Cell Biology: Theory and Lab Skills

Outcome 1: Describe and explain the

structure and function of the cell
membrane.

H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Outcome 1: Describe and explain the structure and function of the cell
membrane.

On completion of this unit you should be able to do the following:

1. Describe the chemical structure and function of membrane lipids:
phospholipids, sphingolipids, and cholesterol.
2. Explain the fluid nature of the cell membrane with relation to the lipid
content.
3. Describe the nature of common membrane proteins.
INTRODUCTION

Cells are the fundamental building blocks of all living organisms. A single cell can
survive on its own (unicellular organism), or it can be part of a large unit of cells
(multicellular organism). Cells require nutrients and oxygen to survive. They also
must remove waste materials efficiently.

To survive then cells must have:

Energy source

Oxygen

Water

Waste products e.g. CO2 removed

A cell can obtain nutrients & get rid of waste by exchanging material with its
environment. It must do this efficiently to survive. If exchange of materials is not
carried out the cell will be deprived of nutrients and oxygen, and will be poisoned
from a build-up of toxic waste materials if these are not removed.
Exchange of materials happens at the cell membrane also known as the
plasma membrane through the processes of diffusion and active transport.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

OUTCOME 1.1: Describe the chemical structure and function of two of the

membrane lipids: phospholipids, sphingolipids, and cholesterol.

Structure of the Cell Membrane

The cell membrane is critical to the survival of the cell as it is the boundary
between the cell & its environment. The membrane keeps cytoplasm in & harmful
particles out, but it is not impenetrable; it is semi-permeable. This is important
because otherwise new nutrients couldnt get in & waste couldnt get out. The cell
membrane is also called the plasma membrane and is more like a tightly woven
fabric as opposed to a solid wall. The cell membrane allows some molecules, for
example respiratory gases, to move freely across it. Other substances, like ions
and glucose, require special mechanisms. The cell membrane also serves as
the attachment site for signalling molecules as well as internal structures, such
as the cytoskeleton. In prokaryotes the cell membrane is crucial in cellular
processes such as ATP production and DNA replication.

Glycoprotein

Phospholipid

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Cholesterol

H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Cell Membrane

The Fluid-mosaic model was first described by Singer & Nicolson in 1972. It is
a term used to describe how parts of the membrane interact (lipid & protein).
Biological membranes are lipid bilayers. These lipids form a viscous solvent into
which proteins can be inserted/attached/integrated.

There are three main types of lipids present in the lipid bilayer:
1. Phospholipids
2. Sphingolipids
3. Cholesterol
Amphipathic is a word used to describe the lipid molecules present in the
plasma membrane. This means that part of the lipid molecule is hydrophilic
(water loving) and part of the lipid is hydrophobic (water fearing). This property
allows the lipids to arrange themselves in such a way as to maximise the
interactions between the hydrophobic regions of the lipids (they interact in the
centre of the lipid bilayer holding the 2 leaves of the membrane together) while
allowing the hydrophilic region of the lipid to interact with the aqueous
environment inside and outside the cell. This is the most energetically favourable
arrangement for these molecules.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

1. Phospholipids
Phospholipids are the most common type of molecule found in biological
membranes and are composed of 2 parts;
a. Negatively charged phosphate based polar head hydrophilic
b. Hydrophobic (water fearing) hydrocarbon chain

Phosphate head
Phospholipid
Lipid tail

A phospholipid
The main class of phospholipids are the phosphoglycerides. They are composed
of a glycerol molecule to which 2 fatty acid chains are added to the first and
second carbon of the glycerol molecule. A polar group is attached to the third
carbon via a phosphodiester linkage.
The phospholipid is named after the polar group if choline is added then the
resulting phospholipid is called phosphatidylcholine (one of the most common
phospholipids in some membranes). The simplest phosphoglyceride is
phosphatidic acid.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

The fatty acid chains in phosphoglycerides can vary in length from 14 24

carbon atoms long. They generally fall into one of two types:
a. Saturated
b. Unsaturated
Fatty acid chains that are saturated have the maximum amount of hydrogen
possible associated with that chain. These molecules have the greatest ability to
pack tightly together. Unsaturated fatty acid chains on the other hand have
double bonds between some of the carbon. When this happens this introduces a
kink into the chains meaning that the phospholipids cannot pack as tightly
together as saturated phospholipids can. A large amount of unsaturated fatty
acids in the phospholipid layer increases membrane fluidity (discussed later).

