Membrane

Structure and
Function
Chapter Concepts
4.1 Plasma Membrane Structure and Function
• The plasma membrane regulates the passage of
molecules into and out of the cell. 68
• The membrane contains lipids and proteins. Each
protein has a specific function. 70

4.2 The Permeability of the Plasma Membrane

• Some substances, particularly small, noncharged
molecules, pass freely across the plasma
membrane. Ions and other types of molecules
need assistance to cross the membrane. 72

4.3 Diffusion and Osmosis

• Molecules spontaneously diffuse (move from an
area of higher concentration to an area of lower
concentration), and some can diffuse across a
plasma membrane. 73
• Water diffuses across the plasma membrane, and
this can affect cell size and shape. 74

4.4 Transport by Carrier Proteins

• Carrier proteins assist the transport of some ions
and molecules across the plasma membrane. 76 A single cell about to be pierced by a fine probe so that DNA can
be removed by the suction tube on the bottom. An intact plasma
4.5 Exocytosis and Endocytosis
membrane is necessary to the life of any cell and if it is ruptured
• Vesicle formation takes other substances the cell cannot continue to exist.
into the cell, and vesicle fusion with the plasma
membrane discharges substances from the
cell. 78

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68 Part 1 Cell Biology 4-2

4.1 Plasma Membrane Structure and

B
anners flying on a castle wall mark off the community
within from the surrounding countryside. Inside, resi-
dents go about their appointed tasks for the good of
the community. Commands passed along from royalty to
knights to workers are obeyed by all. The almost impenetra-
ble wall prevents the enemy without from entering and dis-
turbing the peace within. Only certain small creatures can
pass through the open slitlike windows, and the drawbridge
must be lowered for most needed supplies.
The plasma membrane, which carries markers identify-
ing it as belonging to the individual, can be likened to the cas-
tle wall. Under the command of the nucleus, the organelles
carry out their specific functions and contribute to the work-
ing of the cell as a whole. Very few molecules can freely cross
the membrane, and most nutrients must be transported
across by special carriers. The cell uses these nutrients as a
source of building blocks and energy to maintain the cell. The
operations of the cell will continue only as long as the plasma
membrane selectively permits specific materials to enter and
leave and prevents the passage of others.
Function
The plasma membrane is a phospholipid bilayer in which
protein molecules are either partially or wholly embedded
(Fig. 4.1). The phospholipid bilayer has a fluid consistency,
comparable to that of light oil. The proteins are scattered
throughout the membrane; therefore they form a mosaic pat-
tern. This description of the plasma membrane is called the
fluid-mosaic model of membrane structure.
Phospholipids spontaneously arrange themselves into
a bilayer. The hydrophilic (water loving) polar heads of
the phospholipid molecules face the outside and inside of
the cell where water is found, and the hydrophobic (water
fearing) nonpolar tails face each other (Fig. 4.1). In
addition to phospholipids, there are two other types of
lipids in the plasma membrane. Glycolipids have a struc-
ture similar to phospholipids except that the hydrophilic
head is a variety of sugars joined to form a straight or

Outside cell

hydrophilic
carbohydrate heads
glycolipid glycoprotein chain
hydrophobic
tails

hydrophilic
heads

phospholipid
integral bilayer
protein

cholesterol

peripheral
protein plasma
Inside cell membrane
filaments of
the cytoskeleton

Figure 4.1 Fluid-mosaic model of plasma membrane structure.

The membrane is composed of a phospholipid bilayer in which proteins are embedded. The hydrophilic heads of phospholipids are a part of the
outside surface and the inside surface of the membrane. The hydrophobic tails make up the interior of the membrane. Note the plasma
membrane’s asymmetry—carbohydrate chains are attached to the outside surface and cytoskeleton filaments are attached to the inside surface.
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4-3 Chapter 4 Membrane Structure and Function 69

branching carbohydrate chain. Cholesterol is a lipid that is
found in animal plasma membranes; related steroids are
found in the plasma membrane of plants. Cholesterol re-
duces the permeability of the membrane to most biological
molecules.
The proteins in a membrane may be peripheral proteins
or integral proteins. Peripheral proteins occur either on the
outside or the inside surface of the membrane. Some of these
are anchored to the membrane by covalent bonding. Still
others are held in place by noncovalent interactions that can
be disrupted by gentle shaking or by change in the pH.
Integral proteins are found within the membrane and
have hydrophobic regions embedded within the membrane
and hydrophilic regions that project from both surfaces of
the bilayer:
move through the hydrophobic center of the membrane.)
The fluidity of a phospholipid bilayer means that cells are
pliable. Imagine if they were not—the long nerve fibers in
your neck would crack whenever you nodded your head!
Although some proteins are often held in place by cy-
toskeletal filaments, in general proteins are free to drift lat-
erally in the fluid lipid bilayer. This has been demonstrated
by fusing mouse and human cells, and watching the move-
ment of tagged proteins (Fig. 4.2). Forty minutes after fu-
sion, the proteins are completely mixed. The fluidity of the
membrane is needed for the functioning of some proteins
such as enzymes which become inactive when the mem-
brane solidifies.
The fluidity of the membrane, which is dependent
on its lipid components, is critical to the proper
functioning of the membrane’s proteins.

hydrophobic
region hydrophilic
regions

mouse cell human cell

Many integral proteins are glycoproteins, which have cell fusion

an attached carbohydrate chain. As with glycolipids, the
carbohydrate chain of sugars projects externally. There-
fore it can be said that the plasma membrane is “sugar-
coated.”
The plasma membrane is asymmetrical: the two halves
are not identical. The carbohydrate chains of the glyco-
lipids and proteins occur only on the outside surface and
immediately after fusion
the cytoskeletal filaments attach to proteins only on the inside
surface.

