Membranes are the structures that separate the contents of cells from their environment.

● They also separate the different areas within cells (organelles) from each other and the
cytosol.
● Some organelles are divided further by internal membranes.
● The formation of separate membrane-bound areas in a cell is called
compartmentalisation.
● Compartmentalisation is vital to a cell as metabolism includes many different and often
incompatible reactions.
● Containing reactions in separate parts of the cell allows the specific conditions required
for cellular reactions, such as chemical gradients to be maintained and protects vital cell
components.

Cell membranes are partially permeable barriers

Because cell membranes form a barrier and separate the cell contents from the cell’s exterior
environment, or separate organelles from cytoplasm, they need to allow some molecules
through, into or out of the cell.

Permeability refers to the ability to let substances pass through.

● Some very small molecules simply diffuse through the cell membrane, in between its
structural molecules.
● Some substances dissolve in the lipid layer and pass through.
● Other substances pass through special protein channels or are carried by carrier
proteins.

Because these membranes do not let all types of molecule pass through them, they are
described as partially permeable barriers.
● The properties of the component molecules of the cell membrane determine its
permeability ie. which molecules it allows through.
The fluid mosaic model

Is the theory of cell membrane structure with proteins embedded in a sea of phospholipids.

Membranes were seen for the first time following the invention of electron microscopy.
● Images taken in the 1950s showed the membrane as 2 black parallel lines- supporting
an earlier theory that membranes were composed of a lipid bilayer.

In 1972, Singer and Nicolson proposed a model that allowed the passage of molecules through
the membrane, in which proteins occupy various positions in the membrane.

Their structure explained how cell membranes could be more dynamic and interact more with
the cells’ environment.
● It was called the fluid mosaic model and proposed that the fabric of the membrane
consisted of a phospholipid bilayer (double layer) with proteins floating in it, making up a
mosaic pattern.
● The lipid molecules can change places with each other, and some of the proteins may
move, giving fluidity.
● The phospholipids are free to move
within the layer relative to each other (they
are fluid), giving the membrane flexibility
and because the proteins embedded in the
bilayer vary in shape, size and position (in
the same way as the tiles of a mosaic)
● The fabric of the membrane is the
lipid bilayer made up of 2 layers of
phospholipid molecules. Their hydrophilic
heads are in contact with the watery interior
(cytoplasm).
● The hydrophobic tail regions are in
the centre of the membrane away from
water.
● The fluid mosaic model also helps to explain:
○ Passive and active movement between cells and their surroundings
○ Cell-to-cell interactions
○ Cell signalling
● The fluid mosaic model describes cell membranes as ‘fluid’ because:
○ The phospholipids and proteins can move around via diffusion
○ The phospholipids mainly move sideways, within their own layers
○ The many different types of proteins interspersed throughout the bilayer move
about within it (a bit like icebergs in the sea) although some may be fixed in
position

● The fluid mosaic model of membranes includes four main components:

○ Phospholipids
○ Cholesterol
○ Glycoproteins and glycolipids
○ Transport proteins

● The linkage of phospholipids molecules with molecules of cholesterol contributes

towards keeping the cell membrane cohesive and intact. From here we get the name
‘Fluid Mosaic Structure’.
● Molecules of proteins are embedded between the molecules of these two layers. Some
of the protein molecules work as cell identification sites for different substances, such as
nutrients and hormones, while others work as gates to pass the substances to and from
the cell.
● Some proteins in the membrane are called ‘intrinsic’. This means that they completely
span the bilayer. Others are called ‘extrinsic’ and are partly embedded in the bilayer.
● Some intrinsic proteins are ‘channel proteins’. These are transport proteins that allow the
movement of molecules that are normally too large or too hydrophilic to pass through the
membrane by forming a tube-like structure that goes through the whole membrane.
● Other transport proteins are ‘carrier proteins’. These use energy in the form of ATP to
actively move substances across the membrane. For example, ions in soil are actively
transported into the root hair cells of plants
● Carbohydrates are also a 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).
● Along with peripheral proteins, carbohydrates form specialized sites on the cell’s 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’).
Membrane structure

