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Alexis Omar López

5 Biological Membranes

Lecture Outline
I. Biological membranes are lipid bilayers with associated proteins
A. Phospholipids form bilayers in water
1. Phospholipids have two fatty acid chains linked to a glycerol
2. The hydrophilic portion of the molecule is the phosphate bonded
to the glycerol
3. The fatty acids make up the hydrophobic portion of the molecule
4. The molecule is amphipathic
a) The bilayer forms spontaneously because of the amphipathic
b) Many common detergents are also amphipathic, and so are
able to "solubilize" oil
B. Current data support a fluid mosaic model of membrane structure
1. Davson and Danielli (1935) – phospholipids form a membrane two
molecules thick: the lipid bilayer
a) Singer and Nicholson (1972) – the fluid mosaic model
b) This membrane is only 10 nanometers thick
c) Various types of proteins move around like icebergs in the
lipid sea
C. Biological membranes are two-dimensional fluids
1. Phospholipids act as liquid crystals, and the molecules move
a) Fluidity is affected by temperature, among other factors
2. Molecules rarely move from one side of the membrane to the
3. Frye and Ediden (1970) demonstrated movement of proteins in
the membrane
4. Various bonds may occur between hydrocarbon chains
a) Saturated fats lack double bonds in the fatty acid chains,
causing the membrane to be less fluid
b) Organisms may alter the proportion of saturation in response
to temperature
5. In animal cells, cholesterol, which is slightly amphipathic,
stabilizes the membrane at higher temperatures
6. In plant cells, other steroids function in a similar manner
D. Biological membranes fuse and form closed vesicles
1. Membrane fusion is due to the liquid crystalline state
2. When vesicles and another membrane fuse, their bilayers and
lumens become continuous
3. Endocytosis and exocytosis are products of membrane fusion.
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E. Membrane proteins include integral and peripheral proteins

1. Integral proteins are firmly bound to the membrane, or are bound
to other lipids that are part of the membrane
2. Transmembrane proteins span the entire bilayer
a) Some transmembrane proteins span the membrane only
once, but many weave back and forth a number of times
b) Transmembrane proteins are amphipathic
c) The most common type is an alpha helix
3. Peripheral proteins are bound to the hydrophilic ends of the
integral proteins
F. Proteins are oriented asymmetrically across the bilayer
1. Evidence comes from freeze-fracture electron microscopy
2. Proteins are produced in the ER, then pass to the Golgi, then pass
via a vesicle to the plasma membrane
3. Various chemical modifications occur along the way
G. Membrane proteins function in transport, information transfer, and
as enzymes
1. Membrane proteins are formed by free ribosomes
2. Proteins are involved in transport of small molecules
3. Enzymes may be embedded in the membrane
4. Receptor proteins are embedded in the exterior surface of the
a) Signal molecules convert an extracellular signal into an
intracellular signal via signal transduction
5. Membrane proteins can serve as identification tags functioning in
cell-to-cell recognition; others form junctions between adjacent

II. Cell membranes are selectively permeable

A. Most biological membranes are permeable to small or lipid-soluble
1. Water molecules may pass the lipid bilayer
2. Gases, small polar molecules, and large hydrophobic substances
may also pass
3. Other molecules move through special channels, primarily
through membrane transport membranes
B. Random motion of particles leads to diffusion
1. Atoms and molecules above absolute zero exhibit motion
2. Due to random motion, particles move from an area of higher
concentration to one of lower concentration, ultimately reaching
3. The rate of diffusion depends on temperature, the size of the
molecules, electrical charges, and the concentration gradient
Alexis Omar López

4. Osmosis is the diffusion of water (solvent) across a selectively

permeable membrane
a) The osmotic pressure is the tendency of water to move into
that solution
b) Two solutions may be isotonic to each other, or one may be
relatively hypertonic and the other relatively hypotonic
(1) Human cells are isotonic, with a 0.9% sodium chloride solution
(2) Solutions that are hypertonic have a higher osmotic pressure 
than the cells; solutions that are hypotonic have a lower 
osmotic pressure than the cells
(a) Animal cells placed in a hypertonic solution tend to shrivel 
and die
(b) Plant cells, and others with cell walls, placed in a 
hypertonic solution tend to plasmolyze
(c) Animal cells placed in a hypotonic solution tend to swell 
and burst
(d) Plant cells placed in a hypotonic solution tend to 
become turgid
(3) Turgor pressure is the internal hydrostatic pressure usually 
present in walled cells
(a) Turgor pressure provides structural support in non­
woody plants
C. Carrier-mediated transport of solutes requires special integral
membrane proteins
1. Aquiporins are integral membrane proteins that function as gated
water channels
2. Impermeability of the cell membrane is advantageous so cells do
not lose valuable polar molecules
3. Two forms of carrier-mediated transport are facilitated diffusion
and carrier-mediated active transport
D. Facilitated diffusion occurs down a concentration gradient
1. A membrane may become permeable by a protein that combines
with the material to be transported
2. Glucose transport across erythrocyte membranes is an example
3. Liposomes are artificial vesicles that have been used to study
facilitated diffusion
E. Some carrier-mediated active transport systems "pump" substances
against their concentration gradients
1. Materials are transported from a region of low concentration to
high by carrier-mediated active transport mechanisms
2. ATP is required
3. The sodium-potassium pump is an example
4. Other mechanisms involve the pumping of ions in photosynthesis
and aerobic respiration, which results in a membrane potential
and electrochemical gradient
F. Linked cotransport systems indirectly provide energy for active
1. These transport systems cotransport molecules against their
concentration gradient
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2. ATP is a requirement for linked cotransport systems