The plasma membrane consists of a phospholipid

bilayer. Peripheral proteins are found on the
outside and inside surface of the membrane.
Integral proteins span the lipid bilayer and often
have attached carbohydrate chains.

mixed membrane proteins

The Fluidity of the Plasma Membrane
Figure 4.2 Experiment to demonstrate lateral drifting of
At body temperature, the phospholipid bilayer of the plasma membrane proteins.
plasma membrane has the consistency of olive oil. The After human and mouse cells fuse, the plasma membrane proteins of
greater the concentration of unsaturated fatty acid the mouse (blue circles) and of the human cell (red circles) mix within
residues, the more fluid is the bilayer. In each monolayer, a short time.
the hydrocarbon tails wiggle, and the entire phospholipid
molecule can move sideways at a rate averaging about
2 µm—the length of a prokaryotic cell—per second. (Phos-
pholipid molecules rarely flip-flop from one layer to the
other, because this would require the hydrophilic head to
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70 Part 1 Cell Biology
The Mosaic Quality of the Membrane
The plasma membranes of various cells and the membranes
of various organelles each have their own unique collections
of proteins. The proteins form different patterns according
to the particular membrane and also within the same mem-
brane at different times. When you consider that the plasma
membrane of a red blood cell contains over 50 different
types of proteins, you can see why the membrane is said to
be a mosaic.
The integral proteins largely determine a membrane’s
specific functions. As we will discuss in more detail, certain
plasma membrane proteins are involved in the passage of
molecules through the membrane. Some of these are chan-
nel proteins through which a substance can simply move
across the membrane; others are carrier proteins that com-
bine with a substance and help it to move across the mem-
brane. Still others are receptors; each type of receptor
protein has a shape that allows a specific molecule to bind to
it. The binding of a molecule, such as a hormone (or other
signal molecule), can cause the protein to change its shape
and bring about a cellular response. Some plasma mem-
brane proteins are enzymatic proteins that carry out meta-
bolic reactions directly. The peripheral proteins associated
with the membrane often have a structural role in that they
help stabilize and shape the plasma membrane.
Figure 4.3 depicts the various functions of membrane
proteins.
The mosaic pattern of a membrane is dependent
on the proteins, which vary in structure and function.
Cell–Cell Recognition
The carbohydrate chains of glycolipids and glycoproteins
serve as the “fingerprints” of the cell. The possible diversity
of the chain is enormous; it can vary by the number of sug-
ars (15 is usual, but there can be several hundred), by
whether the chain is branched, and by the sequence of the
particular sugars.
Glycolipids and glycoproteins vary from species to
species, from individual to individual of the same species,
and even from cell to cell in the same individual. Therefore,
they make cell–cell recognition possible. Researchers work-
ing with mouse embryos have shown that as development
proceeds, the different type cells of the embryo develop their
own carbohydrate chains and that these chains allow the tis-
sues and cells of the embryo to sort themselves out.
As you probably know, transplanted tissues are often re-
jected by the body. This is because the immune system is
able to recognize that the foreign tissue’s cells do not have
the same glycolipids and glycoproteins as the rest of the
body’s cells. We also now know that a person’s particular
blood type is due to the presence of particular glycoproteins
in the membrane of red blood cells.
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4-4
Channel Protein
Allows a particular molecule or ion to cross the
plasma membrane freely. Cystic fibrosis, an
inherited disorder, is caused by a faulty chloride
(Cl –) channel; a thick mucus collects in airways
and in pancreatic and liver ducts.
Carrier Protein
Selectively interacts with a specific molecule
or ion so that it can cross the plasma
membrane. The inability of some persons
to use energy for sodium-potassium
(Na+–K+) transport has been suggested as
the cause of their obesity.
Cell Recognition Protein
The MHC (major histocompatibility complex)
glycoproteins are different for each person, so
organ transplants are difficult to achieve. Cells
with foreign MHC glycoproteins are attacked
by blood cells responsible for immunity.
Receptor Protein
Is shaped in such a way that a specific molecule
can bind to it. Pygmies are short, not because
they do not produce enough growth hormone,
but because their plasma membrane growth
hormone receptors are faulty and cannot
interact with growth hormone.
Enzymatic Protein
Catalyzes a specific reaction. The membrane
protein, adenylate cyclase, is involved in ATP
metabolism. Cholera bacteria release a toxin
that interferes with the proper functioning of
adenylate cyclase; sodium ions and water leave
intestinal cells and the individual dies from
severe diarrhea.
Figure 4.3 Membrane protein diversity.
These are some of the functions performed by proteins found in the
plasma membrane.
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 The Growing Field of Tissue Engineering
Tissue culture, the growing of animal cells in laboratory glass- blood to clot.
ware, has been done for quite some time, but researchers al- Certain organs produce chemicals that are needed by other
ways used cancer cells that divide without coaxing. Now cells. Diabetes mellitus occurs when the pancreas is no longer
researchers have learned how to grow all sorts of human cells producing insulin, a molecule that causes all cells to take up
in tissue culture and have hopes that they can even make the glucose and the liver to store glucose as glycogen. Tissue engi-
need for organ transplantation obsolete. Organ transplantation neering can possibly come to the rescue. Insulin-producing pan-
encounters two hurdles that are hard to overcome: (1) there is creatic cells from a pig can be grown in the laboratory. The cells
an overwhelming need, but few human organs are available to are encased in plastic capsules called microreactors, because re-
be transplanted; and (2) immunosuppressive drugs must be ad- actors are typically large vats where chemicals are produced
ministered even if the organs are carefully matched to the (Fig. 4A). These capsules are so small they can be placed into the
recipient, because the body tends to reject foreign organs. To abdomen where they will float freely and produce insulin as
address these problems, some researchers have turned to pigs needed. The membrane of the capsule contains pores that are
as a source of organs for humans. Through genetic engineering, large enough to allow oxygen and nutrients to flow in and
they have crippled the enzymes that produce plasma mem- wastes and insulin to flow out by diffusion. But the membrane
brane carbohydrate chains on pig cells; therefore, the human of a microreactor will prevent immune cells from coming into
body is unable to recognize a pig organ as being foreign. Pigs contact with the enclosed pancreatic cells. Unless immune cells
carry viruses such as the one that causes swine flu, but pig actually come in contact with transplanted cells, they cannot
viruses are not expected to cause infections in humans. There- recognize them as foreign and destroy them. Researchers are
fore, it is predicted that pig-to-human transplants will someday even busily growing implantable liver tissue. They use a spongy
be safely done. material that can be seeded with the patient’s own liver cells.
Tissue engineering offers another possible solution to the Human embryonic cells are grown in tissue culture, and if
transplant problem. Tissue engineering is an endeavor that pro- differentiation can one day be achieved, it may be possible to
duces manufactured bioproducts that can replace normal struc- supply Alzheimer patients with nerve cells, and cardiac pa-
tures in the body. Integra is an artificial skin that consists of a tients with heart cells, and so forth.
porous matrix made of the protein collagen and a derivative of
shark cartilage. This product, which is available in unlim-
ited quantity, will not cause an immune reaction. Integra is
used to cover extensive burns. Once the bottom layer of
skin (the dermis) regrows, a graft of the patient’s own
outer layer of skin (the epidermis) replaces the artificial
matrix.
Researchers have also had success growing human
cartilage for knee operations. In one study, 23 patients
who were experiencing pain because of a lack of cartilage
received a batch of their own chondrocytes (cartilage
cells) grown in the laboratory. All patients reported that
they were doing much better following the procedure.
Other procedures have also been tried. It is possible to
grow tissues to bolster weak ureters that take urine back
to the kidneys instead of to the bladder where it belongs.
And artificial tissue can be stitched into a bladder to in-
crease its capacity. If research continues to be successful,
nearly every human tissue is expected to undergo tissue
engineering. Several groups are working on methods to
reconstruct breast tissue after mastectomy so that one
day women may have an alternative to silicone breast Figure 4A Microreactors.
implants. Epithelial-lined plastic blood vessels are being Microreactors filled with insulin-producing pancreatic cells from pigs
developed because the walls of plastic blood vessels now flourished for 10 weeks in a diabetic mouse without immune system-
used to replace weakened arteries sometimes cause the suppressing drugs.

