BIOLOGICAL

MEMBRANE

 FLUID MOSAIC MODEL

 MEMBRANE TRANSPORT

 GIBBS – DONNAN EQUILIBRIUM

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THE CELL MEMBRANE

The cell membrane (also known as the plasma membrane or

cytoplasmic membrane, or plasmalemma) is a
biological membrane that separates the interior of all cells from the
outside environment (the extracellular space) .The cell membrane is a thin
semi-permeable membrane that surrounds the cytoplasm of a cell.
Animal cells, plant cells, prokaryotic cells, and fungal cells have plasma
membranes. Internal organelles are also encased by membranes.

Functions of plasma membrane

It protects the integrity of the interior of the cell by allowing certain

substances into the cell while keeping other substances out (molecular
traffic). It divide the internal space into discrete compartments to
segregate process and components. It also serves as a base of attachment
for the cytoskeleton in some organisms and the cell wall in others. Thus
the cell membrane also serves to help support the cell and helps to
maintain its shape.

Another function of the membrane is to regulate cell growth through the

balance of endocytosis and exocytosis. In endocytosis, lipids and proteins
are removed from the cell membrane as substances are internalized. In
exocytosis, vesicles containing lipids and proteins fuse with the cell
membrane increasing cell size.

The plasma membrane also plays a role in cell signalling and cell to cell
communication. Receptor proteins on the cell membrane can bind to
molecules of substances produced by other areas of the body, such as
hormones. When a molecule binds to its target receptor on the membrane,
it initiates a signal transduction pathway inside the cell that transmits the
signal to the appropriate molecules. Then, the cell can perform the action

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specified by the signal molecule, such as making or stopping production
of a certain protein.

Fluid Mosaic Model

The fluid mosaic model was first proposed by S.J. Singer and Garth L.
Nicolson in 1972 to explain the structure of the plasma membrane. The
fluid mosaic model describes the structure of the plasma membrane as a
mosaic of components —including phospholipids, cholesterol, proteins,
and carbohydrates—that gives the membrane a fluid character. Plasma
membranes range from 5 to 10 nm in thickness. For comparison, human
red blood cells, visible via light microscopy, are approximately 8 µm wide,
or approximately 1,000 times wider than a plasma membrane. The
proportions of proteins, lipids, and carbohydrates in the plasma
membrane vary with cell type. For example, myelin contains 18% protein
and 76% lipid. The mitochondrial inner membrane contains 76% protein
and 24% lipid.

According to this model, the biological membranes are considered to be

quasi fluid structure. This model is linked to icebergs (membrane
proteins) floating in a sea of phospholipid molecules.

The Components and functions of the Plasma Membrane:

The principal components of a plasma membrane are lipids

(phospholipids and cholesterol), proteins, and carbohydrates attached to

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some of the lipids and some of the proteins. Depending on the
membrane’s location and role in the body, lipids can make up anywhere
from 20 to 80 percent of the membrane, with the remainder being
proteins. While lipids help to give membranes their flexibility, proteins
monitor and maintain the cell's chemical climate and assist in the transfer
of molecules across the membrane.

Diagram depicting The fluid mosaic model of the plasma

membrane: The fluid mosaic model of the plasma membrane describes
the plasma membrane as a fluid combination of phospholipids,
cholesterol, and proteins. Carbohydrates attached to lipids (glycolipids)
and to proteins (glycoproteins) extend from the outward-facing surface
of the membrane.

The main fabric of the membrane is composed of amphiphilic (dual-

loving) phospholipid molecules. The hydrophilic (water-loving) areas of
these molecules are in contact with the aqueous fluid both inside and
outside the cell. Hydrophobic (water-hating molecules) tend to be non-
polar. A phospholipid molecule consists of a three-carbon glycerol
backbone with two fatty acid molecules attached to carbons 1 and 2, and a
phosphate-containing group attached to the third carbon. This
arrangement gives the overall molecule an area described as its head (the
phosphate-containing group), which has a polar character or negative
charge, and an area called the tail (the fatty acids), which has no charge.

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They interact with other non-polar molecules in chemical reactions, but
generally do not interact with polar molecules. When placed in water,
hydrophobic molecules tend to form a ball or cluster. The hydrophilic
regions of the phospholipids tend to form hydrogen bonds with water and
other polar molecules on both the exterior and interior of the cell. Thus,
the membrane surfaces that face the interior and exterior of the cell are
hydrophilic. In contrast, the middle of the cell membrane is hydrophobic
and will not interact with water. Therefore, phospholipids form an
excellent lipid bilayer cell membrane that separates fluid within the cell
from the fluid outside of the cell.

