PoL2e Ch05a Lecture-Cell Membranes and Signaling

5 Cell Membranes and
Signaling
Chapter 5 Cell Membranes and Signaling
Key Concepts
5.1 Biological Membranes Have a Common
Structure and Are Fluid
5.2 Passive Transport across Membranes
Requires No Input of Energy
5.3 Active Transport Moves Solutes against Their
Concentration Gradients
5.4 Large Molecules Cross Membranes via
Vesicles
Chapter 5 Cell Membranes and Signaling
5.5 The Membrane Plays a Key Role in a Cells

Response to Environmental Signals
5.6 Signal Transduction Allows the Cell to
Respond to Its Environment
Chapter 5 Opening Question
What role does the cell membrane play in the

bodys response to caffeine?
Concept 5.1 Biological Membranes Have a Common Structure
and Are Fluid
A membranes structure and functions are

determined by its constituents: lipids,
proteins, and carbohydrates.
The general design of membranes is known as

the fluid mosaic model.
Phospholipids form a continuous bilayer which

is like a lake in which a variety of proteins
float.
Figure 5.1 Membrane Structure
and Are Fluid
The lipid molecules are usually phospholipids

with two regions:
Hydrophilic regionselectrically charged
heads associate with water molecules
Hydrophobic regionsnonpolar fatty acid
tails that do not dissolve in water
Phospholipids
and Are Fluid
A bilayer is formed when the fatty acid tails

associate with each other and the polar heads
face the aqueous environment.
and Are Fluid
Membranes may differ in lipid composition; there

are many types of phospholipids.
Phospholipids may differ in:
Fatty acid chain length
Degree of saturation
Kinds of polar groups present

and Are Fluid
Cholesterol is an important component of animal

cell membranes.
Hydroxyl groups interact with the polar heads of

phospholipids.
Cholesterol is important in modulating

membrane fluidity.
In-Text Art, Chapter 5, p. 84 (2)
and Are Fluid
The fatty acids make the membrane somewhat

fluid. This allows some molecules to move
laterally within the membrane.
Membrane fluidity is influenced by:

Lipid compositionshort, unsaturated
chains increase fluidity
Temperaturefluidity decreases in colder
conditions
and Are Fluid
All biological membranes contain proteins; the

ratio of proteins to phospholipids varies.
Peripheral membrane proteins lack

hydrophobic groups and are not embedded in
the bilayer.
Integral membrane proteins are at least partly

embedded in the phospholipid bilayer.
In-Text Art, Chapter 5, p. 85
and Are Fluid
Anchored membrane proteins have hydrophobic

lipid components that anchor them in the
bilayer.
Transmembrane proteins extend through the

bilayer; they may have domains with different
functions on the inner and outer sides of the
membrane.
and Are Fluid
Some membrane proteins can move within the

phosopholipid bilayer; others are restricted.
Cell fusion experiments illustrate this
migration.
Proteins inside the cell can restrict movement of

membrane proteins, as can attachments to the
cytoskeleton.
Figure 5.2 Rapid Diffusion of Membrane Proteins (Part 1)
Figure 5.2 Rapid Diffusion of Membrane Proteins (Part 2)
and Are Fluid
Diverse carbohydrates are located on the outer

cell membrane and play a role in
communication.
Glycolipidcarbohydrate covalently
bonded to a lipid
Glycoproteinone or more
oligosaccharides covalently bonded to a
protein
Proteoglycanprotein with more and
longer carbohydrates bonded to it
and Are Fluid
Cells can adhere to one another through

interactions between cell surface
carbohydrates and proteins.
and Are Fluid
Membranes are constantly forming, transforming

into other types, fusing, and breaking down.
Though membranes appear similar, there are

major chemical differences among the
membranes of even a single cell.
Concept 5.2 Passive Transport across Membranes Requires
No Input of Energy
Selective permeability: biological membranes

allow some substances, but not others, to
pass
Two processes of transport across membranes:

1. Passive transport - does not require
metabolic energy.
A substance moves down its
concentration gradient.
Concept 5.2 Passive Transport across Membranes Requires
No Input of Energy
2. Active transport - requires input of

metabolic energy.
A substance moves against its
Concept 5.2 Passive Transport across Membranes Requires No
Input of Energy
Passive transport can occur by:

Simple diffusion through the phospholipid
bilayer
Facilitated diffusion through channel
proteins or aided by carrier proteins
Input of Energy
Diffusion is the process of random movement

toward equilibrium; a net movement from
regions of greater concentration to regions of
lesser concentration.
Input of Energy
Speed of diffusion depends on three factors:

Diameter of the moleculessmaller
molecules diffuse faster.
Temperature of the solutionhigher
temperatures lead to faster diffusion.
Concentration gradientthe greater the
concentration gradient, the faster a
substance will diffuse.
Input of Energy
Cell cytoplasm is an aqueous solution, as is the

surrounding environment.
A higher concentration inside the cell causes

the solute to diffuse out; higher concentration
outside causes the solute to diffuse in.
Input of Energy
Some molecules cross the phospholipid bilayer

by simple diffusion:
O2, CO2, and small, nonpolar, lipid-soluble
molecules.
Polar (hydrophilic) molecules do not pass

throughthey are not soluble in the
hydrophobic interior of the membrane.
Amino acids, sugars, ions, water
Input of Energy
Osmosis is the diffusion of water across

membranes through special channels.
It depends on the concentration of water

molecules on either side of the membrane
water moves down its concentration gradient.
The higher the total solute concentration, the

lower the concentration of water molecules.
Input of Energy
When comparing two solutions separated by a

membrane:
A hypertonic solution has a higher solute
concentration.
Isotonic solutions have equal solute
concentrations.
A hypotonic solution has a lower solute
concentration.
Figure 5.3 Osmosis Can Modify the Shapes of Cells
Input of Energy
Facilitated diffusion:
Channel proteins are integral membrane

proteins that form channels across the
membrane through which some substances
can pass.
Substances can also bind to carrier proteins

to speed up diffusion.
Both processes operate in either direction.

Input of Energy
Ion channels: channel proteins that allow

specific ions to pass through
Most are gated channelsthey open when a

stimulus causes the protein to change shape.
Ligand-gatedthe stimulus is a ligand, a
chemical signal.
Voltage-gatedthe stimulus is a change in
electrical charge difference across the
membrane.
Figure 5.4 A Ligand-Gated Channel Protein Opens in Response to a Stimulus
Input of Energy
Water crosses membranes at a faster rate than

simple diffusion.
It may hitchhike with ions such as Na+ as they

pass through ion channels.
Aquaporins are channels that allow large

amounts of water to move along its
Figure 5.5 Aquaporins Increase Membrane Permeability to Water (Part 1)
Figure 5.5 Aquaporins Increase Membrane Permeability to Water (Part 2)
Input of Energy
Carrier proteins in the membrane facilitate

diffusion by binding substances.
Glucose transporters are carrier proteins in

mammalian cells.
Glucose molecules bind to the carrier protein

and cause the protein to change shapeit
releases glucose on the other side of the
membrane.
Figure 5.6 A Carrier Protein Facilitates Diffusion (Part 1)
Input of Energy
Glucose is quickly broken down in the cell, so

there is always a strong concentration
gradient that favors glucose uptake.
But the system can become saturatedwhen

all of the carrier molecules are bound, the rate
of diffusion reaches a maximum.
Figure 5.6 A Carrier Protein Facilitates Diffusion (Part 2)
Concept 5.3 Active Transport Moves Solutes against Their
Cells maintain an internal environment with a

different composition than the outside
environment.
This requires workenergy from ATP is needed

to move substances against their
concentration gradients (active transport).
Specific carrier proteins move substances in

only one direction, either into or out of the cell.
Table 5.1
Two types of active transport:
Primary active transport involves direct

hydrolysis of ATP for energy.
Secondary active transport uses the energy

from an ion concentration gradient or an
electrical gradient. The gradients are
established by primary active transport.
The sodiumpotassium (Na+K+) pump is an

integral membrane protein that pumps Na + out
of a cell and K+ in.
One molecule of ATP moves two K+ and three