71
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72 Part 1 Cell Biology 4-6

Table 4.1 Passage of Molecules into and out of Cells

Name Direction Requirement Examples

DIFFUSION Toward lower Concentration Lipid-soluble molecules,

Transport
Passive

concentration gradient only water, and gases

Means

FACILITATED TRANSPORT Toward lower Carrier and Some sugars and

concentration concentration gradient amino acids

ACTIVE TRANSPORT Toward greater Carrier plus cellular Other sugars, amino acids,
concentration energy and ions
EXOCYTOSIS Toward outside Vesicle fuses with Macromolecules
Transport

plasma membrane
Means
Active

ENDOCYTOSIS
Phagocytosis Toward inside Vacuole formation Cells and subcellular material
Pinocytosis Toward inside Vesicle formation Macromolecules
(includes receptor-
mediated endocytosis)

4.2The Permeability of the Plasma

Membrane
The plasma membrane is differentially (selectively) perme-
able. Some substances can move across the membrane and
some cannot (Fig. 4.4). Macromolecules cannot diffuse across
the membrane because they are too large. Ions and charged
molecules cannot cross the membrane because they are un-
able to enter the hydrophobic phase of the lipid bilayer.
Noncharged molecules such as alcohols and oxygen are
lipid-soluble and therefore can cross the membrane with
ease. They are able to slip between the hydrophilic heads of
the phospholipids and pass through the hydrophobic tails of
noncharged molecule
H2O
macromolecule
+ and ions
–
plasma
membrane
– +
the membrane. Small polar molecules such as carbon diox-
ide and water also have no difficulty crossing through the
membrane. These molecules follow their concentration
gradient which is a gradual decrease in concentration over
distance. To take an example, oxygen is more concentrated
outside the cell than inside the cell because a cell uses
oxygen during aerobic cellular respiration. Therefore oxy-
gen follows its concentration gradient as it enters a cell. Car-
bon dioxide, on the other hand, which is produced when a
cell carries on cellular respiration, is more concentrated
inside the cell than outside the cell, and therefore it moves
down its concentration gradient as it exits a cell.
Special means are sometimes used to get ions and
charged molecules into and out of cells. Macromolecules can
cross a membrane when they are taken in or out by vesicle
formation (Table 4.1). Ions and molecules like amino acids
and sugars are assisted across by one of two classes of trans-
port proteins. Carrier proteins combine with an ion or mole-
cule before transporting it across the membrane. Channel
charged
molecules proteins form a channel that allows an ion or charged mole-
cule to pass through. Our discussion in this chapter is
largely restricted to carrier proteins. Carrier proteins are
specific for the substances they transport across the plasma
membrane.
Ways of crossing a plasma membrane are classified as
passive or active (Table 4.1). Passive ways, which do not use
chemical energy, involve diffusion or facilitated transport.
These passive ways depend on the motion energy of ions
and molecules. Active ways, which do require chemical en-
ergy, include active transport, endocytosis, and exocytosis.

The plasma membrane is differentially permeable.

Figure 4.4 How molecules cross the plasma membrane. Certain substances can freely pass through the
The curved arrows indicate that these substances cannot cross the membrane and others must be transported across
plasma membrane and the back and forth arrows indicate that these either by carrier proteins or by vacuole formation.
substances can cross the plasma membrane.
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4-7 Chapter 4 Membrane Structure and Function 73

water
molecules
(solvent)

dye
molecules
(solute)

a. Crystal of b. Diffusion of c. Equal distribution

dye is placed water and dye of molecules
in water molecules results

Figure 4.5 Process of diffusion.

Diffusion is spontaneous, and no chemical energy is required to bring it about. a. When dye crystals are placed in water, they are concentrated in
one area. b. The dye dissolves in the water, and there is a net movement of dye molecules from higher to lower concentration. There is also a net
movement of water molecules from a higher to a lower concentration. c. Eventually, the water and the dye molecules are equally distributed
throughout the container.