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

The introduction of a double bond into the carbon chain of the fatty acid
introduces a kink (or bend) into the carbon chain. In one group of unnatural fats
called trans fats this kink is not introduced. Membranes with a high content of
trans fats will be less fluid due to the structure of trans fats being more similar to
saturated fats.

2. Sphingolipids

Sphingolipids have a structure similar to phospholipids (polar head, 1 fatty acid

chain) but they contain no glycerol. Instead of glycerol they contain a molecule
called sphingosine. Sphingosine itself is composed of a fatty acid chain and an
attachment sites for the polar head and other fatty acid chain.

Polar
Head

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

There are 3 types of sphingolipids:

a) Ceramide polar head is simply an alcohol group
b) Sphingomyelin phosphocholine polar head
c) Glycosphingolipids sugar molecule for the polar head

Sphingolipids are often found in the outer face of the plasma membrane where
they act as cell surface markers. The glycolipids that make up the ABO blood
group antigens are sphingolipids.
3. Cholesterol

Another type of lipid found in the cell membrane is cholesterol. In some

membranes cholesterol makes up as much as 20% of the lipid content by weight
while others like bacterial membranes have none.

Cholesterol belongs to a group of biomolecules called Steroids. At their most

basic level steroids are composed of 17 carbon atoms arranged into 4 fused ring
structures 3 x 6 carbon rings and 1 x 5 carbon ring. Cholesterol molecules are
short and rigid. It consists of 4 fused carbon rings (3 x 6 carbon rings and 1 x 5
carbon rings) attached to an alcohol group at one end and a hydrocarbon tail at
the other end. Cholesterol is therefore also an amphipathic molecule allowing it
to be embedded into the plasma membrane. Cholesterol plugs any gaps in the

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

plasma membrane and thus decreases the permeability of the membrane to

small water-soluble molecules.

Fused Steroid Rings

C
Polar Head

Hydrocarbon Tail

OUTCOME 1.2: Explain the fluid nature of the cell membrane with relation
to the lipid content.
Properties of the Lipid Bilayer
We have already seen that membrane lipids form a bilayer in aqueous (water
containing) environments. This is due to the amphipathic nature of the
membrane lipids. The hydrophilic polar heads arrange themselves closes to the
water and the hydrophilic tails interact with each other holding the 2 leafs of the
membrane together.

Membrane lipid layers are self healing and will quickly repair any tears in the
membrane in order to exclude water from the hydrophobic core. In the same
way large flat membrane structures will spontaneously fold into a spherical
structure to ensure the membrane layer is continuous forming a boundary
enclosing a space. This property of membrane lipids leads to the
compartmentalisation necessary for complex life to take place.

Each type of membrane has a characteristic set of lipids and proteins associated
with that membrane.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Some lipids are confined to one side of the membrane

The lipid bilayer is a fluid structure. Lipids are prevented from leaving the bilayer
but there is nothing stopping the molecules moving about within the membrane.
The lipid molecules are free to diffuse freely in 2 dimensions, in the same plane
of the membrane but rarely flip from one plane to another. We say that lipids are
laterally mobile. Some enzymes called flippases however, can swap a lipid from
one side of the membrane to the other. Some lipids are found preferentially on
one side of the membrane. Sphingomyelin tends to be found more on the outer
side of the membrane whereas phospholipids are mainly found on the inside of
the membrane. This leads to an asymmetric distribution of lipids in most
membranes.

The fluidity of the membrane depends on its lipid composition.

The longer the hydrocarbon tails the less fluid a membrane is. This is due to the
increased interaction between neighbouring lipids which makes the whole
structure more viscous. Some lipid hydrocarbon tails have double bonds
between the carbon atoms, known as unsaturated fatty acids. This often
introduces a kink (or bend) into the hydrocarbon tail. These lipids do not pack so
tightly together so membranes containing a lot of unsaturated fatty acids are
more fluid than those that are composed of mostly saturated fatty acids.
Cholesterol also plays a role in the fluidity of the membrane. Cholesterol fits in
the gaps between the phospholipid molecules and due to its rigid structure it
stabilises the membrane decreasing the fluidity of the plasma membrane.

Some organisms can change the fluidity of their membrane to suit environmental
conditions, especially changes n temperature. When living at higher
temperatures some bacteria can change the composition of their membranes to
increase the number of longer chain fatty acids present this stabilises the
membrane at higher temperatures.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Membrane proteins diffuse laterally in the membrane

Membrane proteins behave as if they are afloat on a sea of lipids. Like lipids the
membrane associated proteins are free to diffuse laterally in the lipid layer.
Indeed this is where the mosaic nature of the membrane comes from. Some
membrane proteins associate with each other into raft like structures. These
structures termed lipid rafts are important for cell signalling and cellular
adhesion. Lipid rafts often have a high sphingolipid and cholesterol content.
Lipid rafts are more ordered and tightly packed than the surrounding membrane.

