Membranes: Their Structure, Function and Chemistry: Cell Biology

Membranes: their structure,
function and chemistry
Lecture 6
1/11/2020
Cell biology
Chapter 7
Cell Biology, lecture 6, 1/11/2020 1

Functions of membranes Figure 7-2
 Define the boundaries of the cell and its
organelles and act as permeability barrier.
They serve as sites for specific

biochemical functions such as electron
transport during mitochondrial respiration.
 Provide for and regulate transport

processes.
2
Cell Biology, lecture 6, 1/11/2020
Functions of membranes Figure 7-2
Contain protein receptors needed to

detect external signals.
 Provide mechanisms for cell-cell

contact, adhesion and communication.

Functions of membranes; boundaries and
permeability
membrane defines outermost part of living cell
a) Separates physically cell from environment
b) Permeability barrier
c) w/o membranes, important stuff drifts away
d) Must maintain appropriate concentrations of

molecules in smaller volumes
membrane defines outermost part
of living cell
e) Compartmentalize functions, Isolate

incompatible reactions
eg: lysosomes contain hydrolytic enzymes not

compatible with other macromolecules
Some proteins in membranes act as enzymes????

Examples
Membranes are sites of specific
functions
 eg: reactions of oxidative e-transport and
phosphorylation found only
reactions of
photosynthetic
electron transport and
photophosphorylation
found only in the
thylakoid membranes
of chloroplast

role in cell communication and recognition
Plasma membrane interacts with other cells .
 Cells must recognize other cells : immune

response to foreign cells.
 receives communications from other cells
chemical communications = hormones

electrical communications = nerve impulse
mechanical communications = adhesion
 cells transmit communication

Gap junctions (also called communicating junctions) provide
cytoplasmic channels from one cell to an adjacent cell. Gap junctions
consist of special membrane proteins that surround a pore through
which ions, sugars, amino acids, and other small molecules may
pass. Gap junctions are necessary for communication between cells
in many types of tissues, including heart muscle and animal
embryos.
At tight junctions, the membranes of neighboring cells are very tightly
pressed against each other, bound together by specific proteins.
Forming continuous seals around the cells, tight junctions prevent
leakage of extracellular fluid across a layer of epithelial cells.
Desmosomes (also called anchoring junctions) function like
rivets, fastening cells Together into strong sheets. Intermediate
Filaments made of sturdy keratin proteins Anchor desmosomes
in the cytoplasm.
Cell Biology
Biological membranes
Lecture 7
3/11/2020
Chapter 8
Chapter 7, 3/11/2020 membranes

Figure 7-3 Timeline for the Development of the Fluid Mosaic
Model
•The fluid mosaic model of

membrane structure that
Singer and Nicholson
proposed in 1972 was the
culmination of studies that
date back to the 1890s.
•Future developments and

revisions?

Models of membrane structure: An experimental
perspective (Figure 7-3)
 Chalres Overton, 1890, Lipids are

important components of membranes.
 Langmuir, Lipid monolayer with

hydrophilic head and hydrophobic tail.
 Gorter and Grendel, 1925, The basis

of membrane structure is a lipid bilayer.

Models of membrane structure: An experimental
perspective
Previous models didn’t explain how sugars, ions, and other

hydrophilic solutes readily moved into and out of cells even
though pure lipid bilayers are nearly impermeable to water-
soluble substances.
 Davson and Danielli, 1935, Membrane

consist of lipid bilayer coated on both
sides with thin sheets of protein.

Robertson, 1960, Unit membrane (trilaminar)
Trilaminar Appearance of Cellular Membranes.

This TEM of adjacent cells shows their plasma
membranes “railroad track” separated by a small
intercellular space.
Complications of D & D or R models
 Size of membranes under EM is 6-8 nm, lipid

bilayer is 4-5 nm.
distinctiveness of different kinds of membranes

(Protein/ lipid ratios)
(table 7-1 page 181) Mitochonderial inner membrane
3.54, mitochondrial outer membrane 1.22 ....... Why?
Phospholipase extracts 75% of phospholipids
Membrane proteins could not be extracted by water

( organic solvents and detergents).
Composition of several membranes – ratio of
lipids to proteins

Fluid mosaic model of membranes
Proposed by
Singer and Nicolson,
1972
Envisions membrane
as a mosaic of
proteins
discontinuously
embedded in a fluid
lipid bilayer.

Unwin and Henderson: 1975, Most Membrane Proteins Contain
Transmembrane Segments
These transmembrane segments anchor the protein to the

membrane and hold it in proper alignment within the lipid
bilayer.
• Each transmembrane
segment forms an α helix.
• Successive transmembrane
segments are linked to each
other by short loops of
hydrophilic amino acids
that extend into or protrude
from the polar surfaces of
the membrane.

Three main constituents of membranes
 Membrane lipids
 Membrane proteins
 Membrane carbohydrates Chapter 7, 3/11/2020 membranes
Three Types of Membrane Lipid Molecules- all
amphipathic
Glycerol based
phosphoglycerides
 Phospholipids
 Glycolipids Sphingosine based

sphingolipids
 Sterols
Approximately 50% of mass
Lipid bilayers: amphipathic molecules

Examples of polar lipids - Glycerophospholipids and Sphingolipids
• The membrane lipids are composed of the glycerophospholipids and sphingolipids,

which have polar and nonpolar regions.

Three Types of Membrane Lipid Molecules- all amphipathic
Kinds and relative proportions of phospholipids vary

significantly among membranes of various sources.
Sphingomyelin in animal plasma membranes absent

from plants and most bacteria.

Figure 9.8 sphingosine
Sphingolipid
Structure Fatty acid
Polar head
4 major sphingolipids
are:
Found in nerve cell

membranes
brain, nervous system.
Tay-Sachs disease:
accumulation of
ganglioside in brain
and spleen causes
death by age 4.
Three types of membrane lipid molecules- all amphipathic
Phospholipids Sterols
(cholesterol) Glycolipids
serine (sugar lipid)
phosphatidylserine
galactocerebroside
Fat molecules are hydrophobic, whereas phospholipids are
amphipathic.
(A) Triacylglycerol, a fat
molecule, is entirely
hydrophobic.
(B) Phospholipids such as
phosphatidyl-ethanolamine
are amphipathic, containing
both hydrophobic and
hydrophilic portions.
(The third hydrophobic tail of
the triacylglycerol molecule is
drawn here facing upward for
comparison with the
phospholipid, although
normally it is depicted facing
down.)

Figure 11–6 Phosphatidylcholine is the most common phospholipid in cell
membranes
This particular phospholipid is built from five parts: the hydrophilic head, choline, is linked via a phosphate to
glycerol, which in turn is linked to two hydrocarbon chains, forming the hydrophobic tail.
The two hydrocarbon chains originate as fatty acids— that is, hydrocarbon chains with a –COOH group at one end—
which become attached to glycerol via their –COOH groups.
A kink in one of the hydrocarbon chains occurs where there is a double bond between two carbon atoms.

Structure of fatty acids

Factors affecting membrane fluidity
Lecture 9 membranes, 8-3-2020

Degree of saturation of fatty acids

Length of fatty acids & degree of
saturation

Cell Biology
Lecture 8
5/11/2020
Chapter 8

The fluid mosaic model for biological membranes

Amphipathic phospholipids form a bilayer in water.
(A) Schematic drawing of a

phospholipid bilayer in water.
(B) Computer simulation showing the

phospholipid molecules (red heads and
orange tails) and the surrounding water
molecules (blue) in a cross section of a lipid
bilayer.
Phospholipid composition of several kinds of
membranes

Figure 7-13 The Effect of Chain Length and the Number of Double Bonds on the Melting
Point of Fatty Acids
• The melting point of fatty acids:

1. increases with chain length
for saturated fatty acids.
2. decreases dramatically with
the number of double
bonds for fatty acids with a
fixed chain length.
• The 18-carbon fatty acids in
part b are stearate, oleate,
linoleate, and linolenate, with
0, 1, 2, and 3 double bonds,
respectively.

Figure 7-14 The Effect of Unsaturated Fatty Acids on the Packing of Membrane
Lipids
(a) Membrane phospholipids with no unsaturated fatty acids fit together tightly because the fatty acid
chains are parallel to each other. (b) Membrane lipids with one or more unsaturated fatty acids do not
fit together as tightly because the cis double bonds cause bends in the chains, which interfere with
packing. Each of the structures shown is a phosphatidylcholine molecule, with either two 18-carbon
saturated fatty acids (stearate) (part a) or two 18-carbon fatty acids, one saturated (stearate) and the
other with one double bond (oleate) (part b). 8 membranes, 5-11-2020
Lecture
Regulation of Cell Membrane Fluidity
• Most organisms can regulate membrane

fluidity.
• “homoviscous adaptation” in cold-
blooded poikilothermic organisms occurs
by regulating lipid composition:
• Change ratio of 16-carbon to 18-carbon lipids
via hydrolase enzyme (smaller lipids have
lower Tm).
• Change degree of unsaturation (via desaturase
enzyme).
• Important for bacteria, plants, reptiles,
amphibians, hibernating mammals.

Membrane asymmetry

Lipid composition varies in inner & outer
membrane leaflet

Figure 9.16 Distribution of lipids in two monolayers of the human
erythrocyte

Causes of membrane asymmetry

Movement of lipid molecules

Figure 7-11 Demonstration of Lipid Mobility Within Membranes by Fluorescence
Recovery After Photobleaching

The molecular structure common to all steroids
Steroid structures have four fused rings, A, B, C, and D.

Figure 7-6 Structure of the Cholesterol, a Component of Mammalian Cell Membranes
(c) The most common membrane

sterols are cholesterol in animals
and several related phytosterols in
plants.

Figure 11–16 Cholesterol stiffens cell membranes
11_16_Cholesterol.jpg
(A) The structure of cholesterol. (B) How cholesterol fits into the gaps
between phospholipid molecules in a lipid
bilayer. The chemical formula of cholesterol
is shown in Figure 11–7.
Orientation of Cholesterol Molecules in a Lipid Bilayer
(a) Cholesterol molecules are present in both lipid layers

in the plasma membranes of most animal cells, but a
specific molecule is localized to one of the two layers. (b)
Each molecule orients itself in the lipid layer so that its
single hydroxyl group is close to the polar head group of a
neighboring phospholipid molecule, where it forms a
hydrogen bond with the oxygen of the ester bond
between the glycerol backbone and a fatty acid.

Steroids in membranes

Steroids in membranes

The molecular structure common to all steroids
• Cholesterol has a polar head group (OH) and a nonpolar tail.

• Cholesterol and ester derivatives are abundant in
(blood) plasma proteins called lipoproteins.
• Lipoproteins transport cholesterol to tissues for use in
cell membranes and hormone precursors
Cholesterol is traditionally only associated with animal

cells, but derivatives are also identified in plants.
Figure The Structure of Hopanoids
Abundant in petroleum (crude

oil) deposits, suggesting a
prokaryotic (bacterial) role in
formation of oil deposits
(a) A hopanoid, one of a class of

sterol-like molecules that appear
to function in the plasma
membranes of at least some
prokaryotes as sterols do in the
membranes of eukaryotic cells.
(b) The structure of cholesterol, for
comparison. A weakly hydrophilic
side chain (-CH2OH or -OH)
protrudes from each molecule.

Figure 7-9 using thin-layer chromatography to analyze
membrane lipids.
Lipids are extracted from a membrane preparation with a mixture of organic solvents
and separated according to their degree of polarity.
(a) A sample is spotted and dried onto a small area of a glass or metal plate coated
with a thin layer of silica.
(b) Components of the sample are then carried upward by the solvent into which the
TLC plate is placed.
As the solvent moves up the plate by capillary action, the lipids are separated
according to their polarity: Less polar lipids such as cholesterol do not adhere strongly
to the silica and move further up Lecture 8 membranes, 5-11-2020

the plate, while more polar lipids remain closer to the
PDME
phosphatidyldimethylethanolamine
CL cardiolipin
PA phosphatidic acid
PE phosphatidylethanolamine
PG phosphatidylglycerine
PS phosphatidylserine
PI phosphatidylinositol
PC phosphatidylcholine

Human ABO blood groups
involve glycosphingolipids known as A antigen and B
antigen that serve as specific cell surface markers of the
different groups of red blood cells. Cells of blood type
have the A antigen, and cells of blood type B have the B
antigen. Type AB blood cells have both antigen types,
and type O blood cells have neither.

Tay-Sachs disease
GM2 Gangliosidosis
Hexosaminidase A deficiency
Cause: Absence of a lysosomal enzyme, hexosaminidase

A, that is responsible for one of the steps in ganglioside
degradation.
Gangliosides accumulate in the brain and other nervous

tissue, leading to impaired nerve and brain function and
eventually to paralysis, severe mental deterioration, and
death.

Cell Biology
Lecture 9
8/11/2020
Chapter 7
Lecture 9, membranes, 8-11-2020

Figure 7-9 using thin-layer chromatography to analyze
membrane lipids.
Lipids are extracted from a membrane preparation with a mixture of organic solvents
and separated according to their degree of polarity.
(a) A sample is spotted and dried onto a small area of a glass or metal plate coated
with a thin layer of silica.
(b) Components of the sample are then carried upward by the solvent into which the
TLC plate is placed.
As the solvent moves up the plate by capillary action, the lipids are separated
according to their polarity: Less polar lipids such as cholesterol do not adhere strongly
to the silica and move further up Lecture 8 membranes, 5-11-2020

the plate, while more polar lipids remain closer to the
PDME
phosphatidyldimethylethanolamine
CL cardiolipin
PA phosphatidic acid
PE phosphatidylethanolamine
PG phosphatidylglycerine
PS phosphatidylserine
PI phosphatidylinositol
PC phosphatidylcholine

Human ABO blood groups
involve glycosphingolipids known as A antigen and B
antigen that serve as specific cell surface markers of the
different groups of red blood cells. Cells of blood type
have the A antigen, and cells of blood type B have the B
antigen. Type AB blood cells have both antigen types,
and type O blood cells have neither.

Tay-Sachs disease
GM2 Gangliosidosis
Hexosaminidase A deficiency
Cause: Absence of a lysosomal enzyme, hexosaminidase

A, that is responsible for one of the steps in ganglioside
degradation.
Gangliosides accumulate in the brain and other nervous

tissue, leading to impaired nerve and brain function and
eventually to paralysis, severe mental deterioration, and
death.

Freeze-Fracture Analysis of a
Membrane.
Surface view of
monolayers.
This sketch of a freeze-
fractured membrane shows
electron micrographs of
the E and P faces from the
plasma membrane of a
mouse kidney tubule cell.
Individual proteins
imbedded in either face
Figure 7-17 Freeze-fracture electron micrograph of human red blood cells.
Note that the density of intramembrane particles on the

cytosolic (P) face is higher than on the external (E) face.
Fluid mosaic model of membranes
Three classes of membrane proteins:

Protein class Location Interactions
Embedded Held in place by

Integral within lipid the
proteins (Y) bilayer hydrophobic
interactions
Peripheral Located on Linked
Proteins surface of noncovalently
)Z( membrane to the polar
groups of
phospholipids
and proteins
Lipid anchored On the

Proteins periphery but hydrophilic
)X( anchored in the
lipid layer
Membrane proteins can associate with the lipid bilayer in several different
ways
11_21_proteins.associ.jpg
(A) Transmembrane proteins can extend across the bilayer as a single a helix, as multiple a
helices, or as a rolled-up b sheet (called a b barrel).
(B) Some membrane proteins are anchored to the cytosolic surface by an amphipathic a helix.
(C) Others are attached to either side of the bilayer solely by a covalent attachment to a lipid
molecule (red zigzag lines).
(D) Finally, many proteins are attached to the membrane only by relatively weak, noncovalent
interactions with other membrane proteins.
Figure 7-19 The Main Classes of Membrane Proteins
Membrane proteins are classified according to their mode of attachment to the membrane. Integral
membrane proteins contain one or more hydrophobic regions that are embedded within the lipid
bilayer. Peripheral membrane proteins are too hydrophilic to penetrate into the membrane but are
attached to the membrane by electrostatic and hydrogen bonds that link them to adjacent
membrane proteins or to phospholipid head groups. Lipid-anchored proteins are hydrophilic and do
not penetrate into the membrane; they are covalently bound to lipid molecules that are embedded in
the lipid bilayer. (f) Proteins on the inner surface of the membrane are usually anchored by either a
fatty acid or a prenyl group. (g) On the outer membrane surface, the most common lipid anchor is
glycosylphosphatidylinositol (GPI). Lecture 9, membranes, 8-11-2020
Transmembrane protein
Glycophorine one transmembrane span
 Hydrophobic amino acid
residues span membrane
 Hydrophilic domains on both

sides of membrane
 Only outer domain has

covalently attached carbohydrates
Extracted with detergents

Transmembrane proteins span the
bilayer
a-helix
transmembrane
domain
Hydrophobic R groups of
a.a. interact with fatty acid
chains

Integral membrane protein
 Most membrane proteins have multiple

transmembrane spans.
Bacteriorodpsin
More difficult to work with than water soluble

protein
Peripheral protein
 No transmembrane spans.
 Located on surface of membrane .
 Usually bound electrostatically to membrane.

Peripheral protein
 No hydrophobic interactions with interior

of membrane
 bind to other proteins
 bind to lipid head groups.
 Peripheral proteins much easier to isolate

(like water soluble protein)

Lipid anchored proteins
 Hydrophilic proteins that don’t penetrate into the

membranes.
 Covalently bound to lipid molecules that are

embedded in lipid bilayer.

Cell Biology
Lecture 10
8/11/2020
Chapter 7
Functions of membrane proteins
91



Figure 11–20 Plasma membrane proteins have a variety of
functions.

Light transduction
Absorb light:
 Rhodopsin: absorbed light triggers nervous impulse.
 Bacteriorhodopsin: uses light energy to transport H+ across

membrane.
 Light harvesting proteins Transfer light energy

 Reaction center proteins to other protein

Electron transport proteins
Transfer e- from one molecule to another molecule
examples:
 Cytochrome C
 Ferredoxin
 Plastocyanin

Receptor proteins
 Bind other molecules

 Elicit cell response
example: acetylcholine receptor : opens Na+

channels

Membrane carbohydrates
 Approximately 2-10 % of mass
 Confined mainly to the non-cytosolic surface:-
- On the extracellular surface of the cells

- Inward toward the lumen of the compartment


Membrane carbohydrates
 Covalent linkage to
proteins and lipids
Glycoproteins
Proteoglycans
Glycolipids

Externally bound membrane
carbohydrates
Most carbohydrate
attached to protein
 Some carbohydrates
attached to lipid
Glycophorin

In many animal cells, the carbohydrate groups of plasma
membrane glycoproteins and glycolipids protrude from the cell
surface and form a surface coat called the glycocalyx (meaning
“sugar coat”).
they are important components of the recognition sites of

membrane receptors, in antibody-antigen reactions, and in
intercellular adhesion to form tissues.
Glycocalyx surrounding
animal egg cell

Glycocalyx

Glycocalyx of Streptococcus enables it to escape
detection & destruction by immune system

Membrane carbohydrates bound to the
internal surface of lipid bilayer
Covalently bound carbohydrates to the internal surface of
 Golgi vesicles
 Secretion vesicles
 Lysosomes also have

RBC plasma membrane
composition
(by weight)
An erythrocyte is a small, disk-shaped cell with a diameter of
about 7 μm. A mammalian erythrocyte contains no nucleus or
other organelles, which makes it easy to obtain very pure
plasma membrane preparations without contamination by
organelle membranes.
1. 52% protein
2. 40% lipid
3. 8% carbohydrate by weight
Note: Most of the membrane mass IS NOT due to

lipids!!!
(Spectrin and actin are peripheral proteins associated with
membrane, but not in bilayer)
Peripheral membrane proteins

Structural Features of the Erythrocyte Plasma
Membrane
integral proteins
a. Glycophorin
b. Anion
channel
peripheral
proteins
a)Spectrin
b)Ankyrin
c) Actin
d)Band 4.1
FIGURE 7-28 demonstration of the
mobility of membrane proteins by cell
fusion. The mobility of membrane
proteins can be shown experimentally
by the mixing of membrane proteins that
occurs when cells from two different
species (mouse and human) are fused
and the membrane proteins are labeled
with specific fluorescent antibodies.

