CELLULAR TRANSPORT MECHANISMS

A living cell is in a non-stop give and take that never stops unless the cell dies. All cells require nutrients and oxygen
essential to life, and also cells produce wastes that must be removed as fast as possible. Thus, all living cells constantly
take up and give off materials through their cell membrane; i.e. there is a continuous exchange of materials between a cell
and its outer environment: to obtain nutrients, to remove wastes, to generate ionic gradient across membranes and to
maintain suitable or normal internal cell environment. In this regard, the cell membrane plays vital role in controlling the
entry and exit of materials. And all materials must cross the cell membrane so as to get in to the cell or to get out of the
cell. However, not all particles can actually pass through the plasma membrane freely. This is because of largely the lipid
nature of the cell membrane (phospholipid bilayer) which can only allow lipid soluble, nonpolar substances like oxygen
(O2), cholesterol (steroid hormones) and carbon dioxide (CO2), and very small polar substances like water and ammonia.
The diffusion of water through the plasma membrane is of such importance to the cell that it is given a special name
called osmosis, and the lipid bilayers are impermeable to all ions such as K +, Na+, Ca2+, Cl-, HCO3-, H+ and impermeable to
most essential molecules like glucose, amino acids, and macromolecules such as peptides, proteins, polysaacharides.
Thus, if the cell membrane were entirely made up of phospholipid bilayer, larger and polar substances such as sugars,
ions, amino acids and other similar substances could not be allowed to enter in to cells which means starvation of the cell.
However, the cell membrane is also composed of different types of proteins (transport proteins) that serve as windows
or doors through which larger polar substances can enter in to the cell.
The plasma membrane allows some materials to pass while excluding others. This is the property of the cell membrane
known as Selective Permeability. This Selective Permeability influences movement both into and out of the cell, and it is
the key to its function.
The processes by which substances can cross the plasma membrane in to or out of a cell are categorized in to two:
1. Passive transport mechanisms and 2. Active transport mechanisms
1. Passive transport mechanisms
Passive transport mechanisms do not require the cell to do work for the substance to enter or leave the cell. Instead the
energy involved comes from the kinetic energy of the molecules in solution. No cellular energy is required because
substances move across the plasma membrane down their concentration gradient or electrochemical gradient.
Passive transport includes:
A) Simple diffusion B) Facilitated diffusion C) Osmosis

A) SIMPLE DIFFUSION
Diffusion is the net movement of substances (liquid or gas) down their concentration gradient (along their concentration
gradient) - that is from high concentration to low concentration. Since the molecules of any substance (solid, liquid, or
gas) are in motion when that substance is above absolute zero (-273°C), energy is available for movement of the
substances from a higher potential state to a lower potential state. Substances have kinetic energy, which makes them
move about randomly, and substances move spontaneously due to their own kinetic energy.
The majority of substances move from higher to lower concentration, although there will be some that move from low to
high. The overall (or net) movement is thus from high to low concentration. Net directional movement = The net
movement of a substance down a concentration gradient.

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Eventually, if no energy is input into the system, the substances will reach a state of dynamic equilibrium where that
substances become evenly distributed throughout the system, and when there is no net movement of molecules from either
side. And potential energy reaches zero. (There is still plenty of kinetic energy).

The essential characteristics of diffusion are:

 Each molecule or ion move independently of the others and
 Results from the intrinsic kinetic energy of molecules
 Random molecular motion, even though the net movement may be directional
 Diffusion continues until a dynamic equilibrium is reached after which molecules continue to move, but there is
no net directional movement.
 A substance diffuses down its own concentration gradient and is not affected by the gradients of other substances.
Factors that affect diffusion
The rate of diffusion is affected by the following factors:
The steepness of the concentration gradient. (Concentration gradient = Regular, graded concentration change over a
distance in a particular direction or the difference in concentration between two regions). The bigger the difference in
concentration between the two sides of the membrane the quicker the rate of diffusion. high potential gradient.
The surface area. Diffusion is a function of surface area – that is for large amount of substances to diffuse and cross
membranes, there should be enough surface area to take place. Thus, the greater the surface area the faster is the diffusion.
Thickness of membranes- the thinner the membrane the faster the rate of diffusion, or distance that substances move-
efficient diffusion requires small distances
The effects of the above three factors on the rate of diffusion in cells is given by Fick’s law
Rate α Surface Area x Conc. Gradient
Thickness
Temperature. Diffusion is possible at any temperature above absolute zero (-273°C). The higher the temperature the
more rapid the diffusion. Higher temperatures give molecules or ions more kinetic energy. Molecules move around faster,
so diffusion is faster.
The type of molecule or ion diffusing. Large molecules need more energy to get them to move so they tend to diffuse
more slowly. Non-polar molecules diffuse more easily than polar molecules because they are soluble in the non-polar
phospholipid tails.
The rate of diffusion is inversely proportional to the square root of the density of the diffusing substance (Graham’s law
of diffusion).