Membrane fluidity is very important for the following reasons:

a. It allows the diffusion of protein molecules in the membrane which is

important for cell signalling
b. It permits lipids and proteins to diffuse from their site of insertion into the
membrane throughout the entire membrane structure
c. It ensures that membrane molecules are divided evenly when cells divide
d. It allows membranes to fuse and mix with each other e.g. during
endocytosis
e. Membrane fluidity regulates permeability more fluid membranes are
more permeable than less fluid membranes

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

OUTCOME 1.3:

Describe the nature of common membrane proteins.

Membranes as a Mosaic of Proteins

A membrane is a collage of proteins embedded in the fluid lipid membrane.
Whereas the lipids are responsible for the overall structure and integrity of the
plasma membrane it is the proteins that dictate most of the important functions of
the membrane. Each cell type will have its own complement of proteins
associated with that cell type and ensures that that cell will be able to carry out its
necessary functions.
Membrane proteins fall into two broad categories: Integral Membrane proteins
and Peripheral Membrane Proteins. We will not look at both of these types of
proteins in more detail.
Integral Membrane Proteins
Integral proteins are strongly attached to the plasma membrane. They cannot
easily be separated from the membrane. Integral membrane proteins fall into 3
categories:

1) Transmembrane Proteins that become embedded directly lipid bilayer

itself
2) Mono-layer associated -helix containing proteins
3) Lipid linked Proteins

Transmembrane proteins are amphipathic. The hydrophobic region of the

protein (containing amino acids with hydrophobic side chains) is found
embedded in the membrane. These hydrophobic amino acids usually arrange
themselves into either a single -helix, multiple -helices or a structure called a
barrel. The peptide backbone of proteins is usually hydrophilic due to the polar
nature of the peptide bond. When the backbone is arranged into an -helix or
sheet the hydrogen bonding between the protein and water is minimised and
hydrogen bonding within the protein is maximised, thus reducing the hydrophilic

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

nature of the protein in this membrane spanning region. Therefore these

structures allow the protein to become embedded in the membrane. The
hydrophilic regions of the proteins are found either on the cytoplasmic or
exoplasmic side of the membrane. An example of a transmembrane protein is
the G protein coupled receptor which has 7 -helices spanning the plasma
membrane.

1 = single -helix transmembrane protein

2 = Multiple -helices
3 = barrel

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Monolayer associated proteins are can be found either on the cytoplasmic or

exoplasmic side of the plasma membrane. Some proteins use an amphipathic helix or a hydrophobic loop to embed itself in one side of the membrane.
Other proteins become modified with lipid which allows them to become
anchored to the membrane via this attached lipid.

Peripheral proteins are only loosely attached to the plasma membrane usually
via interactions with integral proteins or specialised lipids. Peripheral proteins will
have special domains that allow them to be targeted to the plasma membrane.
E.g. the lipid binding PH domain. Peripheral proteins are often involved in signal
transduction or they serve as anchoring sites either to the extra cellular matrix or
to the cytoskeleton.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Functions of Membrane Proteins

Membrane proteins have a wide variety of functions:

1) Transport
2) Enzymatic Activity
3) Signal Transduction
4) Intercellular Joining
5) Cell-cell Recognition
6) Attachment sites to ECM or Cytoskeleton

1) Transport

There are two basic types of transporters found in the cell membrane. Both are
integral proteins that span the membrane.

Carrier proteins and Ion Channels

One type of transporter is a simple channel protein that facilitates the diffusion of
molecules through the membrane. An example is the Aquaporins whose role in
the cell to conduct water molecules into and out of the cell while preventing ions
and other molecules passing across at the same time. Another example is the
ion channels that play an important role in the generation of the action potential
discussed in Outcome 3. Glucose transporters are also carrier proteins which
facilitate diffusion of glucose into the cell.

Pumps
Another group of proteins responsible for transport across the membrane are the
pumps that actively transport molecules across the plasma membrane usually
against a concentration gradient. Some pumps are powered by the energy
stored in ATP (ATP driven pump) and other pumps are powered by
electrochemical gradients (coupled pumps). The Sodium-Potassium Pump is an
example of an ATP driven pump. The energy released from the hydrolysis of

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

ATP is used to transport both K+ and Na+ against their concentration gradients by
bringing K+ into the cell and transporting Na+ out of the cell.

