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Membrane Structure & Function

Chapter 7

You should be able to:

1. Define the following terms: amphipathic
molecules, aquaporins, diffusion
2. Explain how membrane fluidity is influenced by
temperature and membrane composition
3. Distinguish between the following pairs or sets of
terms: peripheral and integral membrane
proteins; channel and carrier proteins; osmosis,
facilitated diffusion, and active transport;
hypertonic, hypotonic, and isotonic solutions

4. Explain how transport proteins facilitate diffusion

5. Explain how an electrogenic pump creates
voltage across a membrane, and name two
electrogenic pumps
6. Explain how large molecules are transported
across a cell membrane

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Overview: Life at the Edge

o The plasma membrane is the boundary that
separates the living cell from its
surroundings
o Membranes are made of Proteins and Lipids
o The plasma membrane exhibits selective
permeability, allowing some substances to
cross it more easily than others

Cellular membranes are fluid

mosaics of lipids and proteins
o Phospholipids are the most abundant lipid in the
plasma membrane
o Phospholipids are amphipathic molecules,
containing hydrophobic and hydrophilic regions
o The fluid mosaic model states that a membrane
is a fluid structure with a mosaic of various
proteins embedded in it
o Fluid-Substance that flows (vs. locked in place
like a solid)
o Mosaic: Art made of smaller units

Fig. 7-2

A Phospholipid Bilayer

Hydrophilic
head

WATER

Hydrophobic
tail
WATER

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Phospholipid
bilayer

Hydrophobic regions
of protein

Freeze Fracture
TECHNIQUE

Hydrophilic
regions of protein
RESULTS

Extracellular
layer

Proteins Inside of extracellular

layer

Knife

Plasma membrane

Cytoplasmic
layer

Inside of cytoplasmic
layer

The Fluidity of Membranes

o Phospholipids in the plasma membrane can move within
the bilayer
o Most of the lipids, and some proteins, drift laterally
o (Like a person weaving through a crowd)
o Rarely does a molecule flip-flop transversely across the
membrane

Lateral movement
(107 times per second)

Flip-flop
( once per month)

(a) Movement of phospholipids

Fig. 7-6

RESULTS

Membrane proteins

Mouse cell

Mixed proteins
after 1 hour
Human cell

Hybrid cell

oScientists at John Hopkins University labeled the PM proteins of

a mouse cell and a human cell with 2 different markers and fused
the cells
oThe mixing of the mouse and human membrane proteins
indicates that at least some membrane proteins move sideways
within the plane of the PM

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o As temperatures cool, membranes switch

from a fluid state to a more solid state
o Membranes must be fluid to work properly;
they are usually about as fluid as salad oil
o The temperature at which a membrane
solidifies depends on the types of lipids
o Membranes rich in unsaturated fatty acids
are more fluid that those rich in saturated
fatty acids

Fig. 7-5b

Fluid

Unsaturated hydrocarbon
tails with kinks

Viscous

Saturated hydrocarbon tails

(b) Membrane fluidity

Unsaturated hydrocarbon tails of phospholipids have kinks
that keep the molecules from packing together, enhancing
membrane fluidity.

o The steroid cholesterol has different effects on

membrane fluidity at different temperatures
o At warm temperatures (such as 37C), cholesterol
restrains movement of phospholipids
o At cool temperatures, it maintains
fluidity by preventing tight
packing

oCholesterol
o(c) Cholesterol within the animal cell membrane

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Membrane Proteins and Their

Functions
o A membrane is a collage of different
proteins embedded in the fluid matrix of the
lipid bilayer
o Proteins determine most of the membranes
specific functions

Fig. 7-7

Fibers of
extracellular
matrix (ECM)

Glycoprotein

Carbohydrate
Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE

Cholesterol
Microfilaments
of cytoskeleton

Peripheral
proteins
Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE

o Peripheral proteins are bound to the surface

of the membrane
o Integral proteins penetrate into the hydrophobic core

o Integral proteins that span the membrane are called

transmembrane proteins
o Integrins (Chapter 6) connect extracellular matrix to the
cytoskeleton--External forces can change cellular activities
o Membrane proteins- from Rough ER ribosomes

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Fig. 7-8

The hydrophobic
regions of an
integral protein
consist of one or
more stretches of
nonpolar amino
acids, often coiled
into alpha helices

EXTRACELLULAR
SIDE

Nterminus

Cterminus

CYTOPLASMIC
SIDE

Helix

o Six major functions of membrane proteins

o
o
o
o
o
o

Transport
Enzymatic activity
Signal transduction
Cell-cell recognition
Intercellular joining
Attachment to the cytoskeleton and
extracellular matrix (ECM)

