49: Richard Menard;
16.52a: Donald S. Bain; 16.55a:
American Lung Association; 16.55b:
Lewis Lainey; 16.56b: Courtesy of
Utah Valley Regional Medical Center,
Department of Radiology; Clinical
Department or Radiology
Chapter 17
Opener: Barry Slaven/Medichrome;
17.5: CNRI/SPL/Photo Researchers,
Inc.; 17.8c: CNRI/Phototake, Inc.;
17.10a: John D. Cunningham/
Visuals Unlimited; 17.10b: Edwin
A. Reschke; 17.12: John Watney
Photo Library; 17.13: Courtesy of
Eastman Kodak; 17.16: David
Phillips/Visuals Unlimited; 17.18:
McGraw-Hill Higher Higher Education
Group; 17.23: From H. Comroe Jr.,
Physiology of Respiration, 1974,
Yearbook Medical Publishers, Inc.;
17.25: SIU/Photo/Researchers, Inc.;
17.28a: Kent M. Van De Graaff;
17.29: Edward C. Vasquez/R.T.C.R.T./
Dept. of Radiologic Technology,
Los Angeles City College; 17.32a, b:
Martin M. Rotker; 17.33: Martin M.
Rotker/Science Source/Photo Researchers,
Inc.; Clinical Practicum 17.1, 17.2:
Courtesy of University of California, San
Diego, Department of Radiology
Chapter 18
Opener: Courtesy of The Wellcome
Trust Medical Library, London;
18.12: BioPhoto Associates/Photo
Researchers, Inc.; 18.14 Edwin A.
Reschke; 18.15: Courtesy of Utah
Medical Center Department of
Radiation; 18.16a, b: Kent M. Van
De Graaff; 18:18: Edwin A.
Reschke; 18.22: Manfred Kage/Peter
Arnold, Inc.; 18.24: Keith R. Porter/
D. H. Alpers and D. Seetharan New
England of Medicine 296/19771047;
1826: From W. A. Sodeman and T. M.
Watson, Pathologic Physiology, 6/ed
W. B. Saunders Company; 18.27:
Lester V. Bergman & Associates, Inc.;
18.29a: Edwin A. Reschke; 18.29b:
From R. G. Kessel and R. H. Kardon.
Tissues and Organs: A Text-Atlas of
Scanning Electron Microscopy, 1979
W. H. Freeman and Company; 18.31c:
Inc./Rebecca Gray, photographer/Don
Kincaid, dissections; 18.32c: Victor
B. Eichler, PhD.; 18.35a: Carroll M.
Weiss/Camera M.D. Studios; 18.35b:
Dr. Sheril D. Burton; 18.37: Kent
M. Van De Graaff; 18.37d: John Van
De Graaff, DDS & Larry S. Pierce,
DDS, both of Northwestern University;
Clinical Practicums 18.1, 18.2:
Kent M. Van De Graaf
Chapter 19
Opener: John Callan/Shout Pictures;
19.2: SPL/Photo Researchers, Inc.;
19.8b: David M. Phillips/Visuals
Unlimited; 19.9a: Daniel Friend from
William Bloom and Don Fawcett,
Textbook of Histology, 10/ed.
W. B. Saunders Co.; 19.11: Per
H. Kjeldsen, University of Michigan,
Ann Arbor; 19.12: Kent M. Van De
Graaff; 19.14: Bruce Iverson/Visuals
Unlimited; 19.20: SIU/Peter
Arnold, Inc.; Clinical Practicums
19.1, 19.2: Kent M. Van De Graaff
Chapter 20
Opener: Custom Medical Stock;
20.2: Landrum Shettles; 20.7b:
BioPhoto Associates/Photo
Researchers, Inc.; 20.8: Francis
Leroy/Biocosmos/SPL/Photo
Researchers, Inc.; 20.9, 20.10:
Edwin A. Reschke; 20.11, 20.12:
Manfred Kage/Peter Arnold, Inc.,
20.13: Edwin A. Reschke; 20.15:
Lennart Nilsson/A Child Is
Born/Bonniefrlagen AB; 20.20:
Center for Disease Control Atlanta;
Clinical Practicum 20.1: Courtesy of
University of Californa, San Diego,
Chapter 21
Opener: Howard Socherak; 21.4
Edwin A. Reschke; 21.5a: David
M. Phillips/Visuals Unlimited; 21.5b:
Fram R. J. Balandau, A Textbook of
Histology, 10/ed W. B. Saunders Co.;
21.6: Edwin A. Reschke/Peter
Arnold; 21.7: Dr. Landrum Shettles;
21.10, 21.14: Edwin A. Reschke;
21.17: BioPhoto Associates/Photo
Researchers Inc.; 21.20: Courtesy of
Utah Valley Regional Medical Center,
Dept. of Radiology; 21.25: SIU
School of Medicine; 21.26a: McGrawHill Higher Education, Inc./Vincent
Ho, photographer; 21.26bg:
Inc./Bob Coyle, photographer; 21.26h:
Hank Morgan/Photo Researchers,
Inc.; Clinical Practicum 21.2:
Chapter 22
Opener: Custom Medical Stock;
21.1b: David Phillips/Visuals
Unlimited; 22.2: Roy Luciano
Zamboni, from Roy Greep and Leon
Weiss Histology, 3/ed McGraw-Hill
1973; 22.6: Courtesy of Ronan
ORahilly, M.D., Carnegie Institute of
Washington, Department of
Embryology, Davis Division; 22.11,
22.20a: Donald Yeager/Camera
M.D. Studios; 22.20b: Dr. Landrum
Shettles; 22.20c: Petit
Format/Nestle/Science Source/Photo
Researchers, Inc.; 22.20df: Donald
Yeager/Camera M.D. Studios; 22.20g:
Petit Format/Nestle/Science
Source/Photo Researchers, Inc.;
22.21a: A. Tsiaras/Photo
Researchers, Inc.; 22.21b: Courtesy of
Kris Queck; 22.25: Courtesy of Utah
of Radiology; 22.28: J. M. Tanner, The
Physique of the Olympic Athlete, 1964;
22.29: Babu, A. Hirschhorn, K. A.
Guide to Human Chromosomes Reflects
3/ed. White Plains, NY; 22.35:
Lester V. Bergman Associates, Inc.;
22.38: Gregory Dellore, M. D. and
Steven L. Clark, M.D.; Clinical
Practicum 22.1: Courtesy of

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Page numbers followed by f and t indicate figures and tables, respectively.
A bands, 241, 242f, 243f, 245f
Abdomen, 3637
developmental conditions involving, 336
diseases of, 337
internal anatomy of, 315, 317f, 318f
of neonate, 299t
quadrants of, 37, 39f
subdivisions of, 3637, 38f, 38t
surface anatomy of, 312313, 312f
trauma to, 336337
veins of, 579
Abdominal aorta, 566, 567t, 569f, 581f
Abdominal aortic plexus, 441
Abdominal cavity, 40f, 41, 41f, 44f
arteries of, 569f
serous membranes of, 43
Abdominal muscles. See also specific muscles
in respiration, 620
Abdominal organs, 638f
Abdominal reflexes, 430t, 431f
Abdominal thrust maneuver, 608, 627, 628f
Abdominal wall, muscles of, 258, 259f, 260t
Abdominopelvic cavity, 40f, 41, 41f, 42f, 44f
Abducens nerve (VI), 360f, 402f, 404t, 406,
408f, 413t
Abduction, 207, 208f, 211f, 212f
Abductor digiti minimi muscle, 271f, 274f,
275, 275t, 290f, 291f, 325f
Abductor hallucis muscle, 285f, 291f, 329f
Abductor pollicis brevis muscle, 271f, 274f,
275, 275t, 325f
Abductor pollicis longus muscle, 271f, 272,
272f, 273t, 321f, 325f
tendon of, 274f
Abortion, 748, 762
Abrasion, 125f
Abscess, 596
Absorption, 114, 635, 653
Accelerator fibers, 377
Accessory digestive organs, 635
Accessory hemiazygos vein, 576, 577f
Accessory nerves (XI), 309f, 402f, 405t, 412,
412f, 413t, 621t
Accessory nipples, 25f
Accessory olivary nuclei, 377
Accessory pancreatic duct, 654f, 663f
Accessory reproductive glands, 708709
Accessory structures, of eyes, 499502
Accommodation (sensory), 490
Accommodation (visual), 510, 514
Acetabular labrum, 220, 220f
Acetabular notch, 181f, 183
Acetabulum, 181, 181f, 182f
Acetylcholine (ACh), 244, 358, 396, 444,
445, 446f
ACh. See Acetylcholine
Achilles reflex. See Ankle reflex
Achilles tendon. See Tendo calcaneus
Achondroplasia, 169
Acid(s), 51
Acid-base balance, 51, 85
Acid phosphatase, 709
Acinar cells, 664f
Acinar glands, 88f, 89t
Acne, 107t, 128
Acoustic nerve. See Vestibulocochlear nerve
Acquired heart disease, 594
Acquired immune deficiency syndrome
(AIDS), 720, 721t
Acromegaly, 166, 334, 462, 480, 481f
Acromial extremity, 173, 175f
Acromioclavicular joint, 173, 225t
Acromioclavicular ligament, 216f

Acromion, 173, 175f, 311f, 319f, 320f
Acrosomal reaction, 756
Acrosome, 704705, 706f, 756
ACTH. See Adrenocorticotropic hormone
Actin myofilaments, 243f
Action potential, 357, 357f
Active transport, 55t
Acupuncture, 8, 9f
Acute pain, 492
Acute prostatitis, 722
Acute purulent otitis media, 533
Acute renal failure, 693
Adams apple. See Laryngeal prominence
Addisons disease, 483
Adduction, 207, 208f, 211f, 212f
Adductor brevis muscle, 276f, 277, 278f,
281t, 282f
Adductor hallucis muscle, 290f
Adductor hiatus muscle, 278f
Adductor longus muscle, 235f, 276f, 277,
278f, 279f, 281f, 281t, 328f, 331f
Adductor magnus muscle, 235f, 276f, 277,
279f, 280f, 281f, 281t, 282f, 328f
Adductor minimus muscle, 277
Adductor pollicis muscle, 271f, 274f, 275,
275t, 325f
Adenine, 65, 68
Adenohypophysis, 373, 460, 461f, 462f,
464t, 477
Adenoidectomy, 608
Adenoids, 584f, 607
Adenylate cyclase, 459, 459f
ADH. See Antidiuretic hormone
Adipocytes, 93, 445t
Adipose cells. See Adipocytes
Adipose tissue, 9394, 94f, 95t, 107f, 297, 298f
Adolescence, 778779, 779t, 781t
Adrenal cortex, 440, 457t, 471, 473t, 478
Adrenal glands, 456, 456f, 472f, 473f
autonomic nervous system and, 440
description of, 471
development of, 478
disorders of, 483
functions of, 471473
Adrenal hormones, 473t
Adrenaline. See Epinephrine
Adrenal medulla, 440, 457t, 471
autonomic nervous system effects on, 445t
development of, 478
tumors of, 483
Adrenal medullary hormones, 473t
Adrenergic stimulation, 444
Adrenocorticotropic hormone (ACTH),
462, 464t
Adulthood, 779781, 781t
Adventitia. See also Tunica externa
of stomach, 650f
of ureters, 684
of urinary bladder, 687
Afferent glomerular arterioles, 677, 680f, 681f
Afferent lymphatic vessels, 583
Afferent nerves, 356f
Afferent neuron, 352
Afterbirth, 775
Aging, 7374
of muscles, 288289, 292f
of skeletal system, 167168, 167f
of skin, 127, 127f
Aging (senile) atrophy, 72
Agranulocytes, 542, 544t
AIDS. See Acquired immune deficiency
syndrome
Air movements, nonrespiratory, 621, 622t
Air pressure, hearing and, 519
Air spaces, 622t
Airways, 622t
Akinesia, 389
Alar cartilage, 305f, 605
Alar nasal sulcus, 304f, 304t
Alar plates, 390
Albinism, 109, 128, 128f
Albumin, 545, 545t
Alcohol
cytologic effects of, 57
intoxication with, 376
liver effects of, 579
Aldosterone, 457t, 471
Alendronate (Fosamax), 168
Alexandrian Era, and history of anatomy, 1011
Alisphenoid, 142f
Allantois, 667, 764
Alleles, 782783
Alligator, brain of, 26f
Allogenic transplant, 544
Allografts, 102
All-or-none contraction, 437
All-or-none response, 357
Alopecia, 128
Alpha cells, 470, 470f
Alpha globulin, 545, 545t
Alpha waves, 368t
Alveolar bone destruction, 669f
Alveolar ducts, 611, 612, 613f
Alveolar process, of maxilla, 150f,
154, 155f
Alveolar sacs, 611, 612, 613f
Alveolar ventilation, 622t
Alveolus, 136t
Alzheimers disease, 396
Amblyopia, 511, 532
Amelia, 192, 337
Amenorrhea, 747
Amine, 457f, 458, 458t, 459
Amino acids, 51
Amnesia, 392
Amniocentesis, 763, 765f, 787
Amnion, 762763
Amniotic cavity, 80, 760
Amniotic fluid, 762763
Amniotic sac, 762763
Amoebic dysentery, 669
Amphiarthroses, 197
Ampulla
of ductus deferens, 699f, 707, 710f
of inner ear, 526f
of uterine tube, 729f, 734
Ampulla of Vater. See Hepatopancreatic ampulla
Amputation, phantom pain and, 492
Amsler grid, 532
Amylase, 654t
Anabolic effects, of androgens, 703t
Anabolic steroids, 703
Anal canal, 635f, 657, 657f, 658f, 667
Anal column, 657, 658f
Anal membrane, 667
Anal pit, 667
Anal region, pH of, 85
Anal triangle, 39f, 700, 701f
Anaphase, 70, 71f, 703, 704, 704t, 705f
Anastomosis, 555
Anatomical dead space, 622t
Anatomical position, 33, 35f
Anatomical snuffbox, 322, 322f
Anatomical terminology, 30
singular and plural forms of, 33t
standardization of, 17
Anatomy
definition of, 2
developmental, 17
gross, 17
historical documentation of, 6t
history of, 121, 5t

case study, 1, 20
microscopic, 17
prescientific period, 24
radiographic, 1819, 18f, 19f
regional approach to, 29
in scientific period, 419
surface, 17
systematic (systemic) approach to, 29
terminology of, 2
in 17th Century, 1516
in 19th Century, 16
Anatomy Lesson of Dr. Tulp, The
(Rembrandt), 15f
Anconeus muscle, 267f, 268, 269f, 270t,
271f, 272f, 321f
Androgens, 472, 698, 703, 703t
Anemia, 541, 542, 592
Anencephaly, 333, 392
Anesthesia, 157, 391392, 775
Aneurysm, 595, 596f
Angina pectoris, 335, 594
Angiocardiogram, 18, 587
Angiography, 18
Angle
of mandible, 149f, 156f, 307f
of rib, 165, 165f
Angle of Louis. See Sternal angle
Angora hair, 117
Angular gyrus, 371, 371f
Angular movements, 207, 208f, 212f
Animal kingdom, 23, 27t
Ankle, 40
muscles of, 281285, 286t
sprain of, 224
Ankle joint. See Talocrural joint
Ankle reflex, 430t, 431f
Ankylosis, 230
Annular ligament, 217, 218f
ANS. See Autonomic nervous system
Ansa cervicalis, 416t
Antagonistic muscles, 238, 239f
Antebrachium, 38
bones of, 177178
internal anatomy of, 325f
muscles of, 267268, 268272, 270t,
271f, 272f, 273t
Anteflexion, of uterus, 748
Anterior, 36t
Anterior arch, of axis, 160f
Anterior body cavity, 40f, 41, 44f
Anterior border, of tibia, 186f
Anterior cavity, of eye, 505f, 508
Anterior cerebral artery, 563, 563f
Anterior cervical triangle, 307, 307f, 308f, 308t
Anterior chamber, of eye, 500f, 505f, 508, 512f
Anterior commissure, 368, 370f
Anterior communicating artery, 563f
Anterior corneal epithelium, 504
Anterior corticospinal tract, 387, 388t, 389t
Anterior cranial fossa, 149f
Anterior crest, of tibia, 185
Anterior cruciate ligament, 221, 221f, 222f
Anterior crural muscles, 281283, 283f
Anterior division
of brachial plexus, 416
of lumbar plexus, 421
of sacral plexus, 421
Anterior ethmoidal foramina, 150f
Anterior fontanel, 144, 145f
Anterior funiculus, 386, 386f
Anterior gluteal line, 182f, 183
Anterior horn, 365f, 384, 386f, 390

822
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Anterior humeral circumflex artery, 565, 565f

Anterior inferior iliac spine, 181183, 181f, 182f
Anterior intercostal arteries, 567f
Anterior interosseous artery, 565, 565f
Anterior interventricular artery, 548f, 551, 552f
Anterior interventricular vein, 548f, 551
Anterior lobe, of cerebellum, 375f
Anterior median fissure, 384, 386f
Anterior membranous ampulla, 519f
Anterior nasal spine, 148f
Anterior pituitary gland. See Pars distalis
Anterior ramus, of spinal nerves, 414, 414f
Anterior reticulospinal tract, 388t, 389t
Anterior root, of spinal nerves, 386f, 414, 414f
Anterior sacral foramina, 162, 181f
Anterior semicircular canal, 519f
Anterior spinocerebellar tract, 388t, 389t
Anterior spinothalamic tract, 388t, 389t
Anterior sternoclavicular ligament, 215, 215f
Anterior superior iliac spine, 181, 181f,
182f, 312f
Anterior surface
of patella, 186f
of stomach, 650
Anterior talofibular ligament, 222, 223f
Anterior tibial artery, 560f, 570, 570f
Anterior tibial vein, 573f, 576, 578f
Anterior tibiofibular ligament, 223f
Anterior tibiotalar ligament, 223f
Anterior trunk, of internal iliac artery, 569f
Anterior vagal trunk, 441
Anterolateral fontanels, 144, 145f
Anteversion, of uterus, 748
Anthropometry, 780781, 782f
Antibodies, 52
Anticodons, 6869, 69f
Antidiuretic hormone (ADH), 457t, 463,
464t, 481
Antihelix, 517f
Antitragus, 517f
Antrum, 729, 730f, 733f
Anus, 330f, 635f, 657, 658f, 699f, 727f, 737f
Anvil, etymology of term, 30
Anxiety, 396
Aorta, 559, 567t, 568f
abdominal portion of, 566, 567t, 569f, 581f
ascending portion of, 314f, 548f, 550,
552f, 559, 561f, 567t
descending portion of, 549f, 560f
thoracic portion of, 566, 567t
Aortic arch, 24, 314f, 549f, 552f, 559561,
560f, 561f, 567t, 590, 591f
Aortic area, 554, 554f
Aortic bodies, 623
Aortic impression, 615f
Aortic (semilunar) valve, 548t, 549f, 550
Apex
of cochlea, 519f
of heart, 314f, 545, 548f, 553f
of lung, 614, 615f
of nose, 304f, 304t
of patella, 185, 186f
Apical ectodermal ridge, 192
Apical foramen, 645, 645f
Apical turn, of cochlea, 520f
Aplastic anemia, 541
Apnea, 622t, 628
Apneustic area, 374, 374f, 623
Apocrine glands, 87, 88f, 89t, 118
Aponeuroses, 236, 317f
Appendectomy, 313
Appendicitis, 656
Appendicular skeleton
bones of, 132t, 134
case study, 172, 191
clinical considerations, 189191
clinical practicum, 193
development of, 192, 192f
muscles of, 234, 263285
Appendix, 635f, 656, 657f, 658f
Appositional bone growth, 137
Aqueduct of Sylvius. See Mesencephalic
aqueduct
Aqueous humor, 508, 512f
Arachnoid, 378f, 379, 379f, 381f

Arachnoid mater, 385f
Arachnoid villi, 378f
Arbor vitae, 370f, 375, 375f
Arches, of foot, 188, 188f
Arcuate artery, 677, 680f
Arcuate fasciculus, 371, 371f
Arcuate line, 182f
Arcuate popliteal ligament, 221, 221f
Arcuate vein, 678, 680f
Areola, 259f, 312f, 739, 739f
Areolar glands, 739
Areolar tissue. See Loose connective
(areolar) tissue
Argentaffin cells, 651
Aristotle, 7, 910, 11f
Arm. See Brachium; Upper extremity
Armpit. See Axilla
Arrector pili muscle, 107f, 113, 116, 445t
Arrhythmia, 587
Arterial blood, 540
Arterial pressure points, 570, 572f
Arteries, 540, 555, 556f, 559571, 560f, 561f,
590, 591f. See also specific arteries
Arteriography, 587
Arterioles, 540, 555, 556f, 557f
Arteriosclerosis, 595
Arthralgia, 230
Arthritis, 60, 167f, 202, 226, 227f
Arthrolith, 230
Arthrology, 197
Arthrometry, 230
Arthroncus, 230
Arthropathy, 230
Arthroplasty, 200, 230
Arthroscopy, 227229, 227f
Arthrosis, 230
Arthrosteitis, 230
Articular capsule. See Joint capsule
Articular cartilage, 137, 202, 206
Articular disc, 202, 206
Articular surface
of fibula, 186f
of head of rib, 165f
of palatine bone, 156f
of tubercle, 165f
Articulating surface, of bones, 136t
Articulations, 196232. See also Joints
Artificial implantation, 785
Artificial insemination, 714
Artificial ligaments, 200
Artificial respiration, 629f
Arytenoid cartilage, 609
Ascending aorta, 314f, 548f, 550, 552f, 559,
561f, 567t
Ascending colon, 318f, 635f, 638f, 656, 657f
Ascending corticospinal tract, 387f, 388t
Ascending limb, of nephron loop, 680f,
682, 683t
Ascending lumbar veins, 576
Ascending pharyngeal artery, 563
Ascending tracts, of spinal cord, 384, 389t
Aspirin, 358
Association fibers, 368, 369f
Association neurons, 352
Asthma, 611, 628
Astigmatism, 531, 531f
Astrocytes, 350f, 351, 351t
Astrocytomas, 394
Atheroma, 595
Atherosclerosis, 595, 595f
Athletes bradycardia, 592
Athletes foot, 128
Atlantoaxial joint, 225t
Atlanto-occipital joint, 161, 210, 225t
Atlas, 158f, 160f, 161, 204f
Atoms, 28
Atresia, 531
Atretic, 730
Atrial septal defect, 593, 593f
Atrioventricular bundle (AV bundle),
551552, 553f
Atrioventricular node (AV node), 551552, 553f
Atrioventricular valves (AV valves), 547, 548t
Atrium, 547, 589, 590t

Atrophy, 72, 102, 286
aging (senile), 72
disease, 72
disuse, 72, 102
muscle, 102
senescence, 102
Atropine, 445
Audiology, 527
Audiometry, 527
Auditory area, 366f
Auditory impairment, 533
Auditory nerve. See Vestibulocochlear nerve
Auditory ossicles, 24, 132, 132t, 157, 158f,
517, 518f, 529
Auditory tube, 24, 517, 517f, 518f, 529,
605f, 607, 641f
Auerbachs plexus. See Myenteric plexus
Auricle
of ear, 301, 302, 516, 517f, 529
of heart, 547
Auricular abnormalities, 530531
Auricular region, of face, 301, 302303
Auricular surface, 162, 163f, 182f, 183
Auscultation, 35, 297
Australoid (Ngatatjara man), 27f
Autograft, 102, 124
Autonomic dysreflexia, 450451
Autonomic ganglion, 435
Autonomic motor fibers, 354, 356f
Autonomic motor nerve, 345t
Autonomic nervous system (ANS), 344f,
345, 345t, 434453
control of, 448450
dual innervation, 446447
functions of, 444448, 445
neurotransmitters of, 444, 446f
organization of, 435436
parasympathetic division of, 436,
440443, 442f, 443t, 444t
and somatic nervous system, 435, 435t, 436f
structure of, 438443
sympathetic division of, 436, 438f,
439f, 440f, 441f, 442f,
443t, 444t
visceral effector organs of, 436438
Autonomic reflexes. See Visceral reflexes
Autopsy, 1819
Autosomal chromosomes, 782
Autotransplant, 544
AV node. See Atrioventricular node
Avulsion, 125f, 190
AV valves. See Atrioventricular valves
Axial muscles, 265267
Axial skeleton, 132134
bones of, 132t
development of, 141
muscles of, 234, 250263
Axilla, 36, 311f, 312f, 319, 320f
Axillary artery, 560f, 565, 565f
Axillary fossa, 36
Axillary lymph nodes, 584, 584f
Axillary nerve, 324f, 416, 418f, 418t
Axillary process, of breast, 739
Axillary region, 36
Axillary vein, 573f, 575, 575f
Axis, 158f, 160f, 161, 204f
Axon, 101, 348, 349f, 350f, 351, 354f
Axonal transport, 348
Axon plate, 247f
Axon terminal, 244, 247f, 358
Axoplasmic flow, 348
Aye-aye, hands of, 26f
Azygos vein, 549f, 575576, 577f
Babinskis reflex, 387, 430t, 431f
Back, surface anatomy of, 309310, 310f
Backbone. See Vertebral column
Back pain, 226
Bacterial endocarditis, 335336, 594
Bacteriuria, 692
Baculum, 713
Balance, 516527. See also Equilibrium
Ball-and-socket joint, 203, 205f, 205t
Baroreceptors, 490, 561
Bartholins glands. See Vestibular glands
Basal cell carcinoma, 120, 123
Basal metabolic rate (BMR), 480
Basal nuclei, 365f, 368369, 370f
Basal plates, 390
Basal turn, of cochlea, 520f
Base
of heart, 545
of lung, 614
of metacarpals, 178, 179f
of metatarsals, 186, 187f, 188f
of phalanges, 180f
of sacrum, 181f
Basement membrane, 80
Bases, 51, 68
Basilar artery, 360f, 561, 562f, 563f
Basilar membrane, 520, 520f, 521f
Basilar part, of occipital bone, 154f
Basilic vein, 321f, 573f, 575, 575f
Basle Nomina Anatomica (BNA), 17
Basophils, 542, 542f, 544t
Ba spirit, 8
BBB. See Blood-brain barrier
Bedsores, 111, 112f
Behavioral effects, of androgens, 703t
Bells palsy, 250, 409
Belly, of muscle, 236
Benign neoplasms, 73
Benign prostatic hypertrophy (BPH), 709, 722
Benign viral infections, 119
Berengario, Jacopo, 13, 15
Beta cells, 470, 470f
Beta globulin, 545, 545t
Beta waves, 368t
Biaxial joint, 202
Biceps brachii muscle, 235f, 239f, 259f,
264f, 267, 268f, 270f, 270t, 313f,
319f, 320f
Biceps femoris muscle, 235f, 277, 280f, 281,
281f, 282f, 282t, 284f, 287f, 289f,
328f, 328f, 330f, 331f, 333f
tendon of, 328f
Biceps reflex, 430t, 431f
Bicipital aponeurosis, 268f
Bicornuate uterus, 746, 746f
Bicuspid area, 554, 554f
Bicuspids. See Premolars
Bicuspid valve. See Left atrioventricular valve
Bifid spinous process, 160
Bifurcate ligament, 223f
Bile, 662, 663f
Bile canaliculi, 662, 662f, 663f
Bile duct, 663f
Bile ducts, 662
Biliary calculi. See Gallstones
Biliary ductule, 662f, 663f
Binocular vision, 499
Biology, definition of, 2
Biomechanics, 197
of body movements, 210213
Biopsy, 102
Bipedal locomotion, 2527, 183
Bipolar neurons, 352, 356f, 506
Birth, effects of lysosomes on, 59
Birth control. See Contraception
Bitter taste, 497, 498f
Blackhead. See Comedo
Blanching, 592
Blastocoel, 80
Blastocyst, 80, 734, 757, 758f, 760t
Blastocyst cavity, 757
Blastomeres, 757
Bleeding, control of, 596597, 597f
Blind spot, 507, 511f
Blister, 128
Blood, 98, 99f, 540545. See also specific
components
analysis of, 592593
circulatory routes of, 550551

Back Matter
Index
The McGrawHill
Companies, 2001
Index
color of, 541
disorders of, 592593
Blood-brain barrier (BBB), 351, 355f, 383385
Blood-cerebrospinal fluid barrier, 382
Blood clotting, 125126, 126f, 542
Bloodletting, 1
Blood plasma, 98, 540, 541f, 545
Blood platelets, 125126, 126f
Blood pressure, 558559
measurement of, 558f, 559f
variation in, 552
Blood vessels, 445t, 540, 555559. See also
specific types
BMR. See Basal metabolic rate
BNA. See Basle Nomina Anatomica
Body
of axis, 160f
of clavicle, 175f
of corpus callosum, 361f
of epididymis, 707f
of femur, 184, 184f
of fibula, 186f
of humerus, 173, 175f, 176f
of hyoid bone, 157, 157f
of ischium, 181f
of mandible, 156f, 157
of metatarsals, 186, 187f
of nail, 117, 117f
of pancreas, 664f, 668
of penis, 710, 710f, 711, 716
of phalanges, 180f
of pubis, 183
of radius, 177f, 178
of rib, 165, 165f
of sphenoid bone, 152
of sternum, 164, 164f, 311f
of stomach, 649, 649f, 650f
of tibia, 186f
of tongue, 642f
of ulna, 177f
of uterus, 729f, 734
of vertebrae, 159, 160f, 161f, 162f
Body cavities, 40f, 4142, 41f, 44f
Body composition, 781t
Body membranes, 4245, 85
Body organization, 2829, 29f
Body proportions, 777, 777f, 778f, 780781,
780f, 782f
Body regions, 3540, 37f
Body support, muscular system and, 234
Body temperature
variation in, 552
water in, 51
Boil, 128, 334
Bolus, 644
Bone(s). See also Skeletal system; specific bones
demineralization of, 97, 97f
shapes of, 135136, 136f
surface features of, 136t
Bone cells, 138, 138f, 139f
Bone growth, 140144, 140f, 141, 141f
Bone-lining cells, 138
Bone marrow, 134, 137, 584f
Bone marrow transplant, 544
Bone neoplasms, 167
Bone scan, 167, 167f
Bone shaft, 136
Bone tissue, 9598, 139f
Bony callus, 191
Bony labyrinth, 519, 519f, 529
Botulism, 669
Bowmans capsule. See Glomerular capsule
BPH. See Benign prostatic hypertrophy
Brachial artery, 560f, 565, 565f
pressure point for, 321f, 572f
Brachial block, 416
Brachialis muscle, 235f, 239f, 267, 267f,
268f, 269f, 270f, 270t
Brachial plexus, 257f, 309f, 313f, 324f, 385f,
401f, 416419, 417f, 418t
Brachial vein, 573f, 575, 575f
Brachiocephalic trunk, 314f, 549f, 559561,
560f, 561f, 562f, 567t
Brachiocephalic vein, 574

