Membrane Structure and Function: Chapter Concepts
Membrane Structure and Function: Chapter Concepts
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
67
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Cell Biology
4.1
Outside cell
glycolipid
hydrophilic
heads
carbohydrate
chain
glycoprotein
hydrophobic
tails
hydrophilic
heads
phospholipid
bilayer
integral
protein
cholesterol
peripheral
protein
plasma
membrane
Inside cell
filaments of
the cytoskeleton
Figure 4.1
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
membranes asymmetrycarbohydrate chains are attached to the outside surface and cytoskeleton filaments are attached to the inside surface.
4-3
Chapter 4
hydrophobic
region
69
hydrophilic
regions
mouse cell
cell
human cell
fusion
Figure 4.2
After human and mouse cells fuse, the plasma membrane proteins of
the mouse (blue circles) and of the human cell (red circles) mix within
a short time.
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Cell Biology
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.
CellCell Recognition
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
blood to clot.
Certain organs produce chemicals that are needed by other
ware, has been done for quite some time, but researchers alcells. Diabetes mellitus occurs when the pancreas is no longer
ways used cancer cells that divide without coaxing. Now
producing insulin, a molecule that causes all cells to take up
researchers have learned how to grow all sorts of human cells
glucose and the liver to store glucose as glycogen. Tissue engiin tissue culture and have hopes that they can even make the
neering can possibly come to the rescue. Insulin-producing panneed for organ transplantation obsolete. Organ transplantation
creatic cells from a pig can be grown in the laboratory. The cells
encounters two hurdles that are hard to overcome: (1) there is
are encased in plastic capsules called microreactors, because rean overwhelming need, but few human organs are available to
actors are typically large vats where chemicals are produced
be transplanted; and (2) immunosuppressive drugs must be ad(Fig. 4A). These capsules are so small they can be placed into the
ministered even if the organs are carefully matched to the
abdomen where they will float freely and produce insulin as
recipient, because the body tends to reject foreign organs. To
needed. The membrane of the capsule contains pores that are
address these problems, some researchers have turned to pigs
large enough to allow oxygen and nutrients to flow in and
as a source of organs for humans. Through genetic engineering,
wastes and insulin to flow out by diffusion. But the membrane
they have crippled the enzymes that produce plasma memof a microreactor will prevent immune cells from coming into
brane carbohydrate chains on pig cells; therefore, the human
contact with the enclosed pancreatic cells. Unless immune cells
body is unable to recognize a pig organ as being foreign. Pigs
actually come in contact with transplanted cells, they cannot
carry viruses such as the one that causes swine flu, but pig
recognize them as foreign and destroy them. Researchers are
viruses are not expected to cause infections in humans. Thereeven busily growing implantable liver tissue. They use a spongy
fore, it is predicted that pig-to-human transplants will someday
material that can be seeded with the patients own liver cells.
be safely done.
Human embryonic cells are grown in tissue culture, and if
Tissue engineering offers another possible solution to the
differentiation can one day be achieved, it may be possible to
transplant problem. Tissue engineering is an endeavor that prosupply Alzheimer patients with nerve cells, and cardiac paduces manufactured bioproducts that can replace normal structients with heart cells, and so forth.
tures in the body. Integra is an artificial skin that consists of a
porous matrix made of the protein collagen and a derivative of
shark cartilage. This product, which is available in unlimited 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 patients 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 increase 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 systemused to replace weakened arteries sometimes cause the suppressing drugs.
71
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Cell Biology
Active
Transport
Means
Passive
Transport
Means
Direction
Requirement
Examples
DIFFUSION
Toward lower
concentration
Concentration
gradient only
Lipid-soluble molecules,
water, and gases
FACILITATED TRANSPORT
Toward lower
concentration
Carrier and
concentration gradient
ACTIVE TRANSPORT
Toward greater
concentration
EXOCYTOSIS
Toward outside
Macromolecules
Phagocytosis
Toward inside
Vacuole formation
Pinocytosis
(includes receptormediated endocytosis)
Toward inside
Vesicle formation
Macromolecules
ENDOCYTOSIS
4.2
noncharged molecule
H2O
charged
molecules
and ions
macromolecule
plasma
membrane
Figure 4.4
The curved arrows indicate that these substances cannot cross the
plasma membrane and the back and forth arrows indicate that these
substances can cross the plasma membrane.
the membrane. Small polar molecules such as carbon dioxide 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 oxygen follows its concentration gradient as it enters a cell. Carbon 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 transport proteins. Carrier proteins combine with an ion or molecule before transporting it across the membrane. Channel
proteins form a channel that allows an ion or charged molecule 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 energy, include active transport, endocytosis, and exocytosis.
The plasma membrane is differentially permeable.
Certain substances can freely pass through the
membrane and others must be transported across
either by carrier proteins or by vacuole formation.
4-7
Chapter 4
73
water
molecules
(solvent)
dye
molecules
(solute)
a. Crystal of
dye is placed
in water
Figure 4.5
b. Diffusion of
water and dye
molecules
c. Equal distribution
of molecules
results
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
alveoli
capillary
oxygen
Figure 4.6
Oxygen (O2) diffuses into the capillaries of the lungs because there is
a higher concentration of oxygen in the alveoli (air sacs) than in the
capillaries.
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Cell Biology
less water
(higher
percentage
of solute)
solution rises
due to movement
of water toward
lower percentage
of solute
water
more water
(lower
percentage
of solute)
10%
<10%
5%
> 5%
a.
c.
Figure 4.7
b.
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 illustrate osmosis, a thistle tube containing a 10% sugar solution1 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 percentage 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 solution 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.
Tonicity
Tonicity refers to the strength of a solution in relationship to
osmosis. In the laboratory, cells are normally placed in isotonic 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 solutions 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; hemolysis, 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.
