Professional Documents
Culture Documents
Fundamentals
FIFTI{ EDITION
EugeneP.Odum,Ph.D.
oJGeorglaInstitute
LateoJUniversity oJEcologt
GaryW.Barrett,Ph.D.
of Ecolog4UniversityoJGeor$aInstituteof Ecologt
OdumProJessor
-rHorvlsoN
=+-*
BROOKS/COLE
1
TheScopeof Ecology
Ecology:Historyand Relevanceto
Humankind
Levels-of.OrganizationHierarchy
The EmergentPropertyPrinciple
TranscendingFunctionsand Control
Pro@sses
EcologicalIntertacing
About Models
DisciplinaryReductionismto Transdisciplinary
Holism
rl
1 Ecology:
HistoryandRelevance
to llumankind
The word ecologiis derived from rhe Greek oihos,meaning "household,"and logos,
meaning "study."Thus, the study of the environmentalhouseincludes all the organ-
isms in it and all the functional processesthat make the house habitable. Literally,
then, ecology is the study o[ "life at home" with emphasison "the totality or pat-
tern of relationsbetween organismsand their environment," to cite a standarddic-
tionary definition of the word (Merriam-Webster's CollegiateDictionory,1Othedition,
s.v. "ecology").
The word economics is also derived from the Greek root oikos.As nom[s means
"management,"economicstranslatesas "the managementof the household" and, ac-
cordingly, ecology and economicsshould be companion disciplines.Unfortunately,
many people view ecologistsand economistsas advercarieswith antitheticalvisions.
Table 1-1 attempts to illustrate perceiveddifferencesbetween economicsand ecol-
ogy. l-ater, this book will considerthe confrontation that resuhsb€causeeach disci-
pline takesa narrow view of its subjectand, more important, the rapid development
of a new interfacediscipline, ecoiogical economics, that is begnmng to bridge the gap
betweenecologyand economics(Costanza,Cumberland,et al. I997; Barrettand Fa-
rina 2000;L. R. Brown 2001).
Ecologlrwas of practical interestearly in human history. In primitive society,all
individuals neededto know their environment-that is, to understandthe forcesof
nature and the plants and animals around them-to survive. The beginning of civi-
lization, in fact, coincided with the use of fire and other tools to modify the environ-
ment. Becauseo[ technologicalachievements,humans seem to depend less on the
natural environment for their daily needs;many of us lorget our continuing depen-
dence on nature for air, water, and indirectly, food, not to mention wasteassimila-
tion, rccreation,and many other s€rvicessupplied by nature.Also, economicsystems,
of whatever political ideology, value things made by human beings that primarily
benefit the individual, but they placelittle monetary value on the goodsand services
of nature that benefit us as a society.Until there is a cdsis, humans tend to uke nat-
l"
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-
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tn,
ac-
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lap ural goods and servicesfor granted; we assumethey are unlimited or somehow re-
placeableby technologicalinnovations, even though we know rhar life necessities
such as oxygen and water may be recyclablebut nor replaceable.As long as the life-
rlL
support seruicesare consideredfree, they have no value in current market systems
of (seeH. T. Odum and E. P Odum 2000).
ti-
Like all phasesof learning, the scienceof ecologyhas had a gradual if spasmodic
,n-
development during recorded history. The writings of Hippocrates,Aristotle, and
he
other philosophersol ancient Greececlearly contain referencesto ecologicaltopics.
tn-
However, the Greeksdid not have a word for ecology.The word ecolog,,is o[ recenr
la-
origin, having been frrst proposed by the G€rman biologist Ernst Haeckel in 1869.
:ts,
Haeckel defined ecolosl as "the study of the natural environmem including rhe re-
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lations of organismsto one another and to their surroundings" (Haeckel 1869). Be-
ies
fore this, during a biological renaissancein the eighteenthand nineteenthcenturies,
.1t-
many scholarshad contributed to the subject,even though the word ecologrwas not
in use. For example,in the early 1700s,Antoni van Leeuwenhoek,best known as a
I
premier microscopist,also pioneeredthe study of food chains and population regu-
Iation, and the writings of the English botanist Richard Bradley revealedhis under-
standing of biological productivity. All three of thesesubjectsare important areasof
modern ecology.