Sodium Potassium Pump

Coupled pumps work by using the energy generated by the movement of one
ion, usually down its concentration gradient, to move another molecule against its
concentration gradient. One example is the Na+/Glucose transporter which
moves Na+ into the cell, down its concentration gradient, at the same time as it
moves glucose into the cell against its concentration gradient. The Ca2+/Na+
exchanger allows one Ca2+ ion out of the cell for every 3 Na+ ions it allows in.
Coupled pumps can move 2 ions in the same direction (symport) or move the
ions in opposite directions (antiport).
Uniporter

Symport

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Antiport

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

2) Enzymatic Activity

Many enzymes are found embedded in membranes. They can either be part of
signalling pathways, biosynthetic pathways or energy generating pathways. The
enzymes involved in the Electron transport chain are embedded in the inner
mitochondrial membrane. This arrangement allows for the co-ordination of the
task of generating ATP to be split between several enzyme complexes. The
presence of the membrane allows the generation of a Proton (H+) gradient which
is used by an enzyme called ATP synthase to generate ATP in aerobic
respiration. Many enzymes found in the biosynthetic pathway for cholesterol
synthesis are found in membranes also.

3) Signal Transduction

The receptors involved in cell signalling are often found at the cell surface. An
example is the G Protein coupled receptor which is an integral membrane
protein. GPCRs are also called serpentine receptors as the protein spans the
membrane 7 times (crossing the membrane as a hydrophobic -helix) in a shape
similar to a snake. A hydrophilic region of the receptor on the exoplasmic side of
the membrane contains a binding site for a signalling molecule called a ligand.
Once bound to the ligand the GPCR changes shape (this change in shape is
often referred to as a conformational change) which in turn creates a change in
shape of the hydrophilic cytoplasmic portion of the protein. This change in
structure activates a protein on the inside of the cell which can interact with the
cytoplasmic region of the GPCR. This protein is called a G protein. The
activation of the receptor causes an inactive G protein (bound to GDP) to
become active by exchanging the GDP for GTP. The active G protein is now
free to alter the activity of other enzymes in the membrane. The G protein
remains active for only a short time however as it is also a GTPase which breaks
down GTP and turn it back into GDP.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

Other signalling molecules present in the cell membrane include:

Tyrosine Kinase Receptors

JAK/STAT Receptors

4) Intercellular Joining

Cells are held together by proteins present in the cells membrane. Proteins such
as claudins and occludins are present at tight junctions and help to hold two cells
together. Tight junctions are discussed in greater detail in Outcome 3.

5) Cell to Cell Recognition

Many of the proteins, and some of the lipids, surrounding a cell are modified with
carbohydrate. This sugar coating surrounding the cell is called the glycocalyx.
This cell coating serves to protect the cell from mechanical damage the sugar
coating absorbs water making the surface of the cell slimy. A more important
function of the glycocalyx however is that it serves as a major recognition point
for the cell. Each cell type has a specific set of carbohydrates that are found only
on that cell acting like an Identification badge for that type of cell. Certain
proteins called lectins can bind specifically to these carbohydrates allowing

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

important functions that rely on cell recognition to take place. This is very
important in the immune system for example. At a site of infection cells lining
blood vessels can express certain lectins at their cell surface which bind onto
sugars present on the surface of White blood cells thus allowing WBCs to
accumulate at the site of infection.

6) Attachment sites to ECM or Cytoskeleton

Proteins in the cell membrane serve as anchor sites to allow the cell to be
attached to the extracellular matrix and internally to the cytoskeleton. A group of
proteins in the membrane called Integrins serve to anchor the cell to fibres in the
extracellular matrix such as Fibronectin and Collagen. Integrins can also link
cells together. On the inside of the cell the integrin can bind to actin providing
further support to the cell.

Proteins that are involved in cell to cell contact are called Cadherins. These are
important in desmosomes and adherens junctions as discussed in Outcome 3.
They are joined to filaments from the cytoskeleton inside the cell to provide
structure and support to the cell.

The final group of proteins that bind to the ECM are called Selectins. They bind
to carbohydrate in the ECM. They play important roles in targeting White Blood
cells to the correct location in the body.

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H927 34: Cell Biology: Theory and Lab Skills Outcome 1 Student Notes

References
Alberts B, Bray D, Hopkin D, Johnson D, Lewis J, Raff M, Roberts K and Walter
P. Essential Cell Biology 4th Edition New York Garland Science
Campbell NA, Reece JB and Mitchell LG. Biology 5th Edition California
Benjamin/Cummings

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