4.3 Diffusion and Osmosis M

Diffusion is the movement of molecules from a higher to a
lower concentration—that is, down their concentration
gradient—until equilibrium is achieved and they are dis-
tributed equally. Diffusion is a physical process that can be
observed with any type of molecule. For example, when a
crystal of dye is placed in water (Fig. 4.5), the dye and wa-
ter molecules move in various directions, but their net
movement, which is the sum of their motion, is toward the
region of lower concentration. Therefore, the dye is even-
tually dissolved in the water, resulting in a colored solu-
tion. A solution contains both a solute, usually a solid, and
a solvent, usually a liquid. In this case, the solute is the dye
and the solvent is the water molecules. Once the solute and
solvent are evenly distributed, they continue to move
about, but there is no net movement of either one in any
direction.
As discussed, the chemical and physical properties of alveoli
the plasma membrane allow only a few types of molecules
to enter and exit a cell simply by diffusion. Gases can diffuse
through the lipid bilayer; this is the mechanism by which
oxygen enters cells and carbon dioxide exits cells. Also, con-
sider the movement of oxygen from the alveoli (air sacs) of capillary
the lungs to blood in the lung capillaries (Fig. 4.6). After in-
halation (breathing in), the concentration of oxygen in the oxygen
alveoli is higher than that in the blood; therefore, oxygen dif-
fuses into the blood. The principle of diffusion can be em- Figure 4.6 Gas exchange in lungs.
ployed in the treatment of certain human disorders, as is Oxygen (O2) diffuses into the capillaries of the lungs because there is
discussed in the Science Focus on page 71. a higher concentration of oxygen in the alveoli (air sacs) than in the
capillaries.

Molecules diffuse down their concentration

gradients. A few types of small molecules can
simply diffuse through the plasma membrane.
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74 Part 1 Cell Biology 4-8

less water solution rises

(higher due to movement
percentage of water toward
of solute) net movement of water lower percentage
to inside of thistle tube of solute
water solute

more water 10% <10%

(lower
percentage
of solute)
5% > 5%

a. c.

differentially permeable membrane b.

Figure 4.7 Osmosis demonstration.

(Far left) A thistle tube, covered at the broad end by a differentially permeable membrane, contains a 10% sugar solution. The beaker contains a
5% sugar solution. (Middle) The solute (green circles) is unable to pass through the membrane, but the water (blue circles) passes through in
both directions. There is a net movement of water toward the inside of the thistle tube, where there is a lower percentage of water molecules.
(Far right) Due to the incoming water molecules, the level of the solution rises in the thistle tube.

Osmosis
Osmosis is the diffusion of water into and out of cells. To il-
lustrate osmosis, a thistle tube containing a 10% sugar solu-
tion1 is covered at one end by a differentially permeable
membrane and is then placed in a beaker containing a 5%
sugar solution (Fig. 4.7). The beaker contains more water
molecules (lower percentage of solute) per volume, and the
thistle tube contains fewer water molecules (higher percent-
age of solute) per volume. Under these conditions, there is a
net movement of water from the beaker to the inside of the
thistle tube across the membrane. The solute is unable to
pass through the membrane; therefore, the level of the solu-
tion within the thistle tube rises (Fig. 4.7c).
Notice the following in this illustration of osmosis:
1. A differentially permeable membrane separates two
solutions. The membrane does not permit passage of
the solute.
2. The beaker has more water (lower percentage of
solute), and the thistle tube has less water (higher
percentage of solute).
3. The membrane permits passage of water, and there is a
net movement of water from the beaker to the inside of
the thistle tube.
4. In the end, the concentration of solute in the thistle
tube is less than 10%. Why? Because there is now less
solute per volume. And the concentration of solute in
the beaker is greater than 5%. Why? Because there is
now more solute per volume.
Water enters the thistle tube due to the osmotic pressure
of the solution within the thistle tube. Osmotic pressure is
the pressure that develops in a system due to osmosis2. In
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other words, the greater the possible osmotic pressure the
more likely water will diffuse in that direction. Due to
osmotic pressure, water is absorbed from the human large
intestine, is retained by the kidneys, and is taken up by
capillaries from tissue fluid.
Tonicity
Tonicity refers to the strength of a solution in relationship to
osmosis. In the laboratory, cells are normally placed in iso-
tonic solutions; that is, the solute concentration is the same
on both sides of the membrane, and therefore there is no net
gain or loss of water (Fig. 4.8). The prefix iso means the same
as, and the term tonicity refers to the strength of the solution.
A 0.9% solution of the salt sodium chloride (NaCl) is known
to be isotonic to red blood cells. Therefore, intravenous solu-
tions medically administered usually have this tonicity.
Solutions that cause cells to swell, or even to burst, due to
an intake of water are said to be hypotonic solutions. The
prefix hypo means less than, and refers to a solution with a
lower percentage of solute (more water) than the cell. If a cell
is placed in a hypotonic solution, water enters the cell; the net
movement of water is from the outside to the inside of the cell.
Any concentration of a salt solution lower than 0.9% is
hypotonic to red blood cells. Animal cells placed in such a
solution expand and sometimes burst due to the buildup of
pressure. The term lysis is used to refer to disrupted cells; he-
molysis, then, is disrupted red blood cells.
1
Percent solutions are grams of solute per 100 ml of solvent. Therefore, a 10%
solution is 10 g of sugar with water added to make up 100 ml of solution.
2
Osmotic pressure is measured by placing a solution in an osmometer and then
immersing the osmometer in pure water. The pressure that develops is the osmotic
pressure of a solution.

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4-9 Chapter 4 Membrane Structure and Function 75

plasma
membrane

Animal
Cells

Under isotonic In a hypotonic environment, In a hypertonic environment,

conditions, there is no net water enters the cell, which water leaves the cell, which
movement of water. may burst (lysis). shrivels (crenation).

cell
nucleus wall
Plant
Cells plasma membrane

chloroplast

Under isotonic conditions, In a hypotonic environment, In a hypertonic environment,

there is no net movement of vacuoles fill with water, turgor vacuoles lose water, the
water. pressure develops, and cytoplasm shrinks (plasmolysis),
chloroplasts are seen next to the and chloroplasts are seen in the
cell wall. center of the cell.

Figure 4.8 Osmosis in animal and plant cells.

The arrows indicate the net movement of water. In an isotonic solution, a cell neither gains nor loses water; in a hypotonic solution, a cell gains
water; and in a hypertonic solution, a cell loses water.

