CHAPTER 8 MEMBRANE STUCTURE

AND FUNCTION

Section A: Membrane Structure

1. Membrane models have evolved to fit new data
2. Membranes are fluid
3. Membranes are mosaics of structure and function
4. Membrane carbohydrates are important for cell-cell recognition

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Introduction
• The plasma membrane separates the living cell from
its nonliving surroundings.
• This thin barrier, 8 nm thick, controls traffic into and
out of the cell.
• Like other membranes, the plasma membrane is
selectively permeable, allowing some substances to
cross more easily than others.

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• The main macromolecules in membranes are lipids
and proteins, but include some carbohydrates.
• The most abundant lipids are phospholipids.
• Phospholipids and most other membrane
constituents are amphipathic molecules.
• Amphipathic molecules have both hydrophobic regions
and hydrophilic regions.
• The phospholipids and proteins in membranes create
a unique physical environment, described by the
fluid mosaic model.
• A membrane is a fluid structure with proteins embedded
or attached to a double layer of phospholipids.
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1. Membrane modes have evolved to fit new
data
• Models of membranes were developed long before
membranes were first seen with electron
microscopes in the 1950s.
• In 1895, Charles Overton hypothesized that membranes
are made of lipids because substances that dissolve in
lipids enter cells faster than those that are insoluble.
• Twenty years later, chemical analysis confirmed that
membranes isolated from red blood cells are composed of
lipids and proteins.

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• Attempts to build artificial membranes provided
insight into the structure of real membranes.
• In 1917, Irving Langmuir discovered that phosphilipids
dissolved in benzene would form a film on water when
the benzene evaporated.
• The hydrophilic heads were immersed in water.

Fig. 8.1a

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• In 1925, E. Gorter and F. Grendel reasoned that
cell membranes must be a phospholipid bilayer,
two molecules thick.
• The molecules in the bilayer are arranged such that
the hydrophobic fatty acid tails are sheltered from
water while the
hydrophilic phosphate
groups interact
with water.

Fig. 8.1b
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• Actual membranes adhere more strongly to water
than do artificial membranes composed only of
phospholipids.
• One suggestion was that proteins on the surface
increased adhesion.
• In 1935, H. Davson and
J. Danielli proposed a
sandwich model in
which the phospholipid
bilayer lies between two
layers of globular
proteins.
Fig. 8.2a
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• Early images from electron microscopes seemed to
support the Davson-Danielli model and until the
1960s, it was considered the dominant model.
• Further investigation revealed two problems.
• First, not all membranes were alike, but differed in
thickness, appearance when stained, and percentage of
proteins.
• Second, measurements showed that membrane proteins
are actually not very soluble in water.
• Membrane proteins are amphipathic, with hydrophobic
and hydrophilic regions.
• If at the surface, the hydrophobic regions would be in
contact with water.
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• In 1972, S.J. Singer and G. Nicolson presented a
revised model that proposed that the membrane
proteins are dispersed and individually inserted
into the phospholipid bilayer.
• In this fluid mosaic
model, the hydrophilic
regions of proteins
and phospholipids are
in maximum contact
with water and the
hydrophobic regions
are in a nonaqueous
environment.
Fig. 8.2b

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• A specialized
preparation technique,
freeze-fracture, splits a
membrane along the
middle of the
phospholid bilayer
prior to electron
microscopy.
• This shows protein
particles interspersed
with a smooth matrix,
supporting the fluid
mosaic model.
Fig. 8.3
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2. Membranes are fluid
• Membrane molecules are held in place by relatively
weak hydrophobic interactions.
• Most of the lipids and some proteins can drift
laterally in the plane of the membrane, but rarely
flip-flop from one layer to the other.

Fig. 8.4a
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• The lateral movements of phospholipids are rapid,
about 2 microns per second.
• Many larger membrane proteins move more slowly
but do drift.
• Some proteins move in very directed manner, perhaps
guided/driven by the motor proteins attached to the
cytoskeleton.
• Other proteins never move, anchored by the
cytoskeleton.

Fig. 8.5
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• Membrane fluidity is influenced by temperature
and by its constituents.
• As temperatures cool, membranes switch from a
fluid state to a solid state as the phospholipids are
more closely packed.
• Membranes rich in unsaturated fatty acids are more
fluid that those
dominated by saturated
fatty acids because the
kinks in the unsaturated
fatty acid tails prevent
tight packing.
Fig. 8.4b
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• The steroid cholesterol is wedged between
phospholipid molecules in the plasma membrane
of animal cells.
• At warm temperatures, it restrains the movement
of phospholipids and reduces fluidity.
• At cool temperatures, it maintains fluidity by
preventing tight packing.

Fig. 8.4c

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• To work properly with active enzymes and
appropriate permeability, membranes must be
fluid, about as fluid as salad oil.
• Cells can alter the lipid composition of membranes
to compensate for changes in fluidity caused by
changing temperatures.
• For example, cold-adapted organisms, such as winter
wheat, increase the percentage of unsaturated
phospholipids in the autumn.
• This allows these organisms to prevent their membranes
from solidifying during winter.

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3. Membranes are mosaics of structure and
function
• A membrane is a collage of different proteins
embedded in the fluid matrix of the lipid bilayer.

Fig. 8.6
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• Proteins determine most of the membrane’s
specific functions.
• The plasma membrane and the membranes of the
various organelles each have unique collections of
proteins.
• There are two populations of membrane proteins.
• Peripheral proteins are not embedded in the lipid
bilayer at all.
• Instead, they are loosely bounded to the surface of the
protein, often connected to the other population of
membrane proteins.

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 • Integral proteins penetrate the hydrophobic core of the
lipid bilayer, often completely spanning the membrane
(a transmembrane protein).
• Where they contact the core, they have hydrophobic
regions with nonpolar amino acids, often coiled into
alpha helices.
• Where they are in
contact with the
aqueous environment,
they have hydrophilic
regions of amino acids.

Fig. 8.7
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• One role of membrane proteins is to reinforce the
shape of a cell and provide a strong framework.
• On the cytoplasmic side, some membrane proteins
connect to the cytoskeleton.
• On the exterior side, some membrane proteins attach to
the fibers of the extracellular matrix.

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• Membranes have distinctive inside and outside
faces.
• The two layers may differ
in lipid composition, and
proteins in the membrane
have a clear direction.
• The outer surface also has
carbohydrates.
• This asymmetrical
orientation begins during
synthesis of a new membrane
in the endoplasmic
reticulum.
Fig. 8.8

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• The proteins in the plasma membrane may provide
a variety of major cell functions.

Fig. 8.9
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4. Membrane carbohydrates are important
for cell-cell recognition
• The membrane plays the key role in cell-cell
recognition.
• Cell-cell recognition is the ability of a cell to distinguish
one type of neighboring cell from another.
• This attribute is important in cell sorting and organization
as tissues and organs in development.
• It is also the basis for rejection of foreign cells by the
immune system.
• Cells recognize other cells by keying on surface
molecules, often carbohydrates, on the plasma membrane.

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• Membrane carbohydrates are usually branched
oligosaccharides with fewer than 15 sugar units.
• They may be covalently bonded either to lipids,
forming glycolipids, or, more commonly, to
proteins, forming glycoproteins.
• The oligosaccharides on the external side of the
plasma membrane vary from species to species,
individual to individual, and even from cell type to
cell type within the same individual.
• This variation marks each cell type as distinct.
• The four human blood groups (A, B, AB, and O) differ
in the external carbohydrates on red blood cells.
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 CHAPTER 8 MEMBRANE
STRUCTURE AND FUNCTION

Section B: Traffic Across Membranes

1. A membrane’s molecular organization results in selective permeability
2. Passive transport is diffusion across a membrane
3. Osmosis is the passive transport of water
4. Cell survival depends on balancing water uptake and loss
5. Specific proteins facilitate the passive transport of water and selected
solutes: a closer look
6. Active transport is the pumping of solutes against their gradients
7. Some ion pumps generate voltage across membranes
8. In cotransport, a membrane protein couples the transport of two solutes
9. Exocytosis and endocytosis transport large molecules
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1. A membrane’s molecular organization
results in selective permeability
• A steady traffic of small molecules and ions moves
across the plasma membrane in both directions.
• For example, sugars, amino acids, and other nutrients
enter a muscle cell and metabolic waste products leave.
• The cell absorbs oxygen and expels carbon dioxide.
• It also regulates concentrations of inorganic ions, like Na+,
K+, Ca2+, and Cl-, by shuttling them across the membrane.
• However, substances do not move across the barrier
indiscriminately; membranes are selectively
permeable.
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• Permeability of a molecule through a membrane
depends on the interaction of that molecule with
the hydrophobic core of the membrane.
• Hydrophobic molecules, like hydrocarbons, CO2, and
O2, can dissolve in the lipid bilayer and cross easily.
• Ions and polar molecules pass through with difficulty.
• This includes small molecules, like water, and larger
critical molecules, like glucose and other sugars.
• Ions, whether atoms or molecules, and their
surrounding shell of water also have difficulties
penetrating the hydrophobic core.
• Proteins can assist and regulate the transport of ions and
polar molecules.

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• Specific ions and polar molecules can cross the
lipid bilayer by passing through transport
proteins that span the membrane.
• Some transport proteins have a hydrophilic channel that
certain molecules or ions can use as a tunnel through
the membrane.
• Others bind to these molecules and carry their
passengers across the membrane physically.
• Each transport protein is specific as to the
substances that it will translocate (move).
• For example, the glucose transport protein in the liver
will carry glucose from the blood to the cytoplasm, but
not fructose, its structural isomer.
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2. Passive transport is diffusion across a
membrane
• Diffusion is the tendency of molecules of any
substance to spread out in the available space
• Diffusion is driven by the intrinsic kinetic energy (thermal
motion or heat) of molecules.
• Movements of individual molecules are random.
• However, movement of a population of molecules
may be directional.

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• For example, if we start with a permeable membrane
separating a solution with dye molecules from pure
water, dye molecules will cross the barrier
randomly.
• The dye will cross the membrane until both
solutions have equal concentrations of the dye.
• At this dynamic equilibrium as many molecules pass
one way as cross in the other direction.

Fig. 8.10a
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• In the absence of other forces, a substance will
diffuse from where it is more concentrated to
where it is less concentrated, down its
concentration gradient.
• This spontaneous process decreases free energy and
increases entropy by creating a randomized mixture.
• Each substance diffuses down its own
concentration gradient, independent of the
concentration gradients of other substances.

Fig. 8.10b
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• The diffusion of a substance across a biological
membrane is passive transport because it requires
no energy from the cell to make it happen.
• The concentration gradient represents potential energy
and drives diffusion.
• However, because membranes are selectively
permeable, the interactions of the molecules with
the membrane play a role in the diffusion rate.
• Diffusion of molecules with limited permeability
through the lipid bilayer may be assisted by
transport proteins.

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3. Osmosis is the passive transport of water
• Differences in the relative concentration of dissolved
materials in two solutions can lead to the movement
of ions from one to the other.
• The solution with the higher concentration of solutes is
hypertonic.
• The solution with the lower concentration of solutes is
hypotonic.
• These are comparative terms.
• Tap water is hypertonic compared to distilled water but
hypotonic when compared to sea water.
• Solutions with equal solute concentrations are isotonic.
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• Imagine that two sugar solutions differing in
concentration are separated by a membrane that
will allow water through, but not sugar.
• The hypertonic solution has a lower water
concentration than the hypotonic solution.
• More of the water molecules in the hypertonic solution
are bound up in hydration shells around the sugar
molecules, leaving fewer unbound water molecules.

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• Unbound water molecules will move from the
hypotonic solution where they are abundant to the
hypertonic solution where they are rarer.
• This diffusion of water across a selectively
permeable membrane is a special case of passive
transport called osmosis.
• Osmosis continues
until the solutions
are isotonic.

Fig. 8.11
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• The direction of osmosis is determined only by a
difference in total solute concentration.
• The kinds of solutes in the solutions do not matter.
• This makes sense because the total solute concentration
is an indicator of the abundance of bound water
molecules (and therefore of free water molecules).
• When two solutions are isotonic, water molecules
move at equal rates from one to the other, with no
net osmosis.

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4. Cell survival depends on balancing water
uptake and loss
• An animal cell immersed in an isotonic environment
experiences no net movement of water across its
plasma membrane.
• Water flows across the membrane, but at the same rate in
both directions.
• The volume of the cell is stable.

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• The same cell in a hypertonic environment will
loose water, shrivel, and probably die.
• A cell in a hypotonic solution will gain water,
swell, and burst.