The whole bilayer arrangement is held together by noncovalent

interactions, such as van der Waals and hydrophobic interactions.
Because the bilayer is curved, the molecules of inner layer is more tightly
packed. Bulkier molecules such as cerebrosides (any of a group of complex
lipids present in the sheaths of nerve fibres) tend to be located in the outer
layer.

Diagram depicting Phospholipid aggregation: In an aqueous

solution, phospholipids tend to arrange themselves with their polar
heads facing outward and their hydrophobic tails facing inward.
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Diagram depicting The structure of a phospholipid molecule: This
phospholipid molecule is composed of a hydrophilic head and two
hydrophobic tails. The hydrophilic head group consists of a phosphate-
containing group attached to a glycerol molecule. The hydrophobic tails,
each containing either a saturated or an unsaturated fatty acid, are long
hydrocarbon chains.

Four major phospholipids predominate in the plasma membrane of many

mammalian cells

 Phosphatidylcholine
 Phosphatidylethanolamine
 Phosphatidylserine
 sphingomyelin

Cholesterol is another lipid component of animal cell membranes.

Cholesterol molecules are selectively dispersed between membrane
phospholipids. This helps to keep cell membranes from becoming stiff by
preventing phospholipids from being too closely packed together. Instead
of cholesterol plant cells have phytosterol.
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Glycolipids are located on cell membrane surfaces and have a
carbohydrate sugar chain attached to them. They help the cell to recognize
other cells of the body.

Membrane proteins

Proteins make up the second major component of plasma membranes,

which are associated with lipid bilayer in either of the two ways- as
peripheral proteins or as integral proteins.

a)Peripheral membrane proteins (extrinsic proteins) are associated

with membranes surfaces through weak electrostatic forces but do not
penetrate the hydrophobic core of the membrane. They are often found in
association with integral membrane proteins or to the polar head of lipid
molecule. In order to remove peripheral protein from cell membrane, do
not require the disruption of the membrane structure. Salt washes of
membranes are often sufficient to remove peripheral membrane proteins.

An example of peripheral membrane protein is the cytochrome C in the

electron transport chain.

b)Integral proteins or intrinsic proteins (some specialized types are

called integrin) integrated completely into the membrane structure, and
their hydrophobic membrane-spanning regions interact with the
hydrophobic region of the phospholipid bilayer. Single-pass integral
membrane proteins usually have a hydrophobic transmembrane segment
that consists of 20–25 amino acids. Some span only part of the
membrane—associating with a single layer—while others stretch from one
side of the membrane to the other, and are exposed on either side. Those
integral proteins which span entire bilayer is called transmembrane
proteins. They can act as pathways for ions and molecules.

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Polytopic transmembrane proteins cross the membrane several times.
Some are receptor proteins while others form channels.

Some complex proteins are composed of up to 12 segments of a single

protein, which are extensively folded and embedded in the membrane.
This type of protein has a hydrophilic region or regions, and one or several
mildly hydrophobic regions. This arrangement of regions of the protein
tends to orient the protein alongside the phospholipids, with the
hydrophobic region of the protein adjacent to the tails of the
phospholipids and the hydrophilic region or regions of the protein
protruding from the membrane and in contact with the cytosol or
extracellular fluid. Because of the higher affinity of integral protein
towards lipid bilayer, it is difficult to isolate so detergent treatment is often
required to remove integral membrane proteins.

Many hormone receptor molecules enzymes and signalling proteins are

integral proteins and specific polypeptide hormones that bind to these
receptors are peripheral proteins.

Diagram depicting Structure of integral membrane proteins:

Integral membrane proteins may have one or more alpha-helices that
span the membrane (examples 1 and 2), or they may have beta-sheets
that span the membrane (example 3).

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c)Amphitropic proteins are water-soluble channel-forming polypeptide
toxins, they bind weakly (reversibly) to membrane lipids, and this process
regulates their function. Proteins functioning in transduction of signals
generated in cell membranes are commonly regulated by amphitropism.

They have an affinity for both lipid and aqueous environments that is, the
protein has domains (structures) that are embedded in the membrane and
others that extend into the cytoplasm or the extracellular space. Some
pore-forming toxins are a good example.

Amphitropic proteins can be classified into three categories based on the

strategy employed to associate weakly with membranes.

 Lipid clamps – They contain structures (motif) with binding pockets

for lipid monomers.
 Covalent lipid anchors- They contain lipid covalent anchors which
is embedded in the lipid bilayer. A single lipid anchor provides
barely enough binding strength to maintain a membrane
attachment.
 Amphipathic helices- They contain amphipathic helix which binds
to the membrane there by separating the membrane such that it
hydrophobic face is isolated away from water and its polar
(hydrophilic) face can still contact the aqueous phase. It lies parallel
to the lipid bilayer surface. It doesn’t have binding site for lipid
monomer.

