C.

Cell membrane
The cell membrane is also known as the plasma membrane/plasmalemma. It is the outermost
covering of animal cells. It is a semi-permeable membrane composed of lipids, proteins and
carbohydrates. The fundamental structure of the membrane is the phospholipid bilayer, which
forms a stable barrier between two aqueous compartments. The plasma membrane forms the
boundary between the outer environment and living systems. The plasma membrane controls
both the entry and exit of both solute and solvent between the cell and the environment. With
regards to permeability characteristics, a plasma membrane can be semi-permeable,
impermeable, permeable and selectively permeable in nature.

Cell membrane structure

Cell membranes are composed primarily of phospholipids, proteins, cholesterol (sterol), and
carbohydrates.
Phospholipids
Phospholipids are lipids that have two fatty acid tails attached to a glycerol-3-phosphate
molecule, the head. The fatty acid chains are considered the tail of the phospholipid and are
hydrophobic. Hydrophobic is lacking an affinity for water; unable to absorb, or be wetted by
water or water-fearing. The phosphate group is negatively charged, called the "head," and is
hydrophilic. Hydrophilic is having an affinity for water; able to absorb, or be wetted by water or
water loving. Molecules that contain both hydrophobic and hydrophilic parts, are called
amphipathic or amphiphilic. The phospholipids are arranged in a bilayer with the tails pointing in
towards the membrane and the heads facing the extracellular environment and the cytoplasm.
This ensures that the hydrophilic parts of the phospholipids are facing the aqueous parts of the
environment and are interior while the hydrophobic parts are contained within the bilayer.
The phospholipids are packed tightly together and create a seal out of the membrane. They
prevent hydrophilic molecules and large molecules from passing directly through the membrane.
This property of the cell membrane is called selective permeability.
Proteins
Proteins are a type of macromolecule in the cell that is used for many functions, including
structure, support, metabolism and communication. In the cell membrane, there are three types of
proteins. These include integral (intrinsic), peripheral (extrinsic) and transmembrane proteins.
 The intrinsic proteins that are firmly embedded within the phospholipid bilayer and a
portions of these proteins are in Van der Waals contact with the hydrophobic region of the
membrane.These proteins form channels to allow the movement of large molecules and
ions across the hydrophobic layer of the membrane.
 The extrinsic proteins are loosely attached by ionic bonds or calcium bridges to the
electrically charged phosphoryl surface of the bilayer. They carry signals from one
segment of the membrane to another.
 The transmembrane proteins are some integral proteins that span the whole breadth of the
membrane. These proteins can serve as receptors for hormones, neurotransmitters, tissue
specific antigens, growth factors, etc
One of the main functions of proteins in the cell membrane is to help large or hydrophilic
molecules move through the membrane. Some intrinsic proteins that serve this function may act
as pumps, spending energy to move molecules, or they may act as open gates, allowing
molecules to move freely down their concentration gradient. Transport proteins may be always
open, such as aquaporins that regulate the movement of water, or they may be controlled by
small molecules called ligands, such as ligand gated ion channels in neurons. Particles too large
to be diffused or pumped are often swallowed or disgorged whole by an opening and closing of
the membrane.

Proteins of cell membrane

Cholesterol
Cholesterol is a steroid lipid that is small and ring shaped. It fits in between the hydrophobic tails
of the phospholipids and helps regulate fluidity in the membrane. The membrane must be
flexible and able to bend with the cell. Animal cells must move, grow, divide, and perform
functions in the body which require them to change their shape. Thus, the membrane must also
be able to the flex and bend with the cell.

Carbohydrates
Carbohydrates are always found on the exterior surface of cells and are bound either to proteins
(forming glycoproteins) or to lipids (forming glycolipids). These carbohydrate chains may
consist of 2-60 monosaccharide units and can be either straight or branched. Along with
peripheral proteins, carbohydrates form specialized sites on the cell surface that allow cells to
recognize each other. This recognition function is very important to cells, as it allows the
immune system to differentiate between body cells (called “self”) and foreign cells or tissues
(called “non-self”). Similar types of glycoproteins and glycolipids are found on the surfaces of
viruses and may change frequently, preventing immune cells from recognizing and attacking
them. These carbohydrates on the exterior surface of the cell, both glycoproteins and glycolipids
are collectively referred to as the glycocalyx (meaning “sugar coating”). The glycocalyx is
highly hydrophilic and attracts large amounts of water to the surface of the cell. This aids in the
interaction of the cell with its watery environment and in the cell’s ability to obtain substances
dissolved in the water.

Structure of a cell membrane

Molecules that cross cell membrane

i. Small, nonpolar molecules (e.g. oxygen and carbon dioxide): These molecules can pass
through the lipid bilayer and do so by squeezing through the phospholipid bilayers. They
don't need proteins for transport and can diffuse across quickly.
ii. Small, polar molecules (e.g. water): These molecules can also pass through the lipid
bilayer without the help of proteins. Recall that the interior of the phospholipid bilayer is
 made up of the hydrophobic tails. So, it's not easy for water molecules to cross, and it is a
somewhat slower process.
iii. Large, nonpolar molecules (e.g. carbon rings): These rings can pass through but it is also
a slow process.
iv. Large, polar molecules (e.g. simple sugar - glucose): The size and charge of large polar
molecules make it too difficult to pass through the nonpolar region of the phospholipid
membrane without help from transport proteins.
v. Ions (e.g. Na+Na+start text, N, a, end text, start superscript, plus, end superscript):
Similarly, the charge of an ion makes it too difficult to pass through the nonpolar region
of the phospholipid membrane without help from transport proteins.

Functions of cell membrane

 Protecting the integrity of the interior cell.
 Providing support and maintaining the shape of the cell.
 Helps in regulating cell growth through the balance of endocytosis and exocytosis.
 The cell membrane also plays an important role in cell signalling and communication.
 It acts as a selectively permeable membrane by allowing the entry of only selected
substances into the cell.

Biological membranes, structure and function

Biological membranes include cell membrane and internal membranes of organelles. The plasma
membrane usually separates the cell contents from the outer environment. Eukaryotic cells
contain many internal membrane systems that surround the cell organelles. Each internal
membrane system is specialised to assist in the function of organelle it surrounds.

Chemical Composition of biological membranes

Membranes are composed of lipids (lipoproteins and cholesterol), proteins and carbohydrates.
The relative content of these components varies widely from one type of membrane to another,
but typically it contains, 40 per cent of the dry weight is lipids, about 60 per cent proteins and 1
to 10 per cent carbohydrates.

Composition of different membranes: Content of lipid, protein and carbohydrates as

percentage of dry weight
Types of membranes Lipid Protein Carbohydrates
Plasma membrane 43 49 8
Nuclear membrane 35 59 3
Outer mitochondrial membrane 48 52 Trace
Inner mitochondrial membrane 24 76 Trace
Endoplasmic reticulum 44 54 2
Myelin 75 22 3

1. Lipids
Lipids are the basic structural components of cell membranes. Lipid molecules have a ‘polar’ or
ionic head hence hydrophilic and the other end is a ‘nonpolar’ and hydrophobic tail. Hence, they
are amphiphilic or amphipathic.

Basic lipid structure - polar head and nonpolar tails

Types of lipids present in biomembranes
i. Fatty acids
ii. Glycerophospholipids
iii. Sphingolipids
iv. Cholesterol
Fatty acids
They are major components of most membrane lipids. The nonpolar tails of most membrane
lipids are long chain fatty acids attached to polar head groups, such as glycerol-3-Phosphate.
About 50 per cent of the fatty acid groups are saturated, i.e. they contain no double bond. The
most common saturated fatty acid groups in membrane lipids in animals contain 16 to 18 carbon
atoms such as palmitic acid.
The other half of fatty acid molecules contain one or more double bonds, i.e. unsaturated or
polyunsaturated fatty acids. Oleic acid (monounsaturated) is the most abundant unsaturated fatty
acid in animal membrane lipids; others are arachidonic acid (polyunsaturated), linoleic and
linolenic acids. The degree of unsaturation determines the fluidity of the membranes.
Fatty acids and glycerol-3-phosphate
Glycerophospholipids
They are another group of major components of biomembranes. Phosphatidylethanol amine
(cephalin), phosphatidylcholine (Lecithin) and phosphatidylserine are among the most of
common glycerophospholipids.
Sphingolipids
They comprise another group of lipids found in biological membranes specially in the tissues of
nervous system. There are three types of sphingolipids sphingomyelin, cerebrosides and
gangliosides. About 6 per cent of the membrane lipids of grey matter cells in the brain are
gangliosides.
Cholesterol
Cholesterol is another common component of the biomembranes of animals but not of plants and
prokaryotes. It is oriented with its hydrophilic polar heads exposed to water and its hydrophobic
fused ring system and attached hydrocarbon groups buried in the interior. Cholesterol helps to
regulate fluidity of animal membranes.

