1) Where all do you find membranes?

All share common properties

nucleus peroxisome endosome

endoplasmic lysosome
reticulum

The internal membranes form various organelles, including the

endoplasmic reticulum, Golgi apparatus, endosomes, and
mitochondria (Figure 11–3). Although these internal membranes are
constructed on the same principles as the plasma membrane, they
differ subtly in composition, especially in their resident proteins.

Golgi
transport apparatus
vesicle
plasma
membrane
mitochondrion

Figure 11–3 Internal membranes

form many different compartments
in a eukaryotic cell. Some of the main membrane-enclosed organelles in a
typical animal cell are shown here. Note that
the nucleus and mitochondria are each enclosed by two membranes.

Common Features Biological Membranes

1. Membranes are sheetlike structures, only two molecules thick, that form
closed boundaries between different compartments. The thickness of most
membranes is between 60 Å (6 nm) and 100 Å (10 nm).
2. Membranes consist mainly of lipids and proteins. The mass ratio of lipids
to proteins ranges from 1:4 to 4:1. Membranes also contain carbohydrates
that are linked to lipids and proteins.
3. Membrane lipids are small molecules that have both hydrophilic and
hydrophobic moieties. These lipids spontaneously form closed bimolecular
sheets in aqueous media. These lipid bilayers are barriers to the flow of polar
molecules.
4. Specific proteins mediate distinctive functions of membranes. Proteins serve
as pumps, channels, receptors, energy transducers, and enzymes. Membrane
proteins are embedded in lipid bilayers, which create suitable environments
for their action.
5. Membranes are noncovalent assemblies. The constituent protein and lipid
molecules are held together by many noncovalent interactions, which act
cooperatively.
6. Membranes are asymmetric. The two faces of biological membranes
always differ from each other.
7. Membranes are fluid structures because the lipids and many of the
proteins are free to rotate and move sideways in their own half of the
bilayer.
8. Most cell membranes are electrically polarized, such that the inside is
negative [typically 260 millivolts (mV)]. Membrane potential plays a key
role in transport, energy conversion, and excitability.
2)An Overview of Membrane Functions
1. Compartmentalization. Membranes are continuous, unbroken 7. Energy transduction. Membranes are intimately involved in the
sheets and, as such, inevitably enclose compartments. The plasma processes by which one type of energy is converted to another
membrane encloses the contents of the entire cell, whereas the type (energy transduction). The most fundamental energy trans-
nuclear and cytoplasmic membranes enclose diverse intracellular duction occurs during photosynthesis when energy in sunlight is
spaces. The various membrane‐bounded compartments of a cell absorbed by membrane‐bound pigments, converted into chemi-
possess markedly different contents. Membrane compartmen- cal energy, and stored in carbohydrates. Membranes are also
talization allows specialized activities to proceed without exter- involved in the transfer of chemical energy from carbohydrates
nal interference and enables cellular activities to be regulated and fats to ATP. In eukaryotes, the machinery for these energy
independently of one another. conversions is contained within membranes of chloroplasts and
2. Scaffold for biochemical activities. Membranes not only enclose mitochondria.
compartments but are also a distinct compartment themselves.
For reactants floating around in solution, their interactions are
dependent on random collisions. In contrast, components that 8. The mechanical properties of the plasma membrane are
are embedded in a membrane are no longer floating free and can equally impressive. When a cell grows, so does its
be ordered for effective interaction. membrane: this remarkable structure enlarges in area by
adding new membrane without ever losing its continuity,
3. Providing a selectively permeable barrier. Membranes prevent
and it can deform without tearing, allowing the cell to
the unrestricted exchange of molecules from one side to the
move or change shape (Figure 11−2). The membrane is
other. At the same time, membranes provide the means of com-
also self-healing: if it is pierced, it nei-ther collapses like a
munication between the compartments they separate. The
balloon nor remains torn; instead, the membrane quickly
plasma membrane, which encircles a cell, can be compared to a
reseals.
moat around a castle: Both serve as a general barrier, yet both
have gated “bridges” that promote the movement of select
elements into and out of the enclosed living space. 1 receiving
information
macromolecules. The plasma membrane is also able to transport
specific ions, thereby establishing ionic gradients across itself. This
capability is especially critical for nerve and muscle cells.
5. Responding to external stimuli. The plasma membrane plays a
critical role in the response of a cell to external stimuli, a process 3 capacity for
movement and
known as signal transduction. Membranes possess receptors expansion
that combine with specific molecules (ligands) or respond to
other types of stimuli such as light or mechanical tension.
Different types of cells have membranes with different receptors
and are, therefore, capable of recognizing and responding to dif- 2 import and
ferent environmental stimuli. The interaction of a plasma mem- export of
small molecules
brane receptor with an external stimulus may cause the membrane
to generate a signal that stimulates or inhibits internal activities.
For example, signals generated at the plasma membrane may tell
a cell to manufacture more glycogen, to prepare for cell division,
to move toward a higher concentration of a particular com-
pound, to release calcium from internal stores, or possibly to
commit suicide.
6. Intercellular interaction. Situated at the outer edge of every liv-
ing cell, the plasma membrane of multicellular organisms medi-
ates the interactions between a cell and its neighbors. The plasma
membrane allows cells to recognize and signal one another, to
adhere when appropriate, and to exchange materials and infor-
mation. Proteins within the plasma membrane may also facilitate
the interaction between extracellular materials and the intracel-
lular cytoskeleton.
3 What are membranes made up of?
Cell Membrane

consists of

Cholesterol Phospholipids, Sphingolipids Carbohydrates Proteins

together form together form together form

Lipid bilayer Glycolipids Glycoproteins

functions as
whose functions include
Selective barrier between cytosol
and external environment Structural stability Cell recognition Immune response

TABLE 3.1 Composition of Selected Membranes

Membrane Protein Lipid Carbohydrate
Red blood cell membrane 49% 43% 8%

Myelin membrane around nerve cells 18% 79% 3%

Inner mitochondrial membrane 76% 24% 0%

Membrane protein content is highly variable. The amount of proteins equals or exceeds the quantity of lipid in nearly all membranes. The
outstanding exception is myelin, an electri-cal insulator found on many nerve fibers.
 A Typical
Membrane

Plasma membrane

Sphingolipids +
cholesterol Transport
vesicle Cell Membranes of An
Actual Cell

Phosphatidylethanolamine
+ phosphatidylserine

Phosphatidylcholine +
sphingolipids + cholesterol

Trans-Golgi
network

Phosphatidylethanolamine
+ phosphatidylserine

Phosphatidylethanolamine
+ phosphatidylserine

Golgi

Phosphatidylcholine

The distribution of lipids in the membranes of a typical cell. Each

membrane has its own characteristic composition, and the two monolayers of
a given membrane may differ in composition as well.
 4 The Fluid Mosaic Model

Outside of cell Carbohydrates are attached In animal cells, some

to the outer surface of proteins membrane proteins
(forming glycoproteins) or lipids associate with filaments
(forming glycolipids). in the extracellular matrix.

Phospholipid
bilayer

Peripheral membrane Cholesterol molecules interspersed Some membrane Some integral proteins
proteins do not penetrate among phospholipid tails in the proteins interact with the cross the entire phospholipid
Inside of cell the bilayer at all. bilayer influence the fluidity of interior cytoskeleton. bilayer; others penetrate only
fatty acids in the membrane. partially into the bilayer.

6.1 The Fluid Mosaic Model The general molecular structure of bio-
logical membranes is a continuous phospholipid bilayer which has pro-
teins embedded in or associated with it.

• Hydrophilic regions: The phosphorus-containing

“head” of the phospholipid is electrically
charged and therefore associates with polar
water molecules.
• Hydrophobic regions: The long, nonpolar fatty acid
“tails” of the phospholipid associate with other
nonpolar materials, but they do not dissolve in
water or associate with hydrophilic substances.
Because of these properties, one way in which phos-
pholipids can coexist with water is to form a bilayer,
with the fatty acid “tails” of the two layers interact-
ing with each other and the polar “heads” facing
the outside aqueous environment. The thick-ness of
a biological membrane is about 8 nm (0.008 μm),
which is twice the length of a typical phospho-lipid
—another indication that the membrane consists of a
lipid bilayer. This thickness is about 8,000 times
thinner than a piece of paper.
 lateral diffusion 5 The Lipid Bilayer Is a Flexible Two-dimensional Fluid
Phospholipid molecules very rarely tumble from one half of the bilayer,
or monolayer, to the other. Without proteins to facilitate the process, it
flip-flop is estimated that this event, called “flip-flop,” occurs less than once a
(rarely occurs)
month for any individual lipid molecule under conditions similar to
those in a cell. On the other hand, as the result of random thermal
motions, lipid molecules continuously exchange places with their
flexion rotation neighbors within the same monolayer. This exchange leads to rapid
Figure 11–14 Membrane phospholipids lateral diffusion of lipid molecules within the plane of each
move within the lipid bilayer. Because of monolayer.
these motions, the bilayer behaves as a two-
dimensional fluid, in which the individual Individual lipid molecules not only flex their hydrocarbon tails, but they
lipid molecules are able to move in their also rotate rapidly about their long axis—some reaching speeds of
own monolayer. Note that lipid molecules
do not move spontaneously from one
500 revolutions per second. .
monolayer to the other.