As a recognized,distinct field ofscience,ecologydatesfrom about 1900, but only
in the past few decadeshas the word becomepart of the generalvocabulary.At first,
the field was rather sharply divided along taxonomic lines (such asplant ecologyand
animal ecologl), but the biotic community concept of Frederick E. Clementsand
Victor E. Shelford,the food chain and material cycling conceptsof Ra1'rnondLinde-
man and G. Evelyn Hutchinson, and the whole iake studles o[ Edward A. Birge and
ChauncyJuday,amongothers,helped establishbasictheory for a unified 6eld ofgen-
eral ecology.The work of thesepioneerswill be cited often in subsequentchapters.
What can best be describedas a worldwide environmentalawarenessmovement
burst upon the sceneduing two years, 1968 lo 1970, as astronautstook the first
photographsof Earth as seen from outer space.For the first time in human history,
dy we were able to seeEanh as a whole and to realizehow alone and fragile Earth hov-
ers in space(Fig. I-1). Suddenly,during the 1970s,almosteveryonebecamecon-
4 C H A P T E R1 The Scope of Ecology
cemed about pollution, natural areas,population growrh, food and energ/ con-
sumption, and biotic diversity, as indicated by the wide coverageof environmental
concernsin the popular press.The 1970swere frequently referred to as the "decade
of the enyironment,"initiated by the first "Earth Day" on 22 April 1970. Then, in the
1980s and 1990s, environmental issueswere pushed into the political background
by concerns for human relations-problems such as crime, the cold war, govern-
ment budgets,and welfare.As we enter the early stagesof the twenty-first century,
environmental concelns are again coming to rhe forefront becausehuman abuseo[
Earth continuesto escalate.We hope that this time, to usea medical analogy,our em-
phasiswill be on prev€ntionrather than on treatment,and ecologyas outlined in this
book, can contibute a great deal to prevention technology and ecosystemhealth
(Barrett2001).
The increasein public attention had a profound effecton academicecology.Be-
fore the 1970s, ecologywas viewed largely as a subdiscipline of bioiogy. Ecologisrs
were staffedin biology depanmenrc,and ecologycourseswere generallyfound only
in the biological sciencecurricula. Although ecology remains strongly rooted in bi-
ology, it has emergedfrom biologr as an essemiallynew, inregrarivediscipline thar
links physical and biological processesand forms a bridge berweenrhe natural sci-
encesand the social sciences(E. P Odum 1977). Most collegesnow offer campus-
wide coursesand haveseparatemajors,departments,schools,centers,or inslitutes of
ecology.While the scope of ecology is expanding, the study of how individual or-
ganismsand speciesinterfaceand useresourcesintensifies.The multilevel approach,
as outlined in the next section,brings together "evolutionary" and "systems"think-
ing, two approachesthat have tended to divide the field in recentyears.
2 levels-of-0rganizationHierarchy
Perhapsthe best way to delimit modern ecologyis to consider the concept o[ levels
of organization, visualizedas an ecologicalspectrum (Fig. 1-2) and as an extended
ecologicalhierarchy (Fig. 1-3). Hierarchy means "an arrangementinto a graded
setes" (Merriam-Webster's CollegieteDicfionary,10th edition, s.v. "hierarchy"). Inter-
action with the physical environment (energy and matter) at each level produces
characteristicfunctional systems.A system, accordingto a standarddefrmuon, con-
sists of "regularly interacting and interdependent components forming a unified
of
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ts 3 TheEmergent
Propefi Principte
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Al important consequenceof hierarchical organization is that as componems, or
.d subsets,are combined to produce larger functional wholes, new propenies emerge
l;' that were not preseni at the level below Accordingly, an emergent property of in
a ecologicallevel or unit cannot be predicted from the study of the componentsof that
t- level or unit. Another way to expressrhe sameconcept rs nonreducibie propeny-
l- that is, a property ofthe whole not reducibleto the sum ofrhe propeniesof the parts.
a Though findings at any one level aid in the study of the next level, they never com_
i- pletely explain the phenomenaoccurring ar the nexr level,which musr itselfbe stud-
t. ied to complete rhe picrure.
