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© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Chapter 7
Membrane Structure
and Function
Dr. Wendy Sera
Houston Community
College
Biology 1406
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Key Concepts in Chapter 7
1. Cellular membranes are fluid mosaics of lipids and
proteins.
2. Membrane structure results in selective
permeability
3. Passive transport is diffusion of a substance across
a membrane with no energy investment
4. Active transport uses energy to move solutes
against their gradients.
5. Bulk transport across the plasma membrane
occurs by exocytosis and endocytosis.
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Life at the Edge
The plasma membrane is the boundary that
separates the living cell from its surroundings
The plasma membrane exhibits selective
permeability, allowing some substances to cross it
more easily than others
Figure 7.1—How do cell
membrane proteins help
regulate chemical traffic?
2
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Video: Structure of the Cell Membrane
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Concept 7.1: Cellular membranes are fluid mosaics of lipids and proteins
Phospholipids are the most abundant lipid in the
plasma membrane
Phospholipids are amphipathic molecules,
containing both hydrophobic and hydrophilic
regions
A phospholipid bilayer can exist as a stable
boundary between two aqueous compartments
© 2014 Pearson Education, Inc.
Hydrophilic head
Hydrophobic tail
WATER
WATER
Figure 7.2—Phospholipid bilayer (cross section)
3
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Membrane Models: Scientific Inquiry
Membranes have been chemically analyzed and
found to be made of proteins and lipids
Scientists studying the plasma membrane
reasoned that it must be a phospholipid bilayer
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
In 1935, Davson and Danielli proposed a
sandwich model in which the phospholipid bilayer
lies between two layers of globular proteins
Later studies found problems with this model,
particularly the placement of membrane proteins,
which have hydrophilic and hydrophobic regions
In 1972, Singer and Nicolson proposed that the
membrane is a mosaic of proteins dispersed within
the bilayer, with only the hydrophilic regions
exposed to water—called the fluid-mosaic model
The Sandwich Model vs. the Fluid-Mosaic Model
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Phospholipid
bilayer
Hydrophobic regionsof protein
Hydrophilicregions of protein
Figure 7.3—The original fluid mosaic model for membranes
4
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
TECHNIQUE
Extracellularlayer
Knife
Proteins Inside of extracellular layer
RESULTS
Inside of cytoplasmic layerCytoplasmic layerPlasma membrane
• Freeze-fracture studies of the plasma membrane supported the
fluid mosaic model
• Freeze-fracture is a specialized preparation technique that splits
a membrane along the middle of the phospholipid bilayer
Freeze-fracture Studies
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
The Fluidity of Membranes
Phospholipids in the plasma membrane can
move within the bilayer
Most of the lipids, and some proteins, drift
laterally
Rarely, a lipid may flip-flop transversely across
the membrane
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Lateral movement occurs
107 times per second.
Flip-flopping across the membrane
is rare ( once per month).
The fluidity of membranes: The movement of
phospholipids
5
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Membrane proteins
Mouse cellHuman cell
Hybrid cell
Mixed proteins
after 1 hour
Figure 7.4-3—Inquiry: Do membrane proteins move?
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
As temperatures cool, membranes switch from a
fluid state to a solid state
The temperature at which a membrane solidifies
depends on the types of lipids
Membranes rich in unsaturated fatty acids are
more fluid than those rich in saturated fatty acids
Membranes must be fluid to work properly; they
are usually about as fluid as salad oil!
The Fluidity of Membranes, cont.
© 2014 Pearson Education, Inc.
Fluid Viscous
Unsaturated tails
prevent packing.
Saturated tails pack
together.
Cholesterol
Cholesterol reduces
membrane fluidity at
moderate temperatures,
but at low temperatures
hinders solidification.
(a) Unsaturated versus saturated hydrocarbon tails
(b) Cholesterol within the animal cell membrane
Figure
7.5—
Factors
that affect
membran
e fluidity
6
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
The steroid cholesterol has different effects on
membrane fluidity at different temperatures
At warm temperatures (such as 37°C), cholesterol
restrains movement of phospholipids
At cool temperatures, it maintains fluidity by
preventing tight packing
The Role of Cholesterol
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Variations in lipid composition of cell membranes of many
species appear to be adaptations to specific environmental
conditions
Fish in extremely cold environments have high proportion of
unsaturated hydrocarbons tails, enabling their membranes to
remain fluid
Thermophilic bacteria and archaea in hot springs have unusual
lipids that prevent excessive fluidity
Ability to change the lipid compositions in response to
temperature changes has evolved in organisms that live where
temperatures vary
In winter wheat, the % of unsaturated phospholipids increases in
autumn, keeping membranes from solidifying during the winter.