Six major functions of membrane proteins:

Signaling molecule
Enzymes

ATP
(a) Transport
Left: A protein that spans the memb
may provide a hydrophilic channel
across the memb that is selective for
a particular solute. Right: Other
transport proteins shuttle a substance
from 1 side to the other by changing
shape

Receptor

Signal transduction
(b) Enzymatic activity
A protein built into the memb
many be an enzyme with its
active site exposed to
substances in the adjacent
solution. In some cases.
Several enzymes in a memb
are organized as a team that
carries out sequential steps
of a metabolic pathway

(c) Signal transduction

A memb protein (receptor) may
have a binding site with a specific
shape that fits the shape of a
chemical messenger, such as a
hormone. The external
messenger (signaling molecule)
may cause the protein to change
shape, allowing it to relay the
message to the inside of the cell,
usually by binding to a
cytoplasmic protein

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Glycoprotein

(d) Cell-cell recognition

(e) Intercellular joining

Some glycoproteins serve as

identification tags that are
specifically recognized by
memb proteins of other cells.
This type of cell-cell binding is
usually short-lived

Memb proteins of adjacent

cells may hook together in
various kinds of junctions,
such as gap junctions or tight
junctions. This type of binding
is more long lasting.

(f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)
Microfilaments or other
elements of the cytoskeleton
may be noncovalently bound
to the memb proteins, a
function that helps maintain
cell shape and stabilizes the
location of certain membrane
proteins

The Role of Membrane Carbohydrates

in Cell-Cell Recognition
o Cells recognize each other by binding to surface
molecules, often carbohydrates, on the plasma membrane
o Membrane carbohydrates may be covalently bonded to
lipids (forming glycolipids) or more commonly to proteins
(forming glycoproteins)

o Carbohydrates on the external side of the plasma

membrane vary among species, individuals, and even cell
types in an individual

Figure 7.11

HIV

Receptor
(CD4)

Co-receptor
(CCR5)

HIV can infect a cell that

has CCR5 on its surface,
as in most people.

Receptor (CD4)
but no CCR5
Plasma
membrane
HIV cannot infect a cell lacking
CCR5 on its surface, as in
resistant individuals.

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Synthesis and Sidedness of

Membranes
o Membranes have distinct inside and outside
faces
o 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

1. In the ER: syn of memb proteins and lipids.

Carbs may be added to proteins here
2. In the Golgi: Glycoproteins modified and
lipids acquire carbs glycolipids
3. Transmemb proteins, glycolipids, and
secretory proteins are transported by
vesicles to the PM
4. Vesicles fuse with the memb, releasing
secretory proteins

Membrane structure results in

selective permeability
o A cell must exchange materials with its
surroundings, a process controlled by the
plasma membrane
o Plasma membranes are selectively
permeable, regulating the cells molecular
traffic

The Permeability of the Lipid

Bilayer
o Hydrophobic (nonpolar) molecules, such as
hydrocarbons, can dissolve in the lipid
bilayer and pass through the membrane
rapidly
o Polar molecules, such as sugars, do not
cross the membrane easily
Animation: Membrane Selectivity

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Transport Proteins
o Transport proteins allow passage of
hydrophilic substances across the
membrane
o Some transport proteins, called channel
proteins, have a hydrophilic channel that
certain molecules or ions can use as a
tunnel
o varying degrees of specificity

o Channel proteins called aquaporins

facilitate the passage of water

o Other transport proteins, called carrier

proteins, bind to molecules and change
shape to shuttle them across the membrane

o A transport protein is specific for the

substance it moves

Passive transport is diffusion of a substance

across a membrane with no energy
investment
o Diffusion is the tendency for molecules to spread
out evenly into the available space
o Ex. A spilled liquid releasing a smell (molecules into the
air). Eventually, the molecules will spread evenly
throughout the room, and everybody will smell it.

o Although each molecule moves randomly,

diffusion of a population of molecules may exhibit
a net movement in one direction
o Ex. From the place of the spill to the rest of the room

o At dynamic equilibrium, as many molecules cross

one way as cross in the other direction
Animation: Diffusion

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o Substances diffuse down their concentration

gradient, the difference in concentration of a
substance from one area to another
o Higher concentration Lower concentration
o No work (no energy used) must be done to move
substances down the concentration gradient
o The diffusion of a substance across a biological
membrane is passive transport because it
requires no energy from the cell to make it happen