Fluid Mosaic Model

Blood group antigens are
glycosphingolipids

Quiz
The effects of temperature and lipid composition on
membrane fluidity are often studied by using artificial
membranes containing only one or a few kinds of lipids
and no proteins. Assume that you have made the
following artificial membranes:
 Membrane 1: Made entirely from phosphatidylcholine

with saturated 16-carbon fatty acids.
 Membrane 2: Same as membrane 1, except that each

of the 16-carbon fatty acids has a single cis double
bond.

Quiz
Membrane 3: Same as membrane 1, except that each of the
saturated fatty acids has only 14 carbon atoms.
After determining the transition temperatures of samples

representing each of the membranes, you discover that your
lab partner failed to record which membranes the samples
correspond to. The three values you determined are –36°C,
23°C, and 41°C. Assign each of these transition temperatures
to the correct artificial membrane, and explain your
reasoning.

Quiz
For each of the following false statements, change the
statement to make it true and explain your reasoning?
• Because membranes have a hydrophobic interior, polar and

charged molecules cannot pass through membranes.
• Different cellular organelles have membranes with an

identical chemical composition.
• Glycoproteins are proteins contaning oligosaccharide chains

which protrude from the inner membranes.
.

Quiz
• Membranes fluidity is affected by

temperature . When temperature decreases,
membrane fluidity increases, and the
temperature at which this occurs is known
as transition temperature Tm
• You would expect membrane lipids from

tropical plants such as palm and coconut to
have short fatty acids with multiple C=C
double bonds

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Membranes: their structure,

function and chemistry

Cell Biology, lecture 6, 1/11/2020 1

They serve as sites for specific

 Provide for and regulate transport

Contain protein receptors needed to

 Provide mechanisms for cell-cell

Cell Biology, lecture 6, 1/11/2020 3

a) Separates physically cell from environment

c) w/o membranes, important stuff drifts away

d) Must maintain appropriate concentrations of

e) Compartmentalize functions, Isolate

eg: lysosomes contain hydrolytic enzymes not

Some proteins in membranes act as enzymes????

Cell Biology, lecture 6, 1/11/2020 7

Plasma membrane interacts with other cells .

 Cells must recognize other cells : immune

 receives communications from other cells

chemical communications = hormones

 cells transmit communication

Cell Biology, lecture 6, 1/11/2020 8

Chapter 7, 3/11/2020 membranes

•The fluid mosaic model of

•Future developments and

Chapter 7, 3/11/2020 membranes

 Chalres Overton, 1890, Lipids are

 Langmuir, Lipid monolayer with

 Gorter and Grendel, 1925, The basis

Chapter 7, 3/11/2020 membranes

Previous models didn’t explain how sugars, ions, and other

 Davson and Danielli, 1935, Membrane

Chapter 7, 3/11/2020 membranes

Trilaminar Appearance of Cellular Membranes.

 Size of membranes under EM is 6-8 nm, lipid

distinctiveness of different kinds of membranes

Phospholipase extracts 75% of phospholipids

Membrane proteins could not be extracted by water

Chapter 7, 3/11/2020 membranes

Chapter 7, 3/11/2020 membranes

These transmembrane segments anchor the protein to the

Chapter 7, 3/11/2020 membranes

 Glycolipids Sphingosine based

Lipid bilayers: amphipathic molecules

Chapter 7, 3/11/2020 membranes

• The membrane lipids are composed of the glycerophospholipids and sphingolipids,

Chapter 7, 3/11/2020 membranes

Kinds and relative proportions of phospholipids vary

Sphingomyelin in animal plasma membranes absent

Chapter 7, 3/11/2020 membranes

Found in nerve cell

Chapter 7, 3/11/2020 membranes

Chapter 7, 3/11/2020 membranes

Chapter 7, 3/11/2020 membranes

Lecture 9 membranes, 8-3-2020

Lecture 9 membranes, 8-3-2020

Lecture 9 membranes, 8-3-2020

Lecture 8 membranes, 5-11-2020

Lecture 8 membranes, 5-11-2020

(A) Schematic drawing of a