All membranes have the same basic structure. The cell surface membrane which separates the
cell from which separates the cell from its external environment is known as the plasma
membrane.
● Membranes are formed from a phospholipid bilayer. The hydrophilic phosphate heads of
the phospholipids form both the inner and outer surface of a membrane, sandwiching the
fatty acid tails of the phospholipids to form a
hydrophobic core inside the membrane.
● Cells normally exist in aqueous environments. The
inside of cells and organelles are also usually aqueous
environments. Phospholipid bilayers are perfectly
suited as membranes because the outer surfaces of
the hydrophilic phosphate heads can interact with
water.
● Phospholipids bilayers act as a barrier to most water-soluble substances (the non-polar
fatty acid tails prevent polar molecules or ions from passing across the membrane)
● This ensures water-soluble molecules such as sugars, amino acids and proteins cannot
leak out of the cell and unwanted water-soluble molecules cannot get in
● Phospholipids can be chemically modified to act as signalling molecules by:
○ Moving within the bilayer to activate other molecules (eg. enzymes)
○ Being hydrolysed, which releases smaller water-soluble molecules that bind to
specific receptors in the cytoplasm

Membrane proteins

There are 2 types of proteins in the cell-surface membrane- intrinsic and extrinsic proteins.
 Cholesterol
● Cholesterol increases the fluidity of the
membrane, stopping it from becoming too rigid at low
temperatures (allowing cells to survive at lower
temperatures)
● This occurs because cholesterol stops the
phospholipid tails packing too closely together
● Interaction between cholesterol and
phospholipid tails also stabilises the cell membrane at
higher temperatures by stopping the membrane from
becoming too fluid
○ Cholesterol molecules bind to the
hydrophobic tails of phospholipids, stabilising them and
causing phospholipids to pack more closely together
○ The impermeability of the membrane to ions is also affected by cholesterol
● Cholesterol increases the mechanical strength and stability of membranes (without it
membranes would break down and cells burst)

Glycolipids and glycoproteins

● Glycolipids and glycoproteins contain carbohydrate chains that exist on the surface (the
periphery/extrinsically), which enables them to act as receptor molecules
○ The glycolipids and glycoproteins bind with certain substances at the cell’s
surface
● There are three main receptor types:
○ Signalling receptors for hormones and neurotransmitters
○ Receptors involved in endocytosis
○ Receptors involved in cell adhesion and stabilisation (as the carbohydrate part
can form hydrogen bonds with water molecules surrounding the cell
 ● Some glycolipids and glycoproteins act as cell markers or antigens, for cell-to-cell
recognition (eg. the ABO blood group antigens are glycolipids and glycoproteins that
differ slightly in their carbohydrate chains)

Transport proteins
● Transport proteins create hydrophilic channels to allow ions
and polar molecules to travel through the membrane. There
are two types:
○ Channel (pore) proteins
○ Carrier proteins
■ Carrier proteins change shape to transport a
substance across the membrane
● Each transport protein is specific to a particular ion or molecule
● Transport proteins allow the cell to control which substances
enter or leave

Factors affecting membrane structure and permeability

Membranes control the passage of different substances into and out of cells (and organelles).
● If membranes lose their structure, they lose control of this and cell processes will be
disrupted.
● A number of factors affect membrane structure including temperature and the presence
of solvents.

Temperature and kinetic energy

● Increasing the temperature gives all molecules more kinetic energy, and as a result
these molecules move faster.

Phospholipids and changing temperature

● Many organisms do not generate heat to maintain their body temperature and so their
temperature varies with their environment.
 ● At higher temperatures, any diffusion taking
place through the cell membrane will also occur at
a higher speed (due to increased kinetic energy)
● Changes in membrane fluidity are
reversible
○ If temperatures decrease, the lipids
will return to their normal levels of fluidity)
● At a certain temperature (often around
40°C) many proteins (including those in cell
membranes) begin to denature
○ This disrupts the membrane
structure, meaning it no longer forms an effective
barrier
○ As a result, substances can pass
freely through the disrupted membrane
○ This process is irreversible

Solvents

Organic solvents such as acetone and ethanol will

damage cell membranes as they dissolve lipids,
causing the membrane to lose its structure.

Diffusion

Is the movement of molecules

from an area of high concentration of that molecule to
an area of low concentration; it does not involve
metabolic energy (ATP).

The exchange of substances between cells and their environment or between membrane-bound
compartments within cells and the cell cytosol is defined as either active (requiring metabolic
energy) or passive.
 ● All movement requires energy. Passive movement, however, utilises energy from the
natural motion of particles, rather than energy from another source.

The biochemical processes that sustain life go on in cells. Cells therefore need to receive raw
material or reactants for these reactions.
● They respire to make ATP, which provides cellular energy to drive these biochemical
reactions. Therefore they need oxygen and glucose.
● They also need to remove the toxic metabolic waste products, such as carbon dioxide
and they need to export some of the molecules that they make, such as enzymes,
hormones or other signalling molecules.
● Some substances can pass across cell membranes without using any of the cell’s
metabolic energy. These are described as passive processes because they only use the
kinetic energy of the molecules and do not use ATP.

Diffusion is a passive process and will continue until there is a concentration equilibrium
between the 2 areas. Equilibrium means a balance or no difference in concentrations.

Diffusion happens because the particles in a gas or liquid have kinetic energy (they are moving).
● This movement is random and an unequal distribution of particles will eventually become
an equal distribution.
● Equilibrium does not mean the particles stop moving, just that the movements are equal
in both directions.
● Particles move at high speeds and are constantly colliding, which slows down their
overall movement. This means that over short distances, diffusion is fast but as diffusion
distance increases the rate of diffusion slows down because more collisions have taken
place.
● For this reason cells are generally microscopic- the movement of particles within cells
depends on diffusion and a large cell would lead to slow rates of diffusion.
● Reactions would not get the substances they need quickly enough or ATP would be
supplied too slowly to energy-requiring processes.
Water molecules are a special case. Since they are polar and insoluble in lipid the phospholipid
bilayer would seem to be an impenetrable barrier.
● However, water is present in such great concentrations that significant direct diffusion
does happen.
● In membranes where a very high rate of water movement is required there may indeed
be specific water channel proteins known as aquaporins to allow water molecules to
cross the membrane without the challenge of moving through a lipid environment.

How the concentration gradient is maintained

Many molecules entering cells then pass into organelles and are used for metabolic reactions;
this maintains the concentration gradient and keeps more of the molecules entering the cell.
● Oxygen diffusing into the cytoplasm of respiring cells then diffuses into mitochondria and
is used for aerobic respiration.
● Carbon dioxide diffusing into the palisade mesophyll cells of a plant leaf will then diffuse
into chloroplasts and be used for photosynthesis.

Facilitated diffusion

Small molecules that have polarity

(opposite charges at either end of the
molecule), such as ions that have an electrical charge, are insoluble in lipid because they
cannot interact with the hydrophobic tails of the lipid bilayer.
● This means that they diffuse through water-filled protein channels (pores) embedded in
the membrane. These channels are around 0.8nm in diameter.
Channel proteins
● Channel proteins are water-filled pores
● They allow charged substances (eg. ions) to diffuse
through the cell membrane
● The diffusion of these ions does not occur freely, most
channel proteins are ‘gated’, meaning that part of the
channel protein on the inside surface of the membrane
can move in order to close or open the pore
● This allows the channel protein to control the exchange of
ions

Carrier proteins
● Unlike channel proteins which have a fixed shape, carrier proteins can switch between
two shapes
● This causes the binding site of the carrier protein to be open
to one side of the membrane first, and then open to the other
 side of the membrane when the carrier protein switches shape
● The direction of movement of molecules diffusing across the membrane depends on
their relative concentration on each side of the membrane
● Net diffusion of molecules or ions into or out of a cell will occur down a concentration
gradient (from an area containing many of that specific molecule to an area containing
less of that molecule)

Active transport

Is the movement of substances against their concentration gradient from low to high
concentration of that substance.

Many biological processes depend on the presence of a concentration gradient, eg, the
transmission of nerve impulses.
● To maintain this concentration gradient, particles must be moved up it at a rate faster
than the rate of diffusion- active transport.
● This process requires energy and carrier proteins. Energy is needed as the particles are
being moved up a concentration gradient, in the opposite direction to diffusion.
● Metabolic energy is supplied by ATP.
● Carrier proteins span the membranes and act as ‘pumps.’
● Needs more energy than the kinetic energy of the molecules. This energy is provided by
the hydrolysis of ATP.
● Cells or organelles may need to accumulate more of a particular ion than they could do
by simple or facilitated diffusion alone. Process of AT: from inside to outside cell-

Carrier proteins

These membrane proteins have

specific regions, or sites that
combine reversibly with only certain
solute molecules or ions.
● They also have a region that
binds to and allows the hydrolysis of a molecule of ATP, to release energy and in this way they
act as enzymes.
● The energy helps the carrier protein change its conformation (shape) and in doing so it
carries the ion from one side of the cell membrane to the other.

Eg. in guard cells ATP made by chloroplasts provides energy to actively transport potassium
ions from surrounding cells into the guard cells.
● This influx of ions lowers the water potential in the guard cells, so that water enters from
surrounding cells, by osmosis.
● As the guard cells swell, their tips bulge and this opens the stoma between them.

● Active transport is important in:

○ The reabsorption of useful molecules and ions into the blood after filtration into
the kidney tubules
○ The absorption of some products of digestion from the digestive tract
○ The loading of sugar from the photosynthesising cells of leaves into the phloem
tissue for transport around the plant
○ The loading of inorganic ions from the soil into root hairs

Bulk transport

Is another form of active transport

● Large molecules such as enzymes, hormones and whole cells like bacteria are too large
to move through channel or carrier proteins, so they are moved into and out of the cell by
bulk transport.
● Some cells need to transport large molecules and particles that are too large to pass
through the plasma membrane, in or out. They do this by bulk transport, a process that
requires energy from ATP.

● Bulk transport into cells = endocytosis

● Bulk transport out of cells = exocytosis
● These two processes require energy and are therefore forms of active transport

Endocytosis

Is the bulk transport of molecules too large to pass through a cell

membrane even via channel or carrier proteins, into a cell.
● They do not pass through the plasma membrane. Instead, a
segment of the plasma membrane surrounds and encloses the
particle and brings it into the cell, enclosed in a vesicle.
● The cell surface membrane first invaginates (bends inwards)
when it comes into contact with the material to be transported.
● The membrane enfolds the material until eventually the
membrane fuses, forming a vesicle.
 ● The vesicle pinches off and moves into the cytoplasm to transfer the material for further
processing within the cell. Ie. vesicles containing bacteria are moved towards
lysosomes, where the bacteria are digested by enzymes.

○ Phagocytosis:
■ This is the bulk intake of solid material by a cell
■ Cells that specialise in this process are called phagocytes
■ The vacuoles formed are called phagocytic vacuoles
■ An example is the engulfing of bacteria by phagocytic white blood cells
○ Pinocytosis:
■ This is the bulk intake of liquids
■ If the vacuole (or vesicle) that is formed is extremely small then the
process is called micropinocytosis

Exocytosis

Is the bulk transport of molecules, too large to pass through a cell membrane even via channel
or carrier proteins, out of a cell.
● They do not pass through the plasma membrane. Instead,
a vesicle containing them is moved towards and then
fuses with the plasma membrane.
● Ie. at synapses (gaps between neurons) where chemicals
in vesicles are moved, by motor proteins moving along
cytoskeleton threads, to the presynaptic membrane. Here,
the vesicle membranes and plasma membranes fuse and
the neurotransmitter chemicals are released into the
synaptic cleft.
● In all cases, ATP is needed to fuse the membranes
together as well as for moving the vesicles.
● A molecule of ATP is hydrolysed for every step that a
motor protein takes along the cytoskeleton thread as it drags its cargo- the vesicle.
● Energy in the form of ATP is required for movement of vesicles along the cytoskeleton,
changing the shape of cells to engulf materials and the fusion of cell membranes as
vesicles form or as they meet the cell surface membrane.

Osmosis

Is the passage of water molecules down their water potential gradient, across a partially
permeable membrane.

Water potential- is the measure of the tendency of water molecules to diffuse from one region to
another.

In a solution, the liquid in which solute molecules are dissolved in is called the solvent.
● In an aqueous solution, water is the solvent. Water molecules can pass directly through
the phospholipid bilayer.
● Some membranes also have protein channels known as aquaporins which allow water
molecules to cross the membrane more rapidly.
● The inside of cells, the cytoplasm, contains water and the external medium of cells is
also watery, as cells are surrounded by extracellular tissue fluid. Water molecules also
have kinesthetic energy and move randomly but will spread out.

When solute molecules are added to water, the relative number of

water molecules in the resulting solution is changed.
● If the solute molecules dissociate into charged ions, they
exert more effect on the relative number of water molecules than
do larger but non-polar molecules such as glucose.
● This is because as sodium chloride molecules dissociate
into sodium ions and chloride ions, the number of particles in the
solution doubles.

Water potential is the pressure exerted by water molecules as

they collide with a membrane or container.
● The symbol for water potential is the Greek letter psi Ѱ
● Pure water is defined as having a water potential of 0
kPa (at standard temperature (-25 deg C) and (100 kPa))
● All solutions have negative water potentials- the more
concentrated the solution, the more negative the water
potential.

A dilute solution has a high water potential (the right-hand side

of the diagram below) and a concentrated solution has a low
water potential (the left-hand side of the diagram below)

Osmosis in animal and plant cells

The diffusion of water into a solution leads to an increase in

volume of this solution.
● If the solution is in a closed system, such as a cell, this
results in an increase in pressure.
● This pressure is called hydrostatic pressure and has the
same units as water potential, kPa.
● At the cellular level this pressure is relatively large and
potentially damaging.
● Lysis occurs when the cell is in a hypotonic environment
(the solution outside of the cell has a lower solute
concentration than the inside of the cell)
The water potential inside cells is lower than that of pure water, as there are solutes in solution
in the cytoplasm and inside the large vacuole of plant cells.
● When cells are placed in a solution of higher
water potential then water molecules by
osmosis, down the water potential gradient
across the plasma membrane, into the cell.
● In animal cells, if a lot of water molecules
enter, increasing the hydrostatic pressure
inside the cell and the cell will swell and burst
and the plasma membrane breaks.
● All cells have thin-cell surface membranes
(around 7nm) and no cell walls. The cell
surface membrane cannot stretch much and
cannot withstand the increased pressure.
● This is called cytolysis.

Animal cells losing water

● If an animal cell is placed in a solution with a lower water potential than the cell (such as
a concentrated sucrose solution)
● Water will leave the cell through its partially permeable cell surface membrane by
osmosis and the cell will shrink and shrivel up
○ This is crenation (the cell has become crenated), which is usually fatal for the cell
● Crenation occurs when the cell is in a hypertonic environment (the solution outside of the
cell has a higher solute concentration than the inside of the cell)

Animal cells in isotonic environments

● If an animal cell is in an isotonic environment (the solution outside of the cell has the
same solute concentration as the inside of the cell)
● The movement of water molecules into and out of the cell occurs at the same rate (no
net movement of water) and there is no change to the cells

Plant cells

In plant cells, the rigid and strong cellulose cell wall will prevent bursting.
● The cell will swell up to a certain size when its contents push against the cell wall, which
will resist any further swelling. This swollen cell is described as turgid.
● Turgidity of plant cells helps support plants, especially those that are not woody.
● When cells are placed in a solution of lower water potential, water leaves the cells by
osmosis, across the partially permeable plasma membrane
● The cytoplasm of cell plant cells shrinks and the membrane pulls away from the cellulose
cell wall. Water is lost from the cells by osmosis. This leads to a reduction in the volume
of the cytoplasm. The cells are described as plasmolysed.
● The protoplast gradually shrinks and no longer exerts pressure on the cell wall
 ● As the protoplast (all of the contents of a bacterial or plant cell except for the cell wall.
The living parts of a cell) continues to shrink, it begins to pull away from the cell wall
● Plant tissue with plasmolysed cells is described as flaccid.
● Cells that are plasmolysed suffer a degree of dehydration and their metabolism cannot
proceed, as enzyme-catalysed reactions need to be in solution.

Unlike animals, plants are unable to control the water potential, of the fluid around them, ie.
roots are usually surrounded by almost pure water.
● Plant cells have strong cellulose cell walls surrounding the cell-surface membrane.
● When water enters by osmosis the increased hydrostatic pressure pushes the
membrane against the rigid cell walls. This pressure against the cell wall is turgour.
● The expanding protoplast (living part of the cell inside the cell wall) pushes against the
cell wall and pressure builds up inside the cell – the inelastic cell wall prevents the cell
from bursting
● As the turgor pressure increases it resists the entry of further water and the cell is said to
be turgid.
● If plants
do not receive
enough water
the cells cannot
remain rigid and
firm (turgid) and
the plant wilts