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Biology HL

2.4

Membranes

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Biology HL
2.4

Membranes
Features of membrane structure
Phospholipids
The phospholipid bilayer is the backbone of the
Polar hydrophilic
phosphorylated alcohol head
membrane in that it the membrane is held together by
the phospholipids (though not in a rigid structure). This
is because the hydrophobic and hydrophilic regions of
phospholipids in the bilayer of the plasma membrane
cause the phospholipids to always align as a bilayer
(forming a semi-permeable bilayer) if there is water
present and there is a large number of phospholipid
molecules. As the tails of the fatty acids do not
Non-polar hydrophobic fatty acid
strongly attract each other, the membrane is fluid and
(hydrocarbon) tail when packed closely
flexible. This also allows processes called endocytosis
together, they form a semi-permeable
and exocytosis to take place.
bilayer
!
Diffusion across the phospholipid bilayer is only
possible for small, uncharged molecules, e.g. Fig. 2: A model of a phospholipid
substances like O2, water, CO2, urea, ethanol. Large
or charged molecules cannot cross the lipid bilayer, e.g. Na+, K+, Cl-, glucose.
Cholesterol
Cholesterol is a lipid
Phosphorylated alcohol
G
molecule that helps to
l
regulate membrane
y
fluidity and is important
c
for membrane stability.
e
Because membrane
Fatty acid
r
fluidity changes with the
o
temperature, cholesterol
l
molecules are necessary
Fatty acid
because they allow
effective membrane
Fig. 3: Structure of a phospholipid
function at a wider range
of temperatures. For example, at low temperatures, it interferes with solidification of the
phospholipid molecules, and so keeps the structure more fluid. Plant cells do not have
cholesterol molecules so they depend on saturated and unsaturated fatty acids to maintain
proper membrane fluidity.
Membrane proteins
Membrane proteins have many functions. These functions include:
hormone binding sites!!
cell-to-cell communication!
pumps for active transport!

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immobilized enzymes
channels for passive transport
cell adhesion

There are, therefore, many different types of protein found on the plasma membrane to carry
out this diverse range of functions.
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Membranes
Peripheral proteins: Glycoproteins
Glycoproteins are an example of peripheral proteins. They are proteins with a chain of
carbohydrates (glycocalyx) protruding from the surface. They are involved in cell signalling, cellto-cell recognition and in binding cells together. This is because the glycocalyx acts as a
receptor on cells which can bind to hormones, drugs, and other cells.
Integral proteins
There are many types of integral protein, but the main two types are the channel protein and the
carrier protein.
Channel protein:!
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A protein with a specific shape that compliments the shape of a certain


substance to be transported across the plasma membrane. This type of
protein carries out facilitated diffusion of molecules by providing a
continuous gateway for the passive transport of its substrate(s), often ions,
in which case it is called an ion channel.

Carrier protein:!
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A protein with a specific shape that compliments the shape of a certain


substance to be transported across the plasma membrane. Carrier
proteins can carry put facilitated diffusion or active transport. This is done by
providing a binding site in the membrane that can be accessed from within
or outside the cell but only during alternating movements of the carrier
protein. This non-continuous gateway allows for active (co-transport) or
passive transport.
!
In terms of the active transport mechanism, this is done by an
ATP-powered pump, where the protein employs the energy of ATP
hydrolysis to move substrates, often ions, against the concentration
gradient or an electric field.

Transport across membranes


Diffusion is the passive movement of particles from a region of high concentration to a region of
low concentration.
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Osmosis is the passive movement of water molecules, across a partially permeable
membrane, from a region of lower solute concentration to a region of higher solute
concentration.
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Osmosis and diffusion are passive processes that occur along a concentration gradient,
which means that they do not require any energy expenditure.
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Whether a substance can pass across a membrane passively is influenced by two major
factors: size and charge. Substances that are small in size and non-charged can move across
membranes with relative ease. Substances that are large in size, charged or both, cannot: they
rely on membrane transport proteins in order to be able to cross the plasma membrane.
!
Small, uncharged molecules (e.g. oxygen, carbon dioxide, glycerol, urea) can be
transported across the cell membrane (between the phospholipids) by diffusion. They are small
enough to pass through the transient gaps between the phospholipid molecules (Fig. 4).
!
For larger molecules and ions, they can be transported across the plasma membrane by
facilitated diffusion or active transport, but they rely on membrane transport proteins to cross the
plasma membrane. Some larger molecules can pass through channels in specific integral
proteins in the plasma membrane (Fig. 5).
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Biology HL
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Membranes
Passive transport

Phospholipid

Water molecule

Carbon dioxide

Oxygen
Fig. 4: Simple diffusion of small molecules across phospholipid
bilayer via transient gaps between phospholipid molecules

Extracellular space

High concentration

Concentration gradient

Low concentration
Cytoplasm
Fig. 5: Simple diffusion of molecules across
phospholipid bilayer via channels in an integral
protein in the plasma membrane

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Membranes
Facilitated diffusion
Facilitated diffusion is a particular type of diffusion involving a membrane with specific carrier
proteins that are capable of combining with the substance in order to aid in its movement. The
carrier changes shape in order to accomplish this task but this does not require energy. It is
worth noting that the rate of facilitated diffusion will level off when total saturation of available
carriers occurs.
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In facilitated diffusion, the substrate molecule binds to a carrier (integral) protein in the
plasma membrane, where the proteins binding site is initially exposed outside the bilayer. As
the protein can exist in two different conformational states, it then inverts. This means that the
binding site is then exposed inside the cell, so the substrate is released (Fig. 6).
!
Examples of molecules that are transported through facilitated diffusion include glucose,
fructose, non-fat soluble vitamins, urea and ions that the semi-permeable phospholipid bilayer
would normally prevent from diffusing.
Molecule fits into integral protein
Extracellular space
pong

ping

Cytoplasm
Molecule diffuses away from protein
Fig. 6: Facilitated diffusion of molecules across
phospholipid bilayer via carrier proteins
Uniport: !
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Where one solute is simply transported from one side of the membrane to the
other.

Symport:!
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Where the transfer of one solute depends on the simultaneous or sequential


transport of a second solute in the same direction.

Antiport:!
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Where the transfer of one solute depends on the simultaneous or sequential


transport of a second solute in the opposite direction.
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Membranes
Summary of passive transport
Type of transport

Description

Simple diffusion

Substances other than water move between phospholipid molecules


or through proteins which possess channels. The rate of diffusion is
proportional to the concentration gradient (but the rate is also
influenced by the size of the particles, the distance over which
diffusion must occur and the temperature

Facilitated diffusion

Non-channel protein carriers change shape to allow movement of


substances other than water. The rate of facilitated diffusion is
proportional to the concentration gradient and to the number of carrier
proteins that are available.

Osmosis

Only water moves through the membrane using aquaporins which are
proteins with specialised channels for water movement.

Active transport
Active transport is the uptake of substance(s) against the concentration gradient. This requires
ATP (adenosine triphosphate), the form in which usable cellular energy comes and will involve
membrane proteins in this case an ATP-powered pump that employs the energy of ATP
hydrolysis to move substrates (often ions) against a concentration gradient or an electric field.
An example of this is the sodium-potassium-ATP-ase process establishes the most important
gradients across the animal plasma membrane. The sodium gradient is utilised to effectively
absorb glucose from food in the small intestine (see Fig. 12).
Mechanism for active transport 1
1.

On the inside of the cell, a sodium ion binds to an integral protein (ATP pump) on the cell
membrane.

2.

This triggers the breakdown of ATP into ADP and the energy which is released is used to
phosphorylate the protein which forms the pump.

3.

The protein changes shape and the sodium ion is transported to the outside of the cell.

4.

The changed shape of the protein now allows potassium ions on the outside of the cell to
bind to the protein.

5.

This triggers dephosphorylation of the protein (the phosphate group is removed).

6.

This changes the shape of the protein again and the potassium is transported to the inside
of the cell.

In the Na+K+ATPase pump, it is not (strictly speaking) an exchange of one sodium ion for one potassium ion.
Rather, it is an exchange of two sodium ions for three potassium ions.
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Membranes
Extracellular space
Phospholipid
bilayer

Cytoplasm
Integral protein
(ATP pump)

ATP molecule

Fig. 7: On the inside of the cell, an ATP molecule binds to


an integral protein (ATP pump) on the cell membrane.

Extracellular space

Cytoplasm
Sodium ion

Fig. 8: A sodium ion then binds to the ATP pump on the


cell membrane.
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Extracellular space

Cytoplasm
ADP molecule
Fig. 9: This triggers the breakdown of ATP into ADP and the
energy which is released is used to phosphorylate the protein
which forms the pump. The protein changes shape and the
sodium ion is transported to the outside of the cell.

Extracellular
space

Cytoplasm
Potassium ion

Fig. 10: The changed shape of the protein now allows potassium
ions on the outside of the cell to bind to the protein.
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Membranes
Extracellular space

Cytoplasm

Fig. 11: This triggers dephosphorylation of the protein (the phosphate


group is removed). This changes the shape of the protein again and
the potassium is transported to the inside of the cell. The events are
cyclic, so further transport of sodium and potassium ions can occur,
going back to Fig. 7.

(N.B.: There is a tendency for sodium ions to diffuse back into the cell and potassium ions back
out of the cell. Since the membrane is more permeable to potassium than sodium, more ions
leave the cell than enter. This reduces the tendency of water to enter the cell by osmosis thus
the sodium-potassium pump is a method of controlling the volume of the cell.)
In this active transport cycle (Fig. 7 11), the changes in the conformation (ping-pong) of the
integral proteins are analogous to those of integral proteins responsible for facilitated diffusion.
The sodium-potassium-ATP-ase pumps are an example of an antiport. They are particularly
important in animal cells in osmoregulation where they reset the water potential of the cell by
exporting Na+ ions (and thus water), thereby preventing lysis of the cell.
They are also important in the creation of potential differences across membranes which are
crucial in the transmission of nerve impulses.
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Furthermore, sodium-potassium-ATP-ase pumps are important in animal cells in that they drive
the active transport of glucose 2 (via a symport) by generating and maintaining the Na+ gradient
(Fig. 12). It is for this reason that the mechanism for co-transporting glucose and sodium ions is
likely to be very close to the mechanism responsible for the antiport transfer mechanism of
sodium and potassium ions3 .
Na+
Extracellular
space

Cytoplasm

Glucose

Na+

K+

Fig. 12: Diagram showing the glucose symport aided by the


Na+ gradient across the plasma membrane generated and
maintained by the Na+K+ATPase pump
Endocytosis and Exocytosis
Endocytosis and exocytosis are processes that allow larger molecules to move across the
plasma membrane. Endocytosis allows macromolecules to enter the cell, while exocytosis
allows molecules to leave. Both processes depend on the fluidity of the plasma membrane. The
plasma membrane is fluid in consistency because the phospholipid molecules are not closely
packed together due to the rather loose connections between the fatty acid tails. It is also
important to remember why the membrane is stable: the hydrophilic and hydrophobic properties
of the different regions of the phospholipid molecules cause them to form a stable bilayer in an
aqueous environment.
2

Glucose can also be transported via facilitated diffusion, though this depends on the concentration gradient
between the cytoplasm and the extracellular space.
3

Other types of active transport pumps exist, for example Ca2+ pumps in the ER of muscle cells which are
important in muscle contraction and H+ (proton) pumps in mitochondria and chloroplasts which are important in
ATP synthesis.
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Membranes
Endocytosis occurs when a portion of the plasma membrane is pinched off to enclose
macromolecules or other particulates. This pinching off involves a change in the shape of the
membrane. The result is the formation of a vesicle that then enters the cytoplasm of the cell.
The ends of the membrane reattach because of the hydrophobic and hydrophilic properties of
the phospholipids and the presence of water. This could not occur if it were not for the fluid
nature of the plasma membrane.
Examples of endocytosis include phagocytosis the intake of large particulate matter and
pinocytosis the intake of extracellular fluids.
Exocytosis is essentially the reverse of endocytosis where large molecules are released from
the cell. The fluidity of the plasma membrane and the hydrophobic and hydrophilic properties of
its molecules are just as important as in endocytosis. Exocytosis usually begins in the
ribosomes of the rough endoplasmic reticulum (ER) and progresses through a series of four
steps until the substance is secreted to the outside environment of the cell.
1.!

Proteins are manufactured / synthesised / produced (for export) by the ribosomes of the
rough endoplasmic reticulum. They then enter the lumen of the endoplasmic reticulum.

2.!

A vesicle containing the proteins is pinched off. The proteins exit the ER and enter the cis
(receiving) side / face of the Golgi apparatus (Fig. 13).

3.!

The Golgi apparatus packages and processes the proteins, and modifies them. The
modified proteins bud off (come off) the Golgi apparatus, and exits inside a vesicle on the
trans side of the Golgi apparatus, moving towards the plasma membrane.

4.!

The vesicle with the modified proteins moves to and fuses with the plasma membrane.
This results in the secretion of the contents from the cell.

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This is exocytosis, and it is a process that requires energy. An example of exocytosis is synaptic
transmissions between neurons.
Cis side / face (receiving)

Trans side / face (shipping)


(Generally more curved)
Fig. 13: Golgi apparatus
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Biology HL
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Membranes
Appendix:
To explain how the hydrophobic and hydrophilic properties of phospholipids help to maintain the
structure of cell membranes, for example after exocytosis, in which the membrane is split
(temporarily), we must first consider what it means to be hydrophilic and hydrophobic.
Hydrophilic molecules are attracted to water; hydrophobic molecules are repelled by water.
Phospholipids contain both hydrophilic areas and hydrophobic areas.
The phosphate heads are polar and are therefore hydrophilic. The hydrocarbon tails are not
polar, so are hydrophobic.
Membranes are usually in aqueous environments they are surrounded by water molecules.
This means that the heads face towards water, whereas the tails are positioned away from
water.
This forms a spontaneous, self-assembling (and therefore self-reassembling) bilayer, which is
hydrophilic on the outside, and hydrophobic on the inside4.

Membrane proteins will be arranged in accordance with this structure.


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