Importance of diffusion
Simple diffusion is one principle method of movement of substances within cells, as well as the method for essential small
molecules to cross the cell membrane. Some substances can move across the cell membrane by simple diffusion, and
these include small nonpolar lipid soluble molecules, such as 0 2, benzene and small, uncharged but slightly polar
molecules such as H2O, CO2, alcohol and glycerol.
Gas exchange in gills and lungs operates by this process. It is the process involved in transpiration. It is a means of
spreading of ions and other substances throughout the protoplast. It is an effective means of transport of substances, over a

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very short distance. Aroma of flowers is due to diffusion of volatile aromatic compounds and helps to attract pollinating
animals.
B) FACILITATED DIFFUSION

Facilitated diffusion is the passage of molecules or ions across a biological membrane through specific transport
proteins and requires no energy input. Large polar or hydrophilic molecules such as glucose and amino acids, and ions
such as Na+, K+, Ca++ or Cl cannot diffuse across the phospholipid bilayer. The transport of these molecules and ions
across membranes along their concentration gradient is through special transmembrane proteins or integral proteins
that are embedded in the membrane bilayer and facilitate the passage of selected ions and other polar molecules. These
transport proteins are sometimes called permeases, and could be either channels proteins or carrier proteins. Thus,
facilitated diffusion occurs via channels proteins and/or carrier proteins, transport proteins that exhibit specificity for
the transported molecules. Not only is it faster than simple diffusion but it is also responsible for providing a pathway for
ions and large polar molecules to traverse membranes that would otherwise be impermeable to them.
Like normal diffusion, facilitated diffusion is driven by potential energy of a concentration gradient.
Two major types of transport membrane proteins are
1. Channel proteins and 2. Carrier proteins

1. Channel proteins
Channel proteins are multi-pass transmembrane proteins that form small selective aqueous pores (water-filled and
hydrophilic tunnels or pores) across membranes through which ions and some small hydrophilic (water-soluble)
molecules can pass by facilitated diffusion. Channel proteins are especially important in transporting ions, and sometimes
called ion channels. The channels are selective; that is, the structure of the protein admits only certain types of molecules
through. The size of pore and density of surface charges on its interior lining determine its transport specificity.

Channel transport is mainly limited to ions or water. For instance, Na+,

Ca+, K+, etc. But there are some special channels that are used for
passage of other chemicals. For instance, the channels called
Aquaporins are channels designed for the rapid transport of water
across the cell membrane (especially important in the kidney cells for
reabsorption of huge water from the glomerular filtrate in cases of
dehydration of the body).

Porins are proteins that form huge pores in the outer membranes of the plastids, mitochondria and some bacteria allowing
molecules up to the size of small proteins to pass through.

2. Carrier proteins
Carrier proteins are multi-pass transmembrane proteins that undergo reversible conformational changes to transport
specific molecules across the membrane. The so-called "ping-pong" model suggests that the binding of a molecule
causes a reversible conformation change, which allows the transported substance to be released on the other side of the
membrane. That is carrier proteins bind to a substance and physically move it across the membrane. Once the molecule is
released the protein's shape reverts to the original or unoccupied state.
Unlike the channel proteins, carrier proteins function in both passive transport (facilitated diffusion) and active transport.

