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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
Concept 5.1 Biological Membranes Have a Common 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
Concept 5.1 Biological Membranes Have a Common Structure
and Are Fluid

A bilayer is formed when the fatty acid tails


associate with each other and the polar heads
face the aqueous environment.
Concept 5.1 Biological Membranes Have a Common Structure
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


Concept 5.1 Biological Membranes Have a Common Structure
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)
Concept 5.1 Biological Membranes Have a Common Structure
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
Concept 5.1 Biological Membranes Have a Common Structure
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
Concept 5.1 Biological Membranes Have a Common Structure
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.
Concept 5.1 Biological Membranes Have a Common Structure
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)
Concept 5.1 Biological Membranes Have a Common Structure
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
Concept 5.1 Biological Membranes Have a Common Structure
and Are Fluid

Cells can adhere to one another through


interactions between cell surface
carbohydrates and proteins.
Concept 5.1 Biological Membranes Have a Common Structure
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
concentration gradient.
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
Concept 5.2 Passive Transport across Membranes Requires No
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.
Concept 5.2 Passive Transport across Membranes Requires No
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.
Concept 5.2 Passive Transport across Membranes Requires No
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.
Concept 5.2 Passive Transport across Membranes Requires No
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
Concept 5.2 Passive Transport across Membranes Requires No
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.
Concept 5.2 Passive Transport across Membranes Requires No
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
Concept 5.2 Passive Transport across Membranes Requires No
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.


Concept 5.2 Passive Transport across Membranes Requires No
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
Concept 5.2 Passive Transport across Membranes Requires No
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
concentration gradient.
Figure 5.5 Aquaporins Increase Membrane Permeability to Water (Part 1)
Figure 5.5 Aquaporins Increase Membrane Permeability to Water (Part 2)
Concept 5.2 Passive Transport across Membranes Requires No
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)
Concept 5.2 Passive Transport across Membranes Requires No
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
Concentration Gradients

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
Concept 5.3 Active Transport Moves Solutes against Their
Concentration Gradients

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.
Concept 5.3 Active Transport Moves Solutes against Their
Concentration Gradients

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
Concept 5.3 Active Transport Moves Solutes against Their
Concentration Gradients

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.
Concept 5.4 Large Molecules Cross Membranes via Vesicles

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.
Concept 5.4 Large Molecules Cross Membranes via Vesicles

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
Concept 5.4 Large Molecules Cross Membranes via Vesicles

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.
Concept 5.4 Large Molecules Cross Membranes via Vesicles

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
Concept 5.4 Large Molecules Cross Membranes via Vesicles

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.


Concept 5.4 Large Molecules Cross Membranes via Vesicles

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
Concept 5.4 Large Molecules Cross Membranes via Vesicles

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
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

Ligand-receptor binding is noncovalent and


reversible.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cells
Response to Environmental Signals

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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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)
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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
Concept 5.6 Signal Transduction Allows the Cell to Respond to
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