3. Example: glucose transport across microvilli of the intestine
4. More than one transport system may transport one substance
G. Facilitated diffusion is powered by a concentration gradient; active
transport requires another energy source
1. The energy of ATP is used directly or indirectly in transport of
H. The patch-clamp technique has revolutionized the study of ion
1. This was first developed in the mid 1970s at the Max Planck
Institute in Germany
2. With this technique, scientists can study single ion channels of
even very small cells over time
I.In exocytosis and endocytosis, large particles are transported by
vesicles or vacuoles
1. In exocytosis, the cell expels wastes or a secretory product
a) Exocytosis also results in growth of the cell membrane
2. In endocytosis, a cell takes up materials
a) Phagocytosis involves taking in solid materials
b) Pinocytosis involves taking in liquid droplets
3. Receptor-mediated endocytosis involves the bonding of receptor
proteins to the material to be engulfed
a) Molecules bind to receptors, called ligands, concentrated in
coated pits
(1) Coated pits are formed by a protein, clathrin
b) The coated pits move inwards, via endocytosis, and are then
termed coated vesicles
c) The coated vesicle ultimately becomes an endosome when
the coating is lost
d) Cholesterol is taken into cells by this pathway
(1) Brown and Goldstein– 1986 Nobel Prize

III. Cell Signaling is the mechanism by which cells communicate with one
A. An example of a molecule that is a cell signal is cyclic adenosine
monophosphate (cAMP)
1. This is clearly illustrated when studying slime molds
B. Cell signaling involves 6 steps
1. Synthesis and release of the signaling molecules
a) Signaling molecules may be neurotransmitters, hormones, or
other types of molecules
b) Signaling molecules are ligands (often called the first
messengers), which specifically bind to receptors on the
target cells
c) Some ligand-receptor complexes bind to and activate specific
integral membrane proteins: the G proteins
2. Transport to target cells
3. Reception of the information by target cells
a) This is typically composed of a transmembrane protein with a
functional portion on the extracellular surface
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b) Typically there are three domains: an external domain for

docking by the signaling molecule, a second domain
extending through the plasma membrane, and a third domain
resembling a "tail" that extends into the cytoplasm
4. Signal transduction
a) In this process, cells convert and amplify an extracellular
signal into an intracellular one
5. Response by the cell
a) The tail undergoes a conformational change, which then
activates a sequence of proteins that initiate changes within
the cytoplasm
b) The G proteins may pass the message to the second
messenger, often cyclic AMP
c) The second messengers typically activate protein kinases
6. Termination of signaling

IV. Junctions are specialized contacts between cells

A. Anchoring junctions connect epithelial cells
1. A common form of anchoring junction, desmosomes, are points of
attachment between some animal cells
a) Desmosomes hold cells subject to mechanical stresses
b) Desmosomes are composed of intermediate filaments, which
span the gap between two cells
B. Adhering junctions cement cells together with proteins
1. Tight junctions seal off intercellular spaces between some animal
2. Tight junctions seal cells tightly with protein links
3. Tight junctions are important in epithelia
C. Gap junctions permit transfer of small molecules and ions
1. Gap junctions contain pores that connect cells
2. Proteins form the pores
3. Gap junctions allow rapid chemical and electrical communication
between cells
4. The apertures of gap junctions may be controlled
D. Plasmodesmata allow movement of certain molecules and ions
between plant cells
1. Plasmodesmata are connections between plant cells
2. Plasma membranes are continuous through the plasmodesmata
3. A desmotubule may connect the ER of adjacent cells
4. Typically, molecules, but not organelles, pass through
Alexis Omar López

Research and Discussion Topics

• Cystic fibrosis is due to a defect in a membrane pump of the cell

membrane. Discuss the causes and treatments of this disease.
• Discuss the mechanism by which cells use receptor-mediated endocytosis
to take in cholesterol. Investigate the connection to the genetic disorder,
familial hyper-cholesterolemia (FH).
• Compare and contrast the blood-brain barrier to the blood-testes barrier.
How are they similar? How are they different? What are the clinical
applications of these two barriers?