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Diagram depicting the three types of amphitropic proteins

a) Lipid clamps
b) Covalent lipid anchors
c) Amphipathic helices

Cell membrane proteins have a number of different functions.

1) Structural proteins help to give the cell support and shape.

2) Cell membrane receptor proteins help cells communicate with
their external environment through the use of hormones,
neurotransmitters, and other signalling molecules.
3) Transport proteins, such as globular proteins, transport
molecules across cell membranes through facilitated diffusion.
4) Glycoproteins have a carbohydrate chain attached to them. They
are embedded in the cell membrane and help in cell to cell
communications and molecule transport across the membrane.

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Diagram depicting lipid bilayer having membrane proteins, phospholipids
and glycoprotein

Membrane carbohydrates

Carbohydrates are the third major component of plasma membranes.

They sometimes branched chains of sugars attached on the exterior
surface of cells and are bound either to proteins (forming glycoproteins)
or to lipids (forming glycolipids). These carbohydrate chains may consist
of 2-60 monosaccharide units and can be either straight or branched. No
membrane carbohydrate is located at the interior surface.

The oligosaccharide chains of membrane glycoproteins and glycolipids

are formed by various combinations of six principal sugars D-galactose,
D-mannose, L-fucose, N-acetylneuraminic acid (also called sialic acid), N-
acetyl-D-glucosamine, and N-acetyl-D- galactosamine. All of these may be
derived from glucose.

Membrane carbohydrates perform two main functions: participate in cell

recognition and adhesion, either cell-cell signalling or cell-pathogen
interactions, and they have a structural role as physical barrier. For
instance, blood groups are determined by cell surface carbohydrates of
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erythrocytes, and they also have the ability to trigger immunological
responses.

Along with peripheral proteins, carbohydrates form specialized sites on

the cell 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”). Similar types of glycoproteins and glycolipids are
found on the surfaces of viruses and may change frequently, preventing
immune cells from recognizing and attacking them. These carbohydrates
on the exterior surface of the cell—the carbohydrate components of both
glycoproteins and glycolipids—are collectively referred to as the
glycocalyx (meaning “sugar coating”). The glycocalyx is highly hydrophilic
and attracts large amounts of water to the surface of the cell. This aids in
the interaction of the cell with its watery environment and in the cell’s
ability to obtain substances dissolved in the water.

Membrane fluidity

Fluidity of lipid bilayer means the freedom of motion of membrane

components. Lipid molecules are much smaller than proteins and
therefore especially mobile. In a lipid bilayer, the hydrophobic side chain
of fatty acids can be ordered to provide a rather stiff structure. Membrane
fluidity changes with temperature, it decreases as the temperature drops
and increases as temperature rises.

As the temperature increases, the hydrophobic side chains undergo a

transition from the ordered state to a disordered state, taking on a more
fluid arrangement. The temperature at which the structure undergoes the
transition from ordered state to disordered state is called transition

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temperature (Tm). This change of state of membrane is called phase
transition.

The fluidity of membrane primarily depends on the kind of lipids present.

The three most important aspects of lipid composition that affects
membrane fluidity are

 The length of fatty acid chains

 The degree of unsaturation of their side chains (the number of
double bonds present)
 The amount of cholesterol present

Longer and more saturated fatty acid chains interact more strongly with
each other thus cause high values of Tm. In turn, the disordered structure
caused by the presence of unsaturated fatty acids with cis double bonds in
their hydrocarbon chain causes greater fluidity in the bilayer. The lipid
components of a bilayer are always in motion to a greater extend in more
fluid bilayers and to a lesser extend in more rigid ones. The phospholipids
of cellular membranes contain at least one unsaturated fatty acid.

The lipid portion of a plant membrane has a higher percentage of

unsaturated fatty acids, than the lipid portion of an animal membrane. As
a result, animal membranes are less fluid (more rigid) than plant
membranes.

Cholesterol stabilises the extended straight chain arrangement of

saturated fatty acids by van der Waals interactions. It acts as a
bidirectional regulator of membrane fluidity because at high
temperatures, it stabilizes the membrane and raises its melting point,
whereas at low temperatures it intercalates between the phospholipids
and prevents them from clustering together and stiffening.

