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3.

2 BIOLOGICAL MEMBRANES

3.2.1 Properties of Cell Membranes


• Separates living cell from its
nonliving surroundings.
• 8 nm thick.
• Selectively permeable - allows
some substances to cross more
easily than others.

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3.2.2 Fluid Mosaic Model

• Singer and Nicolson (1972) - the


plasma membrane is a mosaic of
proteins dispersed within the lipid
bilayer, with only the hydrophilic
regions exposed to water.

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Hydrophilic region
of protein

Phospholipid
bilayer

Hydrophobic region of protein

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• Plasma membrane is a continuous, fluid,
double layer of phospholipids, the lipid
bilayer.
• Phospholipids & most other membrane
constituents are amphipathic molecules -
have hydrophobic regions & hydrophilic
regions.
Hydrophobic tails face inside of bilayer.
Hydrophilic head faces exterior
(extracellular fluid) and interior
(cytosol).

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WATER

Hydrophilic
head

Hydrophobic
tail

WATER
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• Proteins - embedded in bilayer or
associated with cytoplasmic or
extracellular face.
• Carbohydrates - linked to proteins
(glycoproteins) or lipids (glycolipids)
only on extracellular side.
• Cholesterol - lies within membrane.

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• Membrane molecules held in
place by weak hydrophobic
interactions.
• Most lipids & some proteins drift
laterally.
• Rarely flip-flop from one layer to
the other.

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Lateral movement Flip-flop
(~107 times per second) (~ once per month)
Movement of phospholipids

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• Many larger membrane proteins drift
within the phospholipid bilayer.
• Proteins are much larger than lipids
and move more slowly.
• Other proteins are anchored to
cytoskeleton.

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3.2.3 Membrane Components

a) Membrane proteins

• Amphipathic.
• Determine most of membrane’s
specific functions
• Two groups:
i. Peripheral proteins
 Not embedded but loosely bound to
surface of protein.
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Fibers of
extracellular
matrix (ECM)

Glycoprotein

Carbohydrate

Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE

Cholesterol

Microfilaments Peripheral
of cytoskeleton proteins Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE

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ii. Integral proteins
 Penetrate hydrophobic core, often
completely as transmembrane
proteins.
proteins.
 Hydrophobic segments consist of
stretches of non-
non-polar amino acids,
coiled into α-helices.
 Hydrophilic segments have
hydrophilic non-
non-helical amino
acids.

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EXTRACELLULAR
SIDE
N-terminus

C-terminus
CYTOPLASMIC
SIDE
α Helix
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Six major functions of protein

1. Transport
2. Enzymatic activity
3. Signal transduction
4. Cell-cell recognition
5. Intercellular joining
6. Attachment to the cytoskeleton
and extracellular matrix (ECM)

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Signal
Enzymes

Receptor
ATP

Transport Enzymatic activity Signal transduction

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Glyco-
protein

Cell-cell recognition Intercellular joining Attachment to the


cytoskeleton and extra-
cellular matrix (ECM)

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(b) Carbohydrates

• Branched oligosaccharides with < 15


sugar units.
• Two types:
1. Glycolipids
2. Glycoproteins.
Glycoproteins
• Oligosaccharides on external side of
membrane vary from species to species,
from individual to individual, and from cell
type to cell type within same individual.
• This variation distinguishes each cell
type.
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• Carbohydrates on plasma membrane
surface enables cell-cell recognition
 Ability of a cell to distinguish one
type of neighboring cell from
another by binding to surface
molecules.
• Importance:
1) Sorting and organizing cells into
tissues and organs.
2) Basis for rejection of foreign cells
by immune system.

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(c) Cholesterol

• Interdigitates between phospholipids.