LE Two examples,one from the physical realm and one from rhe ecologicalrealm,
te will sufficeto illusrrate emergentproperries.When hydrogen and oxygen are com-
t- bined in a certain molecular configuration, water is formed-a liquid wirh proper-
ties utterly different from those of its gaseouscomponents.When certain algaeand
l- coelenterateanimals evolvetogetherto produce a coral, an efficientnutrientiycling
(- mechanismis createdthat enablesthe comblned system to maintain a high rate of
l- productivity in watercwith a very low nutrient content. Thus, the fabulousproduc-
8 CHAPTER 1 The Scope of Ecology
tivity and diversity of coral reefsare emergentpropenies only at the level of the reef
community.
Salt (1979) suggestedthat a distinction be made betweenemergempropeties, as
defined previously,and collective properties, which are summationsof the behav-
ior of components.Both are propertiesof the whole, but the collectivepropertiesdo
not lnvolve new or unique characteristicsresuking from the functioning of the whole
lniL Birth rateis an exampleof a population level collectiveproperty, as it is merely
a sum of the individual births in a designaiedtime period, expressedas a lraction or
percentof the total number of individuals in the population. New propertiesemerge
becausethe componentsinteract, not becausethe basic nature of the componentsis
changed.Partsare not "melted down," as it were, but integratedto produce unique
new properties. lt can be demonstratedmathematicallythat integrativehierarchies
evolvemore rapidly from their constituentsthan nonhierarchicalsystemswith the
same number of elements;they are also more resilient in responseto disturbance.
Theoretically,when hierarchiesare decomposedto theirvarious levelsoIsubsysrems,
the latter can still interac[ and reorganizeto achievea higher level of complexity.
Someattributes,obviously,becomemore complex and variableas one proceeds
to higher levelsof organization,but often other attributesbecomelesscomplex and
lessvariableas one goesfrom the smaller to the larger unit. Becausefeedbackmech-
anisms(checksand balances,forcesand counterforces)operatethroughout, the am-
plitude ol oscillations tends to be reduced as smaller units function within larger
units. Statistically,the va ance of the whole system level property is less than the
sum of the varianceof the parts. For example,the rate of photosynthesisof a foresr
community is lessvariablethan that of individual leavesor treeswithin the commu-
nity, becausewhen one component siows down, another component may speedup
to compensate.When one considersboth the emergentpropenies and the increasing
homeostasisthat develop at each level, not all component parts must be known be-
fore the whole can be understood.This is an important point, becausesome contend
that it is uselessto try to work on complex populations and communitieswhen the
smaller units are not yet fully understood. Quite the contrary, one may begin study
at any point in the spectrum, provided that adjacentlevels, as well as the level in
question,are considered,because,as alreadynoted, some attributes are predictable
from parts (collectiveproperties),but others are no[ (emergentproperties).ldeally, a
system-levelstudyis itselfa threefoldhierarchy:system,subsystem(next levelbelow),
and suprasystem(nextlevelabove).Formore on emergenlproperties,seef. F. H. AIIen
and Starr(1982),T. F. H. Allen and Hoekstra(1992),and Ahl and Allen (1996).
Each biosystemlevel has emergentproperties and reduced varianceas well as a
summation of attributes of its subsystemcomponents.The folk wisdom about the
forest being more than just a collection of treesis, indeed, a first working principle
of ecology.Although the philosophy of sciencehas alwaysbeenholistic in seekingto
understandphenomenaasa whole, in recentyearsthe practiceof sciencehasbecome
increasinglyreductionist in seekingto understand phenomenaby detailed study of
smallerand smaller componenls.LaszLoand Margenau(1972) describedwithin the
history o[ sciencean alternation of reductionist and hoiistic thinking (redi{ctiorxism
constructionism and atomism-holism are other pairs of words used to contrast these
philosophicalapproaches).The law of diminishing retums may very well be involved
here, as excessiveellort in any one direction eventuallynecessitates taking the other
(or another) direction.