4-9
Chapter 4
75
plasma
membrane
Animal
Cells
Under isotonic
conditions, there is no net
movement of water.
In a hypotonic environment,
water enters the cell, which
may burst (lysis).
cell
wall
nucleus
Plant
Cells
In a hypertonic environment,
water leaves the cell, which
shrivels (crenation).
plasma membrane
chloroplast
Figure 4.8
In a hypotonic environment,
vacuoles fill with water, turgor
pressure develops, and
chloroplasts are seen next to the
cell wall.
In a hypertonic environment,
vacuoles lose water, the
cytoplasm shrinks (plasmolysis),
and chloroplasts are seen in the
center of the cell.
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 creates turgor pressure. When a plant cell is placed in a hypotonic 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. Turgor pressure in plant cells is extremely important to the
maintenance of the plants erect position. If you forget to water 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
cells are placed in this solution, they shrink. The term crenation 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.
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Cell Biology
Outside
Inside
carrier
Outside
carrier
protein
Inside
protein
2
2
ene
3
Membrane
Figure 4.9
rgy
Membrane
Facilitated transport.
4.4
Facilitated Transport
4-11
Chapter 4
77
Active Transport
During active transport, ions or molecules move through
the plasma membrane, accumulating either inside or outside the cell. For example, iodine collects in the cells of the
thyroid gland; nutrients are completely absorbed from
the gut by the cells lining the digestive tract, and sodium
ions (Na ) can be almost completely withdrawn from
urine by cells lining the kidney tubules. In these instances,
substances have moved to the region of higher concentration, exactly opposite to the process of diffusion. It has
been estimated that up to 40% of a cells energy supply
may be used for active transport of solute across its
membrane.
Both carrier proteins and an expenditure of energy are
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 involved primarily in active transport, such as kidney cells,
have a large number of mitochondria near the membrane
through which active transport is occurring.
Proteins involved in active transport often are called
pumps, because just as a water pump uses energy to move
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 associated with nerve and muscle cells, moves sodium ions
(Na) to the outside of the cell and potassium ions (K) to
the inside of the cell. These two events are presumed to be
linked, and the carrier protein is called a sodiumpotassium pump. A change in carrier shape after the attachment, 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 enzymatically by the carrier.
The passage of salt (NaCl) across a plasma membrane
is of primary importance in cells. The chloride ion (Cl)
usually crosses the plasma membrane because it is attracted by positively charged sodium ions (Na). First,
sodium ions are pumped across a membrane, and then
chloride ions simply diffuse through channels that allow
their passage. As noted in Figure 4.3, the chloride ion
channels malfunction in persons with cystic fibrosis, and
this leads to the symptoms of this inherited (genetic)
disorder.
Outside
Inside
Carrier has a
shape that allows
it to take up three
sodium ions (Na+).
3 Na+
carrier
ATP
ATP is split, and
phosphate group is
transferred to carrier.
3 Na+
ADP
Change in shape
3 Na+
results that
causes carrier to
release three sodium
K+
ions (Na+) outside
the cell. New shape
allows carrier to take up
potassium ions (K+).
P
Phosphate group is
released from carrier.
K+
K+
78
4.5
Part 1
4-12
Cell Biology
plasma
membrane
red blood
cell
Exocytosis
During exocytosis, vesicles often formed by the Golgi apparatus 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.
vacuole
a. Phagocytosis
solute
plasma membrane
vesicle
b. Pinocytosis
Inside
receptor
protein
solute
Endocytosis
During endocytosis, cells take in substances by vesicle formation (Fig. 4.12). A portion of the plasma membrane invaginates 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 phagocytosis. 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 12 m.
c. Receptor-mediated
endocytosis
vesicle
4-13
Chapter 4
receptor
protein
79
solute
endocytosis
2
receptor
protein
3
solutes
removed
exocytosis
a.
b.
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Cell Biology
PD. People who have received such transplants 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
The diffusion of water across a differentially permeable membrane is called osmosis. Water moves across the membrane into the
area of lower water (higher solute) content. When cells are in an isotonic solution, they neither gain nor lose water; when they are in a hypotonic solution, they gain water; and when they are in a hypertonic
solution, they lose water.
4-15
Chapter 4
Testing Yourself
Choose the best answer for each question.
1. Label this diagram of the plasma membrane.
c.
a.
d.
b.
e.
f.
j.
i.
g.
h.
81
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Cell Biology
a.
cell
wall
b.
Thinking Scientifically
1. Considering the movement of molecules across the plasma
membrane:
a. Contrast the manner in which alcohol and water enter a
cell (page 72).
b. Contrast the manner in which sodium ions (Na) and
chloride ions (Cl) exit a cell (Fig. 4.3).
c. Contrast the manner in which amino acids and proteins
enter a cell (page 72).
d. How might the proteins from question c be digested (chapter 3)?
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
constant, how many exocytotic vesicles per endocytotic
vesicles would you expect?
b. Imagine a cell that is moving from left to right. If vesicle
formation is facilitating movement, where would you
expect exocytosis to be occurring? Where would you expect endocytosis to be occurring?
c. Receptor-mediated endocytosis is a process by which a
substance combines with a receptor before endocytosis
brings the entire complex into the cell. Imagine a virus that
enters a cell in this manner. If so, what additional step is
needed for the virus to enter the cell proper?
isotonic solution 74
osmosis 74
osmotic pressure 74
peripheral protein 69
phagocytosis 78
pinocytosis 78
plasmolysis 75
receptor-mediated
endocytosis 79
receptor protein 70
sodium-potassium
pump 77
solute 73
solvent 73
tonicity 74
turgor pressure 75
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4 Diffusion
5 Osmosis