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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 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
Concept 5.1 Biological Membranes Have a Common 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
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.

Bilayer organization helps membranes fuse


during vesicle formation and phagocytosis.
Concept 5.1 Biological Membranes Have a Common Structure
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


Concept 5.1 Biological Membranes Have a Common Structure
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
Concept 5.1 Biological Membranes Have a Common Structure
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.
Concept 5.1 Biological Membranes Have a Common Structure
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.
Concept 5.1 Biological Membranes Have a Common Structure
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
Concept 5.1 Biological Membranes Have a Common Structure
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
Concept 5.1 Biological Membranes Have a Common Structure
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.
Concept 5.2 Some Substances Can Cross the Membrane by
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
Concept 5.2 Some Substances Can Cross the Membrane by
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
Concept 5.2 Some Substances Can Cross the Membrane by
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.
Concept 5.2 Some Substances Can Cross the Membrane by
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.
Concept 5.2 Some Substances Can Cross the Membrane by
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.
Concept 5.2 Some Substances Can Cross the Membrane by
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
Concept 5.2 Some Substances Can Cross the Membrane by
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.
Concept 5.2 Some Substances Can Cross the Membrane by
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.


Concept 5.2 Some Substances Can Cross the Membrane by
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.
Concept 5.2 Some Substances Can Cross the Membrane by
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
Concept 5.2 Some Substances Can Cross the Membrane by
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)
Concept 5.2 Some Substances Can Cross the Membrane by
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)
Concept 5.2 Some Substances Can Cross the Membrane by
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
Concept 5.3 Some Substances Require Energy to Cross the
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.
Concept 5.3 Some Substances Require Energy to Cross the
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.
Concept 5.3 Some Substances Require Energy to Cross the
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
Concept 5.3 Some Substances Require Energy to Cross the
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)
Concept 5.4 Large Molecules Cross the Membrane via Vesicles

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.
Concept 5.4 Large Molecules Cross the Membrane via Vesicles

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.
Concept 5.4 Large Molecules Cross the Membrane via Vesicles

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.
Concept 5.4 Large Molecules Cross the Membrane via Vesicles

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).
Concept 5.4 Large Molecules Cross the Membrane via Vesicles

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)
Concept 5.4 Large Molecules Cross the Membrane via Vesicles

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

Receptors can be classified by their


location in the cell.

This is determined by whether or not their


ligand can diffuse through the
membrane.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

Receptors are also classified by their


activity:

• Ion channel receptors

• Protein kinase receptors

• G protein–linked receptors
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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
Concept 5.5 The Membrane Plays a Key Role in a Cell’s
Response to Environmental Signals

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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to Its
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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to Its
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)
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 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—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.
Concept 5.6 Signal Transduction Allows the Cell to Respond to Its
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)
Concept 5.6 Signal Transduction Allows the Cell to Respond to Its
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.
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 cell’s 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
Concept 5.6 Signal Transduction Allows the Cell to Respond to Its
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)