The reductionistapproachthat hasdominatedscienceand technologysincelsaac
r
SECTION 4 Transcending Functions and Control Processes 9
ref Newton hasmade major contdbutions. For example,researchat the cellular and mo-
lecular levelshas establisheda firm basis for the future cure and prevention of can-
as cersat the level of the organism.However,celllevel sciencewiil contribute very little
to the well-being or survival of human civilization if we understandthe higher levels
lc-t of organization so inadequatelythat we can hnd no solutions to population over-
rle growth, pollution, and other forms ofsocietal and environmentaldisorders.Both ho-
:\' lism and reductionism must be accordedequal value-and simultaneously,not al-
Llf tematively (E. P Odum 1977; Barrett 1994). Ecologyseek slnthesis, not separation.
The revival of the holistic disciplinesmay be due at leastpartly to citizen dissatisfac-
is tion with the specializedscientistwho cannot respond to the large-scaleproblems
ue that need urgent attention. (Histodan L),nn White's 1980 essay"The Ecology of Our
e5 Science"is recommendedreadlng on this viewpoint.) Accordingly, we shall discuss
he ecologicalprinciples at the ecosystemlevel, with appropiate attention to organism,
te. population, and community subsetsand to landscape,biome, and ecospheresupra-
1S, sets.This is the philosophical basisfor the organizationo[ the chaptersin this book.
Fortunately,in the past I0 years,technologicaladvanceshaveallowed humans to
deal quantitativelywith large,complex systemssuch as ecosystemsand landscapes.
rd Tracer methodology,masschemistry (spectrometry,colorimetry, chromatography),
remotesensing,automadcmonitoring, mathematicmodeling, geographicalinforma-
'r't- tion systems(GlS), and computer technologyare providing the tools. Technologyis,
of course,a double-edgedsword: it can be the means of underctandingthe whole-
he nessof humans and nature or of destroyingit.
rp
le-
4 Transcending
Functionsand ContrulProeesses
rd
he Whereaseachlevel in the ecologicalhierarchy can be expectedto haveunique emer-
lv gent and collectiveprope ies, there arc basic functions that operateat a]l levels.Ex-
in amplesof such transcending functions are behavior,development,diversity, ener-
rle getics,evolution, integration,and regulation (seeFig. l-3 for details).Someo[these
.a (energetics,for example)operatethe samethroughout the hierarchy,but others dif-
..), fer in modusoperandiat differentlevels.Natural selectionevolulion, for example,in-
an volvesmutations and other direct geneticinteractionsat the organismlevel but indl-
rect coevolutionaryand group selectionprocessesat higher levels.
ia It is especiallyimpoftant to emphasizethat although positive and negativefeed-
he back controls are universal,from the organismdom, control is sel?oirlf,in that it in-
rle volvesvery exactinggenetic,hormonal, and neural controls on growth and develop-
to ment, Ieadingto what is often called homeostasis. As noted on the right-hand side
ne of Figure 1-4, there are no set-pointcontrolsabovethe organismlevel (no chemostats
oI or thermostatsin nature). Accordingly, feedbackcontrol is much looser,resulting in
ne pulsing rather than sieady states.The term homeorhesis, from the Greek meaning
n- "maintaining the floq" has been suggestedfor this pulsing control. ln other words,
5e there are no equilibriums at the ecosystemand ecospherelevels,but there arepulsing
:d bclances,such asbetweenproduction and respirationor betweenoxygenand carbon
ET dioxide in the atmosphere.Failure to recognizethis differencein cybernetics (the sci-
encedealingwith mechanismsof control or regulation)has resultedin much confu-
sion about the realitiesof the so-called"balanceof nature."