Na+ ions.
Figure 5.7 Primary Active Transport: The SodiumPotassium Pump
Secondary active transport uses energy that is

regained by letting ions move across the
membrane with their concentration gradients.
Example: after the Na+K+ pump
establishes a concentration gradient of
Na+, then passive diffusion of Na+ back into
the cell can provide energy for glucose
transport.
One protein usually moves both the ion and the

transported molecule across the membrane.
Secondary Active Transport
Concept 5.4 Large Molecules Cross Membranes via Vesicles
Macromolecules are too large or too charged to

pass through biological membranes, so
instead they cross within vesicles.
To take up or to secrete macromolecules, cells

must use endocytosis and exocytosis.
Exocytosis moves materials out of the cell in

vesicles.
The vesicle membrane fuses with the cell

membrane and the contents are released into
the environment.
Exocytosis is important in the secretion of

substances made by cells such as digestive
enzymes and neurotransmitters.
Endocytosis brings macromolecules and

particles into eukaryotic cells.
The cell membrane invaginates, or folds

around the particle and forms a vesicle.
The vesicle then separates from the

membrane.
Figure 5.8 Endocytosis and Exocytosis
Endocytosis depends on receptorsproteins

that bind to specific molecules (ligands).
The receptors are integral membrane proteins

on the cell membrane.
The resulting vesicle includes both the receptor

and its ligand, plus other substances present
near the site of invagination.
Phagocytosis (cellular eating): a specialized

cell engulfs a large particle or another cell
A food vesicle (phagosome) forms and
usually fuses with a lysosome, where the
contents are digested.
Pinocytosis (cellular drinking): vesicles are

smaller and bring in fluids and dissolved
substances
Receptor endocytosis brings specific large

molecules into a cell via specific receptors.
This allows cells to control internal processes

by controlling location and abundance of each
type of receptor on the cell membrane.
It also plays a role in cell signaling.

The receptors are located in membrane regions

called coated pits.
The cytoplasmic surface of a pit is coated by

another protein (often clathrin).
When receptors bind to their ligands, the

coated pit invaginates and forms a coated
vesicle.
Clathrin stabilizes the vesicle.

Figure 5.9 Receptor Endocytosis
Once inside, the vesicle loses its clathrin coat

and fuses with a membrane-enclosed
compartment called an endosome.
Receptors may be recycled to the cell

membrane or degraded in a lysosome. This is
an important mechanism for controlling the
abundance of each kind of receptor on the
cell surface.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Cell signaling: cells can process information

from their environment
Signals include physical stimuli, such as heat or

light, and chemicals (ligands). The cell must
have a receptor for the signal in order to
respond.
Following receptor activation by a signal, a signal

transduction pathway is initiateda sequence
of events that lead to a cellular response.
In a multicellular animal, cells are exposed to

many chemical signals:
Autocrine signals affect the same cells that
release them.
Paracrine signals diffuse to and affect
nearby cells.
Juxtacrine signaling requires direct contact
between the signaling and responding cell.
Hormones travel to distant cells.
Figure 5.10 Chemical Signaling Concepts
Only cells with the necessary receptors can

respond to a signalthe target cell must be
able to sense it and respond to it.
A signal transduction pathway involves a

signal, a receptor, and a response.
Figure 5.11 Signal Transduction Concepts
Signal transduction pathways often include

allosteric regulation:
Protein shape changes as a result of a
molecule binding at a site other than the
active site (e.g., a ligand-gated channel).
A signal transduction pathway may produce

short or long term responses.
Receptors can be classified by their location:

Intracellular receptors are located inside
a cell. Their ligands are small or nonpolar
and can diffuse across the membrane.
Membrane receptors located on the cell
surface have large or polar ligands that
cannot diffuse through the membrane.
Membrane receptors:
A chemical ligand fits into a 3-D site on the
receptor protein.
The receptor may have a catalytic domain
on the cytoplasmic side. The ligand is an
allosteric regulatorit exposes the active
site on the catalytic domain.
Figure 5.12 A Signal Binds to Its Receptor
Ligand-receptor binding is noncovalent and

reversible.
Reversible binding is important because cells

need to stop responding to a signal after the
appropriate response has occurred.
Inhibitors, or antagonists, can bind in place of

the normal ligand.
Caffeine binds to receptors in the brain,
preventing binding by the normal ligands.
Ion channel receptors are ligand-gated ion

channels; they change shape when a ligand
binds.
Acetylcholine receptors on skeletal muscle
cells bind acetylcholine to open the
channel and allow Na+ to diffuse into the
cell.
Protein kinase receptors also change shape

when a ligand binds.
The new shape exposes or activates a

cytoplasmic domain that has protein kinase
activityit modifies proteins by adding
phosphate groups.
(Not all protein kinases are receptors.)

Figure 5.13 A Protein Kinase Receptor
G proteincoupled receptors: ligand binding

on the surface exposes a site on the
cytoplasmic side that binds to a mobile
membrane protein, a G protein
The G protein is partially inserted in the lipid

bilayer and partially exposed on the
cytoplasmic surface.
Many G proteins have three subunits and can

bind three different molecules:
The receptor
GDP and GTP, used for energy transfer
An effector protein that causes an effect in
the cell
The activated G proteincoupled receptor

exchanges a GDP nucleotide bound to the G
protein for a higher energy GTP.
The activated G protein activates the effector

protein, leading to signal amplification.
Figure 5.14 A G ProteinLinked Receptor
Concept 5.6 Signal Transduction Allows the Cell to Respond to
Its Environment
Signal activation of a specific receptor leads to

a cellular response, mediated by a signal
transduction pathway.
Signaling can initiate a cascade of protein

interactionsthe initial signal is amplified and
distributed to cause different responses,
ultimately leading to changes in cell function.
Its Environment
There are many ways in which cells respond to

environmental signals:
Opening of ion channelschanges the
balance of ion concentrations between the
outside and inside of the cell and results in
change in the electrical potential across
the membrane.
Its Environment
Alterations in gene expressiongenes

may be switched on (upregulated) or
switched off (downregulated).
Alteration of enzyme activitiesmore rapid

response than those involving change in
gene expression.
Its Environment
The same signal can lead to different

responses in different types of cells.
Example: Heart and digestive tract muscle cells

respond differently to epinephrine (adrenaline)
because the signal transduction pathways
stimulated are different in the two cell types.
Its Environment
Often there is a small molecule intermediary, a

second messenger, between the activated
receptor and the cascade of responses that
ensues.
In the fight-or-flight response, epinephrine

(adrenaline) activates the liver enzyme
glycogen phosphorylase, which catalyzes
breakdown of glycogen for quick energy.
Its Environment
The second messenger was later discovered to be

cyclic AMP (cAMP).
Second messengers regulate target enzymes by

binding to them noncovalently.
They allow the cell to respond to a single

membrane event with many events inside the cell
they distribute the signal.
They amplify the signal by activating more than

one enzyme target.
Figure 5.16 The Formation of Cyclic AMP
Its Environment
Signal transduction pathways involve multiple

steps in which enzymes are either activated
or inhibited by other enzymes.
In liver cells, a signal cascade begins when

epinephrine stimulates a G proteinmediated
protein kinase pathway.
Its Environment
cAMP is produced and activates protein kinase