Fig. 7-11

Molecules of dye

Membrane (cross section)

WATER

Net diffusion

Net diffusion

Equilibrium

(a) Diffusion of one solute

Net diffusion
Net diffusion

Net diffusion
Net diffusion

Equilibrium
Equilibrium

(b) Diffusion of two solutes

Effects of Osmosis on Water Balance

o Osmosis is the diffusion of water across a selectively
permeable membrane (here, only water can pass)
o Water diffuses across a membrane from the region of lower
solute concentration to the region of higher solute
concentration
o Because water molecules H-bond to solute
molecules, they become part of the
solute-water unit.
o Here, only think about the free (unbound) water molecules.
They will be lower on the side with more solute molecules
(higher concentration). So free water will move from the side
with more to the side with less.

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Fig. 7-12

Lower
concentration
of solute (sugar)

Higher
concentration
of sugar

Same concentration
of sugar

H2O
Selectively
permeable
membrane

Osmosis

Water Balance of Cells Without Walls

o Tonicity is the ability of a solution to cause a cell
to gain or lose water
o Isotonic solution: Solute concentration is the
same as that inside the cell; no net water
movement across the plasma membrane
o Hypertonic solution: Solute concentration is
greater than that inside the cell; cell loses water
o Hypotonic solution: Solute concentration is less
than that inside the cell; cell gains water

Fig. 7-13

Hypotonic solution
H2O

Isotonic solution
H2O

H2O

Hypertonic solution
H2O

(a) Animal
cell

Lysed
H2O

Normal
H2O

Shriveled
H2O

H2O

(b) Plant
cell
Turgid (normal)

Flaccid

Plasmolyzed

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o Hypertonic or hypotonic environments

create osmotic problems for organisms
o Your blood is kept isotonic with your cells to prevent water loss or
gain

o Osmoregulation, the control of water

balance, is a necessary adaptation for life in
such environments
o The protist Paramecium, which is hypertonic
to its pond water environment, has a
contractile vacuole that acts as a pump

Fig. 7-14

50 m

Filling vacuole

(a) A contractile vacuole fills with fluid that enters from

a system of canals radiating throughout the cytoplasm.
Contracting vacuole

(b) When full, the vacuole and canals contract, expelling

fluid from the cell.

Water Balance of Cells with Walls

o Cell walls help maintain water balance
o A plant cell in a hypotonic solution swells until the wall
opposes uptake; the cell is now turgid (firm)
o 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 the plant may wilt
o In a hypertonic environment, plant cells lose water;
eventually, the membrane pulls away from the wall, a
usually lethal effect called plasmolysis
Video: Plasmolysis

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Facilitated Diffusion: Passive

Transport Aided by Proteins
o In facilitated diffusion, transport proteins
speed the passive movement of molecules
across the plasma membrane
o Cant go through plasma membrane at all
OR
o Cant go through Fast Enough for cells needs

oChannel proteins provide corridors that allow a

specific molecule or ion to cross the membrane
(selective tube)
oChannel proteins include:
oAquaporins, for facilitated diffusion of water
oIon channels that open or close in response to a
stimulus (gated channels)

oCarrier proteins undergo a subtle change in

shape that translocates the solute-binding site
across the membrane

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Active transport uses energy to

move solutes against their gradients
o Facilitated diffusion is still passive, because the
solute moves down its concentration gradient
o Some transport proteins can move solutes against
their concentration gradients
o Active transport moves substances against their
concentration gradient
o Active transport requires energy, usually in the form
of ATP
o Like using a pump to move water uphill
o Active transport is performed by specific proteins
embedded in the membranes

o Active transport allows cells to maintain

concentration gradients that differ from their
surroundings
o The sodium-potassium pump is one type
of active transport system

Animation: Active Transport

Fig. 7-16-7

EXTRACELLULAR
FLUID

Na+

[Na+] high
[K+] low

Na+
Na+

Na+

Na+

Na+

Na+

Na+

CYTOPLASM

Na+

[Na+] low
[K+] high

1 Cytoplasmic Na binds to
Na/K pump. High affinity
for Na

P
ADP

ATP

2 Binding of Na stimulates

3 Phos causes prot to change

shape, Na affinity decreases,
Na expelled outside

phosphorylation of the
protein by ATP

6 K released, pump has high

affinity for Na again

Loss of phos restores protein

5 to original shape and has

lower affinity for K

New shape has high affinity

for K, phos released

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Passive transport Substances diffuse