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Features of carrier protein mediated transport are:
 Saturation
Due to the specific binding step, a facilitated transporter can saturate if the concentration of the transported substance
becomes high enough. Once this happens, no further increase in the rate of transport can occur, since the binding site on
the transporter is occupied essentially all the time. By contrast, this does not occur with ion channels. As the concentration
of the ion increases, the amount that moves through the pore of the channel increases proportionately, and there is no
process that becomes saturated case of channel protein transport.
 Carriers are highly selective for a particular substrate to be transported.
 A conformational change in the protein is required to transport individual molecules or ions hence the rate of
transport is slower than that of the transport by channel proteins.

Some carrier proteins allow transport, only if two types of molecules move together.
This is called cotransport. When a molecule moves across a membrane independent
of other molecule, the process is called uniport.
Again cotransport can be symport or antiport. In symport, two types of molecules
cross the membrane in the same direction
In antiport, two types of molecules cross the membrane in the opposite direction

Properties of facilitated diffusion

 Movement is still passive (like diffusion), from high concentration to low. Concentration gradient must already be
present for molecules to diffuse. Thus, No ATP is required, and the energy is provided by the concentration
gradient of the substance transported.
 Proteins known as transport proteins in cellular membranes act as carriers or channels/pores to permit flux (influx
or efflux) of substances that cannot diffuse directly through the membrane and thus occurs across cell membranes
only.
 Saturation: A facilitated transporter can saturate if the concentration of the transported substance becomes high
enough. Once this happens, no further increase in the rate of transport can occur, since the binding site on the
transporter is occupied essentially all the time. This is true especially for carrier protein mediated transport.
Maximum rate of transport (the rate when the transporters are fully saturated) is called Tm- the transport
maximum
 Specificity: The transport proteins are specific and able to combine with only certain molecules. However, related
substances can compete for the same carrier or pore.
 Sensitive to inhibitors: The transport proteins are affected by substances called inhibitors which react with the
protein side chains
.
C) OSMOSIS

A cell is a solution of solutes such as salts, sugars, amino acids and peptides/proteins, and water being the solvent– a fairly
concentrated solution, inside a partially permeable membrane (the cell membrane). The salutes (salts, sugars, amino
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acids/peptides) do not freely diffuse out of the cell, but the phospholipid bilayer allows the solvent (water) to pass through
it freely, and water can freely pass through the cell membrane (phospholipid bilayer) from area of high water
concentration to an area of low water concentration. This movement (diffusion) of water through the phospholipid bilayer
is what is called osmosis. Thus, osmosis can be defined as a special kind of diffusion in which water, through a
semipermeable membrane, moves:

What makes osmosis a special type of diffusion?

1. It is diffusion of water (only water)

2. It requires a semipermeable membrane (plasma membrane) that is permeable to water but not to solute,

3. Also there should be two solutions of different concentrations separated by the selectively permeable membrane

When the above criteria are met, water diffuses down its concentration gradient- from an area of high water to an area of
low water (or water diffuses from the hypoosmotic solution (hypotonic solution)-solution with the lower osmotic
concentration, to the hyperosmotic solution (hypertonic solution)- solution with the higher osmotic concentration). For
water to move by osmosis, it is the difference in the free water concentration that is important. The presence of
hydrophilic solutes can reduce the proportion of water molecules that can freely diffuse. Water molecules form a
hydrated shell around hydrophilic solute molecules and this bound water cannot freely diffuse across a membrane. In
dilute solutions, including most biological fluids, it is the different in the proportion of the unbound water that causes
osmosis, rather than the actual difference in water concentration.

Water Potential
Water potential is the fundamental concept for understanding movement of water. The difference between the free energy
of water molecules in pure water, and the energy of water in any other system (e.g., water in a solution or in a cell or
tissue) is termed the water potential. Water potential is the measure of the ability or tendency of water to move out of a
solution by osmosis. It is represented by Greek letter Ѱ (psi) or more accurately Ѱw. The value of Ѱw is measured in bars,
pascals or atmospheres.
The pure water, at normal temperature and pressure has a water potential of zero (0). The presence of solute particles
reduces the free energy of water, and thus decreases the water potential (negative value). Therefore, the water potential of
a solution is always less than zero. Hence, water always moves from the less negative potential (–100 kPa) to more
negative potential (–200 kPa).