2. Proteins
There are three types of proteins found in the membranes. The include:
i. Integral
ii. Peripheral
iii. Transmebrance
Integral membrane proteins
Also known as intrinsic membrane proteins, these proteins are deeply embedded in the
membrane and the portions of these proteins are in Van der Waals contact with the hydrophobic
region of the membrane.
Two major integral proteins are found in red cells membrane. They are:
a) Glycophorin. Glycophorins are glycoproteins. It contains 60 per cent carbohydrates by
weight. The oligosaccharides bound to glycophorin are linked to serine, threonine and
asparagine residues. The polypeptide chain of glycophorin contains 130 amino acid
residues. The function of glycophorin are the M and N blood group antigens.
b) Band-3-Protein. It is dimeric having molecular weight of 93,000. The polypeptide chain
of the dimer is thought to traverse the membrane about a dozen time. Band-3-protein
plays an important role in the function of red blood cells. As red blood cells flow through
the capillaries of the lungs, they exchange bicarbonate anions (HCO 3–.) produced, by the
reaction of CO2 and H2O, for chloride (Cl–) ions. This exchange occurs by way of a
channel in band-3-protein, which forms a Pore through the membrane. Thus band-3-
protein is an example of a membrane transport protein.

Peripheral membrane proteins

Also called extrinsic proteins, these proteins may be weakly bound to the surface of the
membrane by ionic interactions or by hydrogen bonds that form between the proteins and the
‘polar’ heads of the membrane lipids. They may also interact with integral membrane proteins.
They can be removed without disrupting the membrane. The inner face of the red blood cells
membrane is laced with a network of proteins called cytoskeletons that stabilises the membrane
and is responsible for the biconcave shape of the cells. The special peripheral membrane proteins
participate in this stability of red cells are spectrin, actin, and Ankyrin and Band 4, 1 protein.

Transmembrane proteins
Some of the integral proteins span the whole breadth of the membrane and are called as
transmembrane proteins. The hydrophobic side chains of the amino acids are embedded in the
hydrophobic central core of the membrane. These proteins can serve as receptors for hormones,
neurotransmitters, tissue specific antigens, growth factors, etc. Examples include α-helix protein,
transmembrane α-helical protein, and transmembrane β-barrel protein.

3. Carbohydrates
Carbohydrates are the third major component of plasma membranes. They are always found on
the exterior surface of cells and are bound either to proteins (forming glycoproteins) or to lipids
(forming glycolipids). These carbohydrate chains may consist of 2–60 monosaccharide units and
can be either straight or branched. Along with peripheral proteins, carbohydrates form
specialized sites on the cell surface that allow cells to recognize each other. This recognition
function is very important to cells, as it allows the immune system to differentiate between body
cells (called “self”) and foreign cells or tissues (called “non-self”). Glycophorin is a major
integral membrane glycoprotein of human erythrocytes. It has 130 amino acid residues and spans
the lipid membrane, with polypeptide regions outside both the external and internal
(cytoplasmic) surfaces. Carbohydrate chains are attached to the amino terminal portion outside
the external surface. Carbohydrates are also present in apoprotein B of plasma lipoproteins.
Chemical Composition of biological membranes

Components Location Functions

Phospholipid The main fabric of plasma It provides selective permeability to the

membrane cell membrane.

The hydrophilic phosphate side is

outwards and hydrophobic inwards.
Carbohydrates Attached to proteins on outside It helps in cell-to-cell recognition.
membrane layers
Cholesterol Between phospholipids and It helps the plasma membrane to retain its
phospholipid bilayers fluidity.
Proteins Embedded within or on the surface These form channels to allow the
of phospholipid layers movement of molecules.

Formation of Lipid Bilayer

Membrane glycerophospholipids and sphingolipids spontaneously form bilayers, which is the
basis of living biological membranes. Lipid bilayers are oriented with their hydrophobic tails
inside the bilayer while hydrophilic ‘polar’ heads are in contact with the aqueous solution on
each side. Not all lipids can form bilayers. A lipid bilayer can form only when the cross-sectional
areas of the hydrophobic tail and hydrophilic polar head are about equal. Glycerophospholipids
and sphingolipids fulfil this criteria and hence can form bilayer. The lysophospholipids have only
one fatty acyl group, it cannot form the bilayer as the polar heads are too large, similarly
cholesterol also cannot form bilayers as the rigid fused ring systems and additional nonpolar tails
are too large.
The hydrophobic effect and the solvent entropy provide the driving force for the formation of
lipid bilayer. The major interaction responsible for stabilizing plasma membrane is hydrophobic
interactions. A lipid bilayer is about 6 nm across, and this is so thin that it may be regarded as a
two-dimensional fluid.

