CHAPTER
5
MEMBRANE STRUCTURE AND FUNCTION
The complex structure and function of the plasma membrane are described, along with the macromolecules that
comprise the membrane. The mechanisms by which substances move in and out of cells are discussed, as are the
general chemical processes of diffusion and osmosis. Important cell surface modifications and their significance
(e.g., in cellular junctions) are also detailed.
Chapter Outline
5.1 Membrane Models
1. In the early 1900s, researchers noted that lipid-soluble molecules entered cells more rapidly than
water-soluble molecules, suggesting lipids are component of plasma membrane.
2. Later, chemical analysis revealed that the membrane contained phospholipids.
3. Gorter and Grendel (1925) found that the amount of phospholipid extracted from a red blood cell was
just enough to form one bilayer; they also suggested the nonpolar tails were directed inward and polar
heads outward.
4. To account for the permeability of membrane to nonlipid substances, Danielli and Davson (1940s)
proposed the “sandwich” model, with a phospholipid bilayer between layers of protein.
5. Robertson (1950s) proposed that proteins were embedded in an outer membrane and that all
membranes in cells had similar compositions—the “unit membrane” model.
6. Additional research showed great diversity in membrane structure and function.
A. Fluid-Mosaic Model
1. In 1972, Singer and Nicolson introduced the currently accepted fluid-mosaic model.
a. The plasma membrane is a phospholipid bilayer, in which protein molecules are embedded.
b. Embedded proteins are scattered throughout membrane in an irregular pattern; this varies among
membranes.
5.2 Plasma Membrane Structure and Function
1. The plasma membrane is a phospholipid bilayer with embedded proteins.
2. Phospholipids have both hydrophilic and hydrophobic regions; nonpolar tails (hydrophobic) are
directed inward, polar heads (hydrophilic) are directed outward to face both extracellular and
intracellular fluid.
3. The proteins form a mosaic pattern on the membrane.
4. Cholesterol is a lipid found in animal plasma membranes; it stiffens and strengthens the membrane.
5. Glycolipids have a structure similar to phospholipids except the hydrophilic head is a variety of sugar;
they are protective and assist in various functions.
6. Glycoproteins have an attached carbohydrate chain of sugar that projects externally.
7. The plasma membrane is asymmetrical; glycolipids and proteins occur only on outside and
cytoskeletal filaments attach to proteins only on the inside surface.
A. Carbohydrate Chains
1. In animal cells, the glycocalyx is a “sugar coat” of carbohydrate chains; it has several functions.
2. Cells are unique in that they have highly varied carbohydrate chains (a “fingerprint”).
3. The immune system recognizes foreign tissues that have inappropriate carbohydrate chains.
4. Carbohydrate chains are the basis for A, B, and O blood groups in humans.
B. Fluidity of the Plasma Membrane
1. At body temperature, the phospholipid bilayer has the consistency of olive oil.
2. The greater the concentration of unsaturated fatty acid residues, the more fluid the bilayer.
3. In each monolayer, the hydrocarbon tails wiggle, and entire phospholipid molecules can move
sideways.
4. Phospholipid molecules rarely “flip-flop” from one layer to the other.
5. Fluidity of the phospholipid bilayer allows cells to be pliable.
6. Some proteins are held in place by cytoskeletal filaments; most drift in the fluid bilayer.
C. The Functions of the Proteins
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1.
Plasma membrane and organelle membranes have unique proteins; red blood cells (RBC) plasma
membrane contains 50+ types of proteins.
2. Membrane proteins determine most of the membrane’s functions.
3. Channel proteins allow a particular molecule to cross membrane freely (e.g., Cl channels).
4. Carrier proteins selectively interact with a specific molecule so it can cross the plasma membrane
(e.g., Na+-K+ pump).
5. Cell recognition proteins are glycoproteins that allow the body’s immune system to distinguish
between foreign invaders and body cells.