Brachioradialis muscle, 235f, 267, 267f, 268f,
269f, 270t, 271f, 313f, 320f, 321f,
323f, 325f
Brachioradialis reflex. See Supinator reflex
Brachium, 38
bones of, 173176
internal anatomy of, 313f, 323f, 324f
muscles of, 268f
Brachium pontis, 375f
Bradycardia, 587, 592, 592f
Bradykinin, 447
Braille, 491
Brain, 23, 344, 360f361f, 370f
blood supply to, 561563, 563f, 564f
derivation of, 362t
development of, 346347, 346f, 347f
effects of androgens on, 703t
functions of, 362t
general features of, 358359
language and, 369371, 371f
of newborn, 359
respiratory centers of, 623
sections of, 34f
in taxonomy, 24, 25, 26f
trauma to, 367, 377, 450
Braincase. See Cranium
Brain stem, 306f, 373374
Brain waves, 367368
Breast(s), 312f, 738740, 739f, 748
Breast buds, 779
Breast cancer, 264, 335, 740, 745, 748
Breast self-examination (BSE), 319, 335,
745746, 745f
Breathing, 256
abnormal patterns of, 630
mechanics of, 618621, 619f
regulation of, 623, 623f
Breathing reflex, 776
Breech presentation, 775
Bridge, of nose, 303, 304f, 304t
Broad ligament, 728, 729f, 735
Brocas area. See Motor speech area
Bronchial arteries, 566, 567t
Bronchial segments, 614615
Bronchial tree, 610611, 618t
Bronchial wall, 606f
Bronchioles, 445t, 611, 612f, 613f
Bronchitis, 628
Bronchogram, 612, 612f
Bronchopulmonary segments, of lungs, 616f
Bronchoscopy, 335
Bronchus, 604f, 610f, 615f
Brunners glands. See Duodenal glands
Brush border, 655
Buccal cavity. See Oral cavity
Buccal surface, of teeth, 642
Buccinator muscle, 250f, 251f, 251t, 253f, 305f
Bulb, of penis, 686f, 710711, 710f
Bulbar conjunctiva, 500f, 501, 501f
Bulboreticulospinal tract, 388t, 389t
Bulbospongiosus muscle, 260, 260t, 261f,
686f, 701f
Bulbourethral ducts, 686f
Bulbourethral glands, 686f, 687, 698, 699f,
700t, 709
Bulbs of Krause, 489t, 491
Bulbus cordis, 590t
Bulimia nervosa, 645
Bundle of His. See Atrioventricular bundle
Bunion, 202
Burns, 72, 105, 122123, 123f, 124f
Bursae, 202, 203f
Bursitis, 226, 338
Buttock, 37
internal anatomy of, 330, 330f
muscles of, 276277
Butt suture. See Plane suture
Cadavers, dissection of, 2, 15
history of, 4f
during Renaissance, 13, 13f
Caffeine, 358
Calcaneofibular ligament, 222, 223f

Calcaneus muscle, 133f, 174f, 185186,
187f, 188f, 287f, 329f
tendon of, 285f
Calcification, 140
Calcitonin, 457t, 466, 467, 467t, 468
Calcitrol, 114
Calcium, 135
deficiency, 97
Calculus. See Urinary stones
Callus, 109, 323f
Calyces, 677f
Canaliculi, 98, 138f, 140
Cancellous bone. See Spongy bone
Cancer, 73, 73f
of breast, 264, 335, 740, 745, 748
cervical, 119
etymology of term, 30
of lung, 615, 630f
pancreatic, 669
of respiratory system, 628
of skin, 110, 112, 119121
uterine, 747
Cancer cells, 584
Canine teeth, 148f, 154, 304f, 640f, 642,
643f, 644t
Capacitation, 756
CAPD. See Continuous ambulatory
peritoneal dialysis
Capillaries, 540, 555557, 556f, 557f, 563
Capillary network, of respiratory system, 613f
Capitate bone, 178, 179f, 180f
Capitulum, 176, 176f
Capoid (Kalahari bushmen), 27f
Capsular space, 680
Carbohydrates, 51, 52, 52t
Carboxypeptidase, 654t
Carbuncle, 128
Carcinogens, 73
Cardia, 649, 650f
Cardiac arrhythmia, 587
Cardiac catheterization, 550
Cardiac center, of medulla oblongata, 377
Cardiac cycle, 551
Cardiac impression, 614, 615f
Cardiac muscle, 99, 100f, 100t, 234, 435,
436437
Cardiac output, 552
Cardiac plexus, 441
Cardiac septum, congenital defects of, 590
Cardiac tamponade, 335
Cardiac veins, 551, 573f
Cardiac vessels, 547
Cardinal ligament, 735
Cardiogenic area, 589
Cardiovascular assessment, 587592
Cardiovascular regulation, hypothalamus
and, 372373
Cardiovascular system, 540
Carina, 610, 610f
Carotene, 110
Carotid bodies, 561, 623
Carotid canal, 146t, 148f, 151
Carotid hemorrhage, 307
Carotid sheath, 307, 574
Carotid sinus, 560f, 561, 562f
Carotid triangle, 308f, 308t
Carpal bones, 133f, 174f, 179f, 180f, 181t
Carpal extensor retinaculum, 325f
Carpal tunnel syndrome, 338
Carpometacarpal joint, 225t
Carpus, 38, 178
articulation of, 204f
flexion of, 269271
muscles that move, 268272, 273t
surface anatomy of, 323f
Carrier molecules, 54
Cartilage, 95, 97t
alar, 305f, 605
articular, 137, 202, 206
arytenoid, 609
corniculate, 609, 609f
costal, 95, 133f, 164f, 165, 165f, 174f
cricoid, 142f, 306, 306f, 609, 610f
cuneiform, 609, 609f
823
elastic, 95, 97f

epiglottic, 608
fibrocartilage, 95, 96f, 97f
greater alar, 305f, 605f
hyaline, 95, 96f, 97f
hyoid, 142f
lateral, 605, 605f
lateral alar, 305f
Meckels, 142f
septal, 605
styloid, 142f
thyroid, 142f, 306, 306f, 307f, 608, 610f
tracheal, 610f, 611f
zone of calcified, 143
zone of hypertrophic, 143
zone of proliferating, 142
zone of resting, 142
Cartilaginous joints, 197, 199200, 205t
Case study
body organization, 22, 45
clinical
appendicular skeleton, 172, 191
articulations, 196, 229
autonomic nervous system, 434, 451
body organization, 22, 45
central nervous system, 343, 396
circulatory system, 537, 598
cytology, 48, 74
developmental anatomy, 789
digestive system, 634, 671
endocrine system, 454, 483
female reproductive system, 725
growth and development, 754
histology, 77, 103
history of anatomy, 1, 20
integumentary system, 105, 127
male reproductive system, 697, 722
muscular system, 233, 289
peripheral nervous system, 400, 430
regional anatomy, 296, 339
respiratory system, 602, 630
sensory organs, 487, 533
skeletal system, 131, 168
urinary system, 675, 694
Cataracts, 530, 531532
Catecholamines, 472
Catheterization, of heart, 587
CAT scan. See Computerized axial tomography
Cauda equina, 384, 385f
Caudal. See Inferior
Caudate lobe, of liver, 660661, 661f
Caudate nucleus, 365f, 368, 370f
Causasoid, 27f
Cavernous sinus, 574f
Cavities. See Dental caries
CCK. See Cholecystokinin
Cecal diverticulum, 666
Cecum, 635f, 638f, 656, 657f
Celiac plexus, 440, 441
Celiac trunk, 560f, 566, 567t, 568f
Celial ganglia, 440
Cell(s), 28
components of, 53, 53t, 54f
diversity of, 49
as functional units, 49
shapes of, 49, 50f
sizes of, 49
structure of, 5263, 53t, 54f
surfaces of, 49
trauma to, 72
Cell body, 101, 348, 349f, 350f
Cell cycle, 6570, 70f
Cell division, 6970
Cell membrane, 5357, 53t, 54f, 55f
ionic charge of, 54
movement through, 55t, 56f
permeability of, 54
pressure differences between, 54
structure of, 54
Cell theory, 16, 49
Cellular adaptations, 7072
Cellular biology, 17
Cellular chemistry, 5052, 52t
Cellular level, organization at, 28
Celsus, 11, 13

824
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Cementum, 644, 645f

Center, in reflex arc, 427, 427f
Central artery, 505f, 507
Central canal
of bone tissue, 138140
of spinal cord, 358, 381, 382f, 384, 386f
Central (haversian) canal, 9798
Central chemoreceptors, 623
Central nervous system (CNS), 343399,
344f, 345, 345t, 359f. See also
Autonomic nervous system; Nervous
system; Peripheral nervous system
developmental disorders of, 392
injuries to, 392393
neoplasms of, 394
relationship with peripheral nervous
system, 355f
Central nervous system, clinical practicum, 396
Central sulcus, 360f, 363, 364f, 366f
Central vein, 505f, 507, 661, 662f, 663f
Central vessels, of retina, 516
Centrioles, 54f, 61, 61f
Centromere, 65
Centrosome, 53t, 61, 61f
Cephalic. See Superior
Cephalic phase, of gastric secretion, 651t
Cephalic vein, 313f, 321f, 573f, 575, 575f
Cephalization, 624
Cerebellar arteries, 563f
Cerebellar cortex, 375
Cerebellar peduncles, 375, 376f
Cerebellar veins, 574
Cerebellum, 306f, 360f, 361f, 362t, 375,
375f, 376f
and autonomic nervous system, 449, 450f
trauma to, 376
Cerebral angiogram, 391
Cerebral aqueduct. See Mesencephalic
aqueduct
Cerebral arterial circle, 561
Cerebral arteries, 563f
Cerebral cortex, 363, 365f, 367f, 378f
Cerebral embolism, 595
Cerebral gyri, 363, 364f
Cerebral hypophysis, 373. See also Pituitary gland
Cerebral medulla, 365f
Cerebral palsy, 376, 394
Cerebral peduncles, 362t, 374
Cerebral sulci, 363, 364f
Cerebral thrombosis, 395
Cerebral veins, 574
Cerebrospinal fluid (CSF), 345t, 358,
381382, 383f
Cerebrovascular accident (CVA), 376, 395, 595
Cerebrum, 306f, 361f, 362t, 363371, 364f,
365f, 366f
and autonomic nervous system, 449, 450f
lobes of, 363367, 366t
relative sizes of, 26f
structure of, 363
white matter of, 368, 369f
Cerumen, 119, 303, 516
Ceruminous glands, 119, 516
Cervical canal, 734
Cervical cancer, 119
Cervical curve, 158f, 159
Cervical enlargement, of spinal cord, 384, 385f
Cervical lymph nodes, 307, 584, 584f
Cervical muscles, 309f
Cervical nerves, 401f, 413, 621t
Cervical plexus, 385f, 401f, 415416, 415f,
416t, 417f
Cervical region, 35
Cervical rib, 334
Cervical spinal nerve, 385f
Cervical vertebrae, 159, 160161, 160f,
163t, 306f
Cervix
of neck, 306
of uterus, 727f, 729f, 734
Cesarean section, 775
Chalazion, 532
Chancroid, 721t
Cheeks, 640
Chemoreceptors, 489, 490, 561, 623
Chest, of neonate, 299t
Chest cavity. See Thoracic cavity
Cheyne-Stokes breathing, 630
Chief cells. See Principal cells
Childhood, 777778, 781t
Chilitis, 672
Chimpanzee, brain of, 26f
Chin, 303, 304f, 304t
China, and history of anatomy, 8
Chlamydia, 721t
Choana, 604f, 605f, 606
Choking, 608, 627, 628f
Cholecystokinin (CCK), 457t, 476t
Cholera, 9
Cholesterol, 52
Cholinergic, 444
Cholinergic stimulation, 445
Cholinesterase, 358
Chondritis, 230
Chondroblasts, 141
Chondrocranium, 141, 142f
Chondrocytes, 95, 140
Chordae tendineae, 549f, 550
Chorda tympani nerve, in taste sensation,
498f, 499
Chorea, 389
Chorion, 757, 764765
Chorion frondosum, 765
Chorionic villus biopsy, 765
Choroid, 500f, 504505, 505f, 508t, 516
Choroid plexus, 370f, 373, 382
Chromaffin, 471
Chromatids, 64f, 704
Chromatin, 53t, 54f, 63
Chromatophilic substances, 348, 349f
Chromosomal abnormalities, 7273
Chromosomes, 63, 64f, 65
Chronic meningitis, 395
Chronic pain, 492
Chronic prostatitis, 722
Chronic renal failure, 693
Church, influence on study of anatomy, 11, 12
Chyme, 649
Chymotrypsin, 654t
Cilia, 53t, 61, 63f, 83, 606f
Ciliary arteries, 507
Ciliary body, 500f, 504f, 505, 505f, 506f,
508t, 512f, 516
Ciliary ganglion, 441
Ciliary glands, 501
Ciliary muscles, 445t, 503t, 505, 506f
Ciliary processes, 505
Cineradiography, 18
Circle of Willis. See Cerebral arterial circle
Circular fibers, 506f
Circular layer, of pharyngeal muscles, 647
Circular movements, 207209, 209f
Circular muscles. See also Sphincteral muscles
of stomach, 650f
Circulation, fetal, 580581, 581f, 582t
Circulatory routes, 550551
Circulatory system, 32f, 537601, 539f
components of, 540
development of, 588591
relation to lymphatic system, 583f
relation to other body systems, 538f
trauma to, 596598
Circumcision, 711
Circumduction, 209, 209f
Circumflex artery, 551, 552f
Circumflex branches, 548f
Cirrhosis, 579, 670
Cisterna, 57
Cisterna chyli, 582, 584f
Cisternal puncture, 391
Class Mammalia, 24, 27t
Claustrum, 365f, 369, 370f
Clavicle, 133f, 173, 174f, 175f, 181t, 307,
307f, 308f, 309f, 311f
Clavicopectoral triangle, 311f, 319, 319f

Clavicular notch, 164, 164f
Cleavage, 757, 758f, 760t
Cleft lip, 155, 166, 333, 626, 627f
Cleft palate, 155, 166, 333, 624, 626, 627f, 669
Clinical considerations
appendicular skeleton, 189191
autonomic nervous system, 450451
axial skeleton, 165168
central nervous system, 391396
circulatory system, 587598
cytology, 7074
digestive system, 669671
endocrine system, 480483
histology, 101103
integumentary system, 119127
joints, 224229
male reproductive system, 714722, 744751
muscular system, 285289
regional anatomy, 330339
respiratory system, 626630
sensory organs, 527533
urinary system, 688694
Clinical practicum
appendicular skeleton, 193
autonomic nervous system, 451
axial skeleton, 168169
central nervous system, 396
digestive system, 672
endocrine system, 484
female reproductive system, 751
integumentary system, 128
joints, 229230
male reproductive system, 722
muscular system, 292293
peripheral nervous system, 432
regional anatomy, 340
respiratory system, 631
sensory organs, 534
urinary system, 694695
Clinical procedures, terminology of, 3435
Clitoris, 716, 727f, 727t, 737f, 738, 743
Cloaca, 667, 690
Cloacal membrane, 667
Closed fractures, 189
Clostridium botulinum, 669
Clostridium tetani, 250
Clubfoot, 189, 189f, 338
Coarctation, 595
Coccygeal cornua, 163, 163f
Coccygeal nerves, 401f, 413
Coccygeal trauma, 163
Coccygeal vertebrae, 159
Coccygeus muscle, 260, 260t, 261f, 281f
Coccyx, 133f, 158f, 159, 162163, 162f, 163f,
163t, 174f, 181f, 182f, 326f, 330f
Cochlea, 30, 517f, 519f, 520, 520f, 521f,
522f, 529, 529f
Cochlear duct, 519f, 520, 520f, 521f, 529
Cochlear nerve, 410, 410f, 517f, 519f, 520f, 521f
Cochlear window, 158f, 517f, 518f, 520
Codfish, brain of, 26f
Codons, 6869, 68f, 69f
Coelom, 41, 44f
Coitus, 698, 726
Cold receptors, 491492
Cold sore, 128
Colitis, 672
Collagen, 51, 74
Collagenous fibers, 91
Collarbone. See Clavicle
Collateral branches, of axons, 348, 349f
Collateral ligaments, 218, 219f, 222f, 274f
Collecting duct. See Papillary duct
Colles fracture, 178, 190, 338
Colliculi, of midbrain, 361f
Colloid, 467
Colloid substances, 57
Colon, 638f
Coloration
of hair, 116
of skin, 109110, 300
Color blindness, 513, 784785

Color vision, 513
Colostomy, 672
Coma, 377
Comatose, 392
Comedo, 128
Comminuted fractures, 189, 190f
Commissural fibers, 368, 369f
Commissures, of eye, 500
Common bile duct, 635f, 652, 654f, 663f, 664f
Common carotid artery, 307, 314f
pressure point for, 572f
Common cold, 628
Common fibular nerve, 333f, 423, 424t
Common flexor tendon, 272f
Common hepatic artery, 566, 567t, 568f
Common hepatic duct, 663f
Common iliac artery, 560f, 567t, 581f, 736f
Common iliac vein, 573f, 576
Common nerve pathways, 492
Communicating veins, 577f
Communication, skin and, 114
Compact bone, 97, 98f, 136, 138140, 139f
Compensatory hypertrophy, 72
Complete fractures, 189
Compound fractures, 189
Compound glands, 87, 88f, 89t
Compounds, 5051, 51t
Compression fracture, 161
Computerized axial tomography (CT scan/
CAT scan), 18, 19f, 33, 391, 587
Concha, 517f
Conchal crest, 156f
Concussion, 392
Condoms, 750f
Conducting division, of respiratory system, 603
Conducting passages, 604612, 612f
Conduction, of nerve impulse, 488
Conduction aphasia, 371
Conduction deafness, 533
Conduction myofibers, 552, 553f
Conduction system, of heart, 551552, 553f
Conductivity, 357
Condylar process, 147f, 156f, 157
Condyle, 136t
Condyloid canal, 148f, 149f, 151
Condyloid joint, 202, 204f, 205t
Cone cells, 506, 510f, 513
Congenital bilateral absence of ductus
deferens, 722
Congenital cataracts, 530
Congenital deafness, 530
Congenital defects, 789
Congenital diaphragmatic hernia, 336
Congenital dislocation of hip, 338
Congenital heart defects, 593
Congenital megacolon, 336
Congenital syphilis, 334
Congenital uterine abnormalities, 746, 746f
Conjoined twins, 786
Conjugated proteins, 51
Conjunctiva, 302f, 302t, 501, 504f, 505f,
506f, 516
Conjunctival sac, 500f, 501
Conjunctivitis, 532
Connecting stalk, 768
Connective tissue, 78, 8998
characteristics of, 8990
classification of, 8990
dense irregular, 92, 93f, 95t
dense regular, 92, 92f, 95t
elastic, 92, 93f, 95t
embryonic, 90, 90f
loose, 91, 91f, 95t
reticular, 92, 94f, 95t
Connective tissue proper, 9094, 95t
Conoid tubercle, 173, 175f
Constipation, 660
Constriction, of pupil, 510511
Constrictor muscles, of pharynx, 647, 647f
Continuous ambulatory peritoneal dialysis
(CAPD), 693
Continuous capillary, 556f, 557, 557f
Contraception, 475, 749751, 750f
Contraceptive sponge, 750f

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Contractions, 234
of cardiac muscle, 437
of skeletal muscles, 243, 245f, 246f
of smooth muscle, 437
Contrecoup fracture, 334
Conus medullaris, 384, 385f
Convergence, of eyeball, 511
Convergent muscles, 238, 239t
Convolutions, 363, 364f
Convulsion, 293
Copulation, 698
Coracoacromial ligament, 216f
Coracobrachialis muscle, 259f, 264f, 266t,
267, 268f, 270f, 313f
Coracoclavicular ligament, 216f
Coracohumeral ligament, 216, 216f
Coracoid process, 173, 175f
Coranoid fossa, 176f
Corn, 128, 327, 330f
Cornea, 302f, 302t, 500f, 504, 504f, 505f,
506f, 508t, 512f
Corniculate cartilage, 609, 609f
Cornification, 108
Corona glandis, 710f, 711
Coronal plane, 33, 33f, 34f
Coronal suture, 144, 145f, 146, 147f, 148f
Corona radiata, 729, 730f, 733f, 755
Coronary arteries, 560f
Coronary artery bypass surgery, 594f
Coronary circulation, 550551, 552f
Coronary embolism, 595
Coronary sinus, 551, 552f, 589
Coronary sulcus, 547, 549f, 550
Coronoid fossa, 176
Coronoid process, 147f, 149f, 156f, 157,
177, 177f
Corpora cavernosa
of clitoris, 738
of penis, 710f, 711
Corpora quadrigemina, 370f, 374, 376f
Corpus albicans, 732, 733f
Corpus callosum, 306f, 361f, 363, 365f,
368, 370f
Corpus cavernosum penis, 686f
Corpuscles of touch, 489t, 490491
Corpus luteum, 475, 732, 733f
Corpus spongiosum penis, 686f, 710f, 711
Corpus striatum, 368, 370f
Corrugator supercilli muscle, 250f, 251f,
251t, 305f
Cortex, of cerebellum, 370f
Cortex layer, 359
Cortical nephrons, 682, 683f
Corticoids. See Corticosteroids
Corticospinal tracts, 386, 389f
Corticosteroids, 471
Corticotropin-releasing hormone (CRH),
462, 463
Cortisol, 457f, 471, 472
Costal angle, 165, 310, 311f
Costal cartilage, 95, 133f, 164f, 165,
165f, 174f
Costal groove, 165, 165f
Costal margin, of rib cage, 165, 310, 311f, 312
Costal notches, 164, 164f
Costal surface, of lung, 614
Costal tuberosity, 173, 175f
Costocervical trunk, 565
Costochondral joint, 165f
Costoclavicular ligament, 215, 215f
Costovertebral joint, 225t
Coughing, 83, 622t
Cowpers glands. See Bulbourethral glands
Coxal joint, 205f, 220, 220f, 225t
muscles of, 275277, 278t, 281t
Coxarthrosis, 230
Crabs, 721t
Cramped muscle, 286
Cranial. See Superior
Cranial bones, 132t, 144, 145153
Cranial cavity, 40f, 44f, 45f, 144, 149f
Cranial dura mater, 378
septa of, 380t
Cranial encephalocele, 333, 392
Cranial meningocele, 392
Cranial nerves, 401, 401f, 402f, 403413,

404t405t. See also specific nerves
neurological assessment of, 412, 413t
and parasympathetic division
of ANS, 441
Cranial region, 35
Cranial root, of accessory nerves, 412, 412f
Craniosacral division, of autonomic nervous
system. See Parasympathetic
division, of autonomic nervous
system
Craniotomy, 169
Cranium, 35, 301
Cremasteric reflex, 430t
Cremaster muscle, 700, 701f, 710f
Crest, of bones, 136t
Cretinism, 481
Crib death, 628
Cribriform foramina, 146t, 153, 153f
Cribriform plate, of ethmoid bone, 148f,
150f, 153, 153f, 154f
Crick, Francis, 66
Cricoid cartilage, 142f, 306, 306f, 609, 610f
Cricothyroid muscle, 255f
Crista ampullaris, 526, 526f
Cristae, 58, 60f
Crista galli, 148f, 149f, 150f, 153, 153f, 154f
Crossed extensor reflex, 428430, 429f
Cross-sectional plane. See Transverse plane
Crown, of tooth, 644, 645f
Crown-rump length, 773
Crura, 738
Crural muscles, 281285, 285f
Crural region, 40
Crus, of penis, 686f, 710711, 710f
Crutch paralysis, 419
Crying, 502, 622t
Crying reflex, 776
Cryptorchidism, 336, 715, 715f
CSF. See Cerebrospinal fluid
CT scan. See Computerized axial tomography
Cubital fossa, 38, 320, 321f
Cubital lymph nodes, 584
Cubital region, 38
Cuboid bones, 186, 187f, 188f
Cumulus oophorus, 729, 730f, 733f
Cuneiform bones, 186, 188f
Cuneiform cartilage, 609, 609f
Cupola, of lung. See Apex, of lung
Cupula, 526, 526f
Curettage, 102
Cushings syndrome, 483
Cuspids. See Canine teeth
Cutaneous prepatellar bursae, 221, 221f
Cutaneous receptors, 114, 489t
Cuticle. See Eponychium
CVA. See Cerebrovascular accident
Cyanosis, 110, 300, 592, 628
Cyclic AMP, 459, 459f
Cyclopia, 530
Cycloplegic drugs, 527
Cyst(s), dermal, 119
Cystic duct, 663f, 664f
Cystic fibrosis, 626, 672, 790
Cystic vein, 579
Cystitis, 687, 692
Cystoscopy, 692, 692f
Cytokinesis, 70
Cytology, 17, 4876
case study, 48, 74
cell cycle, 6570
cellular chemistry, 5052
cellular structure, 5263
introduction to, 49
Cytoplasm, 53, 53t, 54f, 5761
Cytosine, 65, 68
Cytotrophoblast, 759
Dandruff, 128
Dark Ages, and history of anatomy, 12
Dartos, 700
Daughter cells, 705f
David (Michelangelo), 2, 3f
da Vinci, Leonardo, 13, 14f
Deafness
conduction, 533
congenital, 530
perceptive, 533
Death
autopsy and, 1819
somatic, 102
Decibels (dB), 522
Decidua basalis, 765
Deciduous teeth, 642, 644t
Decubitus ulcers, 111, 112f, 128
Decussation, 386
Deep artery, of penis, 710f
Deep brachial artery, 565, 565f
Deep digital flexor muscle, 271, 272f, 273t
tendon of, 274f, 325f
Deep dorsal vein, of penis, 710f
Deep fascia, 236, 238t, 710f
Deep femoral artery, 560f, 568, 570f
Deep femoral vein, 576, 578f
Deep fibular nerve, 423, 425f
Deep iliac circumflex artery, 568, 569f
Deep infrapatellar bursae, 221, 221f
Deep palmar arch, 575
Deep palmar arch artery, 565f, 566
Deep transversus perinei muscles, 261f
Deep veins, 572, 573f
Defecation, 635, 659t
Defecation reflex, 659
Defibrillation, 592
Definitive hair, 117
Deglutition, 635, 648649, 659t
de Graaf, Regnier, 15
De Humani Corporis Fabrica (Vesalius), 14, 14f
Dehydration, 51
Delirium, 392
Delivery, and molding of fetal head, 144
Delta waves, 368t
Deltoid ligament, 222, 223f
Deltoid muscle, 235f, 259f, 264f, 265f, 266t,
267, 267f, 268f, 269f, 270f, 308f, 310,
311f,313f, 316f, 319f,320f, 323f, 324f
Deltoid region, 38
Deltoid tuberosity, 176, 176f
de Luzzi, Mondino, 13, 15
Demifacet, for rib, 161f
Dendrites, 101, 348, 349f, 350f
Dendritic spinules, 348
Dendritic zone, 348
Dens, of axis, 160f, 161
Dense bone. See Compact bone
Dense irregular connective tissue, 92, 93f, 95t
Dense regular connective tissue, 92, 92f, 95t
Dental alveoli, 154, 199f, 644, 645f
Dental caries, 645, 669670, 669f
Dental cusps, 642, 644
Dental formula, 642
Dental pulp, 645f
Dentate nuclei, 375
Denticulate ligament, 381f
Dentin, 644, 645f
Dentistry, 302
Dentoalveolar joint, 198
Deoxyribose, 67f
Deoxyribose nucleic acid (DNA), 52t, 64f, 67f
replication of, 66f
structure of, 6566, 65f
Depolarization, 357, 357f
Depressed fracture, 190
Depression
of bones, 136t
mental, 396
as type of movement, 209, 210f
Depressor anguli oris muscle, 250f, 251f,
251t, 305f
Depressor labii inferioris muscle, 250f, 251f, 251t
Dermabrasion, 128
Dermal cysts, 119
Dermatitis, 107t, 119, 128
Dermatoglyphics, 110
Dermatology, 128
Dermatomes, 426, 426f
Dermis, 107f, 108f, 110112
Descending aorta, 549f, 560f
Descending colon, 635f, 638f, 656, 657f
825
Descending limb, of nephron loop, 680f,

682, 683t
Descending tracts, of spinal cord, 384, 387f,
388t, 389t
Descriptive terminology, 3335
Detached retina, 506
Detrusor muscle, 687
Development, prenatal
of appendicular skeleton, 192, 192f
of axial skeleton, 141
of brain, 346347, 346f, 347f
case study, 754
of circulatory system, 588591
of digestive system, 665668, 665f
of ear, 528529
of endocrine system, 477478
of eye, 515516, 515f
of female reproductive system, 743
of integumentary system, 120121
of male reproductive system, 716719
of muscular system, 248249, 248f, 249f
of peripheral nervous system, 426, 426f
of spinal cord, 390, 390f
of synovial joints, 206, 206f
of tissues, 7980
of urinary system, 689690
Developmental anatomy, 17
case study, 789
Developmental disorders, 789
of abdomen, 336
of central nervous system, 392
of digestive system, 669
of ears, 527531
of eyes, 527530
of female reproductive system, 746
of head, 333
of hip, 338
of lower extremity, 338
of male reproductive system, 714715
of neck, 333
of shoulder, 337
skeletal, 166, 189
of thoracic region, 334335
of upper extremity, 337
of ureters, 691
of urethra, 691
Diabetes insipidus, 463, 481
Diabetes mellitus, 471, 482
Diapedesis, 542
Diaphragm, 24, 40f, 41, 42f, 43f, 258, 258f,
276f, 314f, 317f, 318f, 618, 620f, 635f
Diaphragm (contraceptive), 750f
Diaphragma sellae, 380t
Diaphragmatic surface, of lung, 615f
Diaphysis, 136
Diarrhea, 639, 660, 670
Diarthroses, 197
Diastole, 552
Diastolic pressure, 559
Diencephalon, 347, 362t, 365f,
372373, 376f
Dieting, 94. See also Nutritional disorders
Diffusion, 56f
facilitated, 55t
simple, 55t
Digastric fossa, 149f
Digastric muscle, 255f, 256, 257f, 257t,
308f, 309f
Digestion, 51, 635
Digestive enzymes, 654t
Digestive system, 32f, 634674, 635f
and circulatory system, 539
introduction to, 635636
Digestive tract. See Gastrointestinal tract

826
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Digit(s), 38, 40
bones of, 178, 179f, 187f
muscles that move, 273t
in taxonomy, 24
Digital arteries, 565f, 566, 570f, 571
Digital rays, 192, 768
Dihybrid cross, 784, 784f
Dilation stage, of labor, 775, 776f
Diphyodont, 642
Diploe, 136, 137f
Diploid chromosomes, 782
Diplopia, 511f, 532
Directional terms, 34, 36t
Disaccharides, 52
Discontinuous capillary, 557, 557f
Disease atrophy, 72
Dislocated articular facet, 226
Dislocation of joints, 225226
Displaced fracture, 190
Dissection, of cadavers, 2, 15
history of, 4f
during Renaissance, 13, 13f
Distal, 36t
Distal convoluted tubule, 679f, 680f, 682, 683t
Distal interphalangeal joints, 322f
Distal phalanx, 179f, 180f, 187f
Distal radioulnar joint, 225t
Distal tibiofibular joint, 225t
Disuse atrophy, 72, 102
Diverticulitis, 671
Diverticulosis, 670671
Dizygotic twins, 785786, 787f
Dizziness, 527
DNA. See Deoxyribose nucleic acid
Dominant alleles, 783, 783t
Dopamine, 351, 390, 463
Dorsal. See Posterior
Dorsal arch, 570
Dorsal artery, of penis, 710f
Dorsal calcaneocuboid ligament, 223f
Dorsal cuneonavicular ligament, 223f
Dorsal hollow nerve cord, 23, 23f
Dorsal interosseous muscle, 271f, 274f, 275,
275t, 290f
Dorsal motor nucleus, 411
Dorsal nerve, of penis, 710f
Dorsal pedal artery, 560f, 570, 570f
pressure point for, 329f, 572f
Dorsal pedal vein, 578f
Dorsal talonavicular ligament, 223f
Dorsal tarsometatarsal ligament, 223f
Dorsal vein, of penis, 710f
Dorsiflexion, 207, 208f, 212f
Dorsum
of foot, 40
of hand, 38
Dorsum sellae, 149f, 153f
Double helix, 65, 65f, 66f
Double-jointed, 200
Double uterus, 746, 746f
Double vagina, 746f
Double vision, 511f, 532
Down syndrome, 73, 763
Dropsy, 1
Drug(s), cytologic effects of, 57, 72
Drug abuse, 391392
Dry skin, 107t
DSR scan. See Dynamic spatial
reconstructor scan
Dual innervation, 446447
antagonistic effects, 446, 447f
complementary effects, 446
cooperative effects, 446
Ductus arteriosus, 581, 581f, 582t, 590
Ductus deferens, 698, 699f, 700t, 701f,
707708, 708f, 710f, 716
Ductus venosus, 580, 581f, 582t
Duodenal glands, 652
Duodenal papilla, 652, 654f, 663f, 664f
Duodenojejunal flexure, 652, 653f
Duodenum, 456, 635f, 638f, 649f, 650f, 652,
653f, 654f, 655f, 663f, 664f
Duplication of ureters, 691
Dural sheath, 379
Dural sinuses, 378, 378f
Dura mater, 360f, 378379, 378f, 379f, 380t,

381f, 385f
Dutch, and history of anatomy, 8
Dutrochet, Ren H., 16
Dwarfism, 166, 462, 480, 481f
Dynamic spatial reconstructor (DSR) scan,
18, 19f, 391
Dysentery, 669
Dyslexia, 394
Dysmenorrhea, 747
Dysplasia, 748
Dyspnea, 621, 622t
Dysuria, 691
Ear(s), 517f. See also Hearing; Inner ear;
Middle ear; Outer ear
of neonate, 299t
Eardrum. See Tympanic membrane
Earlobe, 303, 516, 517f
Earwax. See Cerumen
Eccrine sweat glands, 118, 118f
ECG. See Electrocardiogram
Ectoderm, 80, 80t, 760, 761f
Ectopic pacemakers, 592
Ectopic pregnancy, 734, 746747, 785, 786f
Eczema, 128
Edema, 1, 92, 582, 592
EEG. See Electroencephalogram
Effector, in reflex arc, 427, 427f
Efferent ductules, 702f, 703, 716
Efferent glomerular arterioles, 678, 680f, 681f
Efferent lymphatic vessels, 583
Efferent nerves, 356f
Efferent neuron, 352
Egg cells. See Oocytes
Egypt, and history of anatomy, 58
Ejaculate. See Semen
Ejaculation, 712, 713714, 713f
Ejaculatory ducts, 686f, 687, 698, 699f, 700t,
708, 716
EKG. See Electrocardiogram
Elastic arteries, 555, 556f
Elastic cartilage, 95, 97f
Elastic connective tissue, 92, 93f, 95t
Elastic fibers, 91
Elasticity, of muscle, 234
Elastin, 74
Elbow, surface anatomy of, 321f
Elbow joint, 204f, 213f, 217, 218f, 219f, 225t
Electrical defibrillation, 592
Electrocardiogram (ECG/EKG), 552554,
553f, 587, 595f
Electroencephalogram (EEG), 367368,
368t, 391
Electrolytes, 51, 52t, 373
Electromagnetic spectrum, 511, 513f
Electromyography (EMG), 285
Electronic Monitoring of Fetal Heart Rate
and Uterine Contractions (FHRUC) Monitoring, 787789, 789f
Electron microscope, 17, 17f, 78
Elements, 50
Elevation, 209, 210f
Embalming, 7, 13
Embedding, 102
Embolism, 595
Embryo, 762, 763f, 764f, 773f. See also
Development
bone development in, 141, 141f
changes by week, 767769, 769f, 770f, 771f
chordate, 23f
connective tissue of, 90, 90f
skull of, 142, 142f
Embryoblast, 80, 757
Embryology, 762
Embryonic disc, 760
Embryonic period, 762769
Emetics, 651
EMG. See Electromyography
Emission, 712, 713, 713f
nocturnal, 714
Emmetropia, 531f
Emphysema, 615, 628, 630f
Enamel, 645, 645f
Encephalitis, 394
Encephalomyelitis, 394
Endemic goiter, 481, 481f
Endocarditis, 335, 547
Endocardium, 547, 547t
Endochondral ossification, 95, 140, 141,
141f, 192
Endocrine cells (G cells), 651
Endocrine disorders, 480
Endocrine glands, 455, 455f, 457t
Endocrine organs, 456f
Endocrine system, 31f, 454486. See also
specific components
control of, 373
glands of, 456
and nervous system, 455, 455t
placenta in, 765766, 766t
Endocrinology, 480
Endocytosis, 55t
Endoderm, 80, 80t, 760, 761f
Endolymph, 519, 526f
Endometriosis, 748
Endometrium, 729f, 736
Endomysium, 236, 237f
Endoneurium, 352
Endoplasmic reticulum (ER), 53t
rough, 54f, 57, 58f
smooth, 54f, 57, 58f
Endorphins, 359
Endosteum, 137
Endothelium, 81, 555, 556f
Endothermic animals, 540
Energy deficit, 72
Enkephalins, 359
Entamoeba histolytica, 669
Enteritis, 337, 670
Enterokinase, 654t
Enuresis, 692
Enzymes, 52
digestive, 654t
proteolytic, 757
synthetase, 69
Eosin and hematoxylin stain, 542
Eosinophils, 542, 542f, 544t
Ependymal cells, 350f, 351, 351t
Ependymomas, 394
Epicanthic fold, 501
Epicardium, 546, 547t
Epicondyles, 136t, 185
Epicranius muscle, 251t
Epidermal derivatives, 115119
Epidermis, 106110, 107f, 108f, 109t
Epididymis, 698, 699f, 700t, 701f, 702f, 707,
707f, 710f, 716
Epidural anesthesia, 775
Epidural space, 379, 379f
Epigastric region, 38f, 38t
Epiglottic cartilage, 608
Epiglottis, 604f, 605f, 608, 609f, 641f, 642f
Epilepsy, 363, 393394
Epimysium, 236, 237f
Epinephrine, 246, 440, 446f, 457t, 458, 472, 473t
Epiphyseal line, 137
Epiphyseal plate, 137, 142143, 142f, 143f
Epiphysiolysis, 169
Epiphysis, 137
Epiploic appendages, 657f, 659
Episiotomy, 260, 737
Epispadias, 691, 691f
Epistaxes, 627
Epithalamus, 373
Epithelial tissue, 78, 7989, 86t
Epithelium
glandular, 8789
keratinized stratified squamous, 85, 86t
membranous, 7981
nonkeratinized stratified squamous,
85, 86t
pseudostratified ciliated columnar, 83,

84f, 86t
respiratory, 83
simple, 8183, 86t
simple ciliated columnar, 83, 83f, 86t
simple columnar, 8183, 82f, 86t
simple cuboidal, 81, 82f, 86t
simple squamous, 81, 86t
stratified, 8385, 86t
stratified cuboidal, 85, 85f, 86t
stratified squamous, 8385, 84f, 86t
transitional, 85, 86f, 86t
Epitympanic recess, 517, 518f
Eponychium, 117, 117f
Eponyms, elimination of, 17
Equilibrium, 526f
mechanics of, 425f, 523527
neural pathways for, 526527
ER. See Endoplasmic reticulum
Erasistratus, 11
Erb-Duchenne palsy, 338
Erbs point, 338
Erection
female, 738
male, 711f, 712713, 712f
Erector spinae muscle, 262, 262f, 263t, 310
ERV. See Expiratory reserve volume
Erythema, 110, 128, 592
Erythroblastosis fetalis, 765
Erythrocytes, 98, 99f, 540541, 541f,
542f, 544t
circulation of, 538539
shape of, 50f
Erythropoiesis, 544
Erythropoietin, 457t
Esophageal arteries, 566, 567t
Esophageal atresia, 669
Esophageal hiatus, 648
Esophageal plexus, 441
Esophagus, 306f, 314f, 604f, 605f, 635, 635f,
636t, 641f, 648, 648f, 649f, 650f, 659t
Esotropia, 532
Estradiol-17, 457t
Estriol, during pregnancy, 766
Estrogen, 472, 741, 766t
Estrogens, 474475
Ethmoidal crest, 156f
Ethmoidal sinus, 605f, 607, 607f
Ethmoid arch, 150f
Ethmoid bone, 147f, 153, 153f, 605
Ethmoid sinus, 151f, 153
Ethylene glycol poisoning, 48
Eupnea, 622t
Eustachian tube. See Auditory tube
Eustachius, 15
Evaporative cooling. See Sweating
Eversion, 209, 210f
Examination, 102
Ex anopsia, 511
Excretory lacrimal ductules, 501f
Excretory system, and circulatory system, 539
Exercise, and muscle aging, 289
Exhalation. See Expiration
Exocoelomic membrane, 763
Exocrine glands, 87, 88f, 89t, 455, 455f, 639f
Exocytosis, 55t
Exophthalmos, 482, 482f
Exotropia, 532
Expiration, 256, 618, 619620, 619f, 621t
muscles of, 258f, 620f
Expiratory portion, of rhythmicity area, 623
Expiratory reserve volume (ERV), 621t, 622f
Expulsion stage, of labor, 775, 776f
Exstrophy of urinary bladder, 691
Extensibility, of muscle, 234
Extension, 207, 208f, 211f, 212f
Extensor carpi brevis muscle, tendons of, 272f
Extensor carpi longus muscle, tendons of, 272f
Extensor carpi radialis brevis muscle, 267f,
271, 271f, 273t, 325f
tendon of, 325f
Extensor carpi radialis longus muscle, 267f,
269f, 271, 271f, 273t, 320f, 321f
tendon of, 325f
Extensor carpi radialis muscle, tendons of, 272f

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Extensor carpi ulnaris muscle, 267f, 272,
272f, 273t, 321f, 325f
Extensor digiti minimi muscle, 271f, 272,
273t, 325f
Extensor digitorum brevis muscle, 285, 291f, 329f
Extensor digitorum communis muscle, 271f,
272, 272f, 273t, 325f
tendon of, 325f
Extensor digitorum longus muscle, 235f, 283,
283f, 284f, 286t, 332f
Extensor digitorum muscle, 267f, 272f, 321f
Extensor hallucis brevis muscle, 291f
Extensor hallucis longus muscle, 283, 283f,
284f, 286t
Extensor indicis muscle, 271f, 272f
tendon of, 325f
Extensor pollicis brevis muscle, 271f, 272,
272f, 273t
tendon of, 274f, 322f, 325f
Extensor pollicis longus muscle, 272, 272f, 273t
Extensor retinaculum, 271f
External, 36t
External abdominal oblique muscle, 235f,
258, 258f, 259f, 260t, 264f, 265f,
312f, 313, 313f, 316f, 317f, 323f,
324f, 620f
External acoustic canal, 302, 516, 517f, 529
External acoustic meatus, 147f, 148f, 149f,
151, 152f, 158f, 516
External anal sphincter, 260, 261f, 657, 658f
External carotid artery, 560f, 561, 562f, 563564
External hydrocephalus, 381
External iliac artery, 560f, 566, 567t, 568,
568f, 569f, 581f
External iliac vein, 573f, 576, 578f
External intercostal muscles, 258, 258f, 259f,
264f, 311f, 313f, 618, 620f
External jugular vein, 307, 309f, 324f, 572,
573f, 574
External occipital protuberance, 148f, 152, 315f
External otitis, 533
External respiration, 603
External spermatic fascia, 701f, 710f
External urinary sphincter, 686f, 687
Exteroceptors, 489490
Extraembryonic membranes, 762765, 763f, 764f
Extramedullary tumors, 394
Extrapyramidal tracts, 389, 389f
Extrinsic eye muscles, 502, 503t
Extrinsic laryngeal muscles, 609
Extrinsic ocular muscles, 502f, 514, 516
Extrinsic tongue muscles, 252, 255f, 255t
Eye(s)
accessory structures of, 499502
comparison with camera, 499t
diseases of, 532
diagnosis of, 527
functional impairments of, 531532, 531f
human, comparison with other species, 499
infections of, 532
movements of, 502, 503f, 513514
of neonate, 299t
Eyeball
blood supply to, 507
cavities of, 508
chambers of, 508
convergence of, 511
function of, 509511
structure of, 504508, 504f, 506f, 508t
trauma to, 508
Eyebrows, 302, 302f, 302t, 500, 500f, 501f
Eyelashes, 302, 302f, 302t, 500, 500f, 501, 501f
Eyelids, 302, 302f, 302t, 500, 501f, 516
Eye movement test, 504
Face, surface anatomy of, 301303, 301f

Facet, of bone, 136t, 161, 161f
Facial artery, 305f, 309f, 562f, 563
Facial bones, 132t, 144, 154157
Facial expressions, 252f
muscles of, 301302
Facial muscles, 24, 28, 250, 250f, 251f, 251t
Facial nerve (VII), 250, 360f, 402f,
404t405t, 409, 409f, 410f, 413t,
498f, 499, 517f
Facial region, 35
Facial vein, 305f, 574f
Facilitated diffusion, 55t
Fainting, 392
Falciform ligament, 314f, 317f, 637, 638f, 660
Fallopian tubes. See Uterine tubes
Fallopius, 15
False labor, 775
False pelvis, 180, 183f
False ribs, 164f, 165
False vocal cords. See Vestibular folds
Falx cerebelli, 375, 380t
Falx cerebri, 363, 378f, 380t
Familial cretinism, 790
Family Hominidae, 24, 27t
Farsightedness. See Hyperopia
Fascia, 91, 236, 237f, 238t
Fasciculus, 236, 237f, 352
Fasciculus cuneatus, 388t, 389t
Fasciculus gracilis, 388t, 389t
Fat, 52, 134. See also Adipose tissue
Fat body, 259f
Fauces, 640, 641f
Fecal fistula, 336
Feeding center, 373
Female
comparison with male physique, 780, 780f
distribution of hair in, 115f
growth rate of, 726, 728f
hypodermis of, 297, 298f
infertility in, 747
orgasm in, 738
puberty in, 726
sexual maturation of, 779
Female condoms, 750f
Female infertility, 747
Female reproductive system, 32f, 701f,
725753, 727f, 729f
case study, 725
comparison with male, 743t
development of, 716, 717f, 718f, 743, 743f
diseases of, 720721
disorders of, 747f
Female sterilization, 744
Femoral artery, 331f, 560f, 568, 570f, 571f
Femoral circumflex vein, 578f
Femoral hernia, 293
Femoral nerve, 331f, 401f, 420t, 421, 422f, 571f
Femoral region, 40
Femoral ring, 571f
Femoral triangle, 326, 327f, 568, 571f
Femoral vein, 331f, 571f, 573f, 576, 578f
Femur, 133f, 174f, 184186, 184f, 185f, 188t
Fenestrae, 680, 681f, 682f
Fenestrated capillary, 556f, 557, 557f
Fertilization, 755756, 755f, 757f, 758f
Fetal circulation, 580581, 581f, 582t
Fetal head, molding of, 144
Fetal issue transplantation, 103
Fetal length, 773
Fetal monitoring, 786789
Fetal period, 772774
Fetal skull, 145f
Fetoscopy, 787, 788f
Fetus, 769, 771f, 772, 773f
changes by week, 772774
external appearance of, 772f
Fever blister, 128
FHR-UC. See Electronic Monitoring of Fetal
Heart Rate and Uterine
Contractions Monitoring
Fibrillation, 293, 592, 592f

Fibrils, 53t, 61
Fibrin, 126f
Fibrinogen, 125126, 126f, 545, 545t
Fibroadenoma, 748
Fibroblasts, 91, 126
Fibrocartilage, 95, 96f, 97f
Fibrocystic disease. See Dysplasia
Fibromyositis, 287
Fibrous capsule. See Renal capsule
Fibrous digital sheath, 325f
Fibrous joints, 197198, 205t
Fibrous layer, of vagina, 737
Fibrous pericardium, 546
Fibrous tunic, of eyeball, 504508, 504f, 508t
Fibula, 133f, 174f, 185, 185f, 186f, 188t, 328f
Fibular artery, 570, 570f
Fibular collateral ligament, 221f
Fibular nerve, 425f
Fibular notch, 185
Fight-or-flight response, 444, 473, 474f
Filiform papillae, 497, 497f, 642, 642f
Filtration, 55t, 56f
Filtration slits, 681f, 682f
Filum terminale, 384
Fimbriae, 727f, 729f, 734
Fimbriated fold, 640f
Fingerprints, 110, 110f
Fingers. See also Digits
flexion of, 269271
muscles that move, 268272
First-class lever, 210, 213f
First-degree burns, 122, 123f
First-degree frostbite, 123
First dorsal interosseous muscle, 274f, 321f, 325f
First lumbrical muscle, 274f
First meiotic division, 704, 704t, 705f, 706f
First rib, 164f
First sound, of heartbeat, 554
First trimester, of pregnancy, 785
Fissure, 136t
Fissured fractures, 189, 190f
Fissure of Rolando. See Central sulcus
Fissure of Sylvius. See Lateral sulcus
Fistula, 313
Fixation, 102
Fixed action pattern, 345
Flaccid paralysis, 393
Flagella, 53t, 61, 63f, 704, 706f
Flat bones, 136, 136f
Flatfoot, 188
Flexion, 207, 208f, 211f, 212f, 269271
Flexion creases, 110, 323f
Flexion lines, 110
Flexor carpi radialis muscle, 269, 271f, 273t
tendon of, 321f, 323f, 325f
Flexor carpi ulnaris muscle, 267f, 269, 271f,
273t, 325f
Flexor digiti minimi muscle, 271f, 274f, 275,
275t, 290f, 325f
Flexor digitorum brevis muscle, 290f
Flexor digitorum longus muscle, 284285,
285f, 286t, 287f, 288f
Flexor digitorum profundis muscle, 272f. See
also Deep digital flexor muscle
tendons of, 274f
Flexor digitorum superficialis muscle. See
Superficial digital flexor muscle
Flexor hallucis brevis muscle, 290f
Flexor hallucis longus muscle, 284, 285f,
286t, 288f
Flexor pollicis brevis muscle, 271f, 274f, 275,
275t, 325f
Flexor pollicis longus muscle, 271, 271f,
272f, 273t
tendon of, 274f
Flexor reflex, 428, 429f
Flexor retinaculum, 269, 271f, 274f, 285f, 287f
Flexure, 658f
Floating ribs, 164f, 165
Fluoroscopic analysis, 587
Fold, of buttock, 326, 326f
Folia, 375
Follicles, 729730, 729f, 730f, 733f
827
Follicle-stimulating hormone (FSH), 462,

464t, 741
Follicular cells, 467, 743
Follicular epithelium, 729
Follicular hyperkeratosis, 107t
Fontanels, 144, 300
Food poisoning, 669
Foot, 224. See also Pes
muscles of, 281285, 285, 286t, 290f, 291f
Footdrop, 339
Foramen, 136t
of skull, 144145, 146t
Foramen cecum, 149f, 478
Foramen lacerum, 146t, 148f, 149f, 152
Foramen magnum, 146t, 148f, 149f, 151
Foramen of Magendie. See Median aperture
Foramen of Monro. See Interventricular
foramen
Foramen ovale, 146t, 148f, 149f, 152, 153f,
580581, 581f, 582t, 589
Foramen rotundum, 146t, 149f, 152, 153f
Foramen spinosum, 146t, 149f, 152, 153f
Foramina of Luschka. See Lateral apertures
Forearm. See Antebrachium
Forebrain. See Prosencephalon
Foregut, 624, 665666, 666f
Foreskin. See Prepuce
Formed elements, 98, 99f, 540542,
541f, 544t
Fornix, 500f, 729f, 736
Fosamax. See Alendronate
Fossa, 136t
Fourth ventricle, of brain, 361f, 374, 376,
381, 382f
Fovea capitis femoris, 184, 184f
Fovea centralis, 505f, 506
Fracture hematoma, 190
Fractures, 189191
of clavicle, 173
closed, 189
Colles, 178, 190, 338
comminuted, 189, 190f
complete, 189
compound, 189
compression, 161
contrecoup, 334
depressed, 190
displaced, 190
of fibula, 185
fissured, 189, 190f
greenstick, 189, 190f
impacted, 189
of long bone, 137, 200
nondisplaced, 190
oblique, 189, 190f
open, 189
partial, 189, 190f
pathologic, 189
Potts, 185, 190
of radius, 178
of scaphoid, 178
simple, 189
of skull, 198
of spine, 161
spiral, 189, 190f
stress, 339
transverse, 189, 190f
traumatic, 189
treatment for, 190191, 191f
Fraternal twins. See Dizygotic twins
Freckles, 109
Free border of nail, 117, 117f
Free nerve endings, 489t, 491
Frenulum, 710f, 711
Frequency, of sound, 522
Frog, brain of, 26f
Frontal bone, 133f, 142f, 145146, 145f,
147f, 148f, 149f, 150f, 174f, 306f
Frontal fontanel. See Anterior fontanel
Frontalis muscle, 235f, 250f, 251f, 251t,
305f, 324f
Frontal lobe, 360f, 361f, 363, 364f, 366f, 366t
Frontal plane. See Coronal plane
Frontal process, of maxilla, 154f, 155f

828
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Frontal sinus, 45f, 146, 148f, 151f, 154f,