• Enhances mechanical stability &
flexibility of membrane, and making it
less permeable to water-soluble
substances.
• In animal cell membranes

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3.3 Movement Across Membranes

• Hydrophobic molecules
(hydrocarbons, CO2, & O2) dissolve in
lipid bilayer & cross easily.
• Hydrophobic core of membrane
impedes passage of ions and polar
molecules (water, glucose & other
sugars) - cross membrane with
difficulty.

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• Ions and polar molecules cross bilayer
through transport proteins:
proteins
i. Channel proteins:
proteins Have hydrophilic
channel for passage of certain
molecules or ions.
 Example, passage of water
through membrane facilitated by
channel proteins known as
aquaporins.
aquaporins.
ii. Carrier proteins:
proteins Bind to molecules
& change shape to shuttle them
across membrane.
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3.3.1 Passive Transport

a) Diffusion
 Spontaneous tendency of molecules
of any substance to move down its
concentration gradient from a more
concentrated to a less concentrated
area.

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• Individual molecule moves randomly.
• But diffusion of a population of
molecules exhibit a net movement in
one direction.
• At dynamic equilibrium, as many
molecules cross one way as cross in
the other direction.
• Each substance diffuses independent
of the concentration gradients of other
substances.

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Net diffusion Net diffusion Equilibrium

Net diffusion Net diffusion Equilibrium

Diffusion of two solutes

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• No work done to move substances
down concentration gradient.
• Diffusion of substance across
biological membrane is passive
transport - requires no energy from the
cell.

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Factors determining rate of
diffusion

1) Concentration gradient - the steeper the


gradient, the faster the rate of diffusion.
2) Surface area across which substance is
diffusing - the greater the surface area,
the faster the rate of diffusion.
3) Distance over which substance has to
diffuse (the diffusion distance) - the
greater the diffusion distance, the slower
the rate of diffusion.

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• The rate of diffusion is directly proportional to
the concentration difference and surface
area, and inversely proportional to the
diffusion distance.

Surface x Concentration
Rate of diffusion ∝ area distance
Diffusion distance

• This is known as Fick’s Law.

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• Diffusion through membrane (barrier)
affected by:
i. Nature of membrane, for example,
its permeability.
ii. Size and type of molecule or ion
diffusing through it.

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b) Osmosis

Diffusion of water through a semi-


permeable membrane from a solution
with a low solute concentration (high
water potential) to a solution with a
higher solute concentration (low water
potential) until there is an equal
concentration (water potential) on both
sides of the membrane.
• Direction of osmosis determined only by a
difference in total solute concentration

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• Tendency of water molecules to move
across membrane depends on:
i. Solute concentration.
ii. Pressure on each side of membrane.
• Water potential (ψ
ψ) - combined effect
of solute concentration and pressure
in a solution.
 Units in kilopascals (kPa)

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• Water potential of a solution is the
tendency for water to diffuse out of it.
• Water diffuses from a high water
potential to a low water potential,
down its water potential gradient.
• By definition, pure water at
atmospheric pressure has a water
potential of zero.

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Effect of concentration on water
potential and osmosis:

• Adding solute to pure water will decrease


its water potential - becomes negative.
• The more solute is added, the lower (more
negative) the water potential.
• Example:
• 17 g sucrose/dm3 of water = -130 kPa
• 34 g of sucrose/ dm3 of water = -260 kPa.
• Effect of solute concentration is called
solute potential (Ψs).
• Value of Ψs is always negative.

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Lower Higher Same concentration
concentration concentration of sugar
of solute (sugar) of sugar

H2O

Selectively
permeable mem-
brane: sugar mole-
cules cannot pass
through pores, but
water molecules can

Low solute conc. High solute conc.


High water conc. Low water conc,
High water potential Low water potential
Osmosis
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The effect of pressure on water potential
and osmosis:
• Increasing the pressure would increase
the tendency of water to diffuse out of
the membrane.

High water potential

Low water potential

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Increased pressure

High water potential

Low water potential

There is a substantial
movement of water in
osmosis

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High water potential

Low water potential

Pressure applied
on this side

There is decreased osmosis or


osmosis is stopped or even
completely reversed

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• Effect of pressure on a solution is
called the pressure potential (Ψ Ψp).
• Value of Ψp usually positive.
 In plants, this is the force of the cell
wall pushing inwards on contents of
cells (cytoplasm) when water enters
cell by osmosis.
 In animals this may be due, for
example, to high blood pressure in
glomerulus of the kidney.

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Water potential = Solute potential + Pressure potential

ψ = ψS + ψP
(usually negative) (usually negative) (usually positive)

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Effect of osmosis on plant cells
• Plant cells generally have lower water
potential than that of their surroundings.
Due to presence of solutes in fluid
within vacuole (cell sap).
• Plasma membrane & tonoplast
surrounding vacuole are both partially
permeable, letting water through but not
solutes.
• Cell wall permeable to both water &
solutes.

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• Refer to handout for effect of osmosis
on plant cells.

• Incipient plasmolysis – point when


cytoplasm just starts pulling away from
cell wall.
• Full plasmolysis – when cytoplasm has
completely withdrawn from cell wall
except at plasmodesmata.

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Effect of osmosis on animal cells

• Refer to handout.
• Animals without rigid cell walls have
osmotic problems in environment with
low or high water potential.
• To maintain their internal environment,
they must have adaptations for
osmoregulation.
• Paramecium, which has low water
potential compared to its pond water
environment, has a contractile vacuole
that acts as a pump.
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50 µm
Filling vacuole

50 µm
Contracting vacuole

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Worked examples on the water potential
concept
Example 1
• A plant cell with a water potential of –700 kPa
is immersed in a sucrose solution whose
water potential is –350 kPa. In which
direction will water flow?
Answer
• Water will flow from the sucrose solution into
the cell. This is because the water potential
of the cell is lower than (i.e. more negative
than) the sucrose solution, and there is a net
flow of water from a region of higher water
potential to a region of lower potential, i.e.,
down the water potential gradients.

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Worked examples on the water
potential concept
Example 2
• A plant cell has a solute potential of –240
kPa and a pressure potential of 350 kPa.
What is the water potential of the cell?
Answer
Water potential = Solute potential +
Pressure potential
(Ψ) = (Ψs) + (Ψp)
= -1240 + 350
= -890 kPa

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Worked examples on the water
potential concept
• Example 3:
• A plasmolysed cell is found to have a
solute potential of –960 kPa. What is
the water potential of the cell?
• Answer
• In a plasmolysed cell, pressure
potential (Ψp) is 0.
• Therefore, Ψ = Ψs = -960 kPa

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Worked examples on the water
potential concept
• Example 4:
• Two plant cells, A and B, are next to each
other in a tissue. The water potential of
cell A is –700 kPa, and the water potential
of cell B is –550 kPa. In which direction
will water flow – from A to B, or from B to
A?
• Answer
• Water will flow from B to A. This is
because water flows down a water
potential gradient.

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c) Facilitated Diffusion

• Passive movement of molecules


down their concentration
gradient via transport proteins.

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• Two types of transport proteins:
i. Channel proteins
 Some provide hydrophilic corridors
for passage of specific molecules
or ions.
 Example, aquaporins,, facilitate
diffusion of water.
 Many ion channels function as
gated channels - open or close
depending on presence or absence
of a chemical or physical stimulus.

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EXTRACELLULAR
FLUID

Channel protein Solute


CYTOPLASM

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ii. Carrier proteins
 Some proteins translocate the
solute-binding site and solute
across the membrane as the
transport protein changes shape.