6. Receptor proteins are shaped so a specific molecule (e.g., hormone) can bind to it.
7. Enzymatic proteins carry out specific metabolic reactions.
5.3 Permeability of the Plasma Membrane
1. The plasma membrane is differentially (selectively) permeable; only certain molecules can pass
through.
a. Small non-charged lipid molecules (alcohol, oxygen) pass through the membrane freely.
b. Small polar molecules (carbon dioxide, water) move “down” a concentration gradient, i.e., from
high to low concentration.
c. Ions and charged molecules cannot readily pass through the hydrophobic component of the bilayer
and usually combine with carrier proteins.
2. Both passive and active mechanisms move molecules across membrane.
a. Passive transport moves molecules across membrane without expenditure of energy; includes
diffusion and facilitated transport.
b. Active transport requires a carrier protein and uses energy (ATP) to move molecules across a
plasma membrane; includes active transport, exocytosis, endocytosis, and pinocytosis.
A. Diffusion and Osmosis
1. Diffusion is the movement of molecules from higher to lower concentration (i.e., “down” the
concentration gradient).
a. A solution contains a solute, usually a solid, and a solvent, usually a liquid.
b. In the case of a dye diffusing in water, the dye is a solute and water is the solvent.
c. Once a solute is evenly distributed, random movement continues but with no net change.
d. Membrane chemical and physical properties allow only a few types of molecules to cross by
diffusion.
e. Gases readily diffuse through the lipid bilayer; e.g., the movement of oxygen from air sacs
(alveoli) to the blood in lung capillaries depends on the concentration of oxygen in alveoli.
f. Temperature, pressure, electrical currents, and molecular size influence the rate of diffusion.
2. Osmosis is the diffusion of water across a differentially (selectively) permeable membrane.
a. Osmosis is illustrated by the thistle tube example:
1) A differentially permeable membrane separates two solutions.
2) The beaker has more water (lower percentage of solute) and the thistle tube has less water
(higher percentage of solute).
3) The membrane does not permit passage of the solute; water enters but the solute does not exit.
4) The membrane permits passage of water with a net movement of water from the beaker to the
inside of the thistle tube.
b. Osmotic pressure is the pressure that develops in such a system due to osmosis.
c. Osmotic pressure results in water being absorbed by the kidneys and water being taken up from
tissue fluid.
3. Tonicity is strength of a solution with respect to osmotic pressure.
a. Isotonic solutions occur where the relative solute concentrations of two solutions are equal; a
0.9% salt solution is used in injections because it is isotonic to red blood cells (RBCs).
b. A hypotonic solution has a solute concentration that is less than another solution; when a cell is
placed in a hypotonic solution, water enters the cell and it may undergo cytolysis (“cell bursting”).
c. Swelling of a plant cell in a hypotonic solution creates turgor pressure; this is how plants
maintain an erect position.
d. A hypertonic solution has a solute concentration that is higher than another solution; when a cell
is placed in a hypertonic solution, it shrivels (a condition called crenation).
e. Plasmolysis is shrinking of the cytoplasm due to osmosis in a hypertonic solution; as the central
vacuole loses water, the plasma membrane pulls away from the cell wall.
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B. Transport by Carrier Proteins
1. The plasma membrane impedes passage of most substances but many molecules enter or leave at rapid
rates.
2. Carrier proteins are membrane proteins that combine with and transport only one type of molecule or
ion; they are believed to undergo a change in shape to move the molecule across the membrane.
3. Facilitated transport is the transport of a specific solute “down” or “with” its concentration gradient
(from high to low), facilitated by a carrier protein; glucose and amino acids move across the membrane
in this way.