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Transport across biological membrane

The phospholipid bilayer determines what molecules can move into or out
of the cell, and so is in large part responsible for maintaining the delicate
homeostasis of each cell.
Cell membranes are semipermeable, meaning they have control over what
molecules can or cannot pass through. Some molecules can just drift in
and out, others require special structures to get in and out of a cell, while
some molecules even need an energy boost to get across a cell membrane.
Each cell’s membrane contains the right mix of these structures to help
that cell keep its internal environment just right.
There are two major ways that molecules can be moved across a
membrane, and the distinction has to do with whether or not cell energy
is used. They are
1) Passive transport
 Simple diffusion
 Facilitated diffusion
 osmosis
2) Active transport.
Simple diffusion
Diffusion is the net movement of molecules from a region of its higher
concentration to a region of its lower concentration down
a concentration gradient (across the concentration gradient) until the
concentration reaches equilibrium without the expenditure of energy.
Molecules and ions move freely in gases and liquids, each type of these
particles tends to spread out evenly within the space available by diffusion.
Small non polar mainly gaseous molecules such as oxygen, carbon
dioxide, nitrogen etc. will simply diffuse through cell membrane.
Molecules diffusing through cell membrane doesn’t interact with other
molecules
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Factors That Affect Diffusion
 Extent of the concentration gradient: The greater the difference in
concentration, the more rapid the diffusion. The closer the
distribution of the material gets to equilibrium, the slower the rate of
diffusion becomes.

 Mass of the molecules diffusing: Heavier molecules move more

slowly; therefore, they diffuse more slowly.

 Temperature: Higher temperatures increase the energy and

therefore the movement of the molecules, increasing the rate of
diffusion. Lower temperatures decrease the energy of the molecules,
thus decreasing the rate of diffusion.

 Solvent density: As the density of a solvent increases, the rate of

diffusion decreases. The molecules slow down because they have a
more difficult time getting through the denser medium. If the
medium is less dense, diffusion increases. Because cells primarily use
diffusion to move materials within the cytoplasm, any increase in the
cytoplasm’s density will inhibit the movement of the materials. An
example of this is a person experiencing dehydration. As the body’s
cells lose water, the rate of diffusion decreases in the cytoplasm, and
the cells’ functions deteriorate. Neurons tend to be very sensitive to
this effect. Dehydration frequently leads to unconsciousness and
possibly coma because of the decrease in diffusion rate within the
cells.

 Solubility: nonpolar or lipid-soluble materials pass through plasma

membranes more easily than polar materials, allowing a faster rate
of diffusion.

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  Surface area and thickness of the plasma membrane: Increased
surface area increases the rate of diffusion, whereas a thicker
membrane reduces it.

 Distance travelled: The greater the distance that a substance must

travel, the slower the rate of diffusion.

Examples of diffusion

 Exchange of respiratory gases.

 Transpiration (Release of oxygen out of the stomata of the leaf).
 The distribution of nutrients and water to all the parts of the plant.
 Diffusion of Carbon Dioxide into the leaf of the plants during
photosynthesis.

Diagram depicting transport of different types of molecules across membrane

Facilitated diffusion

Facilitated diffusion is the movement of specific molecules down a

concentration gradient with the aid of special channels or carrier protein
without expending energy. After attaining equilibrium there will be no net
movement of molecules.

Ions or electrically charged molecules are not able to diffuse through

the phospholipid bilayer because they are repelled from the hydrophobic
tails. Large molecules are also unable to move through

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the phospholipid bilayer freely. However, the cell membrane contains
special channel protein that provide hydrophilic passageways for these
special ions and molecules. Diffusion through these channel proteins is
called facilitated diffusion. Each carrier protein has its own shape and
only allows one molecule (or one group of closely related molecules) to
pass through. Selection is by size, shape and charge.

The material being transported is first attached to protein or glycoprotein

receptors on the exterior surface of the plasma membrane. This allows the
material that is needed by the cell to be removed from the extracellular
fluid. The substances are then passed to specific integral proteins that
facilitate their passage. Some of these integral proteins are collections of
beta-pleated sheets that form a channel through the phospholipid bilayer.
Others are carrier proteins which bind with the substance and aid its
diffusion through the membrane. Facilitated diffusion occurs by two ways

a) Channels: - The integral proteins involved in facilitated transport are

collectively referred to as transport proteins; they function as either
channels for the material or carriers. In both cases, they are
transmembrane proteins. Channels are specific for the substance that is
being transported. Channel proteins have hydrophilic domains exposed to
the intracellular and extracellular fluids; they additionally have a
hydrophilic channel through their core that provides a hydrated opening
through the membrane layers. Passage through the channel allows polar
compounds to avoid the nonpolar central layer of the plasma membrane
that would otherwise slow or prevent their entry into the cell.
Aquaporins are channel proteins that allow water to pass through the
membrane at a very high rate.