Active transport Some transport

spontaneously down their concentration gradient,

crossing a membrance with no expenditure of
energy by the cell. The rate of diffusion can be
greatky increased by transport proteins in the
membrane

proteins act as pumps, moving substances

across a membrane against their
concentration (or electrochemical)
gradients. Energy for this work is usually
supplied by ATP

ATP

Diffusion
Hydrophobic
molecs & (at a
slow rate)
very sm
uncharged
polar molecs
can diffuse
through the
lipid bilayer

Facilitated diffusion
Many hydrophilic
substances diffuse
through membs with
the assistance of
transport proteins,
either channel or
carrier proteins

How Ion Pumps Maintain Membrane Potential

o Membrane potential is the voltage difference across a
membrane
o Voltage is created by differences in the distribution of
positive and negative ions
o Areas (around a barrier) do not have a + or - charge
(except for electrons and protons). It has a difference. If
one side of a barrier (membrane or battery terminal) has
more + charges than the other, then the side with more is
+, and the side with less is negative, -, compared to the
side with more.
o Cells are usually Negative compared to outside (fewer
positive ions) (-50-200 mV)

o 2 combined forces, collectively called the

electrochemical gradient, drive the diffusion of
ions across a membrane:
o A chemical force (the ions concentration
gradient)
o An electrical force (the effect of the membrane
potential on the ions movement)
o If the components are opposite, ion flow is
slowed
o If the components (chemical and electrical) are
together, flow is very fast and easy!

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o An electrogenic pump is a transport protein

that generates voltage across a membrane
o The sodium-potassium pump is the major
electrogenic pump of animal cells
o The main electrogenic pump of plants, fungi,
and bacteria is a proton pump
o Electrogenic pumps help store energy that can
be used for cellular work

The main electrogenic pump of plants, fungi, and

bacteria is a Proton (H+) Pump

Cotransport: Coupled Transport by a

Membrane Protein
o Cotransport occurs when active transport of a
solute indirectly drives transport of another
solute
o Plants commonly use the gradient of hydrogen
ions generated by proton pumps to drive active
transport of nutrients into the cell
o You do this with Na+

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Fig. 7-19

H+

ATP

H+

+
H+

Proton pump

H+

H+

H+

+
H+
Sucrose-H+
cotransporter

Diffusion
of H+

H+
Sucrose

+
+

Sucrose

Bulk transport across the plasma membrane

occurs by exocytosis and endocytosis
o Small molecules and water enter or leave
the cell through the lipid bilayer or by
transport proteins
o Large molecules, such as polysaccharides
and proteins, cross the membrane in bulk
via vesicles
o Bulk transport requires energy
Animation: Exocytosis and Endocytosis Introduction

Exocytosis
o In exocytosis, transport vesicles migrate to
the membrane, fuse with it, and release their
contents
o Many secretory cells use exocytosis to
export their products

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Endocytosis
o In endocytosis, the cell takes in
macromolecules by forming vesicles from the
plasma membrane
o Endocytosis is a reversal of exocytosis,
involving different proteins
o There are three types of endocytosis:
o Phagocytosis (cellular eating)
o Pinocytosis (cellular drinking)
o Receptor-mediated endocytosis

o In phagocytosis a cell engulfs a particle in

a vacuole
o The vacuole fuses with a lysosome to digest
the particle
Animation: Phagocytosis

oIn pinocytosis, molecules

are taken up when
extracellular fluid is gulped
into tiny vesicles
oIn receptor-mediated
endocytosis, binding of
ligands to receptors triggers
vesicle formation
oA ligand is any molecule that
binds specifically to a receptor
site of another molecule

Animation: Pinocytosis

Animation: Receptor-Mediated Endocytosis

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Figure 7.22a

Phagocytosis

EXTRACELLULAR
FLUID
Solutes

Pseudopodium
of amoeba

1 m

Bacterium
Food vacuole

Pseudopodium

Food
or other
particle

An amoeba engulfing a bacterium

via phagocytosis (TEM).

Food
vacuole
CYTOPLASM

Figure 7.22b

0.5 m

Pinocytosis

Plasma
membrane

Pinocytosis vesicles forming

in a cell lining a small blood
vessel (TEM).

Vesicle

Figure 7.22c

Receptor-Mediated Endocytosis

Plasma
membrane

Receptor

Coat
proteins

Ligand

0.25 m

Coat proteins

Coated
pit
Coated
vesicle

Top: A coated pit. Bottom: A

coated vesicle forming during
receptor-mediated endocytosis
(TEMs).

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