Carrier protein Solute

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3.3.2 Active Transport – Sodium
Pump and Coupled Transport

 Movement of a substance across a


biological membrane against its
concentration gradient or
electrochemical gradient with the help
of energy input (ATP) and specific
transport proteins.
• Example: sodium-potassium pump

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EXTRACELLULAR [Na+] high Na+
FLUID [K+] low Na+

Na+ Na+ Na+

Na+ Na+

Na+

[Na+] low ATP


Na+ P P
CYTOPLASM [K+] high
ADP
Cytoplasmic Na+ bonds to Na+ binding stimulates Phosphorylation causes
the sodium-potassium pump phosphorylation by ATP. the protein to change its
conformation, expelling Na+
to the outside.

K+

K+

K+
K+
K+

P
P K+

Extracellular K+ binds Loss of the phosphate K+ is released and Na+


to the protein, triggering restores the protein’s sites are receptive again;
release of the phosphate original conformation. the cycle repeats.
group.
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Passive transport Active transport

ATP
Diffusion Facilitated diffusion

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Maintenance of Membrane Potential by
Ion Pumps

• Membrane potential - voltage


difference across a membrane.
• Voltage difference is due to separation
of opposite charges.
• Cytoplasm is negative in charge
compared to extracellular fluid.
Due to unequal distribution of
cations & anions on opposite sides
of membrane.

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• Membrane potential favors passive
transport of cations into cell and
anions out of cell.
• Two combined forces, the
electrochemical gradient,, drive
diffusion of ions across a membrane.
1)Chemical force: an ion’s
concentration gradient.
2)Electrical force: effect of membrane
potential on ion’s movement.

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• Special transport proteins, the
electrogenic pumps,, generate voltage
gradient across a membrane.
• Example:
 Sodium-potassium pump in animal
cells.
 Proton pump in plants, fungi, &
bacteria.

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– EXTRACELLULAR
+
FLUID

ATP –
+ H+

H+
Proton pump
H+

– + H+

H+
– +
CYTOPLASM
H+
– +

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Cotransport
• The coupling of the diffusion of one
substance down its concentration
gradient to the transfer of another against
its concentration gradient.
• Transport protein may move two
substances in the:
Same direction – symport carriers.
Opposite directions – antiport carriers.
carriers

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• Plants commonly use the gradient of
hydrogen ions generated by proton pumps
to drive active transport of nutrients into
the cell.
– +
ATP H+
H+
– +
Proton pump H+
H+
– +
– H+
+
H+ Diffusion
Sucrose-H+ of H+
cotransporter
H+
– +
– + Sucrose
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3.3.3 Bulk Transport

• Small molecules and water enter or


leave the cell through the lipid bilayer
or by transport proteins
• Large molecules, such as
polysaccharides and proteins, cross
the membrane via vesicles

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Exocytosis

 The cellular secretion of


macromolecules by fusion of vesicles
with plasma membrane.
• Transport vesicles migrate to
membrane, fuse with it, and release
their contents
• Many secretory cells use exocytosis to
export their products

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ER

Transmembrane
glycoproteins

Secretory
protein

Glycolipid

Golgi
apparatus

Vesicle

Plasma membrane:
Cytoplasmic face
Extracellular face
Transmembrane
Secreted glycoprotein
protein

Plasma membrane:
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Endocytosis

 The cellular uptake of macromolecules


and particulate substances by
localized regions of plasma membrane
that surround the substances and
pinch off to form an intracellular
vesicle.
• Endocytosis is a reversal of
exocytosis, involving different proteins

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Three types of endocytosis:

1) Phagocytosis (“cellular eating”):


Cell engulfs particle in a vacuole.
2) Pinocytosis (“cellular drinking”):
Cell creates vesicle around fluid.
3) Receptor-mediated endocytosis:
Binding of ligands to receptors
triggers vesicle formation.

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RECEPTOR-MEDIATED ENDOCYTOSIS
Coat protein
Receptor Coated
vesicle

Coated
pit
Ligand

A coated pit
Coat and a coated
protein vesicle formed
during
receptor-
mediated
endocytosis
(TEMs).
Plasma
membrane
0.25 µm

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