Cell Membranes and Signaling: Structure, Transport, 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 Some Substances Can Cross the
Membrane by Diffusion
• 5.3 Some Substances Require Energy to
Cross the Membrane
Chapter 5 Cell Membranes and Signaling
• 5.4 Large Molecules Cross the

Membrane via Vesicles
• 5.5 The Membrane Plays a Key Role in a
Cell’s 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 body’s response to caffeine?
Concept 5.1 Biological Membranes Have a Common Structure
and Are Fluid
A membrane’s structure and functions

are determined by its constituents:
lipids, proteins, and carbohydrates.
The general structure of membranes is

known as the fluid mosaic model.
Phospholipids form a bilayer which is like

a “lake” in which a variety of proteins
“float.”
Figure 5.1 Membrane Molecular Structure
and Are Fluid
Lipids form the hydrophobic core of the

membrane.
Most lipid molecules are phospholipids with two

regions:
• Hydrophilic regions—electrically charged

“heads” that associate with water molecules
• Hydrophobic regions—nonpolar fatty acid

“tails” that do not dissolve in water
and Are Fluid
A bilayer is formed when the fatty acid “tails”

associate with each other and the polar
“heads” face the aqueous environment.
Bilayer organization helps membranes fuse

during vesicle formation and phagocytosis.
and Are Fluid
Membranes may differ in lipid composition as

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
Two important factors in membrane fluidity:
• Lipid composition—types of fatty acids can

increase or decrease fluidity
• Temperature—membrane fluidity decreases

in colder conditions
and Are Fluid
Biological membranes contain proteins, with

varying ratios of phospholipids.
• Peripheral membrane proteins lack

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

embedded in the phospholipid bilayer.
and Are Fluid
Anchored membrane proteins have lipid

components that anchor them in the bilayer.
Proteins are asymmetrically distributed on the

inner and outer membrane surfaces.
A transmembrane protein extends through

the bilayer on both sides, and may have
different functions in its external and
transmembrane domains.
and Are Fluid
Some membrane proteins can move within

the phosopholipid bilayer, while others are
restricted.
Proteins inside the cell can restrict movement

of membrane proteins, as can attachments
to the cytoskeleton.
Figure 5.2 Rapid Diffusion of Membrane Proteins
and Are Fluid
Plasma membrane carbohydrates are

located on the outer membrane and can
serve as recognition sites.
• Glycolipid—a carbohydrate bonded to a

lipid
• Glycoprotein—a carbohydrate bonded

to a protein
and Are Fluid
Membranes are constantly changing by

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 Some Substances Can Cross the Membrane by
Diffusion
Biological membranes allow some

substances, and not others, to pass.
This is known as selective
permeability.
Two processes of transport:
• Passive transport does not require

metabolic energy.
• Active transport requires input of

metabolic energy.
Diffusion
Passive transport of a substance can

occur through two types of diffusion:
• Simple diffusion through the

phospholipid bilayer
• Facilitated diffusion through channel

proteins or aided by carrier proteins
Diffusion
Diffusion is the process of random

movement toward equilibrium.
Speed of diffusion depends on three

factors:
• Diameter of the molecules—smaller

molecules diffuse faster
• Temperature of the solution—higher

temperatures lead to faster diffusion
Diffusion
• The concentration gradient in the

system—the greater the concentration
gradient in a system, the faster a
substance will diffuse
A higher concentration inside the cell

causes the solute to diffuse out, and a
higher concentration outside causes the
solute to diffuse in, for many molecules.
Diffusion
Simple diffusion takes place through the

phospholipid bilayer.
A molecule that is hydrophobic and

soluble in lipids can pass through the
membrane.
Polar molecules do not pass through—

they are not soluble in the hydrophilic
interior and form bonds instead in the
aqueous environment near the
membrane.
Diffusion
Osmosis is the diffusion of water across

membranes.
It depends on the concentration of solute

molecules on either side of the
membrane.
Water passes through special membrane

channels.
Diffusion
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 5.3A Osmosis Can Modify the Shapes of Cells
Figure 5.3B Osmosis Can Modify the Shapes of Cells
Figure 5.3C Osmosis Can Modify the Shapes of Cells
Diffusion
The concentration of solutes in the

environment determines the direction of
osmosis in all animal cells.
In other organisms, cell walls limit the

volume that can be taken up.
Turgor pressure is the internal pressure

against the cell wall—as it builds up, it
prevents more water from entering.
Diffusion
Diffusion may be aided by channel

proteins.
Channel proteins are integral membrane

proteins that form channels across the
membrane.
Substances can also bind to carrier

proteins to speed up diffusion.
Both are forms of facilitated diffusion.

Diffusion
Ion channels are a type of channel

protein—most are gated, and can be
opened or closed to ion passage.
A gated channel opens when a stimulus

causes the channel to change shape.
The stimulus may be a ligand, a

chemical signal.
Diffusion
A ligand-gated channel responds to its

ligand.
A voltage-gated channel opens or closes

in response to a change in the voltage
across the membrane.
Figure 5.4 A Ligand-Gated Channel Protein Opens in Response to a Stimulus
Diffusion
Water crosses membranes at a faster

rate than simple diffusion.
It may “hitchhike” with ions such as Na+

as they pass through channels.
Aquaporins are specific 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)
Diffusion
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
shape—it releases glucose on the other
side of the membrane.
Figure 5.6 A Carrier Protein Facilitates Diffusion (Part 1)
Figure 5.6 A Carrier Protein Facilitates Diffusion (Part 2)
Diffusion
Transport by carrier proteins differs from

simple diffusion, though both are driven
by the concentration gradient.
The facilitated diffusion system can

become saturated—when all of the
carrier molecules are bound, the rate of
diffusion reaches its maximum.
Concept 5.3 Some Substances Require Energy to Cross the
Membrane
Active transport requires the input of

energy to move substances against their
concentration gradients.
Active transport is used to overcome

concentration imbalances that are
maintained by proteins in the
membrane.
Table 5.1 Membrane Transport Mechanisms
Membrane
The energy source for active transport is

often ATP.
Active transport is directional and moves

a substance against its concentration
gradient.
A substance moves in the direction of the

cell’s needs, usually by means of a
specific carrier protein.
Membrane
Two types of active transport:
• Primary active transport involves

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

energy from an ion concentration
gradient, or an electrical gradient.
Membrane
The sodium–potassium (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 Sodium–Potassium Pump
Membrane
Secondary active transport uses energy

that is “regained,” by letting ions move
across the membrane with their
concentration gradients.
Secondary active transport may begin

with passive diffusion of a few ions, or
may involve a carrier protein that
transports both a substance and ions.
Concept 5.4 Large Molecules Cross the Membrane via Vesicles
Macromolecules are too large or too

charged to pass through biological
membranes and instead pass through
vesicles.
To take up or to secrete macromolecules,

cells must use endocytosis or
exocytosis.
Figure 5.8 Endocytosis and Exocytosis (Part 1)
Figure 5.8 Endocytosis and Exocytosis (Part 2)
Three types of endocytosis brings

molecules into the cell: phagocytosis,
pinocytosis, and receptor–mediated
endocytosis.
In all three, the membrane invaginates, or

folds around the molecules and forms a
vesicle.
The vesicle then separates from the

membrane.
In phagocytosis (“cellular eating”), part

of the membrane engulfs a large particle
or cell.
A food vacuole (phagosome) forms and

usually fuses with a lysosome, where
contents are digested.
In pinocytosis (“cellular drinking”),

vesicles also form.
The vesicles are smaller and bring in

fluids and dissolved substances, as in
the endothelium near blood vessels.
Receptor–mediated endocytosis
depends on receptors to bind to
specific molecules (their ligands).
The receptors are integral membrane

proteins located in regions called coated
pits.
The cytoplasmic surface is coated by

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

coated pit invaginates and forms a
coated vesicle.
The clathrin stabilizes the vesicle as it

carries the macromolecules into the
cytoplasm.
Once inside, the vesicle loses its clathrin

coat and the substance is digested.
Figure 5.9 Receptor-Mediated Endocytosis (Part 1)
Figure 5.9 Receptor-Mediated Endocytosis (Part 2)
Exocytosis moves materials out of the

cell in vesicles.
The vesicle membrane fuses with the

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

substances made in the cell.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals
Cells can respond to many signals if they

have a specific receptor for that signal.
A signal transduction pathway is a

sequence of molecular events and
chemical reactions that lead to a cellular
response, following the receptor’s
activation by a signal.
Cells are exposed to many signals and

may have different responses:
• Autocrine signals affect the same cells

that release them.
• Paracrine signals diffuse to and affect

nearby cells.
• Hormones travel to distant cells.

Figure 5.10 Chemical Signaling Concepts
Only cells with the necessary receptors

can respond to a signal—the 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
A common mechanism of signal

transduction is allosteric regulation.
This involves an alteration in a protein’s

shape as a result of a molecule binding
to it.
A signal transduction pathway may

produce short or long term responses.
A signal molecule, or ligand, fits into a

three-dimensional site on the receptor
protein.
Binding of the ligand causes the receptor

to change its three-dimensional shape.
The change in shape initiates a cellular

response.
Figure 5.12 A Signal Binds to Its Receptor
Ligands are generally not metabolized

further, but their binding may expose an
active site on the receptor.
Binding is reversible and the ligand can

be released, to end stimulation.
An inhibitor, or antagonist, can bind in

place of the normal ligand.
Receptors can be classified by their

location in the cell.
This is determined by whether or not their

ligand can diffuse through the
membrane.
Cytoplasmic receptors have ligands, such

as estrogen, that are small or nonpolar
and can diffuse across the membrane.
Membrane receptors have large or polar

ligands, such as insulin, that cannot
diffuse and must bind to a
transmembrane receptor at an
extracellular site.
Receptors are also classified by their

activity:
• Ion channel receptors
• Protein kinase receptors
• G protein–linked receptors
Ion channel receptors, or gated ion

channels, change their three-
dimensional shape when a ligand binds.
The acetylcholine receptor, a ligand-

gated sodium channel, binds
acetylcholine to open the channel and
allow Na+ to diffuse into the cell.
Protein kinase receptors change their

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

cytoplasmic domain that has catalytic
(protein kinase) activity.
Figure 5.13 A Protein Kinase Receptor
Protein kinases catalyze the following

reaction:
ATP + protein  ADP + phosphorylated

protein
Each protein kinase has a specific target

protein, whose activity is changed when
it is phosphorylated.
Ligands binding to G protein–linked

receptors expose a site that can bind to
a 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 molecules:
• The receptor
• GDP and GTP, used for energy transfer
• An effector protein to cause an effect in

the cell
The activated G protein–linked 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 Protein–Linked 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, which is
mediated by a signal transduction
pathway.
Signaling can initiate a cascade of protein

interactions—the signal can then be
amplified and distributed to cause
different responses.
Environment
A second messenger is an intermediary

between the receptor and the cascade
of responses.
In the fight-or-flight response, epinephrine

(adrenaline) activates the liver enzyme
glycogen phosphorylase.
The enzyme catalyzes the breakdown of

glycogen to provide quick energy.
Environment
Researchers found that the cytoplasmic

enzyme could be activated by the
membrane-bound epinephrine in broken
cells, as long as all parts were present.
They discovered that another molecule

delivered the message from the “first
messenger,” epinephrine, to the
enzyme.
Figure 5.15 The Discovery of a Second Messenger (Part 1)
Figure 5.15 The Discovery of a Second Messenger (Part 2)
Environment
The second messenger was later

discovered to be cyclic AMP (cAMP).
Second messengers 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
Environment
Signal transduction pathways involve

multiple steps—enzymes may be either
activated or inhibited by other enzymes.
In liver cells, a signal cascade begins

when epinephrine stimulates a G
protein–mediated protein kinase
pathway.
Environment
Epinephrine binds to its receptor and

activates a G protein.
cAMP is produced and activates protein

kinase A—it phosphorylates two other
enzymes, with opposite effects:
• Inhibition
• Activation
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)
Environment
• Inhibition—protein kinase A inactivates

glycogen synthase through
phosphorylation, and prevents glucose
storage.
• Activation—Phosphorylase kinase is
activated when phosphorylated and is
part of a cascade that results in the
liberation of glucose molecules.
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 cell’s response.
Figure 5.18 Signal Transduction Regulatory Mechanisms
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
Environment
Cell functions change in response to

environmental signals:
• Opening of ion channels
• Alterations in gene expression
• Alteration of enzyme activities

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 it—the result is that
the person remains alert.
Figure 5.19 Caffeine and the Cell Membrane (Part 1)
Figure 5.19 Caffeine and the Cell Membrane (Part 2)

Cell Membranes and Signaling: Structure, Transport, and Signaling

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5 Cell Membranes and

• 5.4 Large Molecules Cross the

What role does the cell membrane play in

A membrane’s structure and functions

The general structure of membranes is

Phospholipids form a bilayer which is like

Lipids form the hydrophobic core of the

Most lipid molecules are phospholipids with two

• Hydrophilic regions—electrically charged

• Hydrophobic regions—nonpolar fatty acid

A bilayer is formed when the fatty acid “tails”

Bilayer organization helps membranes fuse

Membranes may differ in lipid composition as

Phospholipids may differ in:

• Fatty acid chain length

• Kinds of polar groups present

Two important factors in membrane fluidity:

• Lipid composition—types of fatty acids can

• Temperature—membrane fluidity decreases

Biological membranes contain proteins, with

• Peripheral membrane proteins lack

• Integral membrane proteins are partly

Anchored membrane proteins have lipid

Proteins are asymmetrically distributed on the

A transmembrane protein extends through

Some membrane proteins can move within

Proteins inside the cell can restrict movement

Plasma membrane carbohydrates are

• Glycolipid—a carbohydrate bonded to a

• Glycoprotein—a carbohydrate bonded

Membranes are constantly changing by

Though membranes appear similar, there

Biological membranes allow some

Two processes of transport:

• Passive transport does not require

• Active transport requires input of

Passive transport of a substance can

• Simple diffusion through the

• Facilitated diffusion through channel

Diffusion is the process of random

Speed of diffusion depends on three

• Diameter of the molecules—smaller

• Temperature of the solution—higher

• The concentration gradient in the

A higher concentration inside the cell

Simple diffusion takes place through the

A molecule that is hydrophobic and

Polar molecules do not pass through—

Osmosis is the diffusion of water across

It depends on the concentration of solute

Water passes through special membrane

When comparing two solutions separated

• A hypertonic solution has a higher

• Isotonic solutions have equal solute

• A hypotonic solution has a lower solute

The concentration of solutes in the

In other organisms, cell walls limit the

Turgor pressure is the internal pressure

Diffusion may be aided by channel

Channel proteins are integral membrane

Substances can also bind to carrier

Both are forms of facilitated diffusion.