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Channel proteins are either open at all times or they are “gated,” which
controls the opening of the channel. The attachment of a particular ion to
the channel protein may control the opening or other mechanisms or
substances may be involved. In some tissues, sodium and chloride ions
pass freely through open channels, whereas in other tissues, a gate must
be opened to allow passage. An example of this occurs in the kidney, where
both forms of channels are found in different parts of the renal tubules.
Cells involved in the transmission of electrical impulses, such as nerve and
muscle cells, have gated channels for sodium, potassium, and calcium in
their membranes. Opening and closing of these channels changes the
relative concentrations on opposing sides of the membrane of these ions,
resulting in the facilitation of electrical transmission along membranes (in
the case of nerve cells) or in muscle contraction (in the case of muscle
cells).

b) Carrier Proteins: -Another type of protein embedded in the plasma

membrane is a carrier protein. This protein binds a substance and triggers
a change of its own shape, moving the bound molecule from the outside
of the cell to its interior; depending on the gradient, the material may
move in the opposite direction. Carrier proteins are typically specific for a
single substance. This adds to the overall selectivity of the plasma
membrane.

Example -Glucose Transporter (GLUT 4)

When food is digested, there is a high concentration of glucose within

the small intestine. This is transported through the membranes of the cells
of the alimentary canal, towards the endothelial cells lining blood
capillaries. Thereafter, glucose is transported throughout the body by
the circulatory system. When blood flows through tissues that need

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energy, glucose traverses the endothelial cell membranes again and enters
cells with low glucose concentration. Occasionally, when blood sugar
levels drop, the movement can occur in reverse – from body tissues into
blood circulation. For instance, hepatic cells can generate glucose even
from non-carbohydrate sources to maintain a basal blood sugar
concentration and prevent hypoglycemia.

The glucose transporter that facilitates this movement is a carrier protein

that has two major conformational structures. While the exact three-
dimensional structure is not known, the binding of glucose probably
causes a conformational change that makes the binding site face the
interior of the cell. When glucose is released into the cell, the transporter
returns to its original conformation.

Channel and carrier proteins transport material at different rates.

Channel proteins transport much more quickly than do carrier proteins.
Channel proteins facilitate diffusion at a rate of tens of millions of
molecules per second, whereas carrier proteins work at a rate of a
thousand to a million molecules per second.

Mechanism of facilitated transport (PING-PONG MECHANISM)

A ping pong model is put forth to explain the occurrence of facilitated

diffusion. According to this mechanism a transport (carrier) protein exists
in two conformations. In the pong conformation, it is exposed to the side
with high solute concentration. This allows the binding of solute to specific
sites on the carrier protein. The protein then undergoes a conformational
change (ping state) to expose to the side with low solute concentration
where the solute molecule is released. Hormones regulate facilitated

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diffusion. For instance, insulin increase glucose transport in muscles and
adipose tissues; amino acids transport in liver and other tissues.

An example is the movement of glucose into erythrocytes. The

concentration of glucose in the blood is about 5mM. The glucose
concentration in the erythrocyte is less than 5mM. Then glucose
transports through a carrier protein called glucose permease to
erythrocytes.

Diagram depicting
a) Channel proteins
b) Carrier protein

Osmosis
Osmosis is the diffusion of water (solvent) molecules from a region of its
higher concentration (dilute solution) to a region of its lower
concentration (concentrated solution) across a semi permeable

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membrane down a water potential gradient without expending any
energy.
The solute molecules are too large to get through the membrane. Water
molecules carry tiny electrical charges but they are small so, can move
freely through the phospholipid bilayer of most cell membranes and
diffuse across cell membranes.

Tonicity- is directly related to the osmolarity of a solution, affects

osmosis by determining the direction of water flow.

 Osmolarity describes the total solute concentration of a solution;

solutions with a low solute concentration have a low osmolarity,
while those with a high osmolarity have a high solute concentration.

 Water moves from the side of the membrane with lower osmolarity
(and more water) to the side with higher osmolarity (and less water).

 In a hypotonic solution, the extracellular fluid has a lower osmolarity

than the fluid inside the cell; water enters the cell.

 In a hypertonic solution, the extracellular fluid has a higher

osmolarity than the fluid inside the cell; water leaves the cell.

 In an isotonic solution, the extracellular fluid has the same

osmolarity as the cell; there will be no net movement of water into or
out of the cell.
Definition 2- Osmosis is the diffusion of solvent molecules from hypotonic
solution to hypertonic solution across a semi permeable membrane down
a water potential gradient without expending any energy.
How Osmosis Affects Cells
Osmosis affects plant and animal cells differently because plant and
animal cells can tolerate different concentrations of water. In a hypotonic
solution, an animal cell will fill with too much water and lyse, or burst