The two major factors that determine water potential are

1. Solute potential and 2. Pressure potential

1. Solute potential- Ѱs (aka Osmotic potential) = is the decrease in the water potential of a solution over its pure state as
a result of the presence of solutes. Solute potential or osmotic potential ( Ѱs) is always in negative value. The more the
solute molecules, the lower is the solute potential (Ѱs)-which means large negative value.
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Thus, solution with lower solute potential (larger negative value) (Ѱs)-invites water in to it, and such a solution which can
cause an osmotic entry of water in to it, is said to be osmotically active solution. It possesses a low water potential or has
high Osmotic concentration (total solute concentration of a solution) or has high osmotic pressure). Diffusion of water
into the osmotically active solution will continue across the separating membrane until an equilibrium is reached. At
equilibrium, water potential becomes equal on both sides of the membrane.

Osmotic pressure(π ) is the measure of the tendency for a solution to take up water when separated from pure water by a
selectively permeable membrane. The osmotic pressure of pure water is zero (the lowest osmotic pressure). Osmotic
pressure of a solution is proportional to its osmotic concentration (Total solute concentration of a solution). (The greater
the solute concentration, the greater the osmotic pressure), or the osmotic pressure of a solution largely depends upon the
ratio between concentration of solute and solvent particles in a given solution.
For example, a molar solution of sucrose has an osmotic pressure of approximately 22.4 atmospheres at 0°C. However, a
molar solution of sodium chloride has almost twice the osmotic pressure of a molar solution of sucrose. This is because,
sodium chloride is an electrolyte, so it dissociates almost completely into Na+ and Cl– ions (giving twice the particle
number), whereas sucrose or glucose molecules are nonelectrolytes, and do not dissociate in water. Osmotic pressure is
numerically equal to osmotic potential (= solute potential, Ѱ s ) but while osmotic potential has a negative value, osmotic
pressure(π ) has a positive value.

2. Pressure potential-Ѱp (aka Hydrostatic pressure or turgor pressure wall pressure)- is the pressure which develops
in an osmotic system due to osmotic entry or exit of water from it. A positive hydrostatic pressure develops due to the
entry of water into it. A negative hydrostatic pressure develops due to loss of water.

And thus, water potential is the sum of Solute potential and Pressure potential
Water potential= Solute potential + Pressure potential
Ѱw = Ѱs +Ѱp
 In a flaccid or plasmolyzed cell, Pressure potential (Ѱp) =0, thus Ѱw = Ѱs

 In a fully turgid cell, Ѱp = Ѱs and thus Ѱw =0 [no absorption water by the cell in a fully turgid condition.]

The instrument used for measuring osmotic pressure is called osmometer. In one type of osmometer, pure water is
separated from a solution by a selectively permeable membrane that is permeable to water but not solute. The tendency
for water to move into the solution by osmosis is counteracted by applying enough pressure with a piston so the solution's
volume will stay the same. Thus, the amount of pressure required to prevent net movement of water into the solution is the
osmotic pressure.