4. Active transport is transport of a specific solute across plasma membranes “up” or “against” (from
low to high) its concentration gradient through use of cellular energy (ATP).
a. Iodine is concentrated in cells of thyroid gland, glucose is completely absorbed into lining of
digestive tract, and sodium is mostly reabsorbed by kidney tubule lining.
b. Active transport requires both carrier proteins and ATP; therefore cells must have high number of
mitochondria near membranes where active transport occurs.
c. Proteins involved in active transport are often called “pumps”; the sodium-potassium pump is an
important carrier system in nerve and muscle cells.
d. Salt (NaCl) crosses a plasma membrane because sodium ions are pumped across, and the chloride
ion is attracted to the sodium ion and simply diffuses across specific channels in the membrane.
5. Membrane-Assisted Transport
a. In exocytosis, a vesicle formed by the Golgi apparatus fuses with the plasma membrane as
secretion occurs; insulin leaves insulin-secreting cells by this method.
b. During endocytosis, cells take in substances by vesicle formation as plasma membrane pinches
off by either phagocytosis, pinocytosis, or receptor-mediated endocytosis.
c. In phagocytosis, cells engulf large particles (e.g., bacteria), forming an endocytic vesicle.
1) Phagocytosis is commonly performed by ameboid-type cells (e.g., amoebas and
macrophages).
2) When the endocytic vesicle fuses with a lysosome, digestion of the internalized substance
occurs.
d. Pinocytosis occurs when vesicles form around a liquid or very small particles; this is only visible
with electron microscopy.
e. Receptor-mediated endocytosis, a form of pinocytosis, occurs when specific macromolecules
bind to plasma membrane receptors.
1) The receptor proteins are shaped to fit with specific substances (vitamin, hormone, lipoprotein
molecule, etc.), and are found at one location in the plasma membrane.
2) This location is a coated pit with a layer of fibrous protein on the cytoplasmic side; when the
vesicle is uncoated, it may fuse with a lysosome.
3) Pits are associated with exchange of substances between cells (e.g., maternal and fetal blood).
4) This system is selective and more efficient than pinocytosis; it is important in moving
substances from maternal to fetal blood.
5) Cholesterol (transported in a molecule called a low-density lipoprotein, LDL) enters a cell
from the bloodstream via receptors in coated pits; in familial hypocholesterolemia, the LDL
receptor cannot bind to the coated pit and the excess cholesterol accumulates in the
circulatory system.
5.4 Modification of Cell Surfaces
A. Cell Surfaces in Animals
1. Junctions Between Cells are points of contact between cells that allow them to behave in a
coordinated manner.
a. Anchoring junctions mechanically attach adjacent cells.
b. In adhesion junctions, internal cytoplasmic plaques, firmly attached to cytoskeleton within each
cell are joined by intercellular filaments; they hold cells together where tissues stretch (e.g., in
heart, stomach, bladder).
c. In desmosomes, a single point of attachment between adjacent cells connects the cytoskeletons of
adjacent cells.
d. In tight junctions, plasma membrane proteins attach in zipper-like fastenings; they hold cells
together so tightly that the tissues are barriers (e.g., epithelial lining of stomach, kidney tubules,
blood-brain barrier).
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e. A gap junction allows cells to communicate; formed when two identical plasma membrane
channels join.
1) They provide strength to the cells involved and allow the movement of small molecules and
ions from the cytoplasm of one cell to the cytoplasm of the other cell.
2) Gap junctions permit flow of ions for heart muscle and smooth muscle cells to contract.
2. The extracellular matrix is a meshwork of polysaccharides and proteins produced by animal cells.
a. Collagen gives the matrix strength and elastin gives it resilience.
b. Fibronectins and laminins bind to membrane receptors and permit communication between
matrix and cytoplasm; these proteins also form “highways” that direct the migration of cells
during development.
c. Proteoglycans are glycoproteins that provide a packing gel that joins the various proteins in
matrix and most likely regulate signaling proteins that bind to receptors in the plasma protein.
B. Plant Cell Walls
1. Plant cells are surrounded by a porous cell wall; it varies in thickness, depending on the function of
the cell.
2. Plant cells have a primary cell wall composed of cellulose polymers united into threadlike microfibrils
that form fibrils.