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 open. However, plant cells need more water than animal cells, and will not
burst in a hypotonic solution due to their thick cell walls; hypotonic
solutions are ideal for plant cells. The optimal condition for an animal cell
is to be in an isotonic solution, with an equal amount of water and solutes
both inside and outside. When a plant cell is in an isotonic solution, its
cells are no longer turgid and full of water, and the leaves of the plant will
droop. In a hypertonic solution, water will rush out of both animal and
plant cells, and the cells will shrivel. This is why slugs and snails shrivel
and die when salt is sprinkled onto them; water leaves their cells in order
to balance the higher concentration of salt outside the cells.
When a plant cell is placed on hypertonic solution, it became plasmolysed
and the cell is called plasmolysed cell. Plasmolysis means the shrinking of
protoplasm away from the cell wall of a plant due to water loss from
osmosis, thereby resulting in gaps between the cell wall and cell
membrane. This is why slugs and snails shrivel and die when salt is
sprinkled onto them; water leaves their cells in order to balance the higher
concentration of salt outside the cells. By keeping a plasmolysed cell in
water it undergoes deplasmolysis (water re-enters the cell). The solutions
that are isotonic to our cell content are
 0.9% NaCl solution
 5% glucose solution

Hypotonic solution Hypertonic solution Isotonic solution

Animal cell  Endosmosis  Exosmosis No change
(RBC)  Cell swells  Cell shrink
 Become turgid  Become flaccid
 Cell rupture  Become crenate
(RBC)

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Plant cell  Endosmosis  Exosmosis No change
 Cell swells  Cell shrink
 Become turgid  Become flaccid
 Cell never rupture  plasmolysed
due to the
presence of thick
cell wall

This figure shows the effects of osmosis on red blood cells

Examples of Osmosis
 Osmosis is how plants are able to absorb water from soil. The roots
of the plant have a higher solute concentration than the surrounding
soil, so water flows into the roots.
 Opening and closing of stomata.
The opening and closing of stomata is governed by increases or
decreases of solutes in the guard cells, guard cells absorb potassium ions.
So the solute concentration is higher in guard cells then water moves from
subsidiary cells to guard cells. Entering too much water makes guard cells
flaccid and stomata opens, the reverse is for closing the stomata.
 Movement of leaves is Mimosa pudica

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When you touch the leaves of Mimoa pudica, the inner cells the swollen
base of the leaf stalk (called the 'pulvinus') undergo exosmosis. Hence
become flaccid and the cells come closer and leaves folds.
 Germination of seeds
 Osmosis can have adverse effects on animals such as fish. If
freshwater or saltwater fish are put into water that has a different
salt concentration than they are used to, they will die from having
too much water enter or leave their cells.

Summary of passive transport

Active transport across cell membranes

Movement of molecules (mainly ions) across the cell membrane against its
concentration gradient (from lower concentration to higher concentration)
by expending energy usually in the form of adenosine triphosphate (ATP)
E.g. root hair cells in plants take in nitrate ions from the soil. Their
concentration are often higher inside the root hair cell than in the soil, so
the diffusion gradient is from the root hair à the soil. Despite this, the root
hair cells still can take nitrate ions in, by active transport.
The active transport is done using carrier (transporter) proteins in the cell
membrane. These use energy from the breakdown of ATP to move the
ions into the cell. The carrier proteins are ATPases. Each carrier protein

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is specific to just one type of ion or molecule. Cells contain many different
carrier proteins in their membranes.

Diagrams depicting active transport

Active transport mechanisms can be divided into two categories.
1) Primary active transport
2) Secondary active transport(cotransport)
1) Primary active transport
Directly uses a source of chemical energy (e.g., ATP) to move molecules
across a membrane against their gradient. The most important primary
active transport systems are ion-pumps (through the involvement of
pump ATPase or ion transporting ATPase)
The sodium-potassium pump
One of the most important pumps in animal cells is the sodium-
potassium pump, it plays a very important role in nerve cell. The cells

have a high intracellular K+ concentration and a low Na+ concentration,

this is essentially needed for the survival of the cells. High cellular K+ is

required for the optimal glycolysis (pyruvate kinase is dependent on K+)

and for protein biosynthesis.

3 positive Na+ ions are pumped out of the cell for every 2 positive K+ ions

pumped into the cell, with the help of integral plasma membrane protein-

Na+-K+ ATPase. As a result, positive charge builds up outside the cell

compared to inside the cell. The difference in charge between the outside

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 and inside of the cell allows nerve cells to generate electrical impulses
which lead to nerve impulses.
Not only the sodium-potassium pump maintain correct concentrations of
Na+ and K+ in living cells, but it also plays a major role in generating the
voltage across the cell membrane in animal cells. Pumps like this, which
are involved in the establishment and maintenance of membrane voltages,
are known as electrogenic pumps. The sodium-potassium pump
transports sodium out of and potassium into the cell in a repeating cycle
of conformational (shape) changes. In each cycle, three sodium ions exit
the cell, while two potassium ions enter. This process takes place in the
following steps:
1. To begin, the pump is open to the inside of the cell. In this form, the pump
really likes to bind (has a high affinity for) sodium ions, and will take up
three of them.
2. When the sodium ions bind, they trigger the pump to hydrolyse (break
down) ATP which is generated through cellular respiration. One
phosphate group from ATP is attached to the pump, which is then said to
be phosphorylated. ADP is released as a by-product.
3. Phosphorylation makes the pump change shape, re-orienting itself so it
opens towards the extracellular space. In this conformation, the pump no
longer likes to bind to sodium ions (has a low affinity for them), so the
three sodium ions are released outside the cell.
4. In its outward-facing form, the pump likes to bind to (has a high affinity
for) potassium ions. It will bind two of them, and this triggers removal of
the phosphate group attached to the pump in step 2.
5. With the phosphate group gone, the pump will change back to its original
form, opening towards the interior of the cell.
6. In its inward-facing shape, the pump loses its interest in (has a low affinity
for) potassium ions, so the two potassium ions will be released into the