The swelling of a plant cell in a hypotonic solution cre-
ates turgor pressure. When a plant cell is placed in a hypo-
tonic solution, we observe expansion of the cytoplasm
because the large central vacuole gains water and the plasma
membrane pushes against the rigid cell wall. The plant cell
does not burst because the cell wall does not give way. Tur-
gor pressure in plant cells is extremely important to the
maintenance of the plant’s erect position. If you forget to wa-
ter your plants they wilt due to decreased turgor pressure.
Solutions that cause cells to shrink or to shrivel due to a
loss of water are said to be hypertonic solutions. The prefix
hyper means more than, and refers to a solution with a
higher percentage of solute (less water) than the cell. If a cell
is placed in a hypertonic solution, water leaves the cell; the
net movement of water is from the inside to the outside of
the cell.
Any solution with a concentration higher than 0.9%
sodium chloride is hypertonic to red blood cells. If animal
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cells are placed in this solution, they shrink. The term crena-
tion refers to red blood cells in this condition. Meats are
sometimes preserved by salting them. The bacteria are not
killed by the salt but by the lack of water in the meat.
When a plant cell is placed in a hypertonic solution, the
plasma membrane pulls away from the cell wall as the
large central vacuole loses water. This is an example of
plasmolysis, a shrinking of the cytoplasm due to osmosis.
Dead plants you see along a salted roadside after the
winter died because they were exposed to a hypertonic
solution.
In an isotonic solution, a cell neither gains nor
loses water. In a hypotonic solution, a cell gains
water. In a hypertonic solution, a cell loses water
and the cytoplasm shrinks.

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76 Part 1 Cell Biology
Outside Inside
carrier
1 protein
3 3
Membrane
Figure 4.9 Facilitated transport.
A carrier protein speeds the rate at which a solute crosses a membrane
from higher solute concentration to lower solute concentration.
(1) Molecule enters carrier. (2) Molecule is transported across the
membrane and exits on inside. (3) Carrier returns to its former state.
4.4 Transport by Carrier Proteins M
The plasma membrane impedes the passage of all but a few
substances. Yet, biologically useful molecules do enter and
exit the cell at a rapid rate because there are carrier proteins
in the membrane. Carrier proteins are specific; each can
combine with only a certain type of molecule, which is then
transported through the membrane. It is not completely un-
derstood how carrier proteins function; but after a carrier
combines with a molecule, the carrier is believed to undergo
a change in shape that moves the molecule across the mem-
brane. Carrier proteins are required for facilitated and active
transport (see Table 4.1).
Some of the proteins in the plasma membrane
are carriers; they transport biologically useful
molecules into and out of the cell.
Facilitated Transport
Facilitated transport explains the passage of such molecules
as glucose and amino acids across the plasma membrane,
even though they are not lipid soluble. The passage of glu-
cose and amino acids is facilitated by their reversible combi-
nation with carrier proteins, which in some manner
transport them through the plasma membrane. These carrier
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4-10
Outside Inside
carrier
1 protein
2
ene
rgy
Membrane
Figure 4.10 Active transport.
Active transport allows a solute to cross the membrane from lower
solute concentration to higher solute concentration. (1) Molecule enters
carrier. (2) Chemical energy of ATP is needed to transport the molecule
which exits inside of cell. (3) Carrier returns to its former state.
proteins are specific. For example, various sugar molecules
of identical size might be present inside or outside the cell,
but glucose can cross the membrane hundreds of times
faster than the other sugars. This is a good example of the
differential permeability of the membrane.
The carrier for glucose has been isolated and a model
has been developed to explain how it works (Fig. 4.9). It
seems likely that the carrier has two conformations and that
it switches back and forth between the two states. After glu-
cose binds to the open end of a carrier, it closes behind the
glucose molecule. As glucose moves along, the constricted
end of the carrier opens in front of the molecule. After glu-
cose is released into the cytoplasm of the cell, the carrier
changes its conformation so that the binding site for glucose
is again open. This process can occur as often as 100 times
per second. Apparently, the cell has a pool of extra glucose
carriers. When the hormone insulin binds to a plasma mem-
brane receptor, more glucose carriers ordinarily appear in
the plasma membrane. Some forms of diabetes are caused
by insulin insensitivity; that is, the binding of insulin does
not result in extra glucose carriers in the membrane.
The model shows that after a carrier has assisted the
movement of a molecule to the other side of the membrane,
it is free to assist the passage of other similar molecules. Nei-
ther diffusion, explained previously, nor facilitated trans-
port requires an expenditure of chemical energy because the
molecules are moving down their concentration gradient in
the same direction they tend to move anyway.
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4-11 Chapter 4 Membrane Structure and Function 77