3. Cellulose fibrils form a framework whose spaces are filled by non-cellulose molecules.
4. Pectins allow the cell wall to stretch and are abundant in the middle lamella that holds cells together.
5. Non-cellulose polysaccharides harden the wall of mature cells.
6. Lignin adds strength and is a common ingredient of secondary cell walls in woody plants.
7. Plasmodesmata are narrow membrane-lined channels that pass through cell walls of neighboring cells
and connect their cytoplasms, allowing direct exchange of molecules and ions between neighboring
plant cells.
Lecture Enrichment Ideas
Experience Base: This chapter is heavily laden with terminology and chemical concepts that may not be familiar to
many students. Visuals and demonstrations are critical to understanding membrane properties, especially the
discussion of a “fluid” membrane and the concept of “hypotonic” and “hypertonic” as pertains to solutions. Be
aware that students will likely have a negative concept of “cholesterol”; stress that it is necessary for animal life and
is a vital raw material for many important metabolic functions and is the basis of many hormones.
1.
2.
3.
4.
5.
6.
Describe how the sugar residues of glycoproteins and glycolipids are located only on the outside face of the
plasma membrane. Emphasize the importance of these molecules in cellular activity.
Examine scanning electron micrographs of freeze-fractured plasma membrane. Determine which face is the
cytoplasmic and which the external side of the membrane. (The proteins are more frequent in the cytoplasmic
face).
Describe the function of cholesterol in the plasma membranes of animal cells, and discuss why cholesterol is
missing in plant cells.
Describe and discuss the functions of the different kinds of proteins that are located in and attached to the
plasma membrane. Discuss why these membrane-associated proteins are also found in membrane-bound
organelles such as vesicles, vacuoles, mitochondria, and chloroplasts.
Prepare dialysis bags containing different molar solutions of saline, and place them in solutions with different
molarities. Determine which beakers represent a cell in a hypotonic, hypertonic, or isotonic solutions. An egg
with its shell “peeled away” or dissolved away with vinegar can also be used; it can swell to the size of an
orange as water continuously enters. (This experiment should be set up at the beginning of class and analyzed
at the end of class.)
Compare the functions of the different kinds of junctions that hold cells together and with what kinds of cells
they would likely be associated. Discuss the possible effects of various chemical agents that alter or eliminate
the function of each type of junction.
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Critical Thinking
Question 1. If you do not water your houseplants, they will first wilt, then eventually die. Why do wilting and
dying not occur at the same time?
Answer: A plant maintains its rigid shape by holding water inside a large central vacuole; the water pushes against
the plasma membrane, which pushes against the rigid cell wall, and this turgor pressure holds the plant rigid. The
plant wilts when the water level drops to where there is not enough turgor pressure to maintain rigidity, but there is
plenty of water for life processes to continue. Loss of additional water will dehydrate the cell and eventually cause
death of plant tissue.
Question 2. You can “peel” a raw egg without breaking the membrane or “melt” away the shell in vinegar. If you
place it in a glass of distilled, deionized water, it will swell in size until it breaks. Why is the flow one-way?
Answer: The egg white is albumin, a huge protein molecule that cannot pass through the membrane. Water is a
small molecule that can easily pass either way across the membrane. Since the water outside is 100% water and the
albumin cannot leave, the percentage of water inside is always lower than 100 percent; thus the water molecules will
continue to move from higher concentration outside the egg to a lower concentration inside.
Question 3. The DNA of a cell codes for sequences of amino acids in proteins but the main component of a plasma
membrane is phospholipids, a molecule that is not a protein. Where does the “new” plasma membrane come from
when cells reproduce?
Answer: As a lipid, the plasma membrane grows by adding lipid molecules from the cellular environment, a
process called accretion. The unique proteins that are embedded in the membrane would be encoded by DNA but all
non-protein elements of a cell must be developed by other cellular mechanisms.
Technology from McGraw-Hill
Please consult your Course Integration Guide for technology correlations for this chapter.
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