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cytoplasm. The pump is now back to where it was in step 1, and the cycle
can begin again.
This may seem like a complicated cycle, but it just involves the protein
going back and forth between two forms: an inward-facing form with high
affinity for sodium (and low affinity for potassium) and an outward-facing
form with high affinity for potassium (and low affinity for sodium). The
protein can be toggled back and forth between these forms by the addition
or removal of a phosphate group, which is in turn controlled by the
binding of the ions to be transported.

The overall reaction of Na+-K+ pump

3 Na+ (in) + 2 K+ (out) + ATP ---- 3 Na+ (out) + 2 K+ (in) + ADP + Pi

Diagrams depicting the sodium-potassium pump

2) Secondary active transport

Secondary active transport (cotransport) uses an electrochemical
gradient – generated by active transport – as an energy source to move
molecules against their gradient, and thus does not directly require a
chemical source of energy such as ATP.
For example: - In secondary active transport, the movement of the sodium
ions down their gradient is coupled to the uphill transport of a glucose

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molecule up its gradient and into the cell. The carrier protein
(a cotransporter) uses the energy of the sodium gradient to drive the
transport of glucose molecules.

Diagram of a sodium-glucose cotransporter, which uses the energy

stored in a sodium ion gradient to transport glucose "uphill" against its
gradient. The cotransporter accomplishes this by physically coupling the
transport of glucose to the movement of sodium ions down their
concentration gradient.
An electrochemical gradient, created by primary active transport, can
move other substances against their concentration gradients, a process
called secondary active transport or co-transport.
Another example:- galactoside permease in bacteria. The lactose
concentration inside the bacterial cell is higher than outside, so moving
lactose into the cell requires energy. The galactoside permease doesnot
directly hydrolyse ATP. Instead it use the energy generated by letting
hydrogen ions flow through permease into the cell down their
concentration gradient. Active transporters that create hydrogen ion
gradients for the movement of molecules against concentration gradient
are called proton pumps.

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 Co-transport

Cotransport is the name of a process in which two substances are

simultaneously transported across a membrane by one protein, or protein
complex which does not have ATPase activity. It is also called coupled
transport. Different types of co-transport are

a) Symport- When both substances are transported in the same direction

by a transport protein is known as a symport and the carrier protein is
called symporter.

Examples- Sodium-Glucose co-transport mechanism. On its exterior side

the transport protein have 2 binding sites, one for sodium and one for
glucose. When both of these bind to the protein there is a conformational
change allowing the electrochemical gradient to provide the energy
needed to transport both of these molecules into the cell.

 Sodium-Amino acid co-transport occurs in the same manner as for

sodium-glucose co-transport.

b)Antiport- When the substances are transported in opposite directions

by a transport protein is known as antiport and the carrier protein is called
antiporter

Examples- Na+-Ca2+ counter-transport where Na+ binds to the transport

carrier protein on its exterior side, and Ca2+ bound to the same protein on
the membranes interior side. Once both are bound, a conformational
change occurs which releases energy and the sodium ion is transported to
the interior and calcium to the exterior. This transporter is situated on
almost all cell membranes.

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  Na+-H+ counter transport. The mechanism is the same as Na+-
Ca2+ counter-transport. The advantage of this transporter is clearly
seen in the proximal tubules of the kidneys.

Simple diagram of a symporter (carrying two molecules in the same direction) and
an antiporter (carrying two molecules in opposite directions
Note: Transporters: Transporters (also known as carriers) are the
membrane proteins that transport a wide variety of ions and molecules
across the lipid bilayer membrane.
Cotransporters: Cotransporters are proteins that transport two
different solutes such as glucose and amino acids simultaneously across
the cell membrane against a concentration gradient. It mediates coupled
reactions in which an energetically unfavourable reaction (uphill
movement of molecules) is coupled to an energetically favourable
reaction.
C)Uniport : Facilitated diffusion of single solute molecule across plasma
membrane which is carried out by the protein uniporter .
 Example – GLUT 1 glucose transport

Diagram depicting uniport (left) symport (middle) and antiport (right)

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Active transport performs three major functions in cells and
organelles are
 It makes possible the uptake of energy rich molecules and other
essential nutrients from the surrounding fluid even when their
concentrations in the surrounding are very low.
 It allows various substances such as secretory products, waste
materials and sodium ions to be removed from the cell and
organelles even when the concentration outside is greater than
the outside.
 It allows the cell to maintain constant, optimal internal
concentrations of inorganic ions particularly potassium,
calcium and hydrogen ions.