Conditions of osmosis
The three conditions of osmosis are hypertonic, hypotonic and isotonic
1. Hypertonic environment- an environment with a solution that is more concentrated (greater solute concentration) than
the cell inside or it is a solution with lower water potential. In other words, it is a solution that has high osmotic
concentration, hence high osmotic pressure, but more negative solute potential
When a cell is put in hypertonic solution, the cell loses its water- Exosmosis, and the cell shrinks or shrivels, and probably
dies. For example, an increase in the salinity (saltiness) of a lake can kill aquatic animals. If the lake water becomes
hypertonic to the animals’ cells, the cells may shrivel and die. When a plant cell is placed in a hypertonic solution, the net
movement of water by osmosis is from the cell to the outside environment. Thus, the cell losses water and shrinks. This
condition is known as Plasmolysis for animal cells, and for animal cells it is known as Crenation
2. Hypotonic environment- an environment with a solution that is less concentrated (lower solute concentration) than the
cell inside or it is a solution with higher water potential, or Hypotonic solution = a solution that has low osmotic
concentration, hence low osmotic pressure, but less negative solute potential
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 When a cell is put in a hypotonic solution, the cell gains water by
osmosis- Endosmosis.
For plant a cell in a hypotonic solution, the net movement of water by
osmosis is from the hypotonic solution in to the cell. Thus, the cell gains
water and becomes bulged or turgid. This condition is known as
Turgidity. And if an animal cell is put in a hypotonic solution, water
enters in to the cell from the outside by endosmosis. This entry of water
in to an animal cell causes the cell to swell and finally lyses (bursts) like an overfilled water balloon.
A red blood cell placed in a hypotonic solution (e.g., pure water) bursts immediately ("hemolysis") from the influx of
water.
N.B. Plasmolysis and turgor pressure are applied only to plant cells
3. Isotonic environment -an environment with a solution that has equal concentration with that of the cell inside or it is a
solution with equal water potential with that of the cell inside. If a cell is put in an isotonic solution, water diffuses across
the membrane at the same rate in both directions. Thus, there is no net movement of water across the plasma membrane
(or no osmosis).

N.B.
Aquaporins — transmembrane proteins that form hydrophilic channels or pores through which water can pass, and they
greatly accelerate water transport across membranes specially in organs like the kidneys. Each aquaporin allows entry of
as many as 3 billion (109) water molecules per second, passing single file through its central channel, which fits 10 at a
time. Without aquaporin, only a tiny fraction of these water molecules would pass through the same area of the cell
membrane in a second, so the channel protein greatly increases the rate of water movement.

Cell survival depends on the balance between water uptake and loss.
1. Water balance of cells without walls
Organisms without rigid cell walls have osmotic problems in either a hypertonic or a hypotonic environment (due to
excessive osmotic loss of water in hypertonic environment or due to excessive osmotic uptake of water in hypotonic
environment). In a hypertonic environment, an animal cell will lose water by osmosis and crenate (shrivel). In a
hypotonic environment, an animal cell will gain water by osmosis, swell and perhaps lyse (cell destruction).
Water balance is not a problem if such a cell lives in isotonic surroundings. The cells of most terrestrial animals are
bathed in extracellular fluid that is isotonic to the cells. Seawater is isotonic to many marine invertebrates.
However, animals and other organisms without rigid cell walls living in hypertonic or hypotonic environments must have
adaptations for osmoregulation, the control of water balance; that is, they must have mechanisms to prevent excessive loss
or excessive uptake of water by:
 Living in an isotonic environment (e.g., many marine invertebrates are isosmotic with sea water).
 Osmoregulating in a hypo- or hypertonic environment. Organisms can regulate water balance (osmoregulation) by
removing water in a hypotonic environment (e.g., Paramecium with contractile vacuoles in fresh water) or conserving
water and pumping out salts in a hypertonic environment (e.g., bony fish in seawater).

Paramecium is hypertonic to the pond water in which it lives. water

continually enters the Paramecium cell. To solve this problem,
Paramecium and some other single-celled freshwater organisms have a
specialized organelle, the contractile vacuole, which functions to actively
pump out the entering water

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When red blood cells are placed in a physiological saline (0.9% NaCl salt solution), they neither gain nor lose water by
osmosis. Such a solution is said to be isotonic. The extracellular fluid (ECF) of mammalian cells is isotonic to their
cytoplasm. This balance must be actively maintained because of the large number of organic molecules dissolved in the
cytosol but not present in the ECF. These organic molecules exert an osmotic effect that, if not compensated for, would
cause the cell to take in so much water that it would swell and might even burst. This fate is avoided by pumping sodium
ions out of the cell with the Na+/K+ ATPase
The concentration of water can be altered by the active transport of solutes and in this way the movement of water in and
out of the cell can be controlled.
Example: the reabsorption of water from the kidney tubules back into the blood depends on the water following behind
the active transport of Na+.
One of the major functions of blood in animals is to maintain an isotonic internal environment. This eliminates the
problems associated with water loss or excess water gain.

Sea water= hypertonