B2.

1 Membranes and
Membrane Transport
Form and Function - Cells
Guiding questions
How do molecules of lipid and protein assemble into biological membranes?

What determines whether a substance can pass through a biological membrane?

 Syllabus Statements - SL and HL content
B2.1.1—Lipid bilayers as the basis of cell membranes
Phospholipids and other amphipathic lipids naturally form continuous sheet-like bilayers in water.
B2.1.2—Lipid bilayers as barriers
Students should understand that the hydrophobic hydrocarbon chains that form the core of a membrane have low permeability to large molecules and hydrophilic particles,
including ions and polar molecules, so membranes function as effective barriers between aqueous solutions.
B2.1.3—Simple diffusion across membranes
Use movement of oxygen and carbon dioxide molecules between phospholipids as an example of simple diffusion across membranes.
B2.1.4—Integral and peripheral proteins in membranes
Emphasize that membrane proteins have diverse structures, locations and functions. Integral proteins are embedded in one or both of the lipid layers of a membrane. Peripheral
proteins are attached to one or other surface of the bilayer.
B2.1.5—Movement of water molecules across membranes by osmosis and the role of aquaporins
Include an explanation in terms of random movement of particles, impermeability of membranes to solutes and differences in solute concentration.
B2.1.6—Channel proteins for facilitated diffusion
Students should understand how the structure of channel proteins makes membranes selectively permeable by allowing specific ions to diffuse through when channels are open
but not when they are closed.
B2.1.7—Pump proteins for active transport
Students should appreciate that pumps use energy from adenosine triphosphate (ATP) to transfer specific particles across membranes and therefore that they can move particles
against a concentration gradient
B2.1.8—Selectivity in membrane permeability
Facilitated diffusion and active transport allow selective permeability in membranes. Permeability by simple diffusion is not selective and depends only on the size and hydrophilic
or hydrophobic properties of particles..
B.2.1.9—Structure and function of glycoproteins and glycolipids
Limit to carbohydrate structures linked to proteins or lipids in membranes, location of carbohydrates on the extracellular side of membranes, and roles in cell adhesion and cell
recognition.
B2.1.10—Fluid mosaic model of membrane structure
Students should be able to draw a two-dimensional representation of the model and include peripheral and integral proteins, glycoproteins, phospholipids and cholesterol. They
should also be able to indicate hydrophobic and hydrophilic regions.
Syllabus Continued - HL content
B2.1.11—Relationships between fatty acid composition of lipid bilayers and their fluidity
Unsaturated fatty acids in lipid bilayers have lower melting points, so membranes are fluid and therefore flexible at temperatures experienced by a cell.
Saturated fatty acids have higher melting points and make membranes stronger at higher temperatures. Students should be familiar with an example of
adaptations in membrane composition in relation to habitat.

B2.1.12—Cholesterol and membrane fluidity in animal cells

Students should understand the position of cholesterol molecules in membranes and also that cholesterol acts as a modulator (adjustor) of membrane fluidity,
stabilizing membranes at higher temperatures and preventing stiffening at lower temperatures

B2.1.13—Membrane fluidity and the fusion and formation of vesicles

Include the terms “endocytosis” and “exocytosis”, and examples of each process.

B2.1.14—Gated ion channels in neurons

Include nicotinic acetylcholine receptors as an example of a neurotransmitter-gated ion channel and sodium and potassium channels as examples of
voltage-gated channels.

B2.1.15—Sodium–potassium pumps as an example of exchange transporters

Include the importance of these pumps in generating membrane potentials.

B2.1.16—Sodium-dependent glucose cotransporters as an example of indirect active transport

Include the importance of these cotransporters in glucose absorption by cells in the small intestine and glucose reabsorption by cells in the nephron.

B2.1.17—Adhesion of cells to form tissues

Include the term “cell-adhesion molecules” (CAMs) and the understanding that different forms of CAM are used for different types of cell–cell junction. Students
are not required to have detailed knowledge of the different CAMs or junctions.
Drawing & Annotations
Students should be able to draw a two-dimensional representation of the
model and include peripheral and integral proteins, glycoproteins,
phospholipids and cholesterol. They should also be able to indicate
hydrophobic and hydrophilic regions.
 B2.1.1—Lipid bilayers as the basis of cell membranes
Phospholipids and other amphipathic lipids naturally form continuous sheet-like bilayers in water.
Structure of Phospholipids

● Consist of a polar head (hydrophilic) composed of a glycerol and a phosphate

molecule
● Consist of two non-polar tails (hydrophobic) composed of fatty acid
(hydrocarbon) chains
● Because phospholipids contain both hydrophilic (water-loving) and lipophilic
(fat-loving) regions, they are classed as amphipathic
Arrangement in Membranes

● Phospholipids spontaneously arrange into a bilayer (i.e. two parallel layers of

phospholipids)
● The hydrophobic tail regions face inwards and are shielded from the surrounding
polar fluids
● The two hydrophilic head regions face outwards and associate with the cytosolic
and extracellular fluids respectively

Properties of the Bilayer

● The phospholipid bilayer is held together by weak hydrophobic interactions

between the tails
● The presence of hydrophilic and hydrophobic layers restrict the passage of many
substances
● Individual phospholipids can move within the bilayer, allowing for membrane
fluidity and flexibility
● This fluidity allows for the spontaneous breaking and reforming of membranes
(endocytosis / exocytosis)
B2.1.2—Lipid bilayers as barriers
Students should understand that the hydrophobic hydrocarbon chains that form the core of a membrane have low permeability to large
molecules and hydrophilic particles, including ions and polar molecules, so membranes function as effective barriers between aqueous
solutions
 B2.1.3—Simple diffusion across membranes
Use movement of oxygen and carbon dioxide molecules between phospholipids as an
example of simple diffusion across membranes.
● Diffusion is the passive net movement of
particles form an area of high
concentration to an area of low
concentration
○ Down a concentration gradient
● Often through a semipermeable
membrane
● Does not require energy
Factors affecting Rate of Diffusion
B2.1.4—Integral and peripheral proteins in membranes
Emphasize that membrane proteins have diverse structures, locations and functions. Integral proteins are embedded in one or both of
the lipid layers of a membrane. Peripheral proteins are attached to one or other surface of the bilayer.

• Have wide variety of functions

– Junctions ( connect and join two cells together)
– Enzymes (helps catalyze reactions)
– Transport (Responsible for facilitated diffusion/active
transport)
– Recognition (Act as markers for cellular identification)
– Anchorage (Attachment points for cytoskeleton)
– Transduction (Function as receptors for peptide
hormones)
Functions of Membrane Proteins
B2.1.5—Movement of water molecules across membranes by osmosis and the role of aquaporins
Include an explanation in terms of random movement of particles, impermeability of membranes to solutes and differences
in solute concentration.
The Importance of Osmotic Control
B2.1.6—Channel proteins for facilitated diffusion
Students should understand how the structure of channel proteins makes membranes selectively permeable by allowing
specific ions to diffuse through when channels are open but not when they are closed.
B2.1.7—Pump proteins for active transport
Students should appreciate that pumps use energy from adenosine triphosphate (ATP) to transfer specific particles across
membranes and therefore that they can move particles against a concentration gradient.

Active transport uses energy to move molecules against a concentration gradient

This energy may either be generated by:

● The direct hydrolysis of ATP (primary active transport)

● Indirectly coupling transport with another molecule that is moving along its gradient (secondary active transport)

Active transport involves the use of membrane proteins (called protein pumps due to their use of energy)

● A specific solute will bind to the protein pump on one side of the membrane
● The hydrolysis of ATP (to ADP + Pi) causes a conformational change in the protein pump
● The solute molecule is consequently translocated across the membrane (against the gradient) and released
B2.1.8—Selectivity in membrane permeability
Facilitated diffusion and active transport allow selective permeability in membranes. Permeability by simple diffusion is not selective
and depends only on the size and hydrophilic or hydrophobic properties of particles.
B.2.1.9—Structure and function of glycoproteins and glycolipids
Limit to carbohydrate structures linked to proteins or lipids in membranes, location of carbohydrates on the extracellular side
of membranes, and roles in cell adhesion and cell recognition.
Phospholipids and membrane proteins can have carbohydrate chains attached via the process of glycosylation ('glyco’ = sugar)
● Glycosylation of a phospholipid results in a glycolipid while glycosylation of a membrane protein produces a glycoprotein
The carbohydrate chains are located on the extracellular side of the membrane and play important roles in cell adhesion and cell recognition
● The surface carbohydrates can serve as an attachment point for other cells (e.g. sperm binding to an egg is mediated by glycoproteins)
● The surface carbohydrates can also act as a point of recognition between cells (e.g. the ABO blood group antigens are glycolipids)

Glycoproteins and glycolipids also play an important role in maintaining the structural integrity of the extracellular matrix
● The extracellular matrix is a network for external molecules that provide structure and biochemical support to surrounding cells
● The carbohydrate chains can link these extracellular molecules together to help make the matrix a cohesive network

Here we see the Glycocalyx

complex on the exterior of an ovum
important for fertilization by sperm
and for further zygotic development
B2.1.10—Fluid mosaic model of membrane structure
Students should be able to draw a two-dimensional representation of the model and include peripheral and integral proteins,
glycoproteins, phospholipids and cholesterol. They should also be able to indicate hydrophobic and hydrophilic regions.
Drawing the fluid mosaic model example
Cell membranes are represented according to a fluid-mosaic model, due to the fact that they are:

● Fluid – the phospholipid bilayer is viscous and individual phospholipids can move position
● Mosaic – the phospholipid bilayer is embedded with proteins, resulting in a mosaic of
components

When drawing the fluid-mosaic model, it is important to correctly represent the following:

● Phospholipids should be arranged in a bilayer, with the polar and non-polar regions identified
● Glycoproteins and glycolipids should be facing the extracellular side of the membrane
● Integral proteins should be embedded within the bilayer, while peripheral proteins should be
anchored to one side
● Cholesterol may be included in animal cell membranes (it will be interspersed between the
fatty acid tails)
 B2.1.11—Relationships between fatty acid composition of lipid bilayers and their fluidity
Unsaturated fatty acids in lipid bilayers have lower melting points, so membranes are fluid and therefore flexible at
temperatures experienced by a cell. Saturated fatty acids have higher melting points and make membranes stronger at
higher temperatures. Students should be familiar with an example of adaptations in membrane composition in relation to
habitat.

Cell membranes are fluid, meaning they are not fixed in position and can adopt amorphous shapes

● The fluidity of a membrane is affected by the composition of fatty acids within the phospholipid bilayer
Unsaturated fatty acids have double bonds in their lipid chain which results in a kinked hydrocarbon tail

● This means the lipids are harder to pack together, lowering their viscosity (and increasing fluidity)
● Unsaturated fatty acids in lipid bilayers also have lower melting points, so membranes are more fluid and flexible at temperatures experienced
by a cell.

Saturated fatty acids have no double bonds in their lipid chain which results in a straight hydrocarbon tail

● This means the lipids will be easier to pack together, increasing their viscosity (and lowering fluidity)
● Saturated fatty acids have higher melting points and make membranes stronger and more stable at higher temperatures

Many organisms will adjust the composition of lipids in their membranes in order to regulate membrane fluidity (homeoviscous adaptation)

● The general trend is an increase in unsaturated fatty acids at lower temperatures and an increase in saturated fatty acids at higher
temperatures
● Regulating membrane fluidity is particularly important in poikilothermic organisms whose internal body temperatures vary considerably (e.g.
Antarctic fish)
B2.1.12—Cholesterol and membrane fluidity in animal cells
Students should understand the position of cholesterol molecules in membranes and also that cholesterol acts as a
modulator (adjustor) of membrane fluidity, stabilizing membranes at higher temperatures and preventing stiffening at lower
temperatures
 B2.1.13—Membrane fluidity and the fusion and formation of vesicles
Include the terms “endocytosis” and “exocytosis”, and examples of each process.

The membrane is principally held together by weak hydrophobic associations between the fatty acid tails of phospholipids
● This allows for the spontaneous breaking and reforming of the bilayer, as the phospholipids can move around and be
rearranged
● As a consequence, materials can enter or leave the cell without having to cross the membrane (membrane breaks and
reforms around the material)
● This method of transport requires the involvement of vesicles and is an active process (requires ATP hydrolysis but is not
active transport)
Endocytosis
● A vesicle is formed on the inside of a plasma membrane and thus allows material to be take
into the cell
○ Contain water and solutes from outside the cell
○ Often contain larger molecules needed by cell that cannot pass through the plasma membrane
● Process
○ A small region of the membrane is pulled from the rest of the membrane and is pinched off
○ Proteins use energy, ATP, to carry out this process
○ An example is of leukocytes which engulf foreign particles such as bacteria into itself as an
immune response

There are two main types of endocytosis:

● Phagocytosis – The process by which solid substances are ingested (usually to be transported to the lysosome)
● Pinocytosis – The process by which liquids / dissolved substances are ingested (allows faster entry than via
protein channels)

Endocytosis can also be receptor-mediated (involves clathrin-coated pits) – this allows the cell to control the type of
content entering
 Exocytosis

● A vesicle fuses with the plasma membrane and releases its

contents outside the cell
● Vesicles can be used to release materials from the cell
○ After protein made in rER and process in Golgi they can
be released
○ Expel waste products/unwanted materials
○ adds vesicular phospholipids to the cell membrane,
replacing those lost when vesicles are formed via
endocytosis
 B2.1.14—Gated ion channels in neurons
Include nicotinic acetylcholine receptors as an example of a neurotransmitter-gated ion channel and sodium and potassium
channels as examples of voltage-gated channels.

Ion channels are integral membrane proteins which contain a hydrophilic inner pore through which ions may pass

● This allows ions to either enter or exit a cell according to the concentration gradient (facilitated diffusion)
The channel proteins may be ion-selective (only allows passage to specific ions) and may be gated (can control the timing of ion movement)

● Ion channels are essential to the operation of nerve cells as they are used to establish charge differentials across a membrane
(membrane potentials)

Voltage-gated Ion Channels:

● Voltage-gated ion channels cycle between an open and closed conformation according to the transmembrane voltage
● In neurons, voltage-gated sodium channels are used to transport sodium ions into the neuron during depolarisation
● Conversely, voltage-gated potassium channels will transport potassium ions out of the neuron during repolarisation

Ligand-gated Ion Channels:

● Ligand-gated channels change their conformation in response to the binding of a specific chemical (ligand)
● In neurons, acetylcholine is a neurotransmitter released from the nerve cells to stimulate adjacent cells
● Muscles contain nicotinic acetylcholine receptors that will trigger the opening of an ion channel when activated

Binding of acetylcholine to these receptors results in the influx of ions into the muscle, triggering a cascade that results in muscular
contraction
B2.1.15—Sodium–potassium pumps as an example of exchange transporters
Include the importance of these pumps in generating membrane potentials.

Antiport
● Antitransporters move two molecules in opposite directions across the membrane
● An example of antiport is the translocation of sodium and potassium ions by the sodium-potassium pump
● This pump is used by nerve cells (neurons) to establish an electrochemical gradient across the membrane (resting
potential)

○ Three sodium ions bind to intracellular sites on the sodium-potassium pump

○ A phosphate group is transferred to the pump via the hydrolysis of ATP
○ The pump undergoes a conformational change, translocating sodium across the membrane
○ The conformational change exposes two potassium binding sites on the extracellular surface of the pump
○ The phosphate group is released which causes the pump to return to its original conformation
○ This translocates the potassium across the membrane, completing the ion exchange
B2.1.16—Sodium-dependent glucose cotransporters as an example of indirect active transport
Include the importance of these cotransporters in glucose absorption by cells in the small intestine and glucose reabsorption
by cells in the nephron.

Carrier proteins are transmembrane transporters which undergo a conformational change to translocate a material across the bilayer

● They differ from ion channels in their method of transportation (ion channels possess an inert inner pore which is hydrophilic)

Carrier proteins can move two substances simultaneously (cotransport), either in the same direction (symport) or opposite directions
(antiport)

Cotransport
● Cotransporters link the movement of an ion along its concentration gradient to the movement of a solute against its concentration
gradient
● This is a form of secondary active transport, as the electrochemical gradient is used as an energy source (instead of ATP
hydrolysis)
● An example of cotransport is the absorption of glucose in the kidneys and small intestine (contransported with sodium ions)
B2.1.17—Adhesion of cells to form tissues
Include the term “cell-adhesion molecules” (CAMs) and the understanding that different forms of CAM are used for different
types of cell–cell junction. Students are not required to have detailed knowledge of the different CAMs or junctions.

Cell adhesion describes the attachment of cells to other surfaces via specialised membrane proteins called cell adhesion
molecules (CAMs)

● Cells can either be directly attached to other cells or indirectly anchored to the extracellular matrix (a gel-like framework
between cells)
Cell adhesion molecules can play important roles in a variety of cellular processes – including growth, apoptosis, signal
transduction, migration and tissue development

● The targeting of specific cell adhesion molecules can limit the spread of malignant cancers (by inhibiting metastasis)

Different types of cell-cell junctions exist according to the type of cell adhesion molecule involved

● Anchoring junctions – hold cells together to strengthen contact within tissues

● Tight (occluding) junctions – create tight seals that result in an impermeable barrier to diffusion
● Gap junctions – links cells together to allow the movement of material between them
Additional Videos
Membranes and transport: https://www.youtube.com/watch?v=dPKvHrD1eS4
Amoeba Sisters:
https://www.youtube.com/watch?v=Ptmlvtei8hw&pp=ygUYYW1vZWJhIHNpc3RlcnMgdHJhbnNw
b3J0
Fluidity: https://www.youtube.com/watch?v=BWQCAsM-CF4
Gated ion channels: https://www.youtube.com/watch?v=Pl7nzXaVqak