Diagrams depicting passive transport and active transport

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 Endocytosis and exocytosis (bulk transport)
Macromolecules are too large to move with membrane proteins and must
be transported across membranes in vesicles. That is, the mechanism
involve enclosing the substances to be transported in their own small
globes of membrane, which can then bud from or fuse with the membrane
to move the substance across.

Diagrams depicting exocytosis and endocytosis

Exocytosis
Cells release molecules such as signalling proteins and waste products, to
the outside environment by the process called exocytosis. Exocytosis (exo
= external, cytosis = transport mechanism) is a form of bulk transport in
which materials are transported from the inside to the outside of the cell
in membrane-bound vesicles that fuse with the plasma membrane.
Some of these vesicles come from the Golgi apparatus and contain
proteins made specifically by the cell for release outside, such as signalling
molecules. Other vesicles contain wastes that the cell needs to dispose of,
such as the leftovers that remain after a phagocytosed particle has been
digested.
These vesicles are transported to the edge of the cell, where they can fuse
with the plasma membrane and release their contents into the
extracellular space. Some vesicles fuse completely with the membrane and

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are incorporated into it, while others follow the “kiss-and-run” model,
fusing just enough to release their contents (“kissing” the membrane)
before pinching off again and returning to the cell interior

Diagrams depicting mode of exocytosis

Endocytosis

 Endocytosis (endo = internal, cytosis = transport mechanism) is a

general term for the various types of active transport that move
particles into a cell by enclosing them in a vesicle made out of
plasma membrane.
 There are variations of endocytosis, but all follow the same basic
process. First, the plasma membrane of the cell invaginates (folds
inward), forming a pocket around the target particle or particles.
The pocket then pinches off with the help of specialized proteins,
leaving the particle trapped in a newly created vesicle or vacuole
inside the cell.
 Endocytosis can be further subdivided into the following categories:
phagocytosis, pinocytosis, and receptor-mediated endocytosis.

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Endocytosis can be further subdivided into the following categories:
phagocytosis, pinocytosis, and receptor-mediated endocytosis.

 Phagocytosis

Phagocytosis (literally, “cell eating”) is a form of endocytosis in which

large particles, such as cells or cellular debris, are transported into the cell.
(Single-celled eukaryotes called amoebas also use phagocytosis to hunt
and consume their prey).
Once a cell has successfully engulfed a target particle, the pocket
containing the particle will pinch off from the membrane, forming a
membrane-bound compartment called a food vacuole. The food vacuole
will later fuse with an organelle called a lysosome (called the recycling
center of the cell). Lysosomes have enzymes that break the engulfed
particle down into its basic components (such as amino acids and sugars),
which can then be used by the cell.

 Pinocytosis
Pinocytosis (literally, “cell drinking”) is a form of endocytosis in which
a cell takes in small amounts of extracellular fluid. In pinocytosis the cell
engulfs drops of fluid by pinching in and forming vesicles that are smaller
than the phagosomes formed in phagocytosis. Like phagocytosis,
pinocytosis is a non-specific process in which the cell takes in whatever
solutes that are dissolved in the liquid it envelops..

 Receptor-mediated endocytosis

Unlike phagocytosis and pinocytosis, receptor-mediated endocytosis is an

extremely selective process of importing materials into the cell. This
specificity is mediated by receptor proteins (which are transmembrane
proteins) located on depressed areas of the cell membrane called coated

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pits. The cytosolic surface of coated pits is covered by coat proteins. In
receptor-mediated endocytosis, the cell will only take in an extracellular
molecule if it binds to its specific receptor protein on the cell’s surface.
Once bound, the coated pit on which the bound receptor protein is located
then invaginates, or pinches in, to form a coated vesicle. Similar to the
digestive process in non-specific phagocytosis, this coated vesicle then
fuses with a lysosome to digest the engulfed material and release it into
the cytosol. Mammalian cells use receptor-mediated endocytosis to take
cholesterol into cells. Cholesterol in the blood is usually found in lipid-
protein complexes called low-density lipoproteins (LDLs). LDLs bind to
specific receptor proteins on the cell surface, thereby triggering their
uptake by receptor-mediated endocytosis.

Diagrams depicting phagocytosis (left) pinocytosis (middle) and

Receptor-mediated endocytosis (right).

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