Active Transport
During active transport, ions or molecules move through Outside Inside
the plasma membrane, accumulating either inside or out-
side the cell. For example, iodine collects in the cells of the Carrier has a
shape that allows
thyroid gland; nutrients are completely absorbed from 3 Na+
it to take up three
the gut by the cells lining the digestive tract, and sodium sodium ions (Na+).
ions (Na ) can be almost completely withdrawn from carrier
urine by cells lining the kidney tubules. In these instances,
substances have moved to the region of higher concen-
tration, exactly opposite to the process of diffusion. It has
been estimated that up to 40% of a cell’s energy supply ATP
may be used for active transport of solute across its
ATP is split, and P
membrane. phosphate group is 3 Na+
Both carrier proteins and an expenditure of energy are transferred to carrier. ADP
needed to transport molecules against their concentration
gradient (Fig. 4.10). In this case, energy (ATP molecules) is
required for the carrier to combine with the substance to be
transported. Therefore, it is not surprising that cells in- Change in shape
volved primarily in active transport, such as kidney cells, results that 3 Na+
have a large number of mitochondria near the membrane causes carrier to
release three sodium
through which active transport is occurring. ions (Na+) outside K+ P
Proteins involved in active transport often are called the cell. New shape
pumps, because just as a water pump uses energy to move allows carrier to take up
potassium ions (K+).
water against the force of gravity, proteins use energy to
move a substance against its concentration gradient. One
type of pump that is active in all cells, but is especially as-
sociated with nerve and muscle cells, moves sodium ions P
(Na) to the outside of the cell and potassium ions (K) to Phosphate group is
the inside of the cell. These two events are presumed to be released from carrier. K+
linked, and the carrier protein is called a sodium-
potassium pump. A change in carrier shape after the at-
tachment, and again after the detachment, of a phosphate
group allows the carrier to combine alternately with
sodium ions and potassium ions (Fig. 4.11). The phosphate
group is donated by ATP, which is broken down enzymati- Change in shape results
that causes carrier to
cally by the carrier. release potassium ions (K+)
The passage of salt (NaCl) across a plasma membrane inside the cell. New shape is K+
is of primary importance in cells. The chloride ion (Cl) suitable to take up three sodium
ions (Na+) once again.
usually crosses the plasma membrane because it is at-
tracted by positively charged sodium ions (Na). First,
sodium ions are pumped across a membrane, and then
chloride ions simply diffuse through channels that allow Figure 4.11 The sodium-potassium pump.
A carrier protein actively moves three sodium ions (Na) to the
their passage. As noted in Figure 4.3, the chloride ion
outside of the cell for every potassium ion (K) pumped to the inside
channels malfunction in persons with cystic fibrosis, and of the cell. Note that chemical energy of ATP is required.
this leads to the symptoms of this inherited (genetic)
disorder.

During facilitated transport, substances follow their

concentration gradient. During active transport,
substances are moved against their concentration
gradient.
▲
▲

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78 Part 1 Cell Biology 4-12

4.5 Exocytosis and Endocytosis M

red blood plasma
What about the transport of macromolecules such as cell membrane
polypeptides, polysaccharides, or polynucleotides, which
are too large to be transported by carrier proteins? They are
transported in or out of the cell by vesicle formation, thereby
keeping the macromolecules contained so that they do not
mix with those in the cytoplasm.

Exocytosis
vacuole
During exocytosis, vesicles often formed by the Golgi appa- a. Phagocytosis
ratus and carrying a specific molecule, fuse with the plasma
membrane as secretion occurs. This is the way that insulin
leaves insulin-secreting cells, for instance. solute

plasma membrane

vesicle
b. Pinocytosis

Inside solute receptor

protein
Notice that the membrane of the vesicle becomes a part of
the plasma membrane. During cell growth, exocytosis is
probably used as a means to enlarge the plasma membrane,
whether or not secretion is also taking place.

Endocytosis
During endocytosis, cells take in substances by vesicle for-
mation (Fig. 4.12). A portion of the plasma membrane in-
vaginates to envelop the substance, and then the membrane
pinches off to form an intracellular vesicle.
When the material taken in by endocytosis is large, such
as a food particle or another cell, the process is called phago-
cytosis. Phagocytosis is common in unicellular organisms
like amoebas and in amoeboid cells like macrophages,
which are large cells that engulf bacteria and worn-out red
blood cells in mammals. When the endocytic vesicle fuses
with a lysosome, digestion occurs.
Pinocytosis occurs when vesicles form around a liquid
or very small particles. Blood cells, cells that line the kidney
tubules or intestinal wall, and plant root cells all use this
method of ingesting substances. Whereas phagocytosis can
be seen with the light microscope, the electron microscope
must be used to observe pinocytic vesicles, which are no
larger than 1–2 µm.
▲
▲
c. Receptor-mediated vesicle
endocytosis
Figure 4.12 Three methods of endocytosis.
a. Phagocytosis occurs when the substance to be transported into
the cell is large; certain specialized cells in the body can engulf worn-
out red blood cells by phagocytosis. Digestion occurs when the
resulting vacuole fuses with a lysosome. b. Pinocytosis occurs when
a macromolecule such as a polypeptide is to be transported into the
cell. The result is a small vacuole or vesicle. c. Receptor-mediated
endocytosis is a form of pinocytosis. The substance to be taken in (a
ligand) first binds to a specific receptor protein which migrates to a
pit or is already in a pit. The vesicle that forms contains the ligand
and its receptor. Sometimes the receptor is recycled, as shown in
Figure 4.13.

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4-13 Chapter 4 Membrane Structure and Function 79

receptor solute
1
protein

endocytosis

2
receptor
protein

3 solutes
removed

exocytosis

a. b.

Figure 4.13 Receptor-mediated endocytosis.

a. (1) The receptors in the coated pits combine only with a solute. (2) The vesicle that forms is at first coated with a fibrous protein (blue squares),
but soon the vesicle loses its coat. (3) Solutes leave the vesicle. (4) When exocytosis occurs, membrane and therefore receptors are returned to
the plasma membrane. b. Electron micrographs of a coated pit in the process of forming a vesicle.

Receptor-mediated endocytosis is a form of pinocytosis
that is quite specific because it involves the use of a receptor
protein shaped in such a way that a specific molecule such
as vitamins, peptide hormones, and lipoproteins can bind to
it. The binding of a solute to the receptors causes the recep-
tors to gather at one location. This location is called a coated
pit because there is a layer of fibrous protein on the cyto-
plasmic side (see step 1, Fig. 4.13). Once the vesicle is
formed, the fibrous coat is released and the vesicle appears
uncoated (see step 2). The fate of the vesicle and its contents
depends on the kind of solute it contains. Sometimes the
solute simply enters the cytoplasm (step 3). A spent hor-
mone, on the other hand, may be digested when the vesicle
fuses with a lysosome. The membrane of the vesicle and,
therefore, the receptors are returned to the plasma mem-
brane (step 4), or the vesicle can go to other membranous
locations.
Aside from simply allowing substances to enter cells selec-
tively from an extracellular fluid, coated pits are also involved
in the transfer and exchange of substances between cells.
▲
▲
Such exchanges take place when the substances move from
maternal blood into fetal blood at the placenta, for example.
The importance of receptor-mediated endocytosis is
demonstrated by a genetic disorder called familial hyper-
cholesterolemia. Cholesterol is transported in blood by a
complex of lipids and proteins called low-density lipopro-
tein (LDL). These individuals have inherited a gene that
causes them to have a reduced number and/or defective re-
ceptors for LDL in their plasma membranes. Instead of cho-
lesterol entering cells, it accumulates in the walls of arterial
blood vessels, leading to high blood pressure, occluded
(blocked) arteries, and heart attacks.
Substances are secreted from a cell by exocytosis.
Substances enter a cell by endocytosis. Receptor-
mediated endocytosis allows cells to take up
specific kinds of molecules and then sort them
within the cell.

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80 Part 1 Cell Biology 4-14

S uch celebrities as Mohammad Ali, a

former heavyweight boxing champion,
Janet Reno, the attorney general of the
United States, and Michael J. Fox, a
favorite movie actor, have Parkinson dis-
ease. By age 65, Parkinson disease (PD)
affects roughly one of every 100 Ameri-
cans. Due to the death of brain cells that
produce a substance called dopamine, mo-
tor control is not as smooth as it should be.
The three obvious signs of Parkinson are
slowness of movement, tremor, and rigid-
ity. As the disease worsens, patients
become unable to carry out even the sim-
plest activities.
You might think that the condition
could be cured by simply giving a patient
dopamine, but dopamine, like many other
chemical substances, cannot cross the
blood-brain barrier. The blood-brain barrier
is simply due to the impermeability of the
capillaries serving the brain. Nutrients, such
as glucose, and essential amino acids can
only pass through due to facilitated trans-
port. Most drugs can’t get through at all.
Luckily a precursor of dopamine
called L-dopa can get through the blood-
brain barrier, and when L-dopa is given as
a medication, it will be changed into
dopamine until there are few cells left to
do the job. Along the way, physicians and
patients are faced with a wide assortment
of adjunct remedies. Some of these are sur-
gical procedures. Michael J. Fox opted for
pallidotomy, a procedure that kills off cells
that go out of control when the dopamine-
producing cells die off. An experimental
surgical procedure, however, involves the
transplantation of dopamine-producing
fetal tissue into the brains of people with
PD. People who have received such trans-
plants report a lessening of symptoms.
Is it ethical to use tissue from aborted
fetuses for transplants? Is it possible that
women would have abortions just to make
fetal tissue available to loved ones, or for
payment? Should there be governmental
safeguards to prevent such a possibility?
Questions Go To Student OLC
1. Is it ethical to use fetal tissue to prevent
older people from having a debilitating
disorder? Why or why not?
2. Suppose you had a choice between using
fetal tissue (no payment required) and
adult cells bioengineered to produce
dopamine (payment required), which
would you choose and why?
3. Do you favor banning all research using
fetal tissue, or doing such research under
certain circumstances? Explain.

The diffusion of water across a differentially permeable mem-

Summarizing the Concepts
4.1 Plasma Membrane Structure and Function
There are two components of the plasma membrane: lipids and pro-
teins. In the lipid bilayer, phospholipids are arranged with their hy-
drophilic heads at the surfaces and their hydrophobic tails in the
interior. The lipid bilayer has the consistency of oil, and therefore pro-
teins can move laterally in the membrane. Glycolipids and glyco-
proteins are involved in marking the cell as belonging to a particular
individual and tissue.
The hydrophobic portion of an integral protein lies in the lipid bi-
layer of the plasma membrane, and the hydrophilic portion lies at the
surface. Proteins act as receptors, carry on enzymatic reactions, join
cells together, form channels, or act as carriers to move substances
across the membrane.
4.2 The Permeability of the Plasma Membrane
Some substances like gases and water are free to cross a plasma mem-
brane, and others, particularly ions, charged molecules, and macro-
molecules, have to be assisted across. Passive ways of crossing a
plasma membrane (diffusion and facilitated transport) do not require
an expenditure of chemical energy. Active ways of crossing a plasma
membrane (active transport and vesicle formation) do require an ex-
penditure of chemical energy.
4.3 Diffusion and Osmosis
Lipid-soluble compounds, water, and gases simply diffuse across the
membrane from the area of higher concentration to the area of lower
concentration.
▲
▲
brane is called osmosis. Water moves across the membrane into the
area of lower water (higher solute) content. When cells are in an iso-
tonic solution, they neither gain nor lose water; when they are in a hy-
potonic solution, they gain water; and when they are in a hypertonic
solution, they lose water.
4.4 Transport by Carrier Proteins
Some molecules are transported across the membrane by carrier pro-
teins that span the membrane.
During facilitated transport, a carrier protein assists the move-
ment of a molecule down its concentration gradient. No energy is
required.
During active transport, a carrier protein acts as a pump that
causes a substance to move against its concentration gradient. The
sodium-potassium pump carries Na to the outside of the cell and K
to the inside of the cell. Energy in the form of ATP molecules is required
for active transport to occur.
4.5 Exocytosis and Endocytosis
Larger substances can enter and exit a membrane by endocytosis and
exocytosis. Exocytosis involves secretion. Endocytosis includes
phagocytosis and pinocytosis which includes receptor-mediated en-
docytosis. Receptor-mediated endocytosis makes use of receptor
molecules in the plasma membrane. Once specific solutes bind to
their receptors, the coated pit becomes a coated vesicle. After losing
the coat, the vesicle can join with the lysosome, or after freeing the
solute, the receptor-containing vesicle can fuse with the plasma
membrane.