6) Membrane Assembly Begins in the ER

IN THE ER MEMBRANE, PHOSPHOLIPIDS ARE RANDOMLY DISTRIBUTED

But most cell membranes are asymmetric: the two halves of the
bilayer often include strikingly different sets of phospholipids. But if
membranes emerge from the ER with an evenly assorted set of
phospholipids, where does this asymmetry arise? It begins in the Golgi
apparatus.
The Golgi membrane contains another family of phospholipid-handling
transporters, called flippases. Unlike scramblases, which move random
phospholipids from one half of the bilayer to the other, flippases remove
specific phospholipids from the side of the bilayer facing the exterior
space and flip them into the monolayer that faces the cytosol
 7) The Fluidity of a Lipid Bilayer Depends on Its Composition

The fluidity of a cell membrane—the ease with which its lipid molecules move within the
plane of the bilayer—is important for membrane func-tion and has to be maintained within
certain limits. Just how fluid a lipid bilayer is at a given temperature depends on its
phospholipid composition.

CH2 N+(CH3)3
CH2
O
_
O P O

O
CH2 CH CH2
O O
C O C O
CH2 CH2
CH2 CH2
CH2 CH2
CH2 CH2
CH2 CH2
CH2 CH2
CH2 CH2
CH2 CH double
bond
CH2 CH
CH2 CH2
CH2
CH2
CH2
CH2
CH2
CH2 CH2
CH2 CH2
CH2 CH2
CH3
CH2
CH3

For all cells, membrane fluidity is important for a number of reasons. It

enables many membrane proteins to diffuse rapidly in the plane of the
bilayer and to interact with one another, as is crucial, for example, in
cell signaling (discussed in Chapter 16). It permits membrane lipids and
proteins to diffuse from sites where they are inserted into the bilayer after
their synthesis to other regions of the cell. It ensures that membrane mol-
ecules are distributed evenly between daughter cells when a cell divides.
And, under appropriate conditions, it allows membranes to fuse with one
another and mix their molecules (discussed in Chapter 15). If biological
membranes were not fluid, it is hard to imagine how cells could live,
grow, and reproduce.
8) MEMBRANE PROTEINS
Although the lipid bilayer provides the basic structure of all cell mem-
branes and serves as a permeability barrier to the hydrophilic molecules
on either side of it, most membrane functions are carried out by mem-
brane proteins.

TRANSPORTERS AND ANCHORS RECEPTORS ENZYMES Figure 11–20 Plasma membrane

CHANNELS proteins have a variety of functions.
They transport molecules and ions, act
EXTRACELLULAR
SPACE
as anchors, detect signals, or catalyze
reactions.

CYTOSOL

X Y

Each type of cell membrane contains a different set of proteins,

reflecting the specialized functions of the particular membrane.

Membrane proteins largely reflect the functions a cell

can perform.
9 Membrane Proteins Associate with the Lipid Bilayer in Different Ways

(A) (B) (C) (D)

MONOLAYER-
TRANSMEMBRANE ASSOCIATED LIPID-LINKED PROTEIN-ATTACHED

NH2

P P
EXTRACELLULAR SPACE

lipid
bilayer

CYTOSOL

COOH

integral membrane proteins peripheral membrane proteins

Membrane proteins can associate with the lipid bilayer in different ways.
(A) Transmembrane proteins can extend across the bilayer as a single α helix, as multiple α
helices, or as a rolled-up β sheet (called a β barrel).
(B) Some membrane proteins are anchored to the cytosolic half of the lipid bilayer by an
amphipathic α helix.
(C) Others are linked to either side of the bilayer solely by a covalently attached lipid
molecule (red zigzag lines).
(D) Many proteins are attached to the membrane only by relatively weak, noncovalent
interactions with other membrane proteins.
(A−C) are examples of integral membrane proteins;
the proteins shown in (D) are considered peripheral membrane proteins.

Proteins that are directly attached to the lipid bilayer—whether they are transmembrane, associated with the
lipid monolayer, or lipid-linked—can be removed only by disrupting the bilayer with detergents. Such proteins
are known as integral membrane proteins.

The remaining membrane proteins are classified as peripheral membrane proteins; they can be released
from the membrane by more gentle extraction procedures that interfere with protein–protein interactions but
leave the lipid bilayer intact.

Integral membrane proteins are very firmly associ-ated with the lipid bilayer and are removable only by
agents that interfere with hydrophobic interactions, such as detergents, organic solvents, or denaturants.

Peripheral membrane proteins associate with the membrane through electrostatic interac-tions and hydrogen
bonding with the hydrophilic domains of integral proteins and with the polar head groups of membrane lipids.
They can be released by relatively mild treatments that interfere with electrostatic interactions or break
hydrogen bonds; a com-monly used agent is carbonate at high pH.