A, which phosphorylates two other enzymes,
with opposite effects:
Inhibitionglycogen synthase is
inactivated by phosphorylation, which
prevents glucose storage.
Activationphosphorylase kinase is
phosphorylated and starts a cascade that
results in the liberation of glucose
molecules from glycogen.
Figure 5.17 A Cascade of Reactions Leads to Altered Enzyme Activity (Part 1)
Figure 5.17 A Cascade of Reactions Leads to Altered Enzyme Activity (Part 2)
Its Environment
The original signal is amplified at every step in

the cascade.
Each molecule of epinephrine that arrives at

the cell membrane ultimately results in 10,000
molecules of blood glucose.
Its Environment
Signal transduction ends after the cell responds

enzymes convert each transducer back to
its inactive precursor.
The balance between the regulating enzymes

and the signal enzymes determines the cells
ultimate response.
Figure 5.18 Signal Transduction Regulatory Mechanisms
Its Environment
Cells can alter the balance of enzymes in two

ways:
Synthesis or breakdown of the enzyme
Activation or inhibition of the enzymes by
other molecules
Answer to Opening Question
Caffeine is a large, polar molecule that binds to

receptors on nerve cells in the brain.
Its structure is similar to adenosine, which

binds to receptors after activity or stress and
results in drowsiness.
Caffeine binds to the same receptor, but does

not activate itthe result is that the person
remains alert.
Figure 5.19 Caffeine and the Cell Membrane

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5 Cell Membranes and

5.5 The Membrane Plays a Key Role in a Cells

What role does the cell membrane play in the

A membranes structure and functions are

The general design of membranes is known as

Phospholipids form a continuous bilayer which

The lipid molecules are usually phospholipids

A bilayer is formed when the fatty acid tails

Membranes may differ in lipid composition; there

Phospholipids may differ in:

Fatty acid chain length

Kinds of polar groups present

Cholesterol is an important component of animal

Hydroxyl groups interact with the polar heads of

Cholesterol is important in modulating

The fatty acids make the membrane somewhat

Membrane fluidity is influenced by:

All biological membranes contain proteins; the

Peripheral membrane proteins lack

Integral membrane proteins are at least partly

Anchored membrane proteins have hydrophobic

Transmembrane proteins extend through the

Some membrane proteins can move within the

Proteins inside the cell can restrict movement of

Diverse carbohydrates are located on the outer

Cells can adhere to one another through

Membranes are constantly forming, transforming

Though membranes appear similar, there are

Selective permeability: biological membranes

Two processes of transport across membranes:

2. Active transport - requires input of

Passive transport can occur by:

Diffusion is the process of random movement

Speed of diffusion depends on three factors:

Cell cytoplasm is an aqueous solution, as is the

A higher concentration inside the cell causes

Some molecules cross the phospholipid bilayer

Polar (hydrophilic) molecules do not pass

Osmosis is the diffusion of water across

It depends on the concentration of water

The higher the total solute concentration, the

When comparing two solutions separated by a

Channel proteins are integral membrane

Substances can also bind to carrier proteins

Both processes operate in either direction.

Ion channels: channel proteins that allow

Most are gated channelsthey open when a

Water crosses membranes at a faster rate than

It may hitchhike with ions such as Na+ as they

Aquaporins are channels that allow large

Carrier proteins in the membrane facilitate

Glucose transporters are carrier proteins in

Glucose molecules bind to the carrier protein

Glucose is quickly broken down in the cell, so

But the system can become saturatedwhen

Cells maintain an internal environment with a

This requires workenergy from ATP is needed

Specific carrier proteins move substances in

Two types of active transport:

Primary active transport involves direct

Secondary active transport uses the energy

The sodiumpotassium (Na+K+) pump is an

One molecule of ATP moves two K+ and three

Secondary active transport uses energy that is