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4-15 Chapter 4 Membrane Structure and Function 81

3. A phospholipid molecule has a head and two tails. The tails

Studying the Concepts
1. Describe the structure of the plasma membrane, including the
phospholipid bilayer and the various types of proteins. 68–70
2. Why is a plasma membrane called differentially permeable? 72
3. What are the mechanisms by which substances enter and exit
cells? Which are passive ways, and which are active ways? 72
4. Define diffusion, and give an example. 73
5. Define osmosis. Define isotonic, hypertonic, and hypotonic
solutions, and give examples of how these concentrations
affect red blood cells. 74–75
6. Draw a simplified diagram of a red blood cell before and
after being placed in these solutions. What terms are used to
refer to the condition of the red blood cell in a hypertonic
solution and in a hypotonic solution? 75
7. Draw a simplified diagram of a plant cell before and after
being placed in these solutions. Describe the cell contents
under these conditions. 75
8. How does facilitated transport differ from simple diffusion
across the plasma membrane? 76
9. How does active transport differ from facilitated transport?
Give an example. 77
10. Diagram and define endocytosis and exocytosis. Describe
and contrast three methods of endocytosis. 78–79
Testing Yourself
Choose the best answer for each question.
1. Label this diagram of the plasma membrane.
c.
d.
a. b.
e.
f.
j.
i.
h. g.
2. The fluid-mosaic model of membrane structure refers to
a. the fluidity of proteins in the membrane and the pattern of
phospholipids in the membrane.
b. the fluidity of phospholipids and the pattern of proteins in
the membrane.
c. the fluidity of cholesterol and the pattern of sugar chains
outside the membrane.
d. the lack of fluidity of internal membranes compared to the
plasma membrane, and the ability of the proteins to move
laterally in the membrane.
e. the fluidity of hydrophobic regions, proteins and mosaic
pattern of hydrophilic regions.
▲
▲
are found
a. at the surfaces of the membrane.
b. in the interior of the membrane.
c. spanning the membrane.
d. where the environment is hydrophilic.
e. Both a and b are correct.
4. During diffusion,
a. all molecules move only from the area of higher to lower
concentration.
b. solvents move from the area of higher to lower concentra-
tion.
c. there is a net movement of molecules from the area of
higher to lower concentration.
d. a cell must be present for any movement of molecules to
occur.
e. molecules move against their concentration gradient if
they are small and charged.
5. When a cell is placed in a hypotonic solution,
a. solute exits the cell to equalize the concentration on both
sides of the membrane.
b. water exits the cell toward the area of lower solute concen-
tration.
c. water enters the cell toward the area of higher solute
concentration.
d. solute exits and water enters the cell.
e. Both c and d are correct.
6. When a cell is placed in a hypertonic solution,
a. solute exits the cell to equalize the concentration on both
sides of the membrane.
b. water exits the cell toward the area of lower solute concen-
tration.
c. water exits the cell toward the area of higher solute
concentration.
d. solute exits and water enters the cell.
e. Both a and c are correct.
7. Active transport
a. requires a carrier protein.
b. moves a molecule against its concentration gradient.
c. requires a supply of energy.
d. does not occur during facilitated transport.
e. All of these are correct.
8. The sodium-potassium pump
a. helps establish an electrochemical gradient across the
membrane.
b. concentrates sodium on the outside of the membrane.
c. utilizes a carrier protein and energy.
d. is present in the plasma membrane.
e. All of these are correct.
9. A scientist observing a protozoan notices a vacuole discharg-
ing its contents at the plasma membrane. This is an example
of
a. phagocytosis and vacuole formation.
b. endocytosis and active transport.
c. exocytosis and secretion.
d. active transport and vacuole release.
e. Both c and d are correct.

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82 Part 1 Cell Biology 4-16

10. Receptor-mediated endocytosis

a. is no different from phagocytosis. Understanding the Terms
b. brings specific substances into the cell.
active transport 77 isotonic solution 74
c. helps to concentrate proteins in vesicles.
carrier protein 70 osmosis 74
d. All of these are correct.
channel protein 70 osmotic pressure 74
11. Write hypotonic solution or hypertonic solution beneath each
cholesterol 68 peripheral protein 69
cell. Justify your conclusions.
concentration gradient 72 phagocytosis 78
differentially permeable 72 pinocytosis 78
diffusion 73 plasmolysis 75
endocytosis 78 receptor-mediated
enzymatic protein 70 endocytosis 79
exocytosis 78 receptor protein 70
facilitated transport 76 sodium-potassium
fluid-mosaic model 68 pump 77
glycolipid 68 solute 73
glycoprotein 69 solvent 73
a. hypertonic solution 75 tonicity 74
hypotonic solution 74 turgor pressure 75
integral protein 69

cell Match the terms to these definitions:

wall a. Movement of molecules from a region of higher
concentration to a region of lower concentration.
b. Internal pressure that adds to the strength of a cell
and builds up when water moves by osmosis into a plant cell.
c. Solution that contains the same concentration of
solute and water as the cell.
d. Passive transfer of a substance into and out of a
b. cell along a concentration gradient by a process that requires
a carrier.
e. Process in which an intracellular vesicle fuses
Thinking Scientifically with the plasma membrane so that the vesicle’s contents are
released outside the cell.
1. Considering the movement of molecules across the plasma
membrane:
a. Contrast the manner in which alcohol and water enter a
cell (page 72).
Using Technology
b. Contrast the manner in which sodium ions (Na) and Your study of membrane structure and function is
chloride ions (Cl) exit a cell (Fig. 4.3). supported by these available technologies:
c. Contrast the manner in which amino acids and proteins
enter a cell (page 72). Essential Study Partner CD-ROM
d. How might the proteins from question c be digested (chap- Cells £ Cell Membrane
ter 3)?
Visit the Mader web site for related ESP activities.
2. Exocytotic vesicles add plasma membrane to the cell, and
endocytotic vesicles remove plasma membrane.
a. In a cell in which the amount of plasma membrane stays Exploring the Internet
constant, how many exocytotic vesicles per endocytotic
vesicles would you expect? The Mader Home Page provides resources and tools as
b. Imagine a cell that is moving from left to right. If vesicle you study this chapter.
formation is facilitating movement, where would you http://www.mhhe.com/biosci/genbio/mader
expect exocytosis to be occurring? Where would you ex-
pect endocytosis to be occurring?
Virtual Physiology Laboratory CD-ROM
c. Receptor-mediated endocytosis is a process by which a
substance combines with a receptor before endocytosis Diffusion, Osmosis, & Tonicity
brings the entire complex into the cell. Imagine a virus that Enzyme Characteristics
enters a cell in this manner. If so, what additional step is
needed for the virus to enter the cell proper? Life Science Animations 3D Video
4 Diffusion
5 Osmosis
▲
▲

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