Lipid bilayer
Fluid Mosaic Model of Membrane Structure
The fluid mosaic model of membrane structure proposed by Singer and Nicholson in 1972 is
now accepted widely. The fluid mosaic model explains various characteristics regarding the
structure of functional cell membranes as a mosaic of components such as phospholipids,
proteins, cholesterol, and carbohydrates. These components give a fluid character to the
membranes. Each phospholipid has a hydrophilic head pointing outside and a hydrophobic tail
forming the inside of the bilayer. Cholesterol and proteins are embedded in the bilayer that gives
the membrane a mosaic look. Each component has a specific function to perform. The
phospholipid bilayer gives fluidity and elasticity to the membrane. The membrane proteins,
intrinsic proteins (integral) deeply embedded and peripheral proteins loosely attached, float in an
environment of fluid phospholipid bilayers. These molecules are constantly moving in two
dimensions, in a fluid fashion, similar to icebergs floating in the ocean. The movement of the
mosaic of molecules makes it impossible to form a completely impenetrable barrier.
The model demonstrated that it is not only the integral proteins, but the phospholipids also
undergo rapid redistribution in the plane of the membrane. This diffusion within the plane of the
membrane is termed translational diffusion. It can be quite rapid for a phospholipid molecule.
Within the plane of the membrane, one molecule of phospholipid can move several micrometers
per second. The phase changes, and thus the fluidity of the membrane is highly dependent upon
the lipid composition of the membrane.

Proteins in fluid bilayer

Fluid mosaic model of biomembrane
Effects of Fluidity of Membrane
The fluidity of membrane significantly affects its functions:
i. As membrane fluidity increases, its permeability to water and other small hydrophilic
molecules also increases.
ii. As fluidity increases, the lateral mobility of integral proteins also increases.

Factors that influence cell membrane fluidity

i. Temperature: In a lipid bilayer, the hydrophobic chains of the fatty acids can be highly
aligned or ordered to provide a rather stiff structure. As the temperature increases, the
hydrophobic side chains undergo a transition from the ordered state which is more gel
like or crystalline to a disordered state, taking on a more liquid like or fluid arrangements.
The temperature at which the structure undergoes the transition from ordered to
disordered state, i.e. melts, is called the transition temperature (Tm). The longer and more
saturated fatty acid chains interact more strongly with each other via their longer
hydrocarbon chains and thus cause higher values of Tm. Hence, higher temperatures are
required to increase the fluidity of the bilayer.
ii. Cholesterol: The cholesterol molecules are randomly distributed across the phospholipid
bilayer, helping the bilayer stay fluid in different environmental conditions. The
cholesterol holds the phospholipids together so that they don’t separate too far, letting
unwanted substances in, or compact too tightly, restricting movement across the
membrane. Without cholesterol, the phospholipids in the cells will start to get closer
together when exposed to cold, making it more difficult for small molecules, like gases to
squeeze in between the phospholipids like they normally do. Without cholesterol, the
phospholipids start to separate from each other, leaving large gaps.
iii. Saturated and unsaturated fatty acids: Fatty acids are what make up the phospholipid
tails. Saturated fatty acids are chains of carbon atoms that have only single bonds
between them. As a result, the chains are straight and easy to pack tightly. Unsaturated
fats are chains of carbon atoms that have double bonds between some of the carbons. The
double bonds create kinks in the chains, making it harder for the chains to pack tightly.
These kinks increase the fluidity of the membrane.
Anesthetic effects on membrane fluidity
Anaesthesia means "loss of sensation". Medicines that cause anaesthesia such as ketamine,
morphine, benzocaine spray and isoflurane are called anaesthetics. Anaesthetics are used during
tests and surgical operations to numb sensation in certain areas of the body or induce sleep. This
prevents pain and discomfort and enables a wide range of medical procedures to be carried out.
They act by reversibly binding to fast sodium channels from within nerve fibers, thus preventing
propagation of action potential.
Anesthetics increase the membrane fluidity by preferentially acting on the hydrocarbon interior.
This effect is universal at high concentrations above the clinical range, but in some kinds of
membranes low concentrations of drugs have an ordering effect.

Alcohol effects on membrane fluidity

Ethanol treatment leads to increased membrane fluidity and changes in membrane protein
composition. It can also interact directly with membrane proteins, causing conformational
changes and thereby influencing their function. Ethanol can disrupt the physical structure of
almost any type of membrane, including the plasma membrane, membranes of cell organelles. It
can increase membrane fluidity, which contributes directly to its acute toxicity action. Alterations
in the lipid environment of membranes can influence the functions of membrane-associated
proteins, such as receptors, ion channels, and enzymes, and can therefore alter biological
processes, including membrane transport, enzymatic reactions, and signaling pathways.
Moreover, ethanol can interact directly with membrane proteins and bring about conformational
changes, thereby influencing their functions.