Chapter 4

Cell Membrane and Transport

Chapter Learning Outcomes

By the end of this chapter, you should be able to:

1. Describe the fluid mosaic model of membrane
2. Distinguish between the following pairs or
sets of terms: peripheral and integral
membrane proteins; osmosis, facilitated
diffusion, and active transport; hypertonic,
hypotonic, and isotonic solutions
3. Explain how transport proteins facilitate
diffusion
Overview: 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
Cellular membranes are fluid mosaics of lipids and
proteins
• Phospholipids are the most abundant lipid in
the plasma membrane
• Phospholipids are amphipathic molecules,
containing hydrophobic and hydrophilic regions
• The fluid mosaic model states that a
membrane is a fluid structure with a “mosaic” of
various proteins embedded in it
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
 WATER
Hydrophilic
head

Hydrophobic
tail

WATER
• In 1935, Hugh Davson and James 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, J. Singer and G. Nicolson proposed that
the membrane is a mosaic of proteins dispersed
within the bilayer, with only the hydrophilic regions
exposed to water
Phospholipid
bilayer

Hydrophobic regions Hydrophilic

of protein regions of protein
The Fluidity of Membranes

• Phospholipids in the plasma membrane can

move within the bilayer
• Most of the lipids, and some proteins, drift
laterally
• Rarely does a molecule flip-flop transversely
across the membrane
 Lateral movement Flip-flop
(~107 times per second) (~ once per month)

(a) Movement of phospholipids

Fluid Viscous

Unsaturated hydrocarbon Saturated hydro-

tails with kinks carbon tails
(b) Membrane fluidity

Cholesterol

(c) Cholesterol within the animal cell membrane

• 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 that those rich in saturated fatty
acids
• Membranes must be fluid to work properly;
they are usually about as fluid as salad oil
 Fluid Viscous

Unsaturated hydrocarbon Saturated hydro-

tails with kinks carbon tails
(b) Membrane fluidity
• 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
 Cholesterol

(c) Cholesterol within the animal cell membrane

Membrane Proteins and Their Functions

• A membrane is a collage of different proteins

embedded in the fluid matrix of the lipid bilayer
• Proteins determine most of the membrane’s
specific functions
Fig. 7-7

Fibers of
extracellular
matrix (ECM)

Glyco- Carbohydrate
protein
Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE

Cholesterol

Microfilaments Peripheral
of cytoskeleton proteins
Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE
• 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
 N-terminus EXTRACELLULAR
SIDE

C-terminus
CYTOPLASMIC
a Helix SIDE
• Six major functions of membrane proteins:
– Transport
– Enzymatic activity
– Signal transduction
– Cell-cell recognition
– Intercellular joining
– Attachment to the cytoskeleton and
extracellular matrix (ECM)
 Signaling molecule

Enzymes Receptor

ATP
Signal transduction
(a) Transport (b) Enzymatic activity (c) Signal transduction

Glyco-
protein

(d) Cell-cell recognition (e) Intercellular joining (f) Attachment to

the cytoskeleton
and extracellular
matrix (ECM)
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
The Permeability of the Lipid Bilayer

• Hydrophobic (nonpolar) molecules, such as

hydrocarbons, can dissolve in the lipid bilayer
and pass through the membrane rapidly
• Polar molecules, such as sugars, do not cross
the membrane easily
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
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 one way as cross in the other direction
• 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 it
requires no energy from the cell to make it
happen
Molecules of dye Membrane (cross section)

WATER

Net diffusion Net diffusion Equilibrium

(a) Diffusion of one solute

Net diffusion Net diffusion Equilibrium

Net diffusion Net diffusion Equilibrium

(b) Diffusion of two solutes

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
Lower Higher Same concentration
concentration concentration of sugar
of solute (sugar) of sugar

H2O

Selectively
permeable
membrane

Osmosis
Water Balance of Cells Without Walls

• Tonicity is the ability of a 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
 Hypotonic solution Isotonic solution Hypertonic solution

H2O H2O H2O H2O

(a) Animal
cell

Lysed Normal Shriveled

H2O H2O H2O H2O

(b) Plant
cell

Turgid (normal) Flaccid Plasmolyzed

• Hypertonic or hypotonic environments create
osmotic problems for organisms
• Osmoregulation, the control of 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
Water Balance of Cells with 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 the plant
may wilt
• In a hypertonic environment, plant cells lose
water; eventually, the membrane pulls away
from the wall, a usually lethal effect called
plasmolysis
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)
EXTRACELLULAR
FLUID

Channel protein Solute

CYTOPLASM

(a) A channel protein

Carrier protein Solute

(b) A carrier protein

Active transport uses energy to move solutes
against their gradients
• Facilitated diffusion is still passive because the
solute moves down its concentration gradient
• Some transport proteins, however, can move
solutes against their concentration gradients
The Need for Energy in Active Transport

• Active transport moves substances against

their concentration gradient
• Active transport requires energy, usually in the
form of ATP
• Active transport is performed by specific
proteins embedded in the membranes
• Active transport allows cells to maintain
concentration gradients that differ from their
surroundings

• The sodium-potassium pump is one type of

active transport system
EXTRACELLULAR
[Na+] high Na+
FLUID [K+] low Na+

Na+ Na+ Na+

Na+ Na+

Na+

[Na+] low ATP

Na+ P
CYTOPLASM [K+] high P
ADP
1 2 3

K+

K+

+
K+
K
K+

P
K+ P
6 5 4
 Passive transport Active transport

ATP
Diffusion Facilitated diffusion