Fig. 8.12
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• For a cell living in an isotonic environment (for
example, many marine invertebrates) osmosis is
not a problem.
• Similarly, the cells of most land animals are bathed in
an extracellular fluid that is isotonic to the cells.
• Organisms without rigid walls have osmotic
problems in either a hypertonic or hypotonic
environment and must have adaptations for
osmoregulation to maintain their internal
environment.

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• For example, Paramecium, a protist, is hypertonic
when compared to the pond water in which it lives.
• In spite of a cell membrane that is less permeable to
water than other cells, water still continually enters the
Paramecium cell.
• To solve this problem,
Paramecium have a
specialized organelle,
the contractile vacuole,
that functions as a bilge
pump to force water out
of the cell.

Fig. 8.13

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• The cells of plants, prokaryotes, fungi, and some
protists have walls that contribute to the cell’s
water balance.
• An animal cell in a hypotonic solution will swell
until the elastic wall opposes further uptake.
• At this point
the cell is
turgid, a
healthy
state for
most plant
cells.

Fig. 8.12
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• Turgid cells contribute to the mechanical support
of the plant.
• If a cell and its surroundings are isotonic, there is
no movement of water into the cell and the cell is
flaccid and the plant may wilt.

Fig. 8.12
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• In a hypertonic solution, a cell wall has no
advantages.
• As the plant cell loses water, its volume shrinks.
• Eventually, the plasma membrane pulls away from
the wall.
• This
plasmolysis
is usually
lethal.

Fig. 8.12
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5. Specific proteins facilitate passive
transport of water and selected solutes:
a closer look
• Many polar molecules and ions that are normally
impeded by the lipid bilayer of the membrane diffuse
passively with the help of transport proteins that span
the membrane.
• The passive movement of molecules down its
concentration gradient via a transport protein is
called facilitated diffusion.

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• Transport proteins have much in common with
enzymes.
• They may have specific binding sites for the solute.
• Transport proteins can become saturated when they are
translocating passengers as fast as they can.
• Transport proteins can be inhibited by molecules that
resemble the normal “substrate.”
• When these bind to the transport proteins, they
outcompete the normal substrate for transport.
• While transport proteins do not usually catalyze chemical
reactions, they do catalyze a physical process,
transporting a molecule across a membrane that would
otherwise be relatively impermeable to the substrate.
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• Many transport proteins simply provide corridors
allowing a specific molecule or ion to cross the
membrane.
• These channel proteins allow fast transport.
• For example, water channel proteins, aquaprorins,
facilitate massive amounts of diffusion.

Fig. 8.14a
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• Some channel proteins, gated channels, open or
close depending on the presence or absence of a
physical or chemical stimulus.
• The chemical stimulus is usually different from the
transported molecule.
• For example, when neurotransmitters bind to specific
gated channels on the receiving neuron, these
channels open.
• This allows sodium ions into a nerve cell.
• When the neurotransmitters are not present, the
channels are closed.

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• Some transport proteins do not provide channels
but appear to actually translocate the solute-
binding site and solute across the membrane as the
protein changes shape.
• These shape changes could be triggered by the
binding and release of the transported molecule.

Fig. 8.14b

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6. Active transport is the pumping of
solutes against their gradients
• Some facilitated transport proteins can move solutes
against their concentration gradient, from the side
where they are less concentrated to the side where
they are more concentrated.
• This active transport requires the cell to expend its
own metabolic energy.
• Active transport is critical for a cell to maintain its
internal concentrations of small molecules that
would otherwise diffuse across the membrane.
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• Active transport is performed by specific proteins
embedded in the membranes.
• ATP supplies the energy for most active transport.
• Often, ATP powers active transport by shifting a
phosphate group from ATP (forming ADP) to the
transport protein.
• This may induce a conformational change in the
transport protein that translocates the solute across the
membrane.

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• The sodium-potassium pump actively maintains
the gradient of sodium (Na+) and potassium ions
(K+) across the membrane.
• Typically, an animal cell has higher concentrations of
K+ and lower concentrations of Na+ inside the cell.
• The sodium-potassium pump uses the energy of one
ATP to pump three Na+ ions out and two K+ ions in.

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Fig. 8.15

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Fig. 8.16 Both diffusion and facilitated diffusion are forms of passive transport of molecules down
their concentration gradient, while active transport requires an investment of energy to move
molecules against their concentration gradient.
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7. Some ion pumps generate voltage across
membranes
• All cells maintain a voltage across their plasma
membranes.
• The cytoplasm of a cell is negative in charge compared to
the extracellular fluid because of an unequal distribution
of cations and anions on opposite sides of the membrane.
• This voltage, the membrane potential, ranges from -50 to
-200 millivolts.

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• The membrane potential acts like a battery.
• The membrane potential favors the passive
transport of cations into the cell and anions out of
the cell.
• Two combined forces, collectively called the
electrochemical gradient, drive the diffusion of
ions across a membrane:
• A chemical force based on an ion’s concentration
gradient.
• An electrical force based on the effect of the membrane
potential on the ion’s movement.

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• Ions diffuse not simply down their concentration
gradient, but diffuse down their electrochemical
gradient.
• For example, before stimulation there is a higher
concentration of Na+ outside a resting nerve cell.
• When stimulated, a gated channel opens and Na +
diffuses into the cell down the electrochemical gradient.
• Special transport proteins, electrogenic pumps,
generate the voltage gradients across a membrane
• The sodium-potassium pump in animals restores the
electrochemical gradient not only by the active transport
of Na+ and K+, but because it pumps two K+ ions inside
for every three Na+ ions that it moves out.
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• In plants, bacteria, and fungi, a proton pump is
the major electrogenic pump, actively transporting
H+ out of the cell.
• Protons pumps in the cristae of mitochondria and
the thylakoids of chloroplasts, concentrate H+
behind membranes.
• These electrogenic
pumps store energy
that can be accessed
for cellular work.

Fig. 8.17

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8. In cotransport, a membrane protein
couples the transport of two solutes
• A single ATP-powered pump that transports one
solute can indirectly drive the active transport of
several other solutes through cotransport via a
different protein.
• As the solute that has been actively transported
diffuses back passively through a transport protein,
its movement can be coupled with the active
transport of another substance against its
concentration gradient.

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• Plants commonly use the gradient of hydrogen ions
that is generated by proton pumps to drive the
active transport of amino acids, sugars, and other
nutrients into the cell.
• The high concentration of H+ on one side of the
membrane, created by the proton pump, leads to the
facilitated diffusion of
protons back, but only
if another molecule,
like sucrose, travels
with the hydrogen ion.

Fig. 8.18
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9. Exocytosis and endocytosis transport
large molecules
• Small molecules and water enter or leave the cell
through the lipid bilayer or by transport proteins.
• Large molecules, such as polysaccharides and
proteins, cross the membrane via vesicles.
• During exocytosis, a transport vesicle budded from
the Golgi apparatus is moved by the cytoskeleton to
the plasma membrane.
• When the two membranes come in contact, the
bilayers fuse and spill the contents to the outside.
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• During endocytosis, a cell brings in
macromolecules and particulate matter by forming
new vesicles from the plasma membrane.
• Endocytosis is a reversal of exocytosis.
• A small area of the palsma membrane sinks inward to
form a pocket
• As the pocket into the plasma membrane deepens, it
pinches in, forming a vesicle containing the material
that had been outside the cell

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• One type of endocytosis is phagocytosis, “cellular
eating.”
• In phagocytosis, the cell engulfs a particle by
extending pseudopodia around it and packaging it
in a large vacuole.
• The contents of the vacuole are digested when the
vacuole fuses with a lysosome.

Fig. 8.19a
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• In pinocytosis, “cellular drinking,” a cell creates a
vesicle around a droplet of extracellular fluid.
• This is a non-specific process.

Fig. 8.19b

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• Receptor-mediated endocytosis is very specific in
what substances are being transported.
• This process is triggered when extracellular
substances bind to special receptors, ligands, on
the membrane surface, especially near coated pits.
• This triggers the formation of a vesicle

Fig. 8.19c

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• Receptor-mediated endocytosis enables a cell to
acquire bulk quantities of specific materials that
may be in low concentrations in the environment.
• Human cells use this process to absorb cholesterol.
• Cholesterol travels in the blood in low-density
lipoproteins (LDL), complexes of protein and lipid.
• These lipoproteins bind to LDL receptors and enter the
cell by endocytosis.
• In familial hypercholesterolemia, an inherited disease,
the LDL receptors are defective, leading to an
accumulation of LDL and cholesterol in the blood.
• This contributes to early atherosclerosis.
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