Evolution of Differences in Membrane Lipid Composition
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Membrane Proteins and Their Functions
A membrane is a collage of different proteins,
often grouped together, embedded in the fluid
matrix of the lipid bilayer
Proteins determine most of the membrane’s
specific functions
7
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
The fluid mosaic model states that a membrane
is a fluid structure with a “mosaic” of various
proteins embedded in it
Proteins are not randomly distributed in the
membrane
The Fluid Mosaic Model
© 2014 Pearson Education, Inc.
Glyco-
protein Carbohydrate Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE
Microfilaments
of cytoskeleton
Fibers of extra-
cellular matrix (ECM)
Cholesterol
Peripheral
proteins Integral
protein CYTOPLASMIC
SIDE OF
MEMBRANE
Figure 7.3—Updated model of an animal cell’s plasma
membrane
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Peripheral proteins are bound to the surface of
the membrane
Integral proteins penetrate the hydrophobic core
Integral proteins that span the membrane are called
transmembrane proteins
The hydrophobic regions of an integral protein
consist of one or more stretches of nonpolar
amino acids, often coiled into alpha helices
Types of Membrane Proteins
8
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N-terminus
C-terminus
helix
EXTRACELLULAR
SIDE
CYTOPLASMIC
SIDE
Figure 7.6—The structure of a transmembrane protein
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Six major functions of membrane proteins
1. Transport
2. Enzymatic activity
3. Signal transduction
4. Cell-cell recognition
5. Intercellular joining
6. Attachment to the cytoskeleton and extracellular
matrix (ECM)
Functions of Membrane Proteins
© 2014 Pearson Education, Inc.
(a) Transport (b) Enzymatic
activity
(c) Signal
transduction
(d) Cell-cell
recognition(e) Intercellular
joining
(f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)
Enzymes
ATP
Signaling
moleculeReceptor
Signal transduction
Glyco-
protein
Figure
7.7—
Some
functions
of
membrane
proteins
9
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
The Role of Membrane Carbohydrates in Cell-Cell Recognition
Cells recognize each other by binding to
molecules, often containing carbohydrates, on the
extracellular surface of the plasma membrane
Membrane carbohydrates may be covalently
bonded to lipids (forming glycolipids) or more
commonly to proteins (forming glycoproteins)
Carbohydrates on the external side of the plasma
membrane vary among species, individuals, and
even cell types in an individual
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Synthesis and Sidedness of Membranes
Membranes have distinct inside and outside faces
The asymmetrical distribution of proteins, lipids,
and associated carbohydrates in the plasma
membrane is determined when the membrane is
built by the ER and Golgi apparatus
© 2014 Pearson Education, Inc.
Transmembrane
glycoproteinsSecretory
protein
Golgi
apparatus
Vesicle
Attached
carbohydrate
ER
lumen
Glycolipid
Transmembrane
glycoprotein
Plasma membrane:
Cytoplasmic face
Extracellular face
Membrane
glycolipid
Secreted
protein
Figure 7.9—Synthesis of membrane components and their
orientation in the membrane
10
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Concept 7.2: Membrane structure results in selective permeability
A cell must exchange materials with its
surroundings, a process controlled by the
plasma membrane
Plasma membranes are selectively permeable,
regulating the cell’s molecular traffic
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
The Permeability of the Lipid Bilayer
Hydrophobic (nonpolar) molecules, such as
hydrocarbons, can dissolve in the lipid bilayer and
pass through the membrane rapidly
Small molecules, like O2, CO2 (i.e., gases), and
some H2O also can pass between the
phospholipids in the membrane
Hydrophilic molecules including ions and polar
molecules (like sugars) do not cross the
membrane easily
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Membrane Selectivity
11
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Transport Proteins
Transport proteins allow passage of hydrophilic
substances across the membrane
Some transport proteins, called channel proteins,
have a hydrophilic channel that certain molecules or
ions can use as a tunnel
Channel proteins called aquaporins facilitate the
passage of water
Other transport proteins, called carrier proteins,
bind to molecules and change shape to shuttle
them across the membrane
A transport protein is specific for the substance
it moves
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Channel
protein
Carrier
protein
Passive transport:
Facilitated diffusion
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Video: Water Movement Through an Aquaporin
12
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Concept 7.3: Passive transport is diffusion of a substance across a membrane with no energy investment
Diffusion is the tendency for molecules to spread
out evenly into the available space
Although each molecule moves randomly,
diffusion of a population of molecules may exhibit
a net movement in one direction
At dynamic equilibrium, as many molecules cross
the membrane in one direction as in the other
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Diffusion
© 2014 Pearson Education, Inc.