10 CHAPTER1 TheScopeot Ecology
5 Ecologicallnterfacing
6 AboutModels
S E C T I O N6 AboutModels 11
Feedback loop
-
SECTI0N 6 AboutModels 13
--> Negative
Energy source
(Sourceol energyfrom
outsidethesystem)
-\ t-
Slorage
{A compartmentoJ
energy storage)
Y
-Y
----\. =
t--_r lnteraclron
-.L
--:
Heatsink
(Degradedenergy
afterusein work)
(Twoor moreflows
of energyto producea
high-qualityenergy)
Producer
(Converts
and concentrates
sorarener9yl
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$.-.,.4\-- Capitaltransaction
(Fow of moneyto
payfor flowof energy)
7 llisciplinarylieductionismto Tlansdisciptinary
Holism
a gaper entltled "The Emergenceof Ecologr as a New lmegrarive Discipline,,,
ln
E. P Odum (1977) noted that ecologyhad becomea new holistii discipline, having
roots in the biologicai, physical,and social sciences,rather than jusr a iubdiscipline
ofbiology. Thus, a goal of ecoiogyis to link the natural and sociaisciences.It should
be noted that most disciplinesand disciplinary approachesare basedon increased
specializationin isolation (Fig. l-10). The early evoiution and developmentofecol_
to transdisciplinary
holism(af-
)k.
terJantsch 1972).
tI MULTIDISCIPLINARY
no cooperation
CROSSDISCIPLINARY
rigid polarizationtoward
specific monodisciplinary
concepl
INTERDISCIPLINARY
coordination
by higher
levelconcept
TRANSDISCIPLINARY
multi-levelcoordinationof
entire education/ innovation
systern
16 CHAPTER 1 The Scope of Ecology
: "many"), espe-
ogli was frcquently basedon multidisciplinary approaches(multi
cially during the 1960s and 1970s.Unfonunately, the multidisciplinary approaches
lacked cooperation or focus. To achievecooperation and define goals,insliutes or
centerswere establishedon campusesthroughout the world' such as the lnstilute of
Ecologylocatedon the campusof lhe University ofGeorgla Thesecrossdisciplinary
uppro".h", (ctott = "traverse":Fig. 1-10) frequently resultedin polarization toward
a specificmonodisciplinary concept,a poorly funded administrativeunit, or a nar-
row mission. A crossdisciplinaryapproachalso frequently resultedin polarized fac-
uhy reward systems.Institutions of higher leaming, traditiona\ built on disciplinary
structures, have diffrculties in administering programs and addressi.ngenvironmen-
tal problemsaswell astaking advantageofopponunities at greatertemporal and spa-
tial scales.
= "among') were
To addressthe dilemma, interdisciPlinary approaches(inter
employed,resultingin cooperationon a higher-levelconcept,problem, or question'
t'oi example, the process and study of natural ecological successionprovided a
higher-level concept resulting in the successof the SavannahRiver Ecological Labo-
ratory (SREL)during its conception.Researchers theorized that new systemProper-
ties emerge during the course of ecosystem development and that it is theseproper-
ties that largely account for species and growth form changes that occur (E P Odum
1969,1977, seeChapter 8 for details). Today, interdisciplinary approachesare com-
mon when addressing problems at ecosystem, landscape, and global levels'
Much remainsto be done, however. There is an increased need to solve problems,
promote environmental literacy, and manage resources in a transdisciplinary man-
ner. This multilevel, large-scale approach involves entire educalion and innovation
systems(Fig. I-10). Thil integrativeapproach to the need for unlocking cause-and-
#ect explanations across and among disciplines (achieving a transdisciplinary un-
derstanding) has been termed consilience(E. O Wilson 1998), sustainability scien'e
(Kateset al. 2O0l), and integrativescietLce (Barrett2001) Actually, the continued de-
velopmentofthe scienceofecology (the "study of the household"or''place where we
live';) will likely evolve into that much-needed integrative science of the future This
book attempts to provide the knowledge' concepts, principles, and approachesto
underpin this educationalneed and learning process