306f, 605f, 607, 607f
Frontal sulcus, 364f
Frostbite, 72, 123
FSH. See Follicle-stimulating hormone
Functional heart murmur, 554
Functional proteins, 52
Functional syncytium, 437
Fundus
of stomach, 649, 649f, 650f
of uterus, 729f, 734
Fungiform papillae, 497, 497f, 642, 642f
Funiculi, 386
Funnel chest, 312, 334
Furuncle, 128
Gage, Phineas P., 450
Galactosemia, 790
Galea aponeurotica muscle, 250f, 251f, 378f
Galen, 1112
Gallbladder, 317f, 635, 635f, 638f, 661f,
662663, 663f, 664f
Gallstones, 663, 664f
Gametes, 474, 698
Gamma globulin, 545, 545t
Ganglion, 348
Ganglionic gliocytes, 351, 351t
Ganglionic neurons, 506
Gangrene, 102, 123, 128
Gaseter. See Belly, of muscle
Gas exchange, 603
Gastric folds, 650, 650f
Gastric glands, 650, 651f
Gastric inhibitory peptide (GIP), 476t
Gastric juice enzyme, 654t
Gastric phase, of gastric secretion, 651t
Gastric pits, 650, 651f
Gastric secretion, 651t
Gastrin, 457t, 476t
Gastrocnemius muscle, 235f, 280f, 281f, 283,
283f, 284f, 285f, 286t, 287f, 288f,
289f, 328f, 329f, 332f, 333f
Gastrocolic reflex, 659
Gastroileal reflex, 659
Gastrointestinal activity, control of,
hypothalamus and, 373
Gastrointestinal hormones, 476t
Gastrointestinal tract (GI tract), 635, 636t
autonomic nervous system effects
on, 445t
disorders of, 670671, 671f
innervation of, 639640
irritation of, 639
layers of, 637639, 639f
mechanical activity in, 659t
organs of, 635
pathogens of, 669
G cells. See Endocrine cells
General anesthesia, 392
General interpretative area, 366f
General senses, 488
Genes, 782783
Genetics, medical, 7273
Genetic sex, 716
Genetic transcription, 6768, 67f, 68f
Genetic translation, 68, 68f, 69f
Geniculate ganglion, 409
Geniculostriate system, 513
Genioglossus muscle, 252, 255f, 255t
Geniohyoid muscle, 255f, 257t
Genital herpes, 721t
Genitalia. See also Female reproductive
system; Male reproductive system
external
differentiation of, 718f
female, 737f
of neonate, 299t
Genital tract, 743
Genital tubercle, 716, 743
Genitofemoral nerve, 420t
Genotype, 783
Genu, of corpus callosum, 361f, 370f
German Anatomical Society, 17
Germinal epithelium, 704, 728, 733f
Germinal layer, 120
Germ layers, 761f

derivatives of, 80t
formation of, 760
Gestation, 760, 775
GH. See Growth hormone
Gigantism, 166, 462, 480, 481f
Gingiva, 644, 645f
Gingivitis, 672
GIP. See Gastric inhibitory peptide
GI tract. See Gastrointestinal tract
Glabellar reflex, 431f
Glands. See also specific glands
Glandular epithelium, 8789, 89t, 435
Glans clitoris, 738
Glans penis, 686f, 699f, 710, 710f, 711, 716
Glaucoma, 530, 532
Glenohumeral joint, 215217, 216f, 217f,
225t, 264267, 266t
Glenohumeral ligament, 216
Glenoid cavity, 173, 175f
Glenoid labrum, 216, 216f, 217f
Glial cells, 101. See also Neuroglia
Gliding joint, 202, 204f, 205t
Gliomas, 394
Glisson, Francis, 15
Globulin, 545, 545t
Globus pallidus, 365f, 369, 370f
Glomerular capsule, 679f, 680, 680f, 681f, 683t
Glomerular filtration, 682f
Glomerulonephritis, 692
Glomerulus, 678, 680f, 681f, 683t
Glossopalatine arch, 640f, 641
Glossopharyngeal nerve (IX), 402f, 405t,
410411, 410f, 413t, 498f, 499
Glottis, 608, 609f
Glucagon, 457t, 470, 663
Glucocorticoids, 457t, 471, 473t
Gluteal arteries, 569f
Gluteal cleft. See Natal cleft
Gluteal fascia, 265f
Gluteal lines, 183
Gluteal region, 37
muscles of, 265f
Gluteal surface, 183
Gluteal tuberosity, 184f
Gluteus maximus muscle, 235f, 261f, 265f,
276, 277, 278t, 280f, 281f, 316f,
326f, 330f, 331f
Gluteus medius muscle, 235f, 265f, 276, 277,
278t, 279f, 280f, 330f
Gluteus minimus muscle, 276, 277, 278t, 330f
Glycogen, 52
Glycoproteins, 51
GnRH. See Gonadotropin-releasing hormone
Goblet cells, 61, 83, 87, 87f, 610, 637, 650
Goiter, 468, 468f, 481, 482f
Golgi complex (Golgi apparatus), 53t, 54f, 58, 59f
Golgi tendon organs, 493
Gomphoses, 198, 199f, 205t
Gonadal arteries, 567t, 568f
Gonadal ridge, 716, 743
Gonadocorticoids, 472, 473t
Gonadotropin-releasing hormone (GnRH),
462, 741
Gonadotropins, 462
Gonads
differentiation of, 717f
endocrine functions of, 456
female, 474475, 726
male, 474, 698
Gonorrhea, 720721, 721t
Goose, brain of, 26f
Gorilla, hands of, 26f
Gout, 790
Gouty arthritis, 227
G1 phase, 6970, 70f
G2 phase, 6970, 70f
Graafian follicle. See Vesicular ovarian follicle
Gracilis muscle, 235f, 276f, 277, 278f, 279f,
280f, 281f, 281t, 287f, 289f, 328f, 331f
tendon of, 285f
Grand mal epilepsy, 393394
Granular endoplasmic reticulum. See Rough

endoplasmic reticulum
Granulation tissue, 126
Granulocytes, 542, 544t
Granulosa cells, 730f
Graves disease, 481482
Gray commissure, 384, 386f
Gray matter
of brain, 359, 365f
of spinal cord, 384
Gray rami communicantes, 439, 439f
Gray ramus, 414
Great auricular nerve, 315f
Great cardiac vein, 552f
Greater alar cartilage, 305f, 605f
Greater auricular nerve, 416t
Greater cornua, 157, 157f
Greater curvature, of stomach, 650, 650f
Greater multangular. See Trapezium bone
Greater occipital nerve, 309f, 315f
Greater omentum, 42f, 4345, 317f, 637, 638f
Greater palatine foramen, 146t, 148f, 155, 156f
Greater palatine groove, 155f
Greater pelvis, 180, 183f
Greater sciatic notch, 182f, 183
Greater trochanter, of femur, 182f, 184,
184f, 326f, 330f
Greater tubercle, of humerus, 175f, 176, 176f
Greater wing, of sphenoid bone, 147f, 148f,
149f, 150f, 152, 153f
Great saphenous vein, 329f, 571f, 573f, 576,
578f, 579
Grecian period, and history of anatomy, 810
Greek language, and anatomical
terminology, 2
Greenstick fractures, 189, 190f
Gristle, 95
Gross anatomy, 17
Ground substance, 90
Growth, 29, 30f. See also Development
rates of, male vs. female, 726, 728f
Growth hormone (GH), 166, 462,
464t, 480
Growth hormone-inhibiting hormone
(GH-IH), 462
Growth hormone-releasing hormone
(GH-RH), 462
Growth-inhibiting substances, 70
Guanine, 65, 68
Gubernaculum, 716
Gums. See Gingiva
Gustatory cells, 496497, 497f
Gustatory microvilli, 497, 497f
Gustatory pathway, 498f, 499
Gustatory receptors, 496
Gustatory sense, 496499
Gut, 635, 637f
Gynecological examination, 744746
Gynecology, 744
Hair, 107f, 115117, 116f
distribution of, male vs. female, 115f
nutritional deficiencies and, 107t
in taxonomy, 24
Hair bud, 121
Hair bulb, 107f, 115, 116f, 121
Hair cells
of ear, 57, 526f
of saccule, 515, 515f
of spiral organ, 521f
of utricle, 515, 515f
of vestibular organ, 524, 524f
Hair color, 116
Hair cortex, 115116
Hair follicle, 107f, 115, 116f, 121
Hairline, 301
Hair matrix, 121
Hair medulla, 115
Hair papilla, 121
Hair root, 115, 116f
Hair shaft, 115, 116f
Halitosis, 672
Hallux, 188
Hallux valgus, 202
Hamate bone, 178, 179f, 180f
Hamlet (Shakespeare), 2
Hammertoe, 327, 330f
Hamstring injuries, 281
Hamstring muscles, 280, 326f
Hand. See also Manus
bones of, 178, 179f, 180f
extension of, 271272
flexion of, 269271
muscles of, 268272, 273275, 273t,
274f, 275t
pronation of, 268
supination of, 268
surface anatomy of, 321f, 322, 322f, 323f
Hard palate, 604f, 605f, 640f, 641, 641f
Harvey, William, 15, 16f
Haustral churning, 659, 659t
Haustrum, 657, 657f, 658f
Haversian canal. See Central canal
Haversian systems, 138
H bands, 245f
hCG. See Human chorionic gonadotropin
hCS. See Human chorionic
somatomammotropin
Head, 35
arteries of, 307, 561564, 562f
cavities within, 42, 45f
developmental disorders involving, 333
diseases of, 334
internal anatomy of, 303304, 305f,
306f, 324f
surface anatomy of, 301303
trauma to, 333334
veins of, 572574, 574f, 576f
Head (anatomical)
of bones, 136t
of condylar process, 156f
of epididymis, 707f
of femur, 182f, 184, 184f
of fibula, 185, 186f, 328f
of humerus, 173, 175f, 176f
of metatarsals, 186, 187f
of phalanges, 180f
of rib, 165, 165f
of spermatozoa, 704, 706f
of tibia, 185f
of ulna, 177, 177f
Headaches, 392, 564
Hearing, 516527. See also Ear
air pressure in, 519
neural pathways for, 523, 523f
Hearing loss, 533
Heart, 540, 545554
catheterization of, 587
chambers of, 547550, 589f
conduction system of, 551552, 553f
description of, 545546
development of, 588f, 589, 589f, 590t
feedback mechanisms of, 553f
innervation of, 553f
location of, 36, 545546, 546f
serous membranes surrounding, 43f
structure of, 548f549f
in taxonomy, 24
valves of, 547550, 548t, 551f
Heart attack, 551
Heartbeat, variation in, 552
Heartburn, 648, 672
Heart cords, 589
Heart disease, 593595
Heart murmur, 335, 554, 593
Heart sounds, 554, 554f
Heart tube, 589
Heart wall, 546547, 547t
Heat exhaustion, 113
Heat production, muscular system and, 234
Heat receptors, 491492
Heat stroke, 113
Heavy metals, exposure to, 116
Heel, 40

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Height, losses in, 163
Heimlich maneuver, 608, 627, 628f
Helicobacter pylori, 670
Helicotrema, 520
Helix, 303, 516, 517f
Hemarthrosis, 230
Hematoxylin and eosin (H & E) stains, 102
Hematuria, 691692
Heme, 541
Hemiazygos vein, 576, 577f
Hemiplegia, 393
Hemispheres, of cerebellum, 375
Hemivertebrae, 334
Hemocytoblasts, 544
Hemodialysis, 691f, 693
Hemoglobin, 110, 541
Hemophilia, 785
Hemopneumothorax, 602
Hemopoiesis, 134, 135f, 543f, 544
Hemorrhage, 570
control of, 596597, 597f
Hemorrhoidal vessels, 658f
Hemorrhoids, 596, 657, 672
Hemothorax, 335
Heparin, 91
Hepatic artery, 581f, 662f
Hepatic buds, 666
Hepatic ducts, 662, 663f, 664f
Hepatic flexure, 317f, 656, 657f
Hepatic plate, 661, 662f, 663f
Hepatic porphyria, 790
Hepatic portal system, 579, 580f
Hepatic portal vein, 579, 580f, 581f, 662f
Hepatic sinusoids. See Liver sinusoids
Hepatic triad, 662f
Hepatic veins, 573f, 580f
Hepatitis, 670
Hepatitis A, 670
Hepatitis B, 670
Hepatocytes, 661
Hepatomas, 670
Hepatopancreatic ampulla, 652, 654f
Herbs, Chinese beliefs in, 8
Hereditary hemochromatosis, 790
Hereditary leukomelanopathy, 790
Hermaphroditism, 714715
Hernia
congenital diaphragmatic, 336
femoral, 293
hiatal, 293, 336, 670
inguinal, 293
umbilical, 293, 313
Herniated disc, 24, 24f, 226, 423, 425f
Herophilus, 11
Herpes zoster, 414
Hertz (Hz), 522
H & E stains. See Hematoxylin and eosin stains
Heterodont dentition, 24, 642
Heterografts, 102
Heterotransplant, 124
Heterozygous, 783
Hiatal hernia, 293, 336, 670
Hiccuping, 622t
Hidden border of nail, 117, 117f
Hilum, 583, 614, 677, 728
Hindbrain. See Rhombencephalon
Hindgut, 667, 668f
Hinge joint, 202, 204f, 205t
Hip, 40
congenital dislocation of, 338
diseases of, 339
dislocation of, 338, 339
muscles of, 276f
trauma to, 339
Hip joint. See Coxal joint
Hippocrates, 89
Hippocratic Oath, 8, 10f
Hirschsprungs disease, 336
Hirsutism, 115
Histamine, 91
Histology, 77104
case study, 77, 103
connective tissue, 8998

definition of, 17, 28
Histopathology, 102
Histotechnology, 102
Hives, 128
Hodgkins disease lymphomas, 596
Holocrine glands, 88f, 89, 89t
Homer, 8
Hominidae, 24
Homologous chromosomes, 782, 782f, 783f
Homologous structures, 716, 726, 743t
Homo sapiens, 23, 24, 27t
Homotransplant, 102
Homozygous, 783
Homozygous dominant, 783
Homozygous recessive, 783
Homunculus, 16
Hooke, Robert, 16, 49
Hordeola, 532
Horizontal plane. See Transverse plane
Horizontal plates, of palatine bone, 155, 156f
Hormonal disorders, skeletal effects, 166167
Hormonal hypertrophy, 72
Hormones, 52, 456459. See also specific hormone
classes of, 458t
definition of, 455
secretion of, 459460, 460f
Horse, brain of, 26f
Horseshoe kidney, 691
Human chorionic gonadotropin (hCG), 479,
759, 759f, 766t
Human chorionic somatomammotropin
(hCS), 766t
Human Genome Project, 73
Humans
brain of, 26f
characteristics of, 2528, 25f
classification of, 2328, 27t
eyes of, 499
hands of, 26f
races of, 27f
vivisection of, 10, 11
Humeral condyle, 176
Humeroradial joint, 217
Humeroulnar joint, 217
Humerus, 133f, 173176, 174f, 175f, 181t
muscles of, 264267, 267f
Humoral theory, 89
Hunchback. See Kyphosis
Hunger contractions, 659t
Hunger regulation, hypothalamus and, 373
Huntingtons chorea, 790
Hyaline cartilage, 95, 96f, 97f
Hyaline membrane disease, 620, 626
Hyaloid artery, 516
Hyaloid vein, 516
Hyaluronidase, 756
Hydrocele, 722
Hydrocephalism, 382f
Hydrocephalus, 333, 381, 392
Hydrocortisone. See Cortisol
Hydrogen peroxide, 60
Hydrolysis, 51
Hydronephrosis, 693
Hydroregulation, skin and, 112
Hydroureter, 693
Hygiene, 119
Hymen, 736, 737f, 743
Hymenal tag, 300
Hyoglossus muscle, 252, 255f, 255t, 257f
Hyoid bone, 132, 157, 157f, 256, 306, 308f,
309f, 641f
Hyoid cartilage, 142f
Hyperextension, 207, 211f, 212f
Hyperglycemia, 470
Hyperkeratoses
follicular, 107t
seborrheic, 110
Hyperopia, 531, 531f
Hyperpigmentation, 107t
Hyperplasia, 72
Hypertension, 559, 587, 595596
Hyperthermia, 72
Hyperthyroidism, 481482, 482f
Hypertrophic zone, 142143
Hypertrophy, 287
compensatory, 72
hormonal, 72
Hyperventilation, 622t
Hypodermis, 107f, 112
comparison of male and female, 297, 298f
Hypogastric region, 38f, 38t
Hypoglossal canal, 146t, 148f, 151
Hypoglossal nerve (XII), 402f, 405t, 412,
412f, 413t, 498f
Hypoglossal nucleus, 376
Hypoglycemia, reactive, 482
Hyponychium, 117, 117f
Hypophyseal pouch, 478
Hypophysectomy, 480
Hypopituitarism, 480
Hypospadias, 691, 691f
Hypotension, 335
Hypothalamo-hypophyseal portal system,
464, 466f
Hypothalamo-hypophyseal tract, 463
Hypothalamus, 306f, 362t, 370f, 372373,
448449, 449f, 456, 456f, 457t
Hypothenar eminence, 275, 321f, 323f
Hypothenar muscles, 275t
Hypothermia, 72, 113
Hypothyroidism, 334, 481
Hypoventilation, 622t
Hysterectomy, 747
Hysterosalpingogram, 744, 744f
H zone, 241242, 243f
I bands, 241, 242, 242f, 243f, 245f
ICSH. See Interstitial cell-stimulating hormone
Identical twins. See Monozygotic twins
Ileocecal valve, 653, 656, 657f
Ileum, 317f, 638f, 653, 653f, 657f
Iliac crest, 181, 181f, 182f, 276f, 313, 326f, 330f
Iliac fossa, 181f, 183
Iliac tuberosity, 182f, 183
Iliacus muscle, 275, 276f, 278t, 279f, 281f, 331f
Iliad (Homer), 8
Iliococcygeus muscle, 261f
Iliocostalis cervicis muscle, 256f, 262f, 263t
Iliocostalis lumborum muscle, 262f, 263t
Iliocostalis muscle, 262
Iliocostalis thoracis muscle, 262f, 263t, 265f
Iliofemoral ligament, 220, 220f
Iliohypogastric nerve, 420t
Ilioinguinal nerve, 420t
Iliolumbar artery, 568, 569f
Iliopsoas muscle, 235f, 276, 276f, 278f, 279f
Iliotibial tract, 235f, 276, 277, 279f, 280f,
282f, 284f, 328f
Ilium, 133f, 174f, 181183, 181f, 182f
Immunosuppressive drugs, for tissue
transplants, 102
Impacted fractures, 189
Imperforate anus, 669
Imperforate hymen, 743
Impetigo, 129
Implantable contraceptives, 751
Implantation, 757759, 758f, 759f, 760f
abnormal sites of, 785, 786f
artificial, 785
Impotence, 715
Incisive foramen, 146t, 148f, 154, 155f
Incisors, 148f, 154, 304f, 640f, 642, 644t
Incontinence, urinary, 692, 694
Incus, 157, 158f, 517, 518f
Indian subcontinent, people of, 27f
Induced abortion, 748
Induced tissue, 768
Induction, 768
Inductor tissue, 768
Infancy, 777, 781t. See also Development;
Neonate
Infections
benign viral, 119
of eyes, 532
of nasal cavity, 154
of paranasal sinuses, 154
of skin, 119
829
Inferior, 36t
Inferior angle, of scapula, 173, 175f, 310
Inferior articular process, 160, 161f, 162f
Inferior cerebellar peduncles, 375
Inferior colliculi, 362t, 374, 376f
Inferior concha, 501f
Inferior constrictor muscle, 647, 647f
Inferior epigastric artery, 317f, 568, 569f
Inferior extensor retinaculum muscle, 283f,
285f, 291f, 332f
Inferior gemellus muscle, 277, 330f
Inferior gluteal artery, 568, 569f
Inferior gluteal line, 182f, 183
Inferior gluteal nerve, 424t
Inferior horn, 365f
Inferior labial frenulum, 640f
Inferior lacrimal canaliculi, 502
Inferior lacrimal punctum, 501f
Inferior limb buds, 768
Inferior lobe, of lung, 604f, 614, 615f, 616f
Inferior meatus, of palatine bone, 156f
Inferior mesenteric artery, 560f, 566, 567t,
568f, 569f
Inferior mesenteric ganglia, 440
Inferior mesenteric vein, 579, 580f
Inferior nasal concha, 147f, 150f, 154f, 155,
306f, 605f, 606
Inferior nasal meatus, 501f, 605f
Inferior oblique muscle, 252, 254f, 254t, 404,
502, 502f, 503t
Inferior olivary nuclei, 377
Inferior orbital fissure, 146t, 147f, 150f, 155
Inferior parathyroid glands, 469
Inferior peduncle, 376f
Inferior phrenic artery, 566, 567t, 568f
Inferior phrenic vein, 579
Inferior ramus
of ischium, 181f, 182f
of pubis, 181f, 182f, 183
Inferior rectal artery, 568f, 569f
Inferior rectus muscle, 252, 254f, 254t, 404,
502, 502f, 503t, 505f
Inferior sagittal sinus, 574f
Inferior suprarenal arteries, 568f
Inferior surface, of rib, 165f
Inferior thyroid veins, 577f
Inferior trunk, of brachial plexus, 416
Inferior vena cava, 314f, 548f, 549f, 550,
552f, 572, 573f, 578f, 579, 580f, 581f
Inferior vermis, 375f
Inferior vesicular artery, 568, 569f
Infertility
female, 747
male, 700, 715720
Inflammation, 126
Inflammatory skin disorders, 119, 122f
Influenza, 628
Infraglenoid tubercle, 175f
Infrahyoid muscles, 256
Infraorbital foramen, 146t, 147f, 150f, 155, 155f
Infraspinatus muscle, 217, 235f, 265f, 266t,
267, 267f, 310, 315f, 316f, 323f, 324f
Infraspinous fossa, 173, 175f
Infundibulum, 460, 461f, 478, 729f, 732
Ingestion, 635
Ingrown toenail, 327, 330f
Inguinal canal, 708
Inguinal hernia, 293
Inguinal ligament, 279f, 312f, 317f, 318, 578f
Inguinal nodes, 583, 584f
Inhalation. See Inspiration
Inheritance, 58, 782785, 783f, 783t
Inhibitory fibers, 377
Injury. See also Fractures; Trauma
cellular, 72
to knee, 207
to respiratory system, 627628
skeletal, 189191
Inner ear, 517f, 519521, 519f, 528, 528f
Inner-ear cavity, 144
Innervation, 239. See also Nervous system
dual, 446447
of gastrointestinal tract, 639640
of heart, 553f
of ovaries, 728729

830
Back Matter
Index
The McGrawHill
Companies, 2001
Index
of skin, 111
of ureters, 688f
of urethra, 688f
of urinary bladder, 688f
of uterus, 736
Inorganic compounds, 50
Insertion, 236, 236f
Inspection, 34
Inspiration, 256, 618619, 619f, 621t
muscles of, 258f, 620f
Inspiratory portion, of rhythmicity area, 623
Inspiratory reserve volume (IRV), 621t, 622f
Instincts, 345
Insula, 365f, 366t, 367
Insulin, 457t, 470471, 663
Insulin-dependent diabetes mellitus, 482
Integration, 345
Integumentary system, 31f, 105130. See also Skin
Intensity, of sound, 522
Intention tremor, 389
Interatrial septum, 547, 553f
Intercalated discs, 99, 436
Intercarpal joint, 225t
Interclavicular ligament, 215, 215f
Intercondylar eminence, 185, 186f
Intercondylar fossa, 184f, 185
Intercostal arteries, 560f
Intercostal muscles, 258, 259f, 264f
Intercostal nerves, 621t
Intercostal spaces, 164f, 165
Intercostal veins, 576, 577f
Interlobar arteries, 677, 680f
Interlobar veins, 678, 680f
Interlobular arteries, 677, 680f
Interlobular veins, 678, 680f
Intermediate commissure, 370f
Intermediate cuneiform bone, 186, 187f
Intermediate mass, 361f
Intermediate muscles, of hand, 275, 275t
Internal, 36t
Internal abdominal oblique muscle, 258,
258f, 259f, 260t, 264f, 317f, 620f
Internal acoustic meatus, 148f, 149f, 152f
Internal anal sphincter, 657, 658f
Internal anatomy
of abdomen, 315, 317f, 318f
of antebrachium, 325f
of brachium, 313f, 323f, 324f
of buttock, 330, 330f
of eyeball, 511f
of gluteal region, 330f
of hand, 325f
of head, 303304, 305f, 306f, 324f
of hip, 331f
of leg, 332f, 333f
of lower extremity, 330
of neck, 306f, 307, 309f, 324f
of shoulder, 313f, 322, 323f, 324f
of thigh, 331f
of thorax, 313315, 313f, 314f, 324f
of trunk, 313315, 316f
of upper extremity, 322
Internal carotid artery, 307, 360f, 560f, 561,
562f, 563, 563f
Internal hydrocephalus, 381
Internal iliac artery, 560f, 566, 567t, 568,
568f, 569f, 570f, 581f, 736f
Internal iliac vein, 573f, 576, 578f
Internal intercostal muscles, 258, 258f, 259f,
264f, 313f, 618, 620f
Internal jugular vein, 307, 314f, 573f, 574, 574f
Internal obturator nerve, 424t
Internal occipital crest, 149f
Internal pudendal artery, 568, 569f
Internal respiration, 603
Internal spermatic fascia, 701f
Internal surface, of rib, 165f
Internal thoracic artery, 560f, 564565, 567f
Internal thoracic vein, 573f
Internal tunic, of eyeball, 504f, 505507, 508t
Internal urinary sphincter, 686f, 687
International Congress of Anatomists, 17
Interneurons, 352
Interosseous borders, 177f
Interosseous ligaments, 198, 223f
Interosseous membrane, 272f
Interosseous talocalcaneal ligament, 223f
Interphalangeal joints, 218, 219f, 225t
Interphase, 6970, 70f, 71f
Interpretation, of nerve impulse, 488
Intersegmental reflex arc, 428
Interstitial cells, 474, 700t, 702f, 703
Interstitial cell-stimulating hormone
(ICSH), 462, 464t
Interstitial fluid, 92, 540
Interstitial systems, 140
Intertarsal joint, 225t
Intertendinous connections, 325f
Intertrochanteric crest, 184, 184f
Intertrochanteric line, 184, 184f
Intertubercular groove, 176, 176f
Interventricular aqueduct, 382f
Interventricular foramen, 381, 382f
Interventricular septum, 547, 549f,
553f, 589
Interventricular sulci, 547
Intervertebral discs, 24, 24f, 158, 158f, 161f, 162f
compression of, 95, 163
Intervertebral foramen, 161f, 162f
Intervertebral foramina, 158f, 159
Intervertebral joints, 199f, 225t
Intestinal crypt, 655, 655f
Intestinal juice enzyme, 654t
Intestinal phase, of gastric secretion, 651t
Intestinal wall, 653f
Intestine, autonomic nervous system effects
on, 445t
Intracranial tumors, 394
Intraembryonic mesoderm, 768
Intramedullary tumors, 394
Intramembranous ossification, 141
Intraocular pressure, 508
Intraspinal tumors, 394
Intrauterine device (IUD), 750f, 751
Intravenous pyelogram (IVP), 692
Intravertebral tumors, 394
Intrinsic eye muscles, 503t
Intrinsic factor, 651, 652
Intrinsic laryngeal muscles, 609
Intrinsic ocular muscles, 514
Intrinsic tongue muscles, 252
Introns, 68
Inversion, 209, 210f
In vitro fertilization, 785
Iodine, 468
Iodine-deficiency goiter, 468
Ion(s), 51
Ionic charge, of cell membrane, 54
Ipsilateral reflex, 428, 428f
Iris, 302f, 302t, 445t, 447f, 500f, 501f, 504f,
505, 505f, 506f, 508t, 512f, 516
Iron-deficiency anemia, 541
Irregular bones, 136, 136f
Irritability
of muscle, 234
of neurons, 357
IRV. See Inspiratory reserve volume
Ischemia, 594, 594f, 595f
Ischial tuberosity, 182f, 183, 701f
Ischiocavernosus muscle, 260, 260t, 261f, 701f
Ischiofemoral ligament, 220, 220f
Ischium, 133f, 174f, 181f, 182f, 183
Islam, and history of anatomy, 12
Islets of Langerhans. See Pancreatic islets
Isograft, 102
Isometric contractions, 243, 246f
Isotonic contractions, 243, 246f
Isthmus, 466
Isthmus of uterus, 734
IUD. See Intrauterine device
IVP. See Intravenous pyelogram
Janssen, Zacharius, 15
Japan, and history of anatomy, 8
Jaundice, 110, 300, 592, 670
Jaw articulation, in taxonomy, 24
Jawbone. See Mandible
Jejunum, 317f, 638f, 652, 653f

Joint(s), 196232, 225t. See also specific joints
cracking, 202
diseases of, 226227
dislocation of, 225226
disorders of, treatment of, 227229
kinds of, 205t
trauma to, 225226, 229f
Joint capsule, 200, 206
Joint cavity, 200, 206
Joint kinesthetic receptors, 493
Joint prostheses, 228f, 229
Joints
Jugular foramen, 146t, 149f, 151
Jugular fossa, 148f
Jugular notch, 164, 164f, 307, 307f, 311f
Juxtamedullary nephrons, 682, 683f
Kaitai Shinsho, 8
Kalahari bushmen, 27f
Karyotype, 782, 782f
Keratin, 5152, 85, 108
Keratinization, 108
Keratinized stratified squamous epithelium,
85, 86t
Keratinocytes, 108
Keratitis, 532
Keratohyalin, 108
Keratoplasty, 504
Keratosis, 129
Kidneys, 457t, 676f, 677f, 678f, 679f
appearance of, 677
congenital anomalies involving, 691, 691f
gross structure of, 677
microscopic structure of, 677682
position of, 677, 678f
transplantation of, 693
trauma to, 692
Kidney stone. See Renal stone
Kinesiology, 197
Kinesthetic sense, 492493
Kinocilium, 524, 524f
Klinefelters syndrome, 715
Klver-Bucy syndrome, 514
Knee, 40, 185f
injury to, 207
surface anatomy of, 327f, 328f
Kneecap. See Patella
Knee-jerk reflex, 428, 428f, 430t, 431f
Knee joint. See Tibiofemoral joint
Kupffer cells, 661
Kyphosis, 226
Labial frenulum, 640, 640f
Labial surface, of teeth, 642
Labia majora, 686f, 701f, 716, 727f, 727t,
737, 737f, 743
Labia minora, 686f, 701f, 716, 727f, 727t,
737, 737f, 743
Labioscrotal swelling, 716
Labor, 775, 776f
induction of, 463
Labyrinth, 519
Laceration, 125f
Lacrimal apparatus, 501f, 502
Lacrimal bone, 147f, 150f, 154f, 155
Lacrimal canaliculus, 501f
Lacrimal caruncle, 302f, 302t, 500, 501f
Lacrimal duct, 516
Lacrimal fluid, 502
Lacrimal gland, 445t, 501f, 502, 516
Lacrimal groove, 155f
Lacrimal puncta, 502
Lacrimal sac, 501f
Lacrimal secretions, 302
Lacrimal sulcus, 155
Lactase, 654t
Lacteals, 582, 655
Lactic acid, accumulation of, 239
Lactiferous ducts, 739, 739f

Lactiferous sinus, 259f, 739, 739f
Lacuna, 95, 98, 138f, 139f, 140
Lamarck, Jean, 16
Lambdoid suture, 144, 145f, 147f, 148f,
149f, 151
Lamellae, 98, 138
Lamellated corpuscles, 489t, 491
Lamina, 159, 160f, 161f, 162f
Lamina propria, 637
Laminectomy, 160, 169
Langerhans cells. See Nonpigmented granular
dendrocytes
Language, 369371, 371f
Lanugo, 116, 300, 773
Laparoscope, 785, 786f
Laparoscopy, 744
Laparotomy, 313
Lap suture. See Squamous suture
Large artery, 556f
Large intestine, 635, 636t, 656659, 657f,
658f, 659t
mechanical activities of, 659
regions of, 656659
structures of, 656659
Large vein, 556f
Laryngeal muscles, 609
Laryngeal nerve, in taste sensation, 498f
Laryngeal prominence, 306, 608
Laryngitis, 608, 628
Laryngopharynx, 605f, 608, 641f, 647
Laryngotracheal bud, 624
Larynx, 256, 604f, 608609, 608f, 610f, 618t
Lateral, 36t
Lateral alar cartilage, 305f
Lateral angle, of scapula, 173, 175f
Lateral apertures, 381
Lateral border, of scapula, 173, 175f
Lateral cartilage, 605, 605f
Lateral cervical ligament. See Cardinal ligament
Lateral collateral ligament, 222, 222f
Lateral commissure, 302f, 302t, 500, 501f
Lateral condyle, 184, 184f
of tibia, 185, 186f
Lateral cord, of brachial plexus, 416
Lateral corticospinal tract, 387, 388t, 389t
Lateral crural muscles, 283, 284f
Lateral cuneiform bone, 186, 187f
Lateral cutaneous femoral nerve, 420t
Lateral epicondyle
of femur, 184f, 185f, 327f, 328f
of humerus, 176, 176f, 320f, 321f
of tibia, 185
Lateral epicondylitis, 177, 338
Lateral femoral circumflex artery, 568, 570f
Lateral femoral cutaneous nerve, 401f
Lateral flexion, 211f
Lateral funiculus, 386, 386f
Lateral genicular arteries, 570f
Lateral geniculate, 513
Lateral horn, 384, 386f, 390
Lateral ligament, of temporomandibular
joint, 214, 214f
Lateral malleolus, 185, 186f, 284f, 291f, 329f
Lateral membranous ampullae, 519f
Lateral meniscus, 221f, 222f
Lateral patellar retinaculum, 221, 221f
Lateral plantar artery, 570571, 570f
Lateral plantar nerve, 423, 424t, 425f
Lateral plantar vein, 578f
Lateral plate
of pterygoid process, 148f
of sphenoid bone, 154f
Lateral process, of breast, 312f
Lateral pterygoid muscle, 250, 253f, 253t
Lateral pterygoid process, 148f, 153f
Lateral rectus muscle, 252, 254f, 254t, 406,
502, 502f, 503t
Lateral sacral artery, 568
Lateral sacral crest, 163f
Lateral semicircular canal, 519f
Lateral spinothalamic tract, 388t, 389t,
491, 493
Lateral sulcus, 360f, 363, 364f, 366f
Lateral superior artery, 569f

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Lateral ventricle, 306f, 361f, 363, 365f, 381, 382f
Latin, and anatomical terminology, 2
Latissimus dorsi muscle, 235f, 259f, 264265,
264f, 265, 265f, 266t, 267, 268f,
269f, 270f, 310, 311f, 313f, 315f,
316f, 320f, 323f, 324f
Laughing, 622t
Lazy eye, 511
Learning, 345, 363
Lebers hereditary optic neuropathy, 58
Leeuwenhoek, Antoni van, 1516
Left atrioventricular valve, 548t, 549f, 550
Left atrium, 548f, 549f, 550, 552f
Left auricle, 561f
Left brachiocephalic vein, 314f, 561f, 573f, 577f
Left bronchus, 314f
Left bundle branch, 553f
Left colic flexure. See Splenic flexure
Left common carotid artery, 314f, 549f, 559,
560f, 561f, 567t
Left common iliac artery, 566, 568f, 569f
Left coronary artery, 550, 552f, 559, 567t
Left costal margin, of rib cage, 312
Left gastric artery, 566, 567t, 568f
Left gastric vein, 579, 580f
Left gastroepiploic artery, 568f
Left gastroepiploic vein, 579, 580f
Left hemisphere, of brain, 363, 366f
Left hepatic vein, 579
Left hypochondriac region, 38f, 38t
Left inguinal region, 38f, 38t
Left internal jugular vein, 561f
Left lateral region, 38f, 38t
Left lobe, of liver, 314f, 317f, 660, 661f
Left lung, 314f, 604f, 614, 616f
Left ovarian vein, 579
Left principal bronchus, 604f, 610, 610f, 612f
Left pulmonary artery, 548f, 549f, 550, 561f
Left pulmonary vein, 548f, 549f, 561f
Left subclavian artery, 314f, 549f, 559, 560f,
561f, 564, 567t
Left subclavian vein, 572, 573f, 584f
Left suprarenal vein, 579
Left testicular vein, 573f, 579
Left ventricle, 314f, 548f, 549f, 550, 552f
Leg, 40. See also Lower extremity
bones of, 185
internal anatomy of, 332f, 333f
Lens, 500f, 504f, 505, 505f, 506f, 507f, 508t,
510, 512f
Lens capsule, 505, 506f, 512f, 516
Lens fovea, 515
Lens placode, 515
Lens vesicle, 516
Lentiform nucleus, 365f, 368369, 370f
Leonardo da Vinci, 13, 14f
Lesions, 119
Lesser cornua, 157, 157f
Lesser curvature, of stomach, 650, 650f
Lesser multangular. See Trapezoid bone
Lesser occipital nerve, 315f, 416t
Lesser omentum, 42f, 4345, 637, 638f
Lesser palatine foramen, 146t, 155, 156f
Lesser pelvis, 180, 183f
Lesser sciatic notch, 182f, 183
Lesser splanchnic nerve, 440
Lesser trochanter, of femur, 182f, 184, 184f
Lesser tubercle, of humerus, 176, 176f
Lesser wing, of sphenoid bone, 147f, 148f,
149f, 150f, 152, 153f
Leukemia, 542, 593
Leukocytes, 98, 99f, 540, 541f, 542, 544t
circulation of, 539
shape of, 50f
size of, 49
Leukocytosis, 542, 593
Leukopenia, 593
Leukopoiesis, 544
Levator anguli oris muscle, 250f, 251f, 251t
Levator ani muscle, 260, 260t, 261f, 281f,
658f
Levator costarum muscle, 262f
Levator labii superioris muscle, 250f, 251f,
251t, 305f
Levator palpebrae superioris muscle, 252,

254f, 404, 500, 500f, 502f
Levator scapulae muscle, 256f, 257f, 259f, 264,
265f, 266t, 267f, 308f, 309f, 315f
Levers, 210211, 213f
Levodopa, 351, 390
Leydig cells. See Interstitial cells
LH. See Luteinizing hormone
LH surge, 741
Lifting, proper procedure for, 262
Ligaments, 92, 202
Ligamentum arteriosum, 548f, 561f
Ligamentum capitis femoris, 220, 220f
Ligamentum denticulatum, 379
Ligamentum nuchae, 307, 309f
Ligamentum teres, 661, 729f, 735
Light
refraction of, 509510, 512f
transmission of, 509
Light microscope, 17, 17f, 78
Limb buds, 192, 192f
Limbic system, and autonomic nervous
system, 449, 450f
Linea alba, 258, 259f, 311f, 312, 312f,
313, 317f
Linea albicans, 111, 111f
Linea aspera, 184, 184f
Linear bone growth, 137
Linea semilunaris, 312f, 313
Lines of tension, on skin, 110, 111f
Lingual artery, 562f, 563
Lingual frenulum, 305f, 640f, 642
Lingual nerve, in taste sensation, 498f
Lingual surface, of teeth, 642
Lingual tonsils, 605f, 608, 641f, 642
Lipase, 654t
Lipids, 51, 52, 52t
Lipid solubility, 54
Lipoproteins, 51
Lips, 303, 304f, 304t, 640, 640f, 641f
Liver, 314f, 317f, 318f, 635, 635f, 638f,
660662, 661f, 662f
disorders of, 670
effects of alcohol on, 579
location of, 660f
Mesopotamian views on, 7, 7f
Liver cells. See Hepatocytes
Liver lobules, 661, 662f, 663f
Liver sinusoids, 661, 662f, 663f
Liver spots, 110
Lobar bronchus, 604f, 610, 610f
Lobes
of liver, 660
of lungs, 604f, 616f
of mammary glands, 739, 739f
Lobules
of lungs, 616f
of mammary glands, 739, 739f
of testis, 701f, 702
Lobus nervosa, 460, 478
Lobus nervosus, 457t
Local anesthetics, 392
Local effects, 122
Lockjaw, 250
Locomotion, bipedal, 2527
Long bones, 135136, 136f, 137f
growth of, 140f
Longissimus capitis muscle, 255, 256f, 262f,
263t, 309f
Longissimus cervicis muscle, 256f, 262f,
263t, 309f
Longissimus muscle, 262
Longissimus thoracic muscle, 256f, 262f,
263t, 265f
Longitudinal arch, 188, 188f, 327, 329f
Longitudinal cerebral fissure, 363, 364f, 365f
Longitudinal muscle, of stomach, 650f
Long plantar ligament, 223f
Loop of Henle. See Nephron loop
Loose connective (areolar) tissue, 91,
91f, 95t
Lordosis, 226
Lower esophageal sphincter, 648
Lower extremity, 40
arteries of, 566571, 570f
bones of, 132t, 134, 178, 188t
diseases of, 339
internal anatomy of, 330
of neonate, 299t
trauma to, 339
veins of, 576579, 578f
Lower nerve block, 157
Lower respiratory system, 603
Lower spinal nerves, in expiration, 621t
Lumbago, 287
Lumbar arteries, 566, 567t
Lumbar curve, 158f, 159
Lumbar enlargement, of spinal cord, 384, 385f
Lumbar nerves, 401f, 413
Lumbar nodes, 583
Lumbar plexus, 385f, 401f, 420t, 421, 421f
Lumbar puncture, 391, 391f
Lumbar region, 37
Lumbar splanchnic nerve, 440
Lumbar veins, 579
Lumbar vertebrae, 159, 161162, 162f,
163t, 164f
Lumbosacral plexus, 421, 424f
Lumbosacral trunk, 421
Lumbricales muscle, 272f, 274f, 275, 275t,
290f, 325f
Lumen, 556f
Lunate bone, 178, 179f, 180f
Lung cancer, 615, 630f
Lungs, 314f, 614615, 615f, 616f, 618t
autonomic nervous system effects on,
445t, 448t
location of, 36
position of, 614f, 617f
respiratory division of, 612
serous membranes surrounding, 43f
Lung tissue, 613f
Lung volume, 619f
Lunula, 117, 117f
Luteinizing hormone (LH), 462, 464t, 741
Luxation, 225226
Lymph, 540, 582
Lymphadenitis, 596
Lymphangitis, 596
Lymphatic capillaries, 582
Lymphatic nodules, 583
Lymphatic system, 31f, 582586, 584f
disorders of, 596
relation to circulatory system, 540, 583f
Lymphatic vessels, 540, 582, 583f
Lymph ducts, 582
Lymph nodes, 540, 583584, 584f, 585f,
587t, 740, 741f
Lymphoblasts, 544
Lymphocytes, 542, 542f, 544t
Lymphoid organs, 584586, 584f, 587t
Lymphoid tissue, 544
Lymphoma, 596
Lysosomes, 53t, 54f, 59, 60, 60f
Lysozyme, 502
Macrophagic cells, 108
Macula, 515, 515f
Macula lutea, 506507
Macular degeneration, 532
Magnetic resonance imaging (MRI), 18, 19f
Major calyx, 677, 679f, 682
Major vestibular glands, 738
Male
comparison with female physique, 780, 780f
distribution of hair in, 115f
growth rates of, 726, 728f
hypodermis of, 297, 298f
sexual maturation of, 779
Male infertility, 700, 715720
Male pseudohermaphroditism, 714
Male reproductive system, 32f, 697724,
699f, 700t, 701f
clinical considerations, 714722, 744751
831

comparison with female, 743t
diseases of, 720721
Male sterility, 720
Malignant melanoma, 121, 123
Malignant neoplasms, 73
Malleus, 30, 157, 158f, 517, 518f
Malpighi, Marcello, 15
Malposition of teeth, 669f
Maltase, 654t
Mamillary body, 370f
Mammalia, 24
Mammary alveoli, 739
Mammary ducts, 739, 739f
Mammary gland lobule, 259f
Mammary glands, 25f, 119, 119f, 259f, 726,
727t, 738740, 739f, 740f, 741f. See
also Breast(s)
diseases of, 748
lymphatics of, 584f
Mammary lobes, 259f
Mammary region, 3536
Mammary ridge, 25f
Mammary tissue, following weaning, 59
Mammography, 748, 749f
Mandible, 133f, 142f, 145f, 147f, 148f, 156f,
157, 174f, 306f, 308f
Mandibular condyle, 149f
Mandibular foramen, 146t, 149f, 156f, 157
Mandibular fossa, 146, 148f, 149f, 152f
Mandibular margin, 156f
Mandibular nerve, 404t, 405, 406
Mandibular nerve block, 406
Mandibular notch, 147f, 156f, 157
Manic-depressive psychosis, 393
Manometer, 391
Manubrium, 164, 164f, 306f
Manus, 38. See also Hand
articulations of, 204f
bones of, 178, 179f, 180f
Marfans syndrome, 790
Marginal artery, 552f
Massage, 491
Masseteric tuberosity, 156f
Masseter muscle, 235f, 250, 250f, 251f,
253f, 253t
Mass movements, 659, 659t
Mast cells, 91
Mastectomy, 264, 748
Mastication, 635, 659t
muscles of, 250, 253f, 253t
Mastoidal air cells, 517
Mastoid cells, 151
Mastoid fontanels. See Posterolateral
fontanels
Mastoid foramen, 148f, 149f, 151
Mastoiditis, 151, 519
Mastoid part, of temporal bone, 151, 152f
Mastoid process, 147f, 148f, 149f, 151,
152f, 308f
Masturbation, 720
Matrix, 78
Maxilla, 133f, 142f, 145f, 147f, 148f,
150f, 154155, 154f, 155f,
174f, 306f
Maxillary artery, 562f, 563564
Maxillary nerve, 404t, 405, 406
Maxillary nerve block, 406
Maxillary sinus, 150f, 151f, 155, 155f, 605f,
607, 607f
McBurneys point, 312f, 313
Meatal plug, 529
Mechanoreceptors, 489, 490
Meckels cartilage, 142f
Meckels diverticulum, 336
Medial, 36t
Medial border, of scapula, 173, 175f
Medial collateral ligament, 222f
Medial commissure, 302f, 302t, 500, 501f
Medial condyle, 184, 184f, 185, 186f
Medial cord, of brachial plexus, 416
Medial cuneiform bone, 186, 187f

832
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Medial epicondyle
of femur, 184f, 327f, 328f
of humerus, 176, 176f, 320f, 321f
of tibia, 185
Medial femoral circumflex artery, 568, 570f
Medial genicular arteries, 570f
Medial lemniscus, 493
Medial malleolus, 185, 291f, 329f
Medial meniscus, 221f, 222f
Medial patellar retinaculum, 221, 221f
Medial plantar artery, 570571, 570f
Medial plantar nerve, 423, 424t, 425f
Medial plate
of pterygoid process, 148f
of sphenoid bone, 154f
Medial pterygoid muscle, 250, 253f, 253t
Medial pterygoid process, 148f, 153f
Medial rectus muscle, 252, 254f, 254t, 404,
502, 502f, 503t
Medial reticulospinal tract, 388t, 389t
Medial talocalcaneal ligament, 223f
Median aperture, 381
Median cubital vein, 321f, 573f, 575, 575f
Median eminence, 464
Median furrow, 309, 310
Median nerve, 322, 401f, 418t, 419, 420f
Median palatine suture, 148f
Median sacral crest, 162, 163f
Median umbilical ligament, 685, 764
Median vein, 321f
Mediastinal surface, of lungs, 614
Mediastinum, 41, 41f, 44f, 614, 617f
Medical genetics, 7273
Medium-sized artery, 556f
Medium-sized vein, 556f
Medulla oblongata, 360f, 361f, 362t, 370f,
374f, 375f, 376377, 376f, 377f,
385f, 448, 448t
Medullary cavity, 136137
Megakaryoblasts, 544
Megakaryocytes, 542
Meiosis, 703704, 704t, 705f, 757f
Meissners corpuscles. See Corpuscles of touch
Meissners plexus. See Submucosal plexus
Melancholy, 9
Melanin, 108, 109, 121
Melanoblasts, 121
Melanocytes, 108, 109f
Melanocyte-stimulating hormone (MSH),
463, 464t
Melanoma, 112, 129
Melatonin, 457t, 475476
Membranous ampulla, 519f, 520
Membranous epithelia, 7981
Membranous labyrinth, 519, 519f
Membranous part, of urethra, 686f, 687, 709
Memory, 345
Menarche, 726, 779
Menes, 7
Mnieres disease, 533
Meningeal branch, of spinal nerve, 414
Meningeal layer, of dura mater, 378, 378f
Meningeal veins, 574
Meninges, 345t, 358, 378381, 379f, 381f
Meningiomas, 394
Meningitis, 154, 394
Meningomyelocele, 166f
Meniscus, 95, 202, 206
Menopause, 726
Menorrhagia, 747
Menses, 736
Menstruation, 726, 740741, 742f, 742t, 747
Mental artery, 305f
Mental foramen, 146t, 147f, 156f, 157
Mental illness, 393
Mentalis muscle, 250f, 251f, 251t, 305f
Mental nerve, 305f
Mental protuberance, 149f, 156f
Mental retardation, 392
Mental spine, 156f
Mental vein, 305f
Mentolabial sulcus, 304f
Meridional fibers, 506f
Merkel cells. See Tactile cells
Merocrine glands, 87, 88f, 89t
Mesencephalic aqueduct, 361f, 374, 381, 382f

Mesencephalon, 347, 362t, 373374
Mesenchyme, 90, 90f, 120, 141, 768
Mesenteric lymph nodes, 584f
Mesenteric patches, 584, 653
Mesentery, 42f, 45, 318f, 636, 637f, 638f, 652
Mesocolon, 636, 638f, 657f
Mesoderm, 80, 80t, 760, 761f
Mesonephric duct, 716
Mesonephros, 689, 690
Mesopotamia, and history of anatomy, 58
Mesothelium, 81
Mesovarium, 728, 729f
Messenger RNA (mRNA), 67, 69, 69f
Metabolism, 49
Metacarpal arteries, 566
Metacarpal bones, 133f, 174f, 179f, 180f,
181t, 205f
Metacarpophalangeal joints, 204f, 218, 219f,
225t, 322f
Metacarpus, 38, 178
Metanephrogenic mass, 690
Metanephros, 689, 690
Metaphase, 70, 71f, 704t, 705f, 757f
Metaplasia, 72
Metarterioles, 555
Metastases, 73, 394, 584
Metatarsal bones, 133f, 174f, 187f, 188f,
188t, 329f
Metatarsophalangeal joints, 186
Metatarsus, 40, 186
Metencephalon, 347, 362t, 374376
Metopic suture, 145
Metrorrhagia, 747
MI. See Myocardial infarction
Michelangelo, 2
Microcephaly, 333, 392
Microcirculation, 557f
Microglia, 348351, 350f
Micromelia, 192, 337
Microscope and microscopy
and cytology, 49
types of, 17f
Microscopic anatomy, 17
Microtubules, 53t, 54f, 61, 62f
Microvilli, 5457, 56f, 653655, 655f
Micturition, 676, 687688, 688t
Midbrain. See Mesencephalon
Midclavicular line, 310
Middle Ages, and history of anatomy, 12
Middle cerebellar peduncles, 375
Middle cerebral artery, 563, 563f
Middle concha, 501f
Middle constrictor muscle, 647, 647f
Middle ear, 517519, 517f, 528, 530f
Middle-ear cavity, 42, 45f, 144, 158f
Middle lobe, of lung, 604f, 614, 615f, 616f
Middle meatus, of palatine bone, 156f
Middle nasal concha, 147f, 150f, 153, 153f,
154f, 605f, 606
Middle peduncle, 376f
Middle phalanx, 179f, 180f, 187f
Middle rectal artery, 568, 568f, 569f
Middle sacral artery, 566
Middle superior artery, 569f
Middle suprarenal arteries, 568f
Middle trunk, of brachial plexus, 416
Middle turn, of cochlea, 520f
Middle vesicular artery, 568, 569f
Midgut, 666667, 667f
Midgut loop, 666
Midsagittal (median) plane, 33
Migraine headaches, 392
Milia, 300
Milk teeth. See Deciduous teeth
Mineralocorticoids, 471, 473t
Minerals, 52t, 134135
Minor calyx, 677, 677f, 679f, 682
Minor vestibular glands, 738
Miscarriage, 748
Mitochondria, 53t, 54f, 58, 60f, 242f, 247f
Mitosis, 65, 70, 70f, 71f, 706f
Mitral valve. See Left atrioventricular valve
Mixed glands, 456, 663
Mixed nerves, 352, 401, 403

Mnemonic devices, 403
Modified radical mastectomy, 748
Molars, 148f, 154, 640f, 642, 643f, 644t
Molding, of fetal head, 144
Molecules, 28, 50
Mongolian spots, 300
Mongoloid (Thailand), 27f
Monoblasts, 544
Monocytes, 542, 542f, 544t
Monohybrid cross, 784
Mononucleosis, 593
Monosaccharides, 52
Monosomy, 73
Monosynaptic reflex arc, 428
Monozygotic twins, 785786, 788f
Mons pubis, 701f, 737, 737f
Morphogenesis, 756
Morula, 79, 757, 760t
Motor area, of cerebral cortex, 367f, 370f, 371f
Motor end plates, 244, 247f
Motor nerves, 352, 355f, 356f, 401, 403
Motor neuron axon, 247f
Motor neurons, 239, 345t, 349f, 352, 427, 427f
Motor root, of trigeminal nerve, 405
Motor speech area, 364f, 366f, 369371, 371f
Motor unit, 244246
Mounting, 102
Mouth. See Oral cavity
Mouth-to-mouth resuscitation, 629f
Mouth-to-nose resuscitation, 629f
Movement, 211f
muscular system and, 234
skeleton and, 134
M phase, 70
MRI. See Magnetic resonance imaging
mRNA. See Messenger RNA
MS. See Multiple sclerosis
MSH. See Melanocyte-stimulating hormone
Mucocutaneous corpuscles, 491
Mucoid connective tissue, 90, 767
Mucosa, 637, 639f
of stomach, 650f, 651f
of ureters, 684
of uterine tube, 734
Mucosal layer, 736
Mucous cells, 645, 651f
Mucous connective tissue, 90
Mucous glands, autonomic nervous system
effects on, 445t
Mucous membranes, 4243, 85, 303
Mller, Johannes, 16
Mllerian duct. See Paramesonephric duct
Multicellular glands, 8789, 88f, 89t
Multifidus muscle, 262f
Multiple pregnancy, 785786, 787f, 788f
Multiple sclerosis (MS), 395
Multipolar neurons, 352, 356f
Multiunit smooth muscle, 437, 437t
Mummies, Egyptian, 7, 7f
Mumps, 670
Muscarine, 445
Muscle(s). See also Cardiac muscle; Skeletal
muscle; specific muscle
aging of, 288289, 292f
condition of, 285
diseases of, 287288
functional conditions of, 286287
functions of, 234
naming of, 246247
neoplasms of, 288
soreness of, 239
trauma to, 92
Muscle atrophy, 102, 286
Muscle cells, 70
Muscle cramp, 286
Muscle fibers
architecture of, 238
skeletal, 240243, 241f, 242f, 243f
Muscle tissue, 78, 99, 100f, 100t, 234
Muscular arteries, 555
Muscular dystrophy, 287, 339
Muscularis
of ureters, 684
of uterine tube, 734
of vagina, 737
Muscularis mucosa, 637
Muscular system, 31f, 233295. See also
specific muscles
Musculocutaneous nerve, 418t, 419, 419f
Musculotendinous cuff, 217, 267
Mutations, 73
Myalgia, 293
Myasthenia gravis, 288
Myelencephalon, 347, 362t, 376377
Myelin, 349f, 351
Myelination, 351, 352f, 353f
Myelin layer, 351, 352f, 353f
Myeloblasts, 544
Myenteric plexus, 637
Mylohyoid muscle, 255f, 256, 257f, 257t,
308f, 309f
Myoblasts, 248
Myocardial disease, 594
Myocardial infarction (MI), 335, 595
Myocardial ischemia, 335, 594, 594f, 595f
Myocardium, 436437, 546, 547t
Myofibrils, 241, 241f, 242f, 243f, 244f, 247f
Myofilaments, 61, 241, 241f, 243f
Myoglobin, 58
Myokymia, 293
Myoma, 293
Myometrium, 729f, 736
Myoneural junction. See Neuromuscular
junction
Myopathy, 293
Myopia, 531, 531f
Myosin, 241
Myosin myofilaments, 243f
Myotomes, 248
Myotomy, 293
Myotonia, 293
Myringotomy, 533, 533f
Myxedema, 481
Nail(s), 117, 117f, 300, 322f
Nail bed, 117, 117f, 300
Nail fold, 117, 117f
Nail groove, 117, 117f
Nail matrix, 117, 117f
Nares. See Nostrils
Nasal bones, 142f, 145f, 147f, 148f, 150f,
154f, 155, 605, 605f
Nasal cavity, 42, 45f, 144, 154, 154f, 154t,
501f, 604f, 605, 605f, 641f
Nasal crest, 155f, 156f
Nasal fossa, 144, 153, 605
Nasalis muscle, 250f, 251f, 251t
Nasal meatus, 606
Nasal notch, 155f
Nasal region, of face, 301, 303, 304f, 304t
Nasal septum, 144, 501f, 605
Nasal spine, 155f
Nasal vestibule, 605, 605f
Nasolacrimal canal, 146t, 155
Nasolacrimal duct, 501f, 502, 605f
Nasopharynx, 605f, 606, 607, 641f, 647
Natal cleft, 326, 326f
Nausea, 527, 652, 672
Navel. See Umbilicus
Navicular bone, 186, 187f, 188f. See also
Scaphoid bone
Nearsightedness. See Myopia
Neck, 35
arteries of, 307, 561564, 562f
developmental disorders involving, 333
diseases of, 334
of humerus, 176, 176f
internal anatomy of, 306f, 307, 309f, 324f
muscles of, 254256, 257f, 257t, 259f, 265f
of neonate, 299t

Back Matter
Index
The McGrawHill
Companies, 2001
Index
trauma to, 333334
triangles of, 307, 307f, 308f, 308t
Neck (anatomical)
of condylar process, 156f
of femur, 182f, 184, 184f
of fibula, 186f
of humerus, 176, 176f
of radius, 177f
of rib, 165, 165f
of tooth, 645f
veins of, 572574, 574f
Necrosis, 102, 595
Negative feedback system, 459, 459f
Negroid (Africa), 27f
Neisseria gonorrhoeae, 720
Neonatal period, 776, 781t
Neonate. See also Development
brain of, 359
flexion position of, 299, 299f
general appearance of, 299300
palpable structures of, 300
surface anatomy of, 298300, 299t
Neoplasms. See also Cancer; Tumors
benign, 73
of bone, 167
of central nervous system, 394
malignant, 73
metastatic, 394
of muscle, 288
primary, 394
secondary, 394
of skin, 119121
uterine, 747
Nephritis, 692
Nephron, 677, 679f, 680682, 683t
Nephron loop, 679f, 680f, 682, 683t
Nerve(s), 345t. See also specific nerves
classification of, 352354, 356f
structure of, 356f
Nerve block, 157, 391392, 406, 775
Nerve cells. See Neurons
Nerve fiber, 101, 348
Nerve impulse, 357
conduction of, 384, 488
interpretation of, 488
transmission of, 357358
Nerve plexus, 345t, 415423
Nervous layer, of retina, 505
Nervous system, 32f. See also Autonomic
nervous system; Central nervous
system; Peripheral nervous system
degenerative diseases of, 395396
and endocrine system, 455, 455t
functions of, 345
senescence of, 395
Nervous tissue, 78, 100101, 343399
Neural crest, 347
Neural folds, 346
Neural groove, 346
Neural pathways
for equilibrium, 526527
for hearing, 523, 523f
for somatic sensation, 493494, 495f
for visual sense, 513514
Neural plate, 346
Neural tube, 347
Neurocranium, 141, 142f
Neuroendocrine reflexes, 463
Neuroepithelium, 80
Neurofibril(s), 348, 349f
Neurofibril nodes, 349f, 351
Neuroglia, 101, 101f, 346, 348351, 351t
Neurohypophysis, 373, 460, 461f, 463f,
464t, 477
Neurolemmal sheath, 351
Neurolemmocytes, 347, 348, 349f, 351t
Neurological assessment, 391, 412, 413t
Neurology, 344345
Neuromuscular cleft, 244, 247f
Neuromuscular junction, 244, 247f
Neuromuscular spindles, 493
Neurons, 101, 101f, 345, 345t, 348, 350f

damage to, 70
development of, 346
mitotic activity of, 348
myelination of, 351, 352f, 353f
regeneration of, 354f
shape of, 50f
size of, 49
Neuropeptides, 359
Neuropores, 347
Neurosis, 393
Neurosyphilis, 395
Neurotendinous receptors, 493
Neurotransmitters
of autonomic nervous system, 444, 446f
diseases involving, 396
Neutrophils, 542, 542f, 544t
Nevi, 119, 129
Ngatatjara man, 27f
Nicotine, harmful effects of, 559
Nipples, 25f, 259f, 310, 311f, 312f, 739, 739f
Nissl bodies. See Chromatophilic substances
Nitrogenous bases, 6566, 67f
NMR. See Nuclear magnetic resonance
Nociceptors, 490, 491492
Nocturnal emission, 714
Nodes of Ranvier. See Neurofibril nodes
Nonarticulating prominences, of bones, 136t
Nondisplaced fractures, 190
Non-Hodgkins lymphomas, 596
Non-insulin-dependent diabetes mellitus, 482
Nonkeratinized stratified squamous
epithelium, 85, 86t
Nonpigmented granular dendrocytes, 108
Nonrespiratory air movements, 621, 622t
Norepinephrine, 440, 444, 446f, 457f, 458,
472, 473t
Normal amenorrhea, 747
Norplant, 750f
Nose, 605607, 605f, 618t. See also Nasal cavity
Nostrils, 303, 304f, 304t, 604f, 605, 605f, 606
Notochord, 23, 23f, 767
Nucha, 306
Nuchal ligament, 160
Nuclear magnetic resonance (NMR), 18
Nuclear membrane (envelope), 53t, 54f, 63
Nuclear pores, 63
Nuclease, 654t
Nucleic acids, 52t
Nucleolemma cisterna, 63
Nucleolus, 53, 53t, 54f, 63, 349f
Nucleoplasm, 57
Nucleoproteins, 51
Nucleotides, 6566
Nucleus
in brain, 359
of cell, 53, 54f, 63, 64f
of neurons, 348, 349f, 488
Nucleus ambiguus, 376, 411
Nucleus cuneatus, 377
Nucleus gracilis, 377
Nucleus pulposus, 23, 24, 24f
Nurse cells. See Sustentacular cells
Nutrient foramen, 137, 176, 176f
Nutritional disorders. See also specific disorders
skeletal effects, 166167
skin effects of, 107t
Obesity, 636t, 778
Oblique fissure, of lung, 615f
Oblique fractures, 189, 190f
Oblique line, 156f
Oblique muscle(s), 252, 502, 503t
Oblique muscle layer, of stomach, 650, 650f
Oblique popliteal ligament, 221, 221f
Obliquus capitis inferior muscle, 256f, 262f
Obliquus capitis superior muscle, 256f, 262f
Obturator artery, 568, 569f, 570f
Obturator externus muscle, 277, 278f
Obturator foramen, 181f, 182f, 183, 183f
Obturator internus muscle, 261f, 277,
281f, 330f
Obturator nerve, 420t, 421, 422f
Occipital artery, 309f, 315f, 562f, 563
Occipital bone, 133f, 142f, 145f, 147f, 148f,

149f, 151152, 174f, 309f
Occipital condyle, 148f, 149f, 151
Occipital fontanel. See Posterior fontanel
Occipitalis muscle, 235f, 251f, 251t
Occipital lobe, 360f, 361f, 364f, 366367,
366f, 366t
Occipital vein, 574f
Occipitomastoid suture, 149f
Ocular muscles, 252, 254f, 254t, 503f
Ocular region, of face, 301, 302, 302f, 302t
Oculomotor nerve (III), 402f, 403404,
404t, 408f, 413t
Odontoid process. See Dens, of axis
Oily skin, 107t
Olecranon, 177, 177f, 320f, 321f
Olecranon bursa, 217
Olecranon fossa, 176, 176f
Olfaction, 496f
Olfactory bulb, 402f, 403, 406f, 496, 496f
Olfactory hairs, 495, 496f
Olfactory nerve (I), 402f, 403, 404t, 406f,
413t, 496
Olfactory nerve fibers, 496f
Olfactory pathway, 406f
Olfactory pit, 624
Olfactory placode, 624
Olfactory receptor cells, 495, 496f
Olfactory sense, 495496
Olfactory tract, 360f, 402f, 403, 406f, 496
Oligodendrocytes, 348, 350f, 351, 351t
Oligodendrogliomas, 394
Oligospermia, 714
Oliguria, 692
Omoclavicular triangle, 308f, 308t
Omohyoid muscle, 255f, 256, 257f, 257t,
259f, 267f, 308f, 309f
Omos, 38
Omotracheal triangle, 308f, 308t
Ontogeny, 690
Oocytes, 58, 698, 729, 729f, 733f, 757f
primary, 729, 731f, 733f
secondary, 730, 730f, 731f, 732f, 733f
Oogenesis, 733f
Oogonia, 729, 743
Oophoritis, 748
Open fractures, 189
Openings, of bones, 136t
Ophthalmic artery, 562f, 563
Ophthalmic nerve, 404t, 405, 406
Ophthalmic vein, 574
Ophthalmology, 302, 527
Ophthalmoscope, 511f, 527
Opponens digiti minimi muscle, 274f, 275,
275t, 290f, 325f
Opponens pollicis muscle, 274f, 275, 275t
Opposable thumbs, 26f, 27
Optic canal, 147f, 150f, 152, 153f
Optic chiasma, 370f, 402f, 403, 407f, 513
Optic cup, 516
Optic disc, 504f, 507, 511f
Optic fissure, 516
Optic foramen, 146t, 149f
Optic fovea, 529
Optic nerve (II), 360f, 361f, 402f, 403, 404t,
407f, 408f, 413t, 504, 504f, 505f, 516
Optic radiation, 513
Optic tract, 403, 407f, 513
Optic vesicle, 515
Optometrist, 527
Optometry, 302, 527
Oral cavity, 42, 45f, 85, 144, 303, 305f, 306f,
605f, 635, 635f, 636t, 640, 640f,
641f, 659t
Oral contraceptives, 749, 750f
Oral membrane, 665
Oral orifices, 640, 640f
Oral pit, 665
Oral region, of face, 301, 303, 304f, 304t
Oral vestibule, 605f
Ora serrata, 505, 505f
Orbicularis oculi muscle, 235f, 250f, 251f,
251t, 305f, 500, 500f
Orbicularis oris muscle, 235f, 250f, 251f,
251t, 253f, 305f
833
Orbit, of eye, 42, 45f, 144, 145, 150f, 151t,

301, 499
Orbital fat body, 305f
Orbital muscle, 250f
Orbital plate
of ethmoid bone, 147f, 150f, 153f
of frontal bone, 150f
Orbital process, of palatine bone, 156f
Orbital surface, of palatine bone, 156f
Orbitosphenoid, 142f
Orchiectomy, 722
Orchitis, 722
Order Primates, 24, 27t
Organ(s), 28, 42
Organelles, 28, 53, 5761
Organic compounds, 50
Organ of Corti. See Spiral organ
Organs of Ruffini, 489t, 491
Orgasm
female, 738
male, 713714
Origin, of muscles, 236, 236f
Oronasal membrane, 624
Oropharynx, 605f, 608, 641f, 647
Orthopedics, 169
Orthopnea, 622t
Os coxae, 133f, 174f, 180, 182f, 188t
Osmosis, 55t, 56f
Os penis, 713
Ossification, 141, 143t
Ossification zone, 143
Osteitis, 169
Osteoarthritis, 226227
Osteoblastoma, 170
Osteoblasts, 138, 138f, 140
Osteochondritis, 170
Osteoclasts, 138, 138f
Osteocytes, 50f, 9798, 138, 138f, 139f
Osteogenic cells, 138
Osteogenic sarcoma, 167
Osteoids, 140142
Osteology, 132
Osteomalacia, 166
Osteomas, 167
Osteomyelitis, 170, 334
Osteonecrosis, 170
Osteons, 138, 138f, 139f
Osteopathology, 170
Osteoporosis, 167168, 339
Osteotomy, 170
Otic capsule, 529
Otic ganglion, 441
Otocyst, 529
Otolithic membrane. See Statoconial
membrane
Otoliths. See Statoconia
Otorhinolaryngologist, 302
Otorhinolaryngology, 527
Otosclerosis, 533
Otoscope, 527
Outer ear, 516, 517f, 528, 530f
Ova. See Oocytes
Oval window. See Vestibular window
Ovarian arteries, 566, 567t, 733f, 736f
Ovarian cortex, 728
Ovarian cycle, 729732, 758f
Ovarian follicles, 475
Ovarian fossa, 728
Ovarian ligament, 728, 729f
Ovarian medulla, 728
Ovarian tumors, 746
Ovarian veins, 733f
Ovary(ies), 456, 456f, 457t, 474475, 726,
727f, 727t, 729f, 733f
blood supply to, 728729
function of, 728732
innervation of, 728729
palpation of, 729
position of, 728729
problems involving, 746747
Overriding aorta, 594f
Oviducts. See Uterine tubes
Ovulation, 726, 730732, 732f, 733f,
740741, 742t

834
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Oxygen utilization, 603

Oxyphil cells, 469
Oxytocin, 457t, 463, 464t, 775
Pacemaker, 592. See also Sinoatrial node
Pacinian corpuscles. See Lamellated corpuscles
Pagets disease, 166167
Pain receptors, 489t, 490, 491492
Palatal rugae. See Transverse palatine folds
Palate, 641
Palatine bone, 148f, 154f, 155, 156f
Palatine process, of maxilla, 148f, 150f,
154, 155f
Palatine tonsils, 305f, 605f, 608, 640f, 641,
641f, 642f
Palatine uvula, 640f, 641, 641f
Palatoglossal arch, 305f
Palatoglossus muscle, 252, 255f, 255t
Palatopharyngeal arch, 305f
Paleopathology, 34
Palmar arch, 575, 575f
Palmar interossei muscles, 275, 275t
Palmaris longus muscle, 269, 271f, 273t
tendon of, 274f, 321f, 323f, 325f
Palmar ligaments, 218, 219f
Palmar plates, 274f
Palmar region, 38
Palpation, 34, 297, 300
Palpebrae. See Eyelids
Palpebral conjunctiva, 500f, 501
Palpebral fissure, 302f, 302t, 500, 501f
Palpebral muscle, 250f
Pancreas, 456, 456f, 470f, 635, 635f, 638f,
654f, 663669
development of, 478
endocrine function of, 469471
Pancreatic acinar cells, 664f, 668
Pancreatic buds, 666
Pancreatic cancer, 669
Pancreatic duct, 652, 654f, 663f, 664f
Pancreatic islets, 457t, 469470, 470f, 478,
482, 663, 664f, 668
Pancreatic juice, 478, 663, 664f
Pancreatic juice enzyme, 654t
Pancreatic vein, 579
Pancreatoduodenal artery, 568f
Panhypopituitarism, 480
Papanicolaou (Pap) smear, 746
Papilla, 111, 642, 642f
Papillary constrictor muscle, 503t
Papillary dilator muscle, 503t
Papillary duct, 679f, 680f, 682, 683t
Papillary muscle, 549f
Papilloma, 129
Papule, 129
Parafollicular cells, 467
Parallel muscles, 238, 239t
Paralysis, 293, 393
Paralysis agitans. See Parkinsons disease
Paramesonephric duct, 716, 743
Paranasal ducts, 607
Paranasal sinuses, 144, 151f, 154, 605f, 607,
607f, 618t
Paranoia, 393
Paraplegia, 393
Parasympathetic division, of autonomic
nervous system, 436, 440443, 442f,
443t, 444t, 446447
Parathyroid gland, 456, 456f, 457t, 468469,
468f, 469f, 481482
Parathyroid hormone (PTH), 457f, 457t,
469, 469f
Paraventricular nuclei, 463
Parietal bone, 133f, 142f, 145f, 146, 147f,
148f, 149f, 174f
Parietal cells, of stomach, 650, 651f
Parietal lobe, 360f, 361f, 363366, 364f,
366f, 366t
Parietal pericardium, 43, 43f, 545546
Parietal peritoneum, 42f, 43, 317f, 636,
637f, 638f
Parietal pleura, 43, 43f, 615
Parietal portion, of serous membrane, 636
Parkinsons disease, 351, 358, 390, 396

Parotid duct, 305f, 645
Parotid gland, 635f, 645, 646f, 647t
Pars distalis, 457t, 460, 461f, 462f, 478
Pars intermedia, 460, 478
Pars tuberalis, 460, 461f, 478
Partial fractures, 189, 190f
Parturition, 775, 776f
Patella, 40, 133f, 174f, 185, 185f, 186f, 188t,
327f, 328f, 332f
Patellar ligament, 220, 221f, 278f, 279f,
283f, 327f, 332f
Patellar reflex. See Knee-jerk reflex
Patellar region, 40
Patellar surface, 184f, 185
Patellar tendon, 276f, 278f, 279f, 283f, 327f
Patellofemoral joint, 220
Patellofemoral stress syndrome, 207
Patent foramen ovale, 334, 593
Patent urachus, 336, 690
Pathogens, cytologic effects of, 72
Pathologic fractures, 189
Pathology, 78
PBI test. See Protein-bound iodine test
Pectineus muscle, 235f, 276f, 277, 278f,
279f, 281f, 281t
Pectoral girdle, 133f, 173, 174f
bones of, 132t, 134, 181t
muscles acting on, 266t
Pectoralis major muscle, 259f, 264265,
264f, 265, 266t, 268f, 270f, 308f,
311f, 312f, 313f, 319f, 320f, 324f
Pectoralis minor muscle, 264, 264f, 266t,
268f, 313f, 324f
Pectoralis muscle, 235f
Pectus carinatum. See Pigeon breast
Pectus excavatum. See Funnel chest
Pedicels, 681f, 682, 682f
Pedicle, 159, 160f, 161f, 162f
Pellagrous dermatitis, 107t
Pelvic brim, 180, 182f, 183f
Pelvic cavity, 40f, 4142, 41f, 44f
Pelvic curve, 158f, 159
Pelvic diaphragm, 260
Pelvic examination, 744746
Pelvic foramina, 162, 163f
Pelvic girdle, 133f, 174f, 178, 180183
bones of, 132t, 134, 181f, 182f, 188t
female, 183, 183f, 183t
male, 183, 183f, 183t
Pelvic inflammatory disease (PID), 637,
734, 748
Pelvic inlet, 180, 182f, 183f
Pelvic muscles, 276f
Pelvic outlet, 180
female, 261f
male, 261f
muscles of, 260, 260t, 261f
Pelvic region, 37
Pelvic surface, of sacrum, 181f
Pelvimetry, 183
Pelvis, 180
female, 685f
male, 685f
regions of, 38t
trauma to, 692
Pendular movements, 656, 659t
Penis, 698, 699f, 700t, 710711, 710f, 711f
trauma to, 711
Pennate muscles, 238, 239t, 246
Pepsin, 654t
Peptic ulcers, 337, 651, 670
Peptidase, 654t
Perceptive deafness, 533
Percussion, 34, 297
Percutaneous muscle biopsy, 102
Perforating canals, 140
Perforating cutaneous nerve, 424t
Perforating fibers, 137
Pericapsulitis, 217
Pericardial arteries, 566, 567t
Pericardial cavity, 41, 41f, 43, 43f, 44f, 546
Pericardial fluid, 546

Pericardial membranes, 43
Pericarditis, 335, 546, 594
Pericardium, 314f
Perichondrium, 95, 206
Periderm, 120
Perikaryon, 101
Perilymph, 519, 529
Perilymphatic space, 529
Perimetrium, 729f, 735
Perimysium, 236, 237f
Perineal muscles, 260
Perineal raphe, 700, 701f
Perineum, 37, 309, 700
female, 39f, 701f, 737, 737f
male, 39f, 701f
Perineurium, 352
Periodontal disease, 198, 645, 670
Periodontal membrane, 644, 645f
Periodontitis, 669f
Periosteal bone collar, 142
Periosteal bud, 142
Periosteal layer, of dura mater, 378, 378f
Periosteum, 97, 137, 251f
Peripheral chemoreceptors, 623
Peripheral ganglia, 439440
Peripheral nervous system (PNS), 344f, 345,
345t, 355f, 400433, 401f
cranial nerves, 403413
nerve plexuses, 415423
relationship with central nervous
system, 355f
spinal nerves, 413415
Peripheral vision, 513
Peristalsis, 635, 656, 659t
Peristaltic contraction, 648649, 649f
Peristaltic movements, 659
Peritoneal cavity, 42f, 43, 636, 637, 637f
Peritoneal lavage, 22
Peritoneal membranes, 43, 636, 638f
Peritoneum, 636
Peritonitis, 337, 637, 656, 671
Peritubular capillaries, 678, 680f
Permanent teeth, 642, 643f, 644t
Pernicious anemia, 541, 652
Peroneal artery, 570
Peroneal nerve. See Common fibular nerve
Peroneus brevis muscle, 283, 283f, 284f,
286t, 287f, 288f, 333f
Peroneus longus muscle, 235f, 283, 283f, 284f,
286t, 287f, 288f, 290f, 332f, 333f
tendon of, 329f
Peroneus tertius muscle, 283, 283f, 284f,
286t, 291f, 332f
tendon of, 332f
Peroxisomes, 53t, 6061
Perpendicular plate, of ethmoid bone, 147f,
148f, 150f, 153, 153f
Pes, 40. See also Foot
bones of, 185188, 187f
Pes planus, 188
Pestinate muscles, 547
Petit mal epilepsy, 393
Petrous part, of temporal bone, 149f, 151, 152f
PET scan. See Positron emission
tomography scan
Peyers patches. See Mesenteric patches
pH, 51, 85
Phagocytic cells, 59, 92
Phagocytosis, 55t, 61, 62f
Phalanges, 38, 133f, 174f, 178, 179f, 180f,
181t, 187f, 188, 188f, 188t
Phallus, 743
Phantom pain, 492
Pharyngeal pouches, 23, 23f
Pharyngeal tonsils, 605f, 607, 641f
Pharyngopalatine arch, 640f, 641
Pharynx, 24, 604f, 607608, 618t, 635, 635f,
636t, 640, 641f, 645648, 647f, 659t
Phasic receptors, 490
Phenobarbital, cytologic effects of, 57

Phenotype, 783
Phenylketonuria (PKU), 790
Pheochromocytomas, 483
Philtrum, 304f, 304t
Phimosis, 711
Phlebitis, 596
Phlegm, 9
Phobias, 393
Phosphate group, 65
Phospholipids, 52
Phosphorus, and bones, 135
Photoreceptors, 57, 489
Photorefractive keratectomy, 531
Phrenic nerves, 314f, 416, 416t, 621t
Phylogeny, 23, 690
Phylum Chordata, 23, 23f, 27t
Physical protection, of skin, 112
Physiological contracture, 286
Physiological dead space, 622t
Physiological jaundice of the newborn, 670
Physiology, definition of, 2
Pia mater, 378f, 379, 379f, 381f
PID. See Pelvic inflammatory disease
Pigeon breast, 312, 334
Pigmented layer, of retina, 505
Pigmented moles (nevi), 119, 129
Piles, 657
Pineal gland, 370f, 373, 456, 456f, 457t,
475476
Pinna, 301
Pinocytosis, 55t, 61, 62f
Piriformis muscle, 261f, 265f, 277, 281f, 330f
Piriformis nerve, 424t
Pisiform bone, 178, 179f, 180f, 325f
Pitch, of sound, 522, 523
Pituitary cachexia, 480
Pituitary dwarfism, 480, 481f
Pituitary gland, 360f, 362t, 370f, 372f, 373,
456, 456f, 457t, 460464, 461f
blood supply to, 563, 563f
Pituitary hormones, 462, 464t, 465f
Pituitary stalk, 370f, 460, 461f
Pivot joint, 202, 204f, 205t
PKU. See Phenylketonuria
Placenta, 456, 478479, 580, 764f
circulation of blood within, 766f
endocrine functions of, 765766, 766t
exchange of molecules across, 765
in taxonomy, 24
Placental stage, of labor, 775, 776f
Placenta previa, 785
Placode, 529
Plagiocephaly, 333
Planes of reference, 33, 33f
Plane suture, 198
Plantar aponeurosis muscle, 269, 290f
Plantar calcaneonavicular ligament, 223f
Plantar flexion, 207, 208f, 211f, 212f
Plantar interosseous muscle, 290f
Plantaris muscle, 284, 284f, 286t, 287f,
288f, 289f
Plantar ligaments, 223f, 290f
Plantar reflex, 430t, 431f
Plantar surface, 40
Plasma membrane. See Cell membrane
Plasma proteins, 545t
Platelets, 98, 99f, 540, 541f, 542, 542f, 544t
Platysma, 250f, 251f, 251t, 257f, 264f, 324f
Pleurae, 43, 615, 617f, 618t
Pleural cavity, 41, 41f, 43, 43f, 44f, 615
Pleural membranes, 617f
Pleurisy, 615, 628
Plicae circulares, 653, 653f
PMN leukocytes. See Polymorphonuclear
leukocytes
Pneuma, 603
Pneumonia, 628
Pneumotaxic area, of brain, 374, 374f, 623
Pneumothorax, 335, 622t, 627, 627f
Podocytes, 681f, 682

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Poisons
gastrointestinal effects, 669
Polar body, 730
Polarization, 357
Poliomyelitis, 288, 339, 394
Pollex, 178
Polycythemia, 542, 593
Polydactyly, 189, 189f, 337, 338
Polymorphonuclear (PMN) leukocytes, 542
Polypeptide, formation of, 69
Polyribosomes, 68
Polysaccharides, 52
Polysomes, 68
Polysynaptic reflex arc, 428, 429f
Polythelia, 25f
Polyuria, 692
Pons, 360f, 361f, 362t, 370f, 374, 374f, 375f
Popliteal artery, 560f, 568, 570f
Popliteal bursae, 221f, 222f
Popliteal fossa, 40, 221, 284, 289f, 326f,
327, 328f
Popliteal ligaments, 221f
Popliteal nodes, 583
Popliteal vein, 573f, 576, 578f
Popliteus muscle, 284, 286t, 288f
Portal system, 464, 563, 579
Positron emission tomography (PET) scan,
18, 19f, 359, 362f
Postcentral gyrus, 363, 364f, 493
Posterior, 36t
Posterior arch, of atlas, 160f
Posterior auricular artery, 562f, 563
Posterior cardiac vein, 549f, 551
Posterior cavity, 40f, 41, 44f
of eye, 505f, 508
Posterior cerebellar artery, 563f
Posterior cerebral artery, 561, 563f
Posterior cervical triangle, 307, 307f, 308f, 308t
Posterior chamber, of eye, 500f, 505f, 508, 512f
Posterior commissure, 370f, 373
Posterior communicating artery, 561, 563f
Posterior cranial fossa, 149f
Posterior cruciate ligament, 221, 221f, 222f
Posterior crural muscles, 283285, 287f, 288f
Posterior division
of brachial plexus, 416
of lumbar plexus, 421
Posterior ethmoidal foramina, 150f
Posterior fontanel, 144, 145f
Posterior funiculus, 386, 386f
Posterior gluteal line, 182f, 183
Posterior horns, 384, 386f, 390
Posterior humeral circumflex artery, 565, 565f
Posterior inferior iliac spine, 182f, 183
Posterior intercostal arteries, 566, 567f, 567t
Posterior interventricular artery, 551, 552f
Posterior lobe, of cerebellum, 375f
Posterior median sulcus, 384, 386f
Posterior membranous ampullae, 519f
Posterior pituitary gland. See Lobus nervosa
Posterior ramus, of spinal nerves, 414, 414f
Posterior root, of spinal nerves, 386f, 414, 414f
Posterior sacral foramen, 162, 163f
Posterior semicircular canal, 519f
Posterior spinocerebellar tract, 388t, 389t
Posterior sternoclavicular ligament, 215, 215f
Posterior superior iliac spine, 182f, 183
Posterior surface, of stomach, 650
Posterior talocalcaneal ligament, 223f
Posterior talofibular ligament, 222, 223f
Posterior tibial artery, 560f, 570, 570f
Posterior tibial vein, 573f, 576, 578f
Posterior tibiofibular ligament, 223f
Posterior tibiotalar ligament, 223f
Posterior trunk, of internal iliac artery, 569f
Posterior tubercle, 161
Posterior vagal trunk, 441
Posterolateral fontanels, 144, 145f
Postganglionic neuron, 435436
Postmature infant, 777
Postnatal growth, periods of, 775781, 781t
Postsynaptic neuron, 358

Posture, muscular system and, 234
Potts fracture, 185, 190
Pouch of Douglas. See Rectouterine pouch
Precapillary sphincter muscles, 555557
Precentral gyrus, 363, 364f
Precursor messenger RNA (pre-mRNA), 67
Preembryonic period, 757762, 760t
Preganglionic neuron, 435
Pregnancy
ectopic, 734, 746747, 785, 786f
erector spinae muscle in, 262
multiple, 785786, 787f, 788f
termination of, 748
trimesters of, 785
uterus in, 735f
vulva in, 738
Pregnancy tests, 759
Prehensile hands, 24, 26f
Premature infant, 777
Premature synostosis, 333
Premolars, 148f, 154, 304f, 640f, 642,
643f, 644t
pre-mRNA. See Precursor messenger RNA
Prepuce, 699f, 710f, 711, 737
Presbyopia, 531
Prescientific period, anatomy in, 24
Pressure cuff, 558559, 558f
Pressure points, 321f, 323f, 329f, 570, 572f
Pressure receptors, 490491
Presynaptic neuron, 358
Primary amenorrhea, 747
Primary curves, 159
Primary fissure, 375f
Primary follicles, 729, 730f, 733f
Primary germ layers, 80, 760
Primary growth, 73
Primary neoplasms, 394
Primary oocytes, 729, 731f, 733f
Primary ossification center, 142
Primary sex cords, 716, 743
Primary sex organs, 698, 726
Primary spermatocytes, 703, 704, 706f
Primates, 24
Primitive gut, 665
Primitive line/streak, 767, 768f
Primitive node, 767, 768f
Primordial follicles, 729, 730f, 743
Principal artery, of thumb, 565f
Principal bronchus, 604f
Principal cells
of parathyroid gland, 469
of stomach, 650, 651f
P-R interval, 553
Process, 136t
Proctodeum, 667
Proerythroblasts, 544
Profunda vein, 640f
Progesterone, 457t, 472, 474475, 741, 766t
Projection fibers, 368, 369f
Prolactin, 463, 464t
Prolactin-inhibiting hormone (PIH), 463
Prolapse, of uterus, 748
Proliferation zone, 142
Proliferative phase, of ovulation, 741, 742f
Promontory, 158f
Pronation, 209, 268
Pronator quadratus muscle, 268, 272f, 273t,
274f, 325f
Pronator teres muscle, 268, 271f, 272f, 273t
Pronephric duct, 690
Pronephros, 689, 690
Prophase, 70, 71f, 703, 704t, 705f
Proprioceptors, 375, 490, 492493, 494f
Prosencephalon, 347, 362t
Prostaglandins, 775
Prostate, 686f, 687, 698, 699f, 700t,
708709, 709f, 710f
disorders of, 722
palpation of, 709
Prostatectomy, 722
Prostatic carcinoma, 722
Prostatic part, of urethra, 686f, 687, 709
Prostatitis, 722
Prostheses, joint, 228f, 229
Prostroatory vestibular nystagmus, 527

Protection, in circulatory system, 539
Protein-bound iodine (PBI) test, 480
Protein hormones, 457f, 458, 458t, 459, 459f
Proteins, 52t
in adults, 51
functional, 52
plasma, 545t
structural, 5152
synthesis of, 68, 68f
Proteolytic enzymes, 757
Protoplasm, 57
Protraction, 209, 210f
Proximal, 36t
Proximal convoluted tubule, 679f, 680f,
681f, 682, 683t
Proximal interphalangeal joints, 322f
Proximal phalanx, 179f, 180f, 187f
Proximal radioulnar joint, 217, 225t
Proximal tibiofibular joint, 222f, 225t
Pruritus, 129
Pseudohermaphroditism, 714
Pseudohypertrophic muscular dystrophy, 790
Pseudopolar neurons, 352
Pseudostratified ciliated columnar
epithelium, 83, 84f, 86t
Pseudounipolar neurons, 356f
Psoas major muscle, 276, 276f, 278t, 279f, 281f
Psoas minor muscle, 276f, 279f, 281f
Psoriasis, 129
Psychomotor epilepsy, 393
Psychosis, 393
Pterygoid canal, 153f
Pterygoid hamulus, 153f
Pterygoid plexus, 574f
Pterygoid processes, 148f, 152, 153f
Pterygoid tuberosity, 156f
Pterygopalatine ganglion, 441
PTH. See Parathyroid hormone
Puberty, 29, 726, 778779
Pubescence, 778
Pubic arch, 183f
Pubic region, 37, 38f, 38t
Pubic tubercle, 181f, 182f, 183
Pubis, 133f, 174f, 181f, 182f, 183
Pubococcygeus muscle, 261f
Pubofemoral ligament, 220, 220f
Pudendal cleft, 727t, 737
Pudendal nerve, 424t
Pudendal nerve block, 775
Pulmonary alveoli, 612614, 613f, 618t, 624
Pulmonary arteriole, 613f
Pulmonary artery, 581f, 615f
Pulmonary capillary, 613f
Pulmonary circulation, 550
Pulmonary disorders, 621
Pulmonary embolism, 595
Pulmonary ligament, 615, 615f
Pulmonary plexus, 441
Pulmonary respiration, 256
Pulmonary semilunar valve. See Pulmonary
valve
Pulmonary stenosis, 593, 594f
Pulmonary trunk, 548f, 549f, 550, 552f, 561f
Pulmonary valve, 548t, 549f, 550
Pulmonary vein, 550, 581f, 615f
Pulmonary ventilation. See Breathing
Pulmonary venule, 613f
Pulmonic area, 554, 554f
Pulp, 645
Pulp cavity, 645
Pulse pressure, 559
Puncture wound, 125f, 126f
Punnett square, 784, 784f
Pupil, of eye, 302f, 302t, 501f, 505, 505f,
509f, 510511
Pupillary reflex, 514
Purkinje fibers. See Conduction myofibers
Pustule, 129
Putamen, 365f, 369, 370f
P wave, 553
Pyelitis, 687
Pyelonephritis, 687
Pyloric sphincter, 649, 650f
Pyloric stenosis, 336, 669
835
Pylorus, 649650, 649f, 650f

Pyogenic granuloma, 669f
Pyorrhea, 672
Pyramid, 518f
Pyramidalis muscle, 317f
Pyramidal process, 156f
Pyramidals, 264f
Pyramidal tracts. See Corticospinal tracts
Pyramids, of medulla oblongata, 376
Pyuria, 692
QRS complex, 553554
Quadrate lobe, of liver, 661, 661f
Quadratus femoris muscle, 277, 278f, 330f
Quadratus femoris nerve, 424t
Quadratus lumborum muscle, 262f, 263,
263t, 276f
Quadratus plantae, 290f
Quadriceps femoris muscle, 280, 282t, 327f
Quadriplegia, 393
Quickening, 206, 248, 773
Racial groups, body proportion differences
among, 780781, 782f
Radial artery, 560f, 565, 565f
pressure point for, 321f, 323f, 572f
Radial collateral ligament, 217, 219f
Radial fossa, 176f
Radial keratotomy, 531
Radial nerve, 324f, 401f, 416, 418f, 418t
Radial notch, 177, 177f
Radial recurrent artery, 565
Radial vein, 321f, 573f, 575, 575f
Radiate sternocostal ligament, 215f
Radiation, cytologic effects of, 72
Radical mastectomy, 264, 748
Radiocarpal joint, 225t
Radiograms, 587
Radiographic anatomy, 1819, 18f, 19f
Radioimmunoassay (RIA), 480
Radius, 133f, 174f, 177f, 178, 181t, 198f, 325f
Rami communicantes, 414, 439, 439f
Ramus
definition of, 136t
of ischium, 183
of mandible, 149f, 156f, 157
Range of motion, of synovial joints, 200
RAS. See Reticular activating system
Rathkes pouch. See Hypophyseal pouch
Reactive hypoglycemia, 482
Receptive relaxation, 659t
Receptors
cutaneous, 488, 489t
in reflex arc, 427, 427f
Recessive alleles, 783, 783t
Reciprocal inhibition, 430
Rectal arteries, 568f, 569f
Rectouterine folds, 735
Rectouterine pouch, 727f, 735
Rectum, 635f, 638f, 657, 657f, 658f, 685f, 727f
Rectus abdominis muscle, 235f, 258, 258f,
259f, 260t, 264f, 311f, 312f, 313,
313f, 317f, 620f
Rectus capitis posterior major muscle, 256f, 262f
Rectus capitis posterior minor muscle, 256f, 262f
Rectus femoris muscle, 276f, 278f, 279f, 280,
281f, 282f, 282t, 327f, 328f, 331f
tendon of, 331f
Rectus muscles, 252, 502, 503t
Rectus sheath, 259f, 264f
Recurrent laryngeal nerve, 411
Red blood cells. See Erythrocytes
Red bone marrow, 134, 137
Red-green color blindness, 513, 784785
Red nucleus, 374
Red pulp, 585, 586f
Referred pain, 491492, 492f, 493f
Reflex arc, 427, 427f
Reflexes
kinds of, 427430, 430t, 431f
neonatal, 776
spinal cord and, 384
testing of, 35, 430
Refraction, of light, 509510, 512f
Regeneration, of neurons, 354f

836
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Regeneration tube, 351

Regional anatomy
Regional approach, to anatomy, 29
Regulatory system, and circulatory
system, 539
Regurgitation. See Vomiting
Relaxin, 775
Releasing factors, 459460
Rembrandt, 15f
Renaissance, and history of anatomy, 1315
Renal adipose capsule, 677
Renal agenesis, 691
Renal arteries, 566, 567t, 581f, 676, 677,
678f, 679f, 680f
Renal biopsy, 692
Renal blood vessels, 677679, 680f
Renal calyx, 678f, 679f
Renal capsule, 677, 678f, 679f
Renal columns, 677, 678f, 679f
Renal corpuscle, 680, 681f, 683t
Renal cortex, 677, 678f, 679f, 680f
Renal ectopia, 691
Renal failure, 693
Renal fascia, 677
Renal medulla, 677, 678f, 679f, 680f
Renal papilla, 677, 679f, 682
Renal pelvis, 677, 678f, 679f, 680f, 682
Renal pelvises, 677f
Renal ptosis, 677
Renal pyramids, 677, 678f, 679f
Renal stone, 684, 684f
Renal tubules, 679f
Renal vein, 573f, 579, 581f, 676, 677, 678,
678f, 679f, 680f
Repolarization, 357
Reproductive structures, categories of, 698
Reproductive system
female (See Female reproductive system)
male (See Male reproductive system)
Reserve zone, 142
Residual volume (RV), 621t, 622f
Resorption zone, 143
Respiration. See also Respiratory system
definition of, 603
Respiratory bronchioles, 611, 613f
Respiratory capacities, 620, 621t, 622f
Respiratory centers, of brain, 374, 374f, 377, 623
Respiratory distress syndrome, 620
Respiratory division, of respiratory system,
603, 612, 613f
Respiratory epithelium, 83
Respiratory system, 32f, 602633, 604f
basic structure of, 603
conducting passages, 604612, 612f
control of, 628630
disorders of, 628
epithelial tissue of, 606f
functions of, 603
injury/trauma to, 627628
lower, 603, 624, 626f
physical requirements of, 603
respiratory division of, 613f
upper, 603, 605f, 624, 625f
Respiratory volumes, 620, 621t, 622f
Resting potential, 357
Resting tremor, 390
Rete testis, 701f, 702f, 703
Reticular activating system (RAS), 377, 377f
Reticular connective tissue, 92, 94f, 95t
Reticular fibers, 91
Reticular formation, 377
Reticulospinal tracts, 389
Retina, 505507, 505f, 508t, 511f
damage to, 532
development of, 516
layers of, 510f

photoreceptor cells of, 510f
Retinaculum, 236
Retinal detachment, 532
Retinitis pigmentosa, 790
Retraction, 209, 210f
Retroflexion, 735
Retroflexion, of uterus, 748
Retromandibular vein, 305f
Retroperitoneal organs, 636
Retroversion, of uterus, 748
Rhabdomyosarcoma, 288
Rheumatic fever, 336
Rheumatism, 287
Rheumatoid arthritis, 60, 226, 227f
Rheumatology, 230
Rhinencephalon, 449
Rhombencephalon, 347, 362t
Rhomboideus major muscle, 256f, 265f,
266t, 267f, 315f, 316f, 323f
Rhomboideus minor muscle, 256f, 265f,
266t, 267f, 315f
Rhomboideus muscle, 235f, 264
Rhythmicity area, of brain, 374f, 623
Rhythmic segmentation, 656, 659t
Rhythm method, of contraception, 749
RIA. See Radioimmunoassay
Rib(s), 133f, 165, 165f, 166f, 174f, 311f
Rib cage, 133f, 164165, 164f, 174f
abnormalities of, 312
bones of, 132t, 134
Ribonucleic acid (RNA), 52t, 67f
structure of, 6668
synthesis of, 6668, 67f
Ribonucleotide, 67
Ribose, 67f
Ribosomal RNA (rRNA), 67
Ribosomes, 53t, 54f, 57, 58f, 69f
Rickets, 97, 114, 114f, 135, 166
Right atrioventricular valve, 314f, 548t,
549f, 550
Right atrium, 548f, 549f, 550, 552f
Right auricle, 561f
Right axillary vein, 574f
Right brachiocephalic vein, 314f, 561f, 573f,
574f, 577f
Right bundle branch, 553f
Right colic flexure. See Hepatic flexure
Right common carotid artery, 561, 561f,
562f, 567t
Right common iliac artery, 566, 570f
Right common iliac vein, 578f
Right coronary artery, 548f, 550, 552f, 559, 567t
Right costal margin, of rib cage, 312
Right external iliac artery, 570f
Right external jugular vein, 574f
Right gastric artery, 568f
Right gastric vein, 579, 580f
Right gastroepiploic vein, 579, 580f
Right hemisphere, of brain, 363
Right hepatic vein, 579
Right hypochondriac region, 38f, 38t
Right inguinal region, 38f, 38t
Right internal jugular vein, 561f
Right lateral region, 38f, 38t
Right lobe, of liver, 314f, 317f, 660, 661f
Right lung, 314f, 604f, 614, 616f
Right lymphatic duct, 582, 584f
Right marginal artery, 551
Right marginal vein, 552f
Right ovarian vein, 579
Right principal bronchi, 610, 612f
Right principal bronchus, 610f
Right pulmonary artery, 548f, 549f, 550, 561f
Right pulmonary veins, 548f, 549f, 561f
Right renal artery, 568f
Right subclavian artery, 560f, 561, 561f,
562f, 564, 567t
Right subclavian vein, 314f, 324f, 572, 573f,
574f, 584f
Right suprarenal vein, 579
Right testicular vein, 573f, 579
Right ventricle, 548f, 549f, 550, 552f
Right ventricular hypertrophy, 594f
Right vertebral artery, 560f
Rigor mortis, 102, 286

Risorius muscle, 250f, 251f, 251t, 305f
Rivlins ducts, 645
RNA. See Ribonucleic acid
RNA polymerase, 67
Rods, of eye, 506, 510f
Roentgen, Wilhelm Konrad, 18
Roentgenograph, 18
Roman Era
and history of anatomy, 1112
surgical instruments in, 12f
Root
of nose, 303, 304f, 304t
of penis, 710711, 710f
of tongue, 642f
of tooth, 644, 645f
Root canal, 645, 645f
Root hair plexus, 489t, 491
Rooting reflex, 776
Rotation, 209, 209f, 211f, 212f
Rotator cuff, 217, 267
Rough endoplasmic reticulum, 54f, 57, 58f
Round ligaments. See Ligamentum teres
Round window. See Cochlear window
rRNA. See Ribosomal RNA
Rubella virus, 527530
Rubrospinal tract, 388t, 389, 389t
Rugae, of urinary bladder, 685
Rule of nines, 123, 124f
Runners knee, 207
RV. See Residual volume
Saccadic eye movements, 514
Saccular portion, of otocyst, 529
Sacculations. See Haustrum
Saccule, 515, 519f, 520, 524
Sacral arteries, 569f
Sacral canal, 162, 163f
Sacral cornua, 163f
Sacral curve, 159
Sacral nerves, 401f, 413
Sacral plexus, 385f, 401f, 421423, 423f, 424t
Sacral promontory, 162, 163f, 182f
Sacral region, 37
Sacral tuberosity, 162, 163f
Sacral vertebrae, 159
Sacroiliac articulation, 181f
Sacroiliac joint, 162, 182f, 225t
Sacrospinous ligament, 281f
Sacrotuberous ligament, 330f
Sacrum, 133f, 158f, 159, 162, 162f, 163f,
163t, 174f, 181f, 182f, 330f
Sacs, of eye, 57
Saddle joint, 202203, 205f, 205t
Sagittal plane, 33, 33f, 34f
Sagittal suture, 144, 145f, 146151
Saliva, 645
Salivary ducts, 645
Salivary enzyme, 654t
Salivary glands, 635, 645, 646f, 647t
clinical problems of, 669670
Salmonella, 669
Salpingitis, 734, 746, 748
Salpingography, 734
Salpingolysis, 734
Salpinx, 734
Salts, 51
Salty taste, 497, 498f
Sanguine, 9
SA node. See Sinoatrial node
Saphenous nerve, 420t
Sarcolemma, 237f, 241, 241f, 242f, 243f, 247f
Sarcomeres, 241, 243f, 244f
Sarcoplasm, 237f, 241, 241f, 243f
Sarcoplasmic reticulum, 241, 242f
Sartorius muscle, 235f, 276f, 278f, 279, 279f,
280f, 281f, 282f, 282t, 285f, 287f,
289f, 327f, 328f, 331f
Satiety center, 373
Scab, 126f
Scala tympani, 520, 520f, 521f, 529
Scala vestibuli, 520, 520f, 521f, 529
Scalene muscles, 258, 258f, 619, 620f
Scalenus anterior muscle, 257f, 308f

Scalenus medius muscle, 256f, 257f, 259f, 308f
Scalenus posterior muscle, 256f
Scalp, 301, 306f
Scaphocephaly, 333
Scaphoid bone, 178, 179f, 180f. See also
Navicular bone
Scapula, 133f, 173, 174f, 175f, 181t,
309310
Scapular notch, 173, 175f
Scar tissue, 98, 126, 127f
Schizophrenia, 393, 396
Schleiden, Matthias, 16, 49
Schwann, Theodor, 16, 49
Schwann cells. See Neurolemmocytes
Sciatica, 423
Sciatic nerve, 331f, 333f, 401f, 421, 423, 424t
Scientific period, anatomy in, 419
Sclera, 302f, 302t, 501f, 504, 505f, 506f,
508t, 516
Scleral venous sinus, 512f
Sclerotomes, 248
Scoliosis, 226, 226f
Scrotal dermatitis, 107t
Scrotal septum, 700, 701f
Scrotum, 698, 699f, 700, 700t, 701f, 710f, 716
disorders of, 722
trauma to, 711
Sebaceous glands, 107f, 118, 121, 301
Seborrhea, 129
Seborrheic hyperkeratoses, 110
Sebum, 118
Secondary amenorrhea, 747
Secondary curves, 159
Secondary follicles, 729, 730f, 733f
Secondary growth, 73
Secondary neoplasms, 394
Secondary oocytes, 730, 730f, 731f, 732f, 733f
Secondary ossification center, 142
Secondary sex characteristics, 698, 703t, 726
Secondary sex organs, 698, 726, 732738
Secondary spermatocytes, 703, 704, 706f
Second-class lever, 210211, 213f
Second-degree burns, 122, 123f
Second-degree frostbite, 123
Second-division nerve block, 157, 406
Second meiotic division, 704, 704t, 705f, 706f
Second sound, of heartbeat, 554
Second trimester, of pregnancy, 785
Secretin, 457t, 476t
Secretion granule, 54f
Secretory phase, of ovulation, 741, 742f
Sectioning, 102
Segmental bronchus, 604f, 610, 610f
Sella, etymology of term, 30
Sella turcica, 148f, 149f, 152, 153f, 154f
Semen, 708, 712, 713714, 714, 714t
Semicircular canals, 519f, 520, 526, 529
Semicircular ducts, 517f, 519f, 520, 524
Semilunar valves, 547
Semimembranosus bursae, 221f, 222f
Semimembranosus muscle, 235f, 277, 280f,
281, 281f, 282f, 282t, 287f, 289f,
328f, 331f
tendon of, 285f
Seminal fluid. See Semen
Seminal vesicles, 698, 699f, 700t, 708, 708f,
710f, 716
Seminiferous tubules, 700t, 702, 702f, 706f, 716
Semispinalis capitis muscle, 255, 256f, 257f,
262f, 263t, 265f, 308f, 309f
Semispinalis cervicis muscle, 262f, 263t
Semispinalis thoracic muscle, 262f, 263t
Semitendinosus muscle, 235f, 277, 280f, 281,
281f, 282f, 282t, 287f, 289f, 328f,
330f, 331f, 333f
tendon of, 285f
Senescence, 781t
atrophy, 102
of nervous system, 395
Senile atrophy, 72
Sensation, 488
Senses. See also specific sense

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Sensorineural deafness, 533
Sensory adaptation, 490
Sensory area, of cerebral cortex, 367f
Sensory hair cells, 493
Sensory homunculus, 494
Sensory nerves, 352, 355f, 356f, 401, 403
Sensory neurons, 239, 345t, 349f, 352, 427, 427f
Sensory organs, 487536. See also specific
organs
Sensory perception, 488
Sensory receptors, 111, 114
Sensory root, of trigeminal nerve, 405
Septa, of testis, 701f
Septal cartilage, 605
Septal defect, 593, 593f, 594f
Septate uterus, 746f
Septicemia, 596
Septum pellucidum, 370f
Septum penis, 710f, 711
Septum secundum, 589
Serosa, 639, 639f
Serotonin, and blood clotting, 542
Serous cells, 645
Serous fluid, 43
Serous layer, of uterine tube, 734
Serous membranes, 42, 42f, 43, 43f, 85,
636637, 637f
Serous pericardium, 546
Serrate suture, 197
Serratus anterior muscle, 235f, 259f, 264,
264f, 266t, 311f, 312f, 313f
Serratus posterior inferior muscle, 265f
Serratus posterior muscle, 309f
Serratus posterior superior muscle, 256f
Sertoli cells. See Sustentacular cells
Sesamoid bones, 132, 180f
Sex cells. See Gametes
Sex chromosomes, 782
Sex determination, 716
Sex differentiation, 703t, 717f
Sex hormones, 458, 472
female, 741
male, 698
Sex-linked inheritance, 784785
Sexual center, 373
Sexual dysfunction, male, 715720
Sexual intercourse, 698, 726
Sexually dimorphic characteristics, 779
Sexually transmitted diseases (STDs),
720721, 721t
Sexual reproduction, 698
Sexual response, hypothalamus and, 373
Shaft, of penis. See Body, of penis
Shakespeare, William, 2
Sharpeys fibers. See Perforating fibers
Shin, 40
Shingles, 414
Shinsplints, 293, 339
Shivering, 113
Shock
recognition of, 597
treatment of, 597598, 597f
Shoes, improperly fitted, 202
Short bones, 136, 136f
Short head, of biceps brachii muscle, 267
Shoulder, 38. See also Pectoral girdle
arteries of, 564566
diseases of, 338
internal anatomy of, 313f, 322, 323f, 324f
muscles of, 259f, 264f, 265f, 268f
surface anatomy of, 319, 319f, 320f
trauma to, 338
Shoulder joint. See Glenohumeral joint
Siamese twins. See Conjoined twins
Sickle-cell anemia, 593
SIDS. See Sudden infant death syndrome
Sighing, 622t
Sight. See Eye(s); Visual sense
Sigmoid colon, 317f, 318f, 635f, 638f, 656, 657f
Sigmoid sinus, 574f
Simmonds disease. See Pituitary cachexia
Simple ciliated columnar epithelium, 83,

83f, 86t
Simple columnar epithelium, 8183, 82f, 86t
Simple cuboidal epithelium, 81, 82f, 86t
Simple diffusion, 55t
Simple epithelia, 8183
Simple epithelium, 86t
Simple fractures, 189
Simple glands, 87, 88f, 89t
Simple mastectomy, 748
Simple squamous epithelium, 81, 81f, 86t
Single-unit smooth muscle, 437, 437t
Sinoatrial node (SA node), 551552, 553f
Sinus. See also Paranasal sinuses
of bones, 136t
Sinus bradycardia, 592f
Sinusitis, 628
Sinusoids, 557, 579, 580f
Sinus rhythm, 592f
Sinus tachycardia, 592f
Sinus venosus, 589, 590t
Skeletal developmental disorders, 166
Skeletal muscle, 99, 100f, 100t
architecture of, 238, 239t
associated connective tissue, 236, 237f
attachments of, 235236, 236f
contractions of, 243, 245f, 246f
groups of, 236238
nerve supply to, 239, 240f
Skeletal muscle cells, shape of, 50f
Skeletal muscle fibers, 240243, 241f, 242f, 243f
Skeletal muscle pump, 558
Skeletal system, 31f, 131171. See also
Appendicular skeleton; Axial
skeleton
aging of, 167168, 167f
Skeleton, 174f. See also Appendicular
skeleton; Axial skeleton
bones of, 132t, 133f
and fat storage, 134
and hemopoiesis, 134, 135f
and mineral storage, 134135
movement and, 134
protection of, 134
support of, 134
Skin, 78f, 107f
absorption by, 114
aging of, 127, 127f
coloration of, 109110, 300
communication and, 114
hydroregulation of, 112
innervation of, 111
layers of, 106112
of neonate, 299t
nutritional deficiencies and, 107t
as organ, 106
pH of, 85
physical protection of, 112
sensory reception in, 114
surface patterns of, 110, 110f
synthesis in, 114
thermoregulation and, 112113, 113f
vascular supply of, 111
Skin cancer, 110, 112, 119121
Skin creases, 300, 300f
Skin disorders
infectious, 119
inflammatory, 119, 122f
Skin glands, 117119, 118f
Skin grafts, 124, 125f
Skull, 132, 133f, 144158, 174f
anterior view of, 147f
bones of, 132t
fetal, 145f
foramina of, 146t
fracture of, 198
frontal section of, 150f

growth of, 777, 778f
inferior view of, 148f
inferolateral view of, 149f
lateral view of, 147f
of neonate, 299t
sagittal view of, 148f
sutures of, 198f
Skull joints, 225t
Sleep apnea, 628
Sleep center, 373
Sleep response, 377
Slipped disc, 24
Slit pores, 681f, 682
Small cardiac vein, 552f
Small intestine, 318f, 457t, 476t, 478, 635,
635f, 636t, 639f, 652656, 653f, 659t
mechanical activities of, 655656
regions of, 652653
structural modifications of, 653655
Small saphenous vein, 578f, 579
Smegma, 711
Smooth chorion, 764
Smooth endoplasmic reticulum, 54f, 57, 58f
Smooth muscle, 99, 100f, 234
autonomic nervous system and, 435,
437438
multiunit, 437, 437t
of respiratory system, 613f
single-unit, 437, 437t
Smooth muscle cell, shape of, 50f
Smooth pursuit movements, 514
Sneezing, 83, 496, 622t
Snellens chart, 527
Soft corns, 327
Soft palate, 305f, 604f, 605f, 640f, 641, 641f
Soft spots. See Fontanels
Solar plexus, 440
Sole, of foot, 40
Sole creases, 300, 300f
Soleus muscle, 235f, 283f, 284, 284f, 285f,
286t, 287f, 288f, 329f, 332f, 333f
Solutes, 51
Solutions, 51
Solvents, 51
Somatic death, 102
Somatic motor nerve, 345t
Somatic motor neurons, 354, 356f
Somatic nervous system, and autonomic
nervous system, 435, 435t, 436f
Somatic pain, 491
Somatic reflexes, 428430
Somatic senses, 488489, 490494, 495f
Somatic sensory neurons, 354, 356f
Somatomammotropin, 479
Somatotropin, 464t. See also Growth
hormone
Soul, 7
Sound waves, 522523, 522f
Sour taste, 497, 498f
Spallanzani, Lazzaro, 15
Spastic paralysis, 393
Specialized cells, 70
Special movements, 209, 210f
Special senses, 488
Speculum, 744
Spermatic cord, 264f, 317f, 700, 701f, 702f,
707708, 710f, 716
Spermatic ducts, 707708
Spermatids, 703, 704, 706f
Spermatocytes, 703, 704, 706f
Spermatogenesis, 702, 703704, 703t, 706f
Spermatogonia, 702703, 706f
Spermatozoa, 63f, 698, 702f, 706f, 707f,
756, 756f
movement of, 83
shape of, 50f
Spermiogenesis, 704
S phase, 6970, 70f
Sphenofrontal suture, 148f
Sphenoidal process, 156f
Sphenoidal sinus, 45f, 148f, 151f, 154f, 306f,
605f, 607, 607f
Sphenoidal sinuses, 152
837
Sphenoid bone, 145f, 147f, 148f, 149f, 152,

153f, 154f
Sphenoid fontanels. See Anterolateral
fontanels
Sphenomandibular ligament, 214f, 215
Sphenopalatine notch, 156f
Sphenoparietal suture, 148f
Sphincteral muscles, 238, 239t
Sphincter of ampulla, 652, 654f, 662
Sphincter of Oddi, 652
Sphygmomanometer, 558559, 558f, 587
Spina bifida, 166, 166f, 392
Spina bifida cystica, 392
Spina bifida occulta, 392
Spinal arteries, 563f
Spinal cavity, 44f
Spinal column, 159
Spinal cord, 23, 159, 360f, 381f, 384390,
385f, 386f
structure of, 384
tracts of, 384389
Spinal curvature, 2527
Spinal dura mater, 378
Spinal ganglion, 386f, 414, 414f
Spinalis muscle, 262
Spinalis thoracic muscle, 262f, 263t, 265f
Spinal meningitis, 394
Spinal nerves, 24, 379f, 381f, 384, 385f,
386f, 401, 401f, 402f, 413415
Spinal plexus, 385f
Spinal root, of accessory nerves, 412, 412f
Spindle fibers, 65
Spine
of bones, 136t
of ischium, 181f, 182f, 183
of scapula, 173, 175f, 315f, 324f
Spinous process, 160, 160f, 161, 161f, 162f, 310
Spiral fractures, 189, 190f
Spiral organ, 410, 520f, 521, 521f, 529, 529f
Spiritus, 603
Spirometer, 620, 621f
Splanchnic nerves, 439
Splanchnocranium, 141
Spleen, 445t, 584f, 585, 586f, 587t
Splenectomy, 337, 585
Splenic artery, 566, 567t, 568f, 586f
Splenic flexure, 317f, 656, 657f
Splenic vein, 579, 580f, 586f
Splenium, of corpus callosum, 361f, 370f
Splenius capitis muscle, 254, 256f, 257f,
259f, 265f, 308f, 315f
Splenius cervicis muscle, 256f, 265f
Split brain, 514
Spondylitis, 230
Spongy bone, 97, 136, 138
Spongy part, of urethra, 686f, 687, 709
Spontaneous abortion, 748
Spoon nails, 107t, 117
Sprain, 92, 203
Sprains, 224, 225, 339
Sprengels deformity, 337
Squamosal-dentary jaw articulation, in
taxonomy, 24
Squamous cell carcinoma, 121, 123
Squamous epithelial cell, shape of, 50f
Squamous part, of temporal bone, 146, 152f
Squamous suture, 144, 145f, 146, 147f, 148f,
149f, 197198
Staining, 102
Stapedius muscle, 518f, 519
tendon of, 518f
Stapes, 30, 157, 158f, 517, 518f
Statoacoustic nerve. See Vestibulocochlear
nerve
Statoconia, 525, 525f
Statoconial membrane, 525
STDs. See Sexually transmitted diseases
Stelluti, Francisco, 15
Stenosis, 335, 554
Stensen, Niels, 15
Stensens duct. See Parotid duct
Stereocilia, 524f
Stereoscopic vision, 27
Sterility, male, 720

838
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Sterilization, 744, 749

Sternal angle, 164f, 165, 310
Sternal extremity, 173, 175f
Sternal joint, 225t
Sternal region, 36
Sternoclavicular joint, 173, 215, 215f, 225t
Sternocleidomastoid muscle, 235f, 250f,
251f, 254, 256f, 257f, 257t, 258,
258f, 259f, 264f, 265f, 307, 307f,
309f, 315f, 619, 620f
Sternocostal joint, 225t
Sternohyoid muscle, 255f, 256, 257f, 257t,
259f, 308f, 309f
Sternothyroid muscle, 255f, 256, 257f,
257t, 308f
Sternum, 133f, 164165, 164f, 174f
Steroid hormones, 457f, 458, 458f, 458t, 459
Steroids, 246, 703
Stethoscope, 587
Stimulus, 345, 488
Stomach, 445t, 456, 457t, 476t, 478, 635,
635f, 636t, 649652, 649f, 650f, 659t
Stomodeum, 665
Strabismus, 532
Straight sinus, 574f
Strains, 92, 225, 226, 286
Stratified cuboidal epithelium, 85, 85f, 86t
Stratified epithelium, 8385, 86t
Stratified squamous epithelium, 8385, 84f, 86t
Stratum basale, 106108, 107f, 109t, 736
Stratum corneum, 107f, 108109, 109t
Stratum functionale, 736
Stratum germinativum, 108
Stratum granulosum, 107f, 108, 109t
Stratum lucidum, 108, 109t
Stratum papillarosum, 111
Stratum reticularosum, 111
Stratum spinosum, 107f, 109t
Stress, 473, 474f
Stress fractures, 339
Stress lines, 143, 143f
Stretch marks, 111, 111f
Stretch reflex, 428
Striate cortex, 513
Striations, 237f
Stroke, 376, 395, 595
Stroma, 91, 728
Structural proteins, 5152
Strychnine, 358
S-T segment, 554
Sty, 532
Styloglossus muscle, 252, 255f, 255t
Stylohyoid muscle, 255f, 256, 257f, 257t, 308f
Styloid cartilage, 142f
Styloid process
of radius, 177f, 178, 321f, 322f
of temporal bone, 147f, 148f, 149f, 151, 152f
of ulna, 177, 177f, 321f, 322f
Stylomandibular joint, 214f
Stylomandibular ligament, 215
Stylomastoid foramen, 148f, 151
Stylopharyngeus muscle, 255f
Subacromial bursae, 216f, 217
Subarachnoid space, 358, 378f, 379, 379f,
381, 381f
Subclavian artery, 562f, 565f, 566f
Subclavian vein, 575f
Subclavius muscle, 264, 264f, 266t, 313f
Subcoracoid bursae, 216f, 217
Subcutaneous adipose tissue, surface anatomy
and, 297, 298f
Subcutaneous injections, 112
Subcutaneous prepatellar bursae, 221, 221f
Subcutaneous tissue. See Hypodermis
Subdeltoid bursae, 216f, 217
Subdural hemorrhage, 333334
Sublingual duct, 640f
Sublingual fold, 640f
Sublingual gland, 635f, 645, 646f, 647t
Subluxation, 226
Submandibular duct, 305f, 640f, 645
Submandibular ganglion, 441
Submandibular gland, 305f, 309f, 635f, 645,
646f, 647t
Submandibular triangle, 308f, 308t

Submental triangle, 308f, 308t
Submucosa
of gastrointestinal tract, 637, 639f
of stomach, 650f, 651f
Submucosal plexus, 637
Subphylum Vertebrata, 27t
Subscapular bursae, 216f, 217
Subscapular fossa, 173, 175f
Subscapularis muscle, 217, 259f, 264f, 266t,
267, 268f
Subserous fascia, 238t
Substantia nigra, 374
Subtarsal sulcus, 302f, 302t
Suckling, 641, 776
Sucrase, 654t
Suction lipectomy, 94
Sudden infant death syndrome (SIDS), 628
Sudoriferous glands, 118119
Sugars, 52
Sulcus, 136t
Sulcus limitans, 390
Superficial circumflex iliac artery, 571f
Superficial digital flexor muscle, 269, 271f,
273t, 325f
tendon of, 272f, 274f, 321f, 323f, 325f
Superficial epigastric artery, 571f
Superficial epigastric vein, 578f
Superficial fascia, 236, 238t, 701f
Superficial fibular nerve, 423, 425f
Superficial muscles, 235f
Superficial palmar arch, 575
Superficial palmar arch artery, 565f, 566
Superficial temporal artery, 305f, 562f, 563564
Superficial temporal vein, 574f
Superficial transversus perinei muscles,
260, 261f
Superficial veins, 572, 573f
Superior, 36t
Superior angle, of scapula, 173, 175f
Superior articular facet, 160f
Superior articular process, 160, 160f, 161f,
162, 162f, 163f, 181f
Superior articular surface, 161
Superior border, of scapula, 173, 175f
Superior cerebellar arteries, 563f
Superior cerebellar peduncles, 375
Superior colliculi, 362t, 374, 376f, 513514
Superior constrictor muscle, 647, 647f
Superior extensor retinaculum muscle, 283f,
291f, 332f
Superior gemellus muscle, 277, 330f
Superior gluteal artery, 568, 569f
Superior gluteal nerve, 424t
Superior labial frenulum, 640f
Superior lacrimal canaliculi, 502
Superior lacrimal punctum, 501f
Superior limb buds, 768
Superior lobe, of lung, 604f, 614, 615f, 616f
Superior mesenteric artery, 318f, 560f, 566,
567t, 568f
Superior mesenteric ganglia, 440
Superior mesenteric vein, 318f, 579, 580f
Superior nasal concha, 153, 153f, 154f, 606
Superior nasal meatus, 605f
Superior nuchal line, 148f, 152, 301
Superior oblique muscles, 252, 254f, 254t,
404, 502, 502f, 503t
Superior ophthalmic vein, 574f
Superior orbital fissure, 146t, 147f, 150f,
152, 153f
Superior parathyroid glands, 468469
Superior peduncle, 376f
Superior peroneal retinaculum, 287f
Superior phrenic arteries, 566, 567t
Superior ramus, of pubis, 181f, 182f, 183
Superior rectal artery, 568f, 569f
Superior rectus muscle, 252, 254f, 254t, 404,
500f, 502, 502f, 503t, 505f
Superior sagittal sinus, 574f
Superior suprarenal arteries, 568f
Superior surface, of rib, 165f
Superior tarsal plate, 500f
Superior thyroid artery, 562f, 563

Superior thyroid vein, 574f
Superior trunk, of brachial plexus, 416
Superior vena cava, 314f, 548f, 549f, 550,
552f, 561f, 572, 573f, 574, 577f, 581f
Superior vermis, 375f
Superior vesicular artery, 568, 569f
Supination, 209, 268
Supinator muscle, 272f, 273t
Supinator reflex, 430t, 431f
Supplementary motor cerebral cortex, 371f
Supporting cells, 497, 497f
Supraclavicular fossa, 311f, 319f
Supraclavicular nerve, 309f, 416t
Supraclavicular triangle, 308f, 308t
Suprahyoid muscles, 256
Supraoptic nuclei, 463
Supraorbital foramen, 146, 146t, 147f, 150f
Supraorbital margin, 145146, 147f, 149f, 305f
Supraorbital ridge, 301
Suprapatellar bursae, 221, 221f
Suprarenal arteries, 566, 567t, 568f
Suprarenal glands. See Adrenal glands
Supraspinatus muscle, 217, 265f, 266t, 267,
267f, 315f, 316f, 324f
Supraspinous fossa, 173, 175f
Supratrochlear artery, 305f
Surface anatomy, 17. See also specific body region
of neonate, 298300
Surfactant, 612, 620
Surgical biopsy, 102
Surgical instruments, in Roman Era, 12f
Surgical neck, of humerus, 176, 176f
Suspensory ligament
of breasts, 739
of lens, 500f, 505, 506f
of ovaries, 728, 729f
of penis, 710f
zonular fibers of, 505f
Sustentacular cells, 703, 704
Sutural bones, 132, 152
Sutures, 197198, 198f, 205t
Swallowing, 608, 648649
Swayback. See Lordosis
Sweat glands, 107f, 108f, 118, 118f, 121,
301, 445t
Sweating, water in, 51
Sweat pore, 107f
Sweet taste, 497, 498f
Swimmers muscle, 265
Sympathetic division, of autonomic nervous
system, 436, 438f, 439f, 440f, 441f,
442f, 443t, 444t, 446447
Sympathetic rami, 414f
Sympathetic trunk ganglion, 414, 414f
Symphyses, 199, 199f, 205t
Symphysial surface, 182f
Symphysis pubis, 180, 181f, 182f, 183f, 199f,
225t, 312, 685f, 699f, 701f, 727f
Synapse, 358
Synaptic cleft, 358
Synaptic knob. See Axon terminal
Synaptic transmission, 358, 358f
Synaptic vesicles, 244, 247f, 358
Synarthroses, 197
Synchondroses, 200, 200f, 205t
Syncytial myotubes, 248
Syncytiotrophoblast, 759, 759f
Syndactyly, 189, 337, 338
Syndesmoses, 198, 198f, 205t
Synergistic muscles, 238, 239f
Synostosis, 198, 200
Synovial fluid, 200202
Synovial joints, 197, 200203, 200f, 205t
kinds of, 202203
movements at, 207213
range of motion, 200
trauma to, 203
Synovial membrane, 202, 206
Synovitis, 225, 230
Synthetase enzymes, 69
Synthetic skin, 125f
Syphilis, 334, 394395, 720, 721f, 721t

Syringomyelia, 395
Systematic (systemic) approach,
to anatomy, 29
Systemic circulation, 550
Systemic effects, 122
System level, organization at, 28, 31f32f
Systole, 552
Systolic pressure, 559
Tabes dorsalis, 395
Tachycardia, 587, 592, 592f
Tactile cells, 108
Tactile receptors, 489t, 490491
Taeniae coli, 657, 657f
Tail
of epididymis, 707f
Tailbone. See Coccyx
Tailors muscle, 279
Talipes, 189, 189f, 338
Talocrural joint, 222224, 223f, 225t
Talus, 187f, 188f
Tanning
of animal hides, 111
of skin, 109, 110
Target cells, 455
Tarsal bones, 133f, 174f, 187f, 188t
Tarsal glands, 501
Tarsal plates, 501
Tarsier, hands of, 26f
Tarsometatarsal joints, 186, 225t
Tarsus, 40, 185186
Taste, 496499
Taste buds, 409, 496497, 497f, 498f
Taste pore, 497, 497f
Tattooing, 109
Taxonomy, 2328
Tay-Sachs disease, 395, 763, 790
Tears. See Lacrimal fluid; Lacrimal secretions
Tectal system, 513
Tectorial membrane, 521, 521f
Tectospinal tract, 388t
Teeth, 304f, 635, 635f, 642645, 643f, 645f.
See also specific tooth
clinical problems of, 669670
in taxonomy, 24, 28
Telencephalon, 347, 362t
Telophase, 70, 71f, 704, 704t, 705f
TEM. See Transmission electron microscope
Temporal bone, 133f, 145f, 147f, 148f, 149f,
152f, 174f, 517f
Temporal fascia, 251f
Temporalis muscle, 235f, 250, 250f, 251f,
253f, 253t, 301, 305f, 324f
Temporal lobe, 360f, 361f, 364f, 366,
366f, 366t
Temporal process, of zygomatic bone, 155
Temporal region, 301
Temporomandibular joint, 146, 214215,
214f, 225t, 305f
Temporomandibular joint syndrome, 215
Tendinous inscriptions, 258, 259f, 311f, 312f
Tendo calcaneus, 223f, 235f, 283284, 284f,
287f, 288f, 327, 329f, 332f, 333f
Tendonitis, 226
Tendons, 92
Tendon sheath, 202, 203f, 236
Tennis elbow, 177, 217, 338
Tenosynovitis, 338
Tensor fasciae latae muscle, 235f, 276277,
276f, 278t, 279f, 280f, 328f, 331f
Tensor tympani muscle, 518f, 519
tendon of, 518f
Tentorium cerebelli, 367, 375, 380t
Teres major muscle, 235f, 259f, 264f,
265f, 266t, 267, 267f, 268f, 316f,
323f, 324f
Teres minor muscle, 217, 265f, 266t, 267,
267f, 316f, 323f, 324f
Terminal bronchioles, 611, 613f
Terminal cisternae, of sarcoplasmic
reticulum, 242f
Terminal ganglia, 441
Terminal hair, 116117

Back Matter
Index
The McGrawHill
Companies, 2001
Index
Testes, 456, 457t, 474, 699f, 700t, 701f,
702705
descent of, 716, 719f
disorders of, 722
of neonate, 300
removal of, 722
trauma to, 711
Testicular artery, 560f, 566, 567t, 569f
Testicular lobules, 702
Testosterone, 115, 457t, 474, 703
Tetanus, 250
Tetralogy, 73, 789
Tetralogy of Fallot, 335, 593, 594f
Thalamus, 306f, 362t, 365f, 370f, 372
Thalidomide, 192
Theca interna, 730, 730f
Thenar eminence, 275, 321f, 323f
Thenar muscles, 275, 275t
Therapeutic touch, 491
Thermogram, 113f
Thermoreceptors, 489t, 490, 491492
Thermoregulation
dual innervation and, 447
hypothalamus and, 373
skin and, 112113, 113f
Theta waves, 368t
Thickened skin, 107t
Thick filaments, 241
Thigh, 40
anterior muscles of, 275276, 279280, 282t
bones of, 183185
muscles of, 275277, 276f, 278f, 278t,
279281, 279f, 280f, 281t, 282f
surface anatomy of, 326327, 326f,
327f, 328f
Thin filaments, 241
Third-class lever, 211, 213f
Third-degree burns, 122, 123f
Third-degree frostbite, 123
Third-division nerve block, 157, 406
Third molars, 642, 643f, 644t
Third trimester, of pregnancy, 785
Third ventricle, 365f, 381, 382f
Thoracic aorta, 566, 567t
Thoracic cavity, 40f, 41, 41f, 44f, 314f, 617f
Thoracic curve, 158f, 159
Thoracic duct, 582, 584f
Thoracic lymph nodes, 584
Thoracic nerves, 401f, 413, 621t
Thoracic region, 3536
developmental conditions involving,
334335
surface anatomy of, 311f
Thoracic spinal nerves, 385f, 414f
Thoracic vertebrae, 159, 161, 161f, 163t, 164f
Thoracic wall, arteries of, 567f
Thoracolumbar division, of autonomic
nervous system. See Sympathetic
division, of autonomic nervous
system
Thorax, 3536
diseases of, 335336
internal anatomy of, 313315, 313f,
314f, 324f
serous membranes of, 43f
trauma to, 335
veins of, 575576, 577f
Thoroughfare channels, 555
Thrombocytes. See Platelets
Thrombus, 595
Thumb, 178, 565f
Thymine, 65, 67f
Thymosin, 457t, 476
Thymus, 456, 456f, 457t, 476, 476f, 584f,
586, 586f, 587t
Thymus dependent cells, 476
Thyrocalcitonin. See Calcitonin
Thyrocervical trunk, 562f, 564
Thyroglossal duct, 478
Thyrohyoid muscle, 255f, 256, 257f, 257t, 308f
Thyroid, etymology of term, 30

Thyroid cartilage, 142f, 306, 306f, 307f,
608, 610f
Thyroid diverticulum, 478
Thyroid follicles, 467
Thyroid gland, 306307, 456, 456f, 457t,
466468, 467f, 468f
functions of, 468
Thyroid hormones, 467t
Thyroid-stimulating hormone (TSH), 462,
464t, 468
Thyrotropin, 462, 464t
Thyrotropin-releasing hormone (TRH), 462
Thyroxine, 457t, 458, 466, 467t, 468
Tibia, 133f, 174f, 185, 185f, 186f, 188t,
328f, 329f, 332f
Tibial collateral ligament, 221f
Tibialis anterior muscle, 235f, 283, 283f,
284f, 285f, 286t, 328f, 329f, 332f
tendon of, 329f
Tibialis posterior muscle, 285, 285f, 286t, 288f
Tibial nerve, 333f, 423, 424t, 425f
Tibial tuberosity, 185, 186f, 327f
Tibiocalcaneal ligament, 223f
Tibiofemoral joint, 200f, 220222, 221f,
222f, 225t
Tibionavicular ligament, 223f
Tidal volume (TV), 621t, 622f
Tinea pedis, 128
Tinnitus, 533
Tissue(s). See also specific type
definition of, 78
Tissue analysis, 102
Tissue composition, changes in, 102
Tissue fluid, 540
Tissue level, organization at, 28
Tissue-rejection reaction, 102
Tissue repair, 98
Tissue transplantation, 102103
TLC. See Total lung capacity
Toes. See also Digits
Tongue, 306f, 605f, 635, 635f, 640f, 641f,
642, 642f
extrinsic muscles of, 252, 255f, 255t
intrinsic muscles of, 252
Tonic contracture, 659t
Tonic receptors, 490
Tonometer, 527
Tonsillectomy, 584, 608
Tonsillitis, 628
Tonsils, 584, 584f, 587t, 607, 608
Tooth. See Teeth
Torso. See Trunk
Torticollis, 286
Total lung capacity (TLC), 621t, 622f
Touch. See Tactile receptors
Toxic goiter, 481
Trabeculae, 138, 583
Trabeculae carneae, 549f, 550
Trace minerals, 52t
Trachea, 306, 306f, 314f, 604f, 609610,
610f, 611f, 612f, 618t, 641f
Tracheal cartilage, 610f, 611f
Trachealis muscle, 611f
Tracheobronchitis, 628
Tracheostomy, 306, 610, 611f
Tracheotomy, 610
Trachoma, 532
Tragus, 303, 517f
Transfer RNA (tRNA), 67, 68, 69, 69f
Transitional epithelium, 85, 86f, 86t
Transmission, of light rays, 509
Transmission electron microscope (TEM), 57f
Transplantation
allogenic, 544
autotransplant, 544
bone marrow, 544
corneal, 504
fetal tissue, 103
heterotransplant, 124
homotransplant, 102
kidney, 693
tissue, 102103
Transportation, in circulatory system, 538539
Transurethral prostatic resection, 709
Transversalis fascia, 259f
Transverse abdominis muscle, 317f
Transverse acetabular ligament, 220
Transverse arch, 188, 188f
Transverse cervical nerve, 309f, 416t
Transverse colon, 317f, 318f, 635f, 638f, 656
Transverse fissure, 375
Transverse flexure, 657f
Transverse foramen, 160, 160f
Transverse fractures, 189, 190f
Transverse humeral retinaculum, 216
Transverse ligament, 221f, 222f
Transverse lines, 162, 163f
Transverse palatine folds, 640f, 641
Transverse plane, 33, 33f, 34f
Transverse process, 160, 160f, 161f, 162f, 163f
Transverse sinus, 574f
Transverse tubules, 241, 242f
Transversus abdominis muscle, 258, 258f,
259f, 260t, 264f, 317f, 620f
Transversus perinei muscle, 260, 260t
Trapezium bone, 178, 179f, 180f, 205f
Trapezius muscle, 235f, 254, 257f, 259f, 264,
264f, 265f, 266t, 267f, 307, 307f, 308f,
309f, 310, 311f, 315f, 316f, 319f, 323f
Trapezoid bone, 178, 179f, 180f
Trauma
to abdomen, 336337
to brachial plexus, 416
to brain, 367, 377, 450
to cells, 72
to cerebellum, 376
to circulatory system, 596598
to coccyx, 163
to connective tissue, 92
to eyeball, 508
to head, 333334
to hip, 339
to joints, 225226, 229f
to kidneys, 692
to ligaments, 92
to lower extremity, 339
to muscles, 92
to neck, 333334
to pelvis, 692
to penis, 711
to respiratory system, 627628
to scrotum, 711
to shoulder, 338
skeletal, 189191
to sphenoid bone, 152
to synovial joints, 203
to tendons, 92
to testis, 711
to thorax, 335
to upper extremity, 338
to urethra, 692
to urinary bladder, 692
to urinary organs, 692693
Traumatic fractures, 189
Trench mouth, 669f, 672
Trepanation, 3, 4f
Treponema pallidum, 721
TRH. See Thyrotropin-releasing hormone
Triangle of auscultation, 310, 315f, 323f
Triangular fossa, 517f
Triceps brachii muscle, 235f, 259f, 267f,
268, 268f, 269f, 270f, 270t, 316f,
319f, 320f, 321f, 323f, 324f
Triceps reflex, 430t, 431f
Triceps surae muscle, 284
Tricuspid area, 554, 554f
Tricuspid valve. See Right atrioventricular
valve
Trigeminal ganglion, 405
Trigeminal nerve (V), 157, 360f, 402f, 404t,
405406, 408f, 413t, 496, 498f
839
Trigone, 685, 686f

Triiodothyronine, 457t, 466, 467t, 468
Triplets, 786
Triquetrum bone, 178, 179f, 180f
Trisomy, 73
Trisomy 21, 763
tRNA. See Transfer RNA
Trochanter, 136t
Trochlea, 176, 176f, 252, 502, 502f
Trochlear nerve (IV), 402f, 404405, 404t,
408f, 413t
Trochlear notch, 177, 177f
Trophic hormones, 457t, 462
Trophoblast, 80, 757
True hermaphroditism, 714
True labor, 775
True pelvis, 180, 183f
True ribs, 164f, 165
True vocal cords. See Vocal folds
Truncus arteriosus, 589, 590t
Trunk, 3537
cavities of, 41f
internal anatomy of, 313315, 316f
muscles of, 259f, 264f, 265f
Trypsin, 654t
TSH. See Thyroid-stimulating hormone
T tubules. See Transverse tubules
Tubal ligation, 744, 744f, 749
Tubal pregnancy, 747, 785
Tubercle, 136t
of rib, 165, 165f
Tuberculosis, 628
Tuberosity, 136t
of calcaneus, 186, 187f, 290f
of tibia, 283f
of ulna, 177f
Tubular glands, 87, 88f, 89t
Tumors, 73, 394, 483, 746
Tunic(s)
of gastrointestinal tract, 637639, 639f
of rectum, 658f
Tunica albuginea, 701f, 702, 710f, 728
Tunica externa, 555, 556f
Tunica interna, 555, 556f
Tunica media, 555, 556f
Tunica muscularis, 637, 639f
Tunica vaginalis, 701f, 702
Tuning fork tests, 527
Tunnel vision, 513
Turbinates, 153, 606
Turners syndrome, 73, 715
TV. See Tidal volume
T wave, 554
Twelfth rib, 164f
Tympanic cavity, 517, 517f, 518f, 529
Tympanic membrane, 158f, 516, 517, 518f,
529, 533
Tympanic part, of temporal bone, 151, 152f
Tympanum, etymology of term, 30
Tyrosinase, 109
Tyrosine, 109
Ulcers
decubitus, 111, 112f, 128
peptic, 337, 651, 670
Ulna, 133f, 174f, 177, 177f, 181t, 198f, 325f
Ulnar artery, 560f, 565, 565f
Ulnar collateral ligament, 217, 218f, 219f
Ulnar nerve, 320f, 401f, 418t, 419, 419f
Ulnar notch, 177f, 178
Ulnar recurrent artery, 565f, 566
Ulnar sulcus, 176, 176f
Ulnar vein, 321f, 573f, 575, 575f
Ultrasonography, 587, 772, 774f, 786787, 788f
Ultrastructure, 17
Ultraviolet light, protection from, 112
Umbilical arteries, 580, 581f, 582t, 765, 767
Umbilical cord, 300, 580, 766767, 767f
Umbilical hernia, 293, 313
Umbilical region, 38f, 38t
Umbilical vein, 580, 581f, 582t, 765, 767
Umbilicus, 36, 259f, 312313, 312f, 317f
Unicellular glands, 87, 87f, 89t

840
Back Matter
Index
The McGrawHill
Companies, 2001
Index
Unicornuate uterus, 746, 746f

Upper arm birth palsy, 338
Upper back, muscles of, 256f
Upper extremity, 38
bones of, 132t, 134, 181t
diseases of, 338
internal anatomy of, 322
of neonate, 299t
surface anatomy of, 319322, 319f, 320f
trauma to, 338
veins of, 574575, 575f, 576f
Upper leg, 40
Upper lip, 605f
Upper respiratory system, 603, 605f, 624, 625f
Upright posture, 2527, 183
Urachus, 690, 764
Uracil, 67, 67f, 68
Uremia, 692
Ureteric bud, 690
Ureters, 676, 676f, 677, 677f, 679f, 684,
684f, 685f, 699f, 710f
Urethra, 676, 676f, 685f, 686f, 687, 698, 699f
female, 727f
male, 709, 709f, 710f
trauma to, 692
Urethral folds, 716, 743
Urethral glands, 687, 709
Urethral groove, 716, 743
Urethral orifice, 686f, 687, 701f, 711, 716, 737f
Urethritis, 692, 721t
Urinalysis, 480, 682, 692
Urinary bladder, 638f, 676, 676f, 677f,
685687, 685f, 699f, 710f, 727f
female, 686f
male, 686f
trauma to, 692
Urinary incontinence, 694
Urinary retention, 688
Urinary stones, 684, 693
Urinary system, 31f, 675696, 676f
functions of, 676
infections of, 692
obstruction of, 693694
trauma to, 692693
Urinary tract infections (UTI), 692
Urination. See Micturition
Urine, 682, 683t
Urogenital diaphragm, 260, 261f, 686f
Urogenital membrane, 667
Urogenital ridge, 689
Urogenital sinus, 667
Urogenital triangle, 39f, 700, 701f
Urorectal septum, 667
Urticaria, 128
Uterine artery, 568, 736f
Uterine cancer, 747
Uterine cavity, 734, 744f
Uterine displacement, 748
Uterine neoplasms, 747
Uterine ostium, 734
Uterine prolapse, 735, 748
Uterine tubes, 716, 726, 727f, 727t, 729f,
732734, 734f, 744f, 746747
Uterine wall, 735736, 735f
Uterus, 685f, 726, 727f, 727t, 735f
abnormalities of, 746
atrophy of, following delivery, 59
development of, 716
innervation of, 736

problems involving, 747748
size of, 736
structure of, 734
support of, 735
UTI. See Urinary tract infections
Utricle, 515, 519f, 520, 524
Utricular portion, of otocyst, 529
UVA rays, 110
UVB rays, 110
Uvea, 504505, 508t
Uvula, 30, 305f, 605f, 607
Vacuoles, 53t, 61
Vagal reflexes, 448t
Vagina, 638f, 685f, 716, 726, 727f, 727t,
729f, 736737, 737f, 748
Vaginal artery, 568, 736f
Vaginal fornix, 727f
Vaginal lining, pH of, 85
Vaginal orifice, 701f, 727f, 736, 737f
Vaginal rugae, 736737
Vaginal spermicides, 750f
Vaginal vestibule, 727t, 738
Vaginitis, 721t, 748
Vagotomy, 672
Vagus nerve (X), 314f, 402f, 405t,
411412, 411f
assessment of, 413t
protection of, 307
in taste sensation, 498f
Vagus nuclei, 377
Vallate papillae, 497, 497f, 642, 642f
Valsalvas maneuver, 574
Valves, of heart, 547550, 548t, 551f
Valvular auscultatory areas, 554, 554f
Valvular incompetence, 335
Valvular insufficiency, 554
Valvular stenosis, 335
Varicocele, 703
Varicose veins, 327, 558, 595596
Vasa recta, 678, 680f
Vascular disorders, 595596
Vascular processes, 351
Vascular shunts, 555
Vascular tissue, 98. See also Blood
Vascular tunic, of eyeball, 504505, 504f, 508t
Vas deferens, 707
Vasectomy, 720, 720f
Vasomotor center, of medulla oblongata, 377
Vastus intermedius muscle, 276f, 278f, 279f,
280, 282f, 282t
Vastus lateralis muscle, 235f, 276f, 277,
278f, 279f, 280, 280f, 282f, 282t,
284f, 289f, 327f, 328f, 331f
Vastus medialis muscle, 235f, 276f, 278f, 279f,
280, 281f, 282f, 282t, 327f, 328f, 331f
VC. See Vital capacity
Veins, 540, 555, 556f, 558, 561f, 571579,
573f. See also specific veins
Vein stripping, 596
Vellus, 116
Venereal warts, 119, 721t
Venous blood, 540
Venous sinuses, 574
Venous sinus thrombosis, 334
Venous valves, 558, 558f
Ventilation, 603
Ventral. See Anterior
Ventricles
of brain, 381, 382f
of heart, 547, 589, 590t
Ventricular fibrillation, 592, 592f
Ventricular septal defect, 334, 593,
593f, 594f
Ventricular tachycardia, 592, 592f
Venules, 540, 555, 556f, 557f
Vermilion, 640, 641f
Vermis, 375
Vernix caseosa, 121, 299300, 773
Vertebrae, 158, 158f, 159
regional characteristics of, 160163
in taxonomy, 28
Vertebral arch, 159, 160f

Vertebral artery, 360f, 561, 562f, 563f, 564
Vertebral cavity, 40f
Vertebral column, 24f, 132134, 133f,
158163, 158f, 174f, 262f
articulations of, 226
bones of, 132t
curvature disorders of, 226
functions of, 159
injuries to, 392393
regions of, 163t
Vertebral curves, 158f, 159, 159f
Vertebral foramen, 159, 160f, 161f, 162f
Vertebral region, 36
Vertebral vein, 574f
Vertebra prominens, 265f
Vertex position, of fetus, 774, 774f
Vertigo, 527, 533
Vesalius, Andreas, 1415, 14f
Vesicouterine pouch, 727f, 735
Vesicular artery, 569f
Vesicular ovarian follicle, 730, 731f, 733f
Vestibular bulbs, 686f, 738
Vestibular folds, 609, 609f
Vestibular ganglion, 409
Vestibular glands, 727t, 738
Vestibular membrane, 520, 520f, 521f
Vestibular nerve, 409, 410f, 517f, 520f
Vestibular nuclei, 410
Vestibular organs, 409
hair cells of, 524, 524f
Vestibular window, 517, 518f, 520, 520f
Vestibule
of inner ear, 519f, 520
of oral cavity, 640, 641f
of reproductive system, 716
Vestibulocochlear nerve (VIII), 360f, 402f,
405t, 409410, 410f, 413t, 520f, 523
Vestibulospinal tract, 388t, 389
Vibrissae, 606f
Villi, intestinal, 653, 655f
Villous chorion, 765
Vincula longa, 274f
Virchow, Rudolf, 16
Viscera, 41, 635
Visceral effector organs, and autonomic
nervous system, 436438
Visceral motor neurons, 354, 356f
Visceral organs, 41, 42f
Visceral pain, 491
Visceral pericardium, 43, 43f, 546
Visceral peritoneum, 42f, 43, 636, 637f, 638f
Visceral pleura, 43, 43f, 615
Visceral portion, of serous membrane, 636
Visceral reflexes, 427
Visceral senses, 489
Visceral sensory neurons, 354, 356f
Visceroceptors, 490
Viscerocranium, 141, 142f
Visible light, 511
Vision. See also Eye(s); Visual sense
binocular, 499
color, 513
double, 511f, 532
stereoscopic, 27
tunnel, 513
Visual cortex, 403
Visual fields, 514f
Visual information, processing of, 514
Visual inspection, 297
Visual pathways, 407f
Visual portion, of retina, 505
Visual sense. See also Eye(s)
accessory structures of eye and, 499502
function of eyeball in, 509511
neural pathways for, 513514
structure of eyeball and, 504508
Visual spectrum, 511
Vital capacity (VC), 621t, 622f
Vitamin(s), 52t, 114
Vitamin B12, 652
Vitamin D deficiency, 135, 166
Vitiligo, 110
Vitreous humor, 508, 512f, 516
Vivisection, human, 10, 11

Vocal cords. See Vocal folds
Vocal folds, 609, 609f
Vocal structures, in taxonomy, 27
Volkmanns canals, 140
Vomer, 147f, 148f, 150f, 157, 605
Vomiting, 639, 651652, 672
Vomiting center, 651
Vulva, 737738, 748
Vulval dermatitis, 107t
Vulvovaginitis, 748
Wakefulness center, 373
Warm-blooded animals, 540
Warts, 119, 129
Water
as component of body, 51, 52t
hypothalamus regulation of, 373
Watson, James, 66
Wernickes aphasia, 371
Wernickes area, 371, 371f
Wharton, Thomas, 15
Whartons duct. See Submandibular duct
Whartons jelly. See Mucoid connective tissue
Whiplash, 161, 334, 393, 393f
White blood cells. See Leukocytes
Whitehead. See Comedo
White matter
of brain, 351, 359, 363, 365f, 368, 369f
of spinal cord, 384
White pulp, 585, 586f
White rami communicantes, 439, 439f
White ramus, 414
William of Saliceto, 13
Willis, Thomas, 15
Wing, of nose, 303, 304f, 304t
Wisdom teeth. See Third molars
Withdrawal reflex, 428, 429f
Womb. See Uterus
Wormian bones. See Sutural bones
Wounds, 125126, 125f, 126f. See also Trauma
Wrist. See Carpus
Wristdrop, 338, 419
Wryneck, 286
Xenograft, 102, 124
Xiphisternal joint, 310
Xiphoid process, 164, 164f, 311f, 312f
Xiphos, etymology of term, 30
X-rays, 18
Yang, ancient Chinese concept of, 8
Yawning, 622t
Yellow bone marrow, 134, 137
Yin, ancient Chinese concept of, 8
Yolk sac, 763764
Z lines, 241, 242, 242f, 243f, 245f
Zona fasciculata, 471
Zona glomerulosa, 471
Zona pellucida, 729, 730f, 733f, 755
Zona reticularis, 471
Zone of calcified cartilage, 143
Zone of dechondrification, 143
Zone of hypertrophic cartilage, 143
Zone of proliferating cartilage, 142
Zone of resting cartilage, 142
Zonular fibers, of suspensory ligament, 505,
505f, 506f
Zygomatic arch, 146, 149f, 150f, 155, 251f
Zygomatic bone, 133f, 142f, 145f, 147f,
148f, 150f, 155, 174f
Zygomaticofacial foramen, 146t, 155
Zygomatic process, 146, 147f, 148f, 152f,
155, 155f
Zygomaticus major muscle, 250f, 251f
Zygomaticus minor muscle, 250f, 251f
Zygomaticus muscle, 235f, 251t, 305f
Zygote, 79, 732, 760t

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Van De Graaff: Human

Anatomy, Sixth Edition

uman Anatomy was written to serve as a foundation and

Van De Graaff: Human

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Van De Graaff: Human

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