Molecules of dye Membrane (cross section)
WATER
(a) Diffusion of one solute
(b) Diffusion of two solutes
Net diffusion Net diffusion
Net diffusionNet diffusion
Net diffusion Net diffusion
Equilibrium
Equilibrium
Equilibrium
Figure
7.10—The
diffusion
of solutes
across a
synthetic
membrane
13
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Substances diffuse down their concentration
gradient, the difference in concentration of a
substance from one area to another
No work must be done to move substances down
the concentration gradient
The diffusion of a substance across a biological
membrane is passive transport because no
energy is expended by the cell to make it happen
Passive Transport, cont.
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Effects of Osmosis on Water Balance
Osmosis is the diffusion of water across a
selectively permeable membrane
Water diffuses across a membrane from the
region of lower solute concentration to the region
of higher solute concentration until the solute
concentration is equal on both sides
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Osmosis
14
© 2014 Pearson Education, Inc.
Lower concentration
of solute (sugar)
Higher concentration
of soluteMore similar
concentrations of solute
Sugar
moleculeH2O
Selectively
permeable
membrane
OsmosisFigure 7.11—Osmosis
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Water Balance in Cells with Cell Walls
Tonicity is the ability of a surrounding solution to
cause a cell to gain or lose water
Isotonic solution: Solute concentration is the
same as that inside the cell; no net water
movement across the plasma membrane
Hypertonic solution: Solute concentration is
greater than that inside the cell; cell loses water
Hypotonic solution: Solute concentration is less
than that inside the cell; cell gains water
© 2014 Pearson Education, Inc.
Hypotonic Isotonic Hypertonic
H2O
Lysed Normal Shriveled
Plasma
membrane
Cell
wall
Turgid (normal) Flaccid Plasmolyzed
(a)
An
imal cell
(b)
Pla
nt
cell
Plasma
membrane
H2OH2O
H2OH2O
H2OH2O
H2O
Figure 7.12—The water balance of living cells
15
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Video: Turgid Elodea
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Video: Plasmolysis
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Hypertonic or hypotonic environments create
osmotic problems for organisms
Osmoregulation, the control of solute
concentrations and water balance, is a necessary
adaptation for life in such environments
The protist Paramecium, which is hypertonic to its
pond water environment, has a contractile
vacuole that acts as a pump
Osmoregulation
16
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Contractile vacuole50 μm
Figure 7.13—The contractile vacuole of Paramecium
caudatum
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Video: Paramecium Vacuole
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Water Balance of Cells with Cell Walls
Cell walls help maintain water balance
A plant cell in a hypotonic solution swells until the wall
opposes uptake; the cell is now turgid (firm)
If a plant cell and its surroundings are isotonic, there is no
net movement of water into the cell;
the cell becomes flaccid (limp) and if enough cells are
flaccid, the plant will wilt.
In a hypertonic environment, plant cells lose water
The membrane pulls away from the cell wall causing the
plant to wilt, a usually lethal effect called plasmolysis
17
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Facilitated Diffusion: Passive Transport Aided by Proteins
In facilitated diffusion, transport proteins speed the
passive movement of molecules across the plasma
membrane
Channel proteins provide corridors that allow a specific
molecule or ion to cross the membrane
Channel proteins include
Aquaporins for facilitated diffusion of water
Ion channels that open or close in response to a
stimulus (gated channels)
Carrier proteins undergo a subtle change in shape that
translocates the solute-binding site across the membrane
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(a) A channel
protein
(b) A carrier protein
Carrier protein
Channel protein Solute
Solute
EXTRACELLULAR
FLUID
CYTOPLASM
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Concept 7.4: Active transport uses energy to move solutes against their gradients
Facilitated diffusion is still passive because the
solute moves down its concentration gradient, and
the transport requires no energy
Some transport proteins, however, can move
solutes against their concentration gradients
18
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The Need for Energy in Active Transport
Active transport moves substances against their
concentration gradients
Active transport requires energy, usually in the
form of ATP
Active transport is performed by specific proteins
embedded in the membranes
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Active transport allows cells to maintain
concentration gradients that differ from their
surroundings
The sodium-potassium pump is one type of
active transport system
Active Transport: The Sodium-Potassium Pump
© 2014 Pearson Education, Inc.
EXTRACELLULAR
FLUID
CYTOPLASM
1 2
5
6
4
3
[Na+] low
[K+] high
[Na+] high
[K+] low
Na+
K+
K+
K+
K+
K+
K+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
Na+
ATP
ADP
P
PP
i
P
Figure 7.15—The sodium-potassium pump: a specific case of
active transport
19
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Active Transport
© 2014 Pearson Education, Inc.
ATP
Passive transport Active transport
Diffusion Facilitated diffusion
Figure 7.16—Review: passive
and active transport
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
BioFlix: Membrane Transport
20
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How Ion Pumps Maintain Membrane Potential
Membrane potential is the voltage difference
across a membrane
Voltage is created by differences in the distribution
of positive and negative ions across a membrane
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Two combined forces, collectively called the
electrochemical gradient, drive the diffusion of
ions across a membrane
A chemical force (the ion’s concentration gradient)
An electrical force (the effect of the membrane
potential on the ion’s movement)
Electrochemical Gradient
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
An electrogenic pump is a transport protein that
generates voltage across a membrane
The sodium-potassium pump is the major
electrogenic pump of animal cells
The main electrogenic pump of plants, fungi, and
bacteria is a proton pump
Electrogenic pumps help store energy that can be
used for cellular work
Electrogenic Pump
21
© 2014 Pearson Education, Inc.
EXTRACELLULAR
FLUID
ATP
CYTOPLASM
H+
H+
H+
H+
H+
H+
Proton pump
−
−
−
−
+
+
+
+
Figure 7.17—A proton pump: An electrogenic
pump in plant, fungi, and bacterial cells
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Cotransport: Coupled Transport by a Membrane Protein
Cotransport occurs when active transport of a
solute indirectly drives transport of other
substances
EXAMPLE: Plants commonly use the gradient of
hydrogen ions generated by proton pumps to drive
active transport of nutrients into the cell
© 2014 Pearson Education, Inc.
Sucrose
ATP
H+
H+
H+
H+
H+
Proton pump
−
−
−
−
+
+
+
+
Sucrose-H+
cotransporter
H+
H+
H+
H+
Sucrose
Diffusion of H+
Figure 7.18—Cotransport: active transport driven by a
concentration gradient
22
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Concept 7.5: Bulk transport across theplasma membrane occurs by exocytosis and endocytosis
Small molecules and water enter or leave the
cell through the lipid bilayer or via transport
proteins
Large molecules, such as polysaccharides and
proteins, cross the membrane in bulk via vesicles
Bulk transport requires energy
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Exocytosis and Endocytosis Introduction
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Exocytosis
In exocytosis, transport vesicles migrate to the
membrane, fuse with it, and release their contents
outside the cell
Many secretory cells use exocytosis to export
their products
23
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Exocytosis
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Endocytosis
In endocytosis, the cell takes in macromolecules
by forming vesicles from the plasma membrane
Endocytosis is a reversal of exocytosis, involving
different proteins
There are three types of endocytosis:
Phagocytosis (“cellular eating”)
Pinocytosis (“cellular drinking”)
Receptor-mediated endocytosis
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
In phagocytosis a cell engulfs a particle in a
vacuole
The vacuole fuses with a lysosome to digest the
particle
Endocytosis, cont.
24
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Phagocytosis
Solutes
Pseudopodium
“Food”
or
other
particle
Food
vacuole
CYTOPLASM
EXTRACELLULAR
FLUIDPseudopodium
of amoeba
Bacterium
Food vacuole
An amoeba engulfing a
bacterium via phago-
cytosis (TEM)
1 μm
Figure 7.19a—Exploring
endocytosis in animal cells
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Phagocytosis
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In pinocytosis, molecules dissolved in droplets
are taken up when extracellular fluid is “gulped”
into tiny vesicles
Endocytosis, cont.
25
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Pinocytosis
Plasma
membrane
Coated
pit
Coated
vesicle
Coat
proteinPinocytotic vesicles
forming (TEMs)
0.2
5 μm
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Animation: Pinocytosis
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In receptor-mediated endocytosis, binding of
ligands to receptors triggers vesicle formation
A ligand is any molecule that binds specifically
to a receptor site of another molecule
Endocytosis, cont.
26
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Receptor-Mediated
Endocytosis
ReceptorCoat
protein
Plasma
membrane
Top: A coated pit.
Bottom: A coated vesicle
forming during receptor-
mediated endocytosis (TEMs)
0.2
5 μm
Figure 7.19c—Exploring endocytosis in animal cells
© 2014 Pearson Education, Inc.© 2014 Pearson Education, Inc.
Animation: Receptor-Mediated Endocytosis
© 2014 Pearson Education, Inc.
Phagocytosis Pinocytosis
Receptor-Mediated
Endocytosis
Solutes
Pseudopodium
“Food”
or
other
particle
Food
vacuole
CYTOPLASM
EXTRACELLULAR
FLUID
Plasma
membrane
Coated
pit
Coated
vesicle
Coat
protein
Receptor
Figure 7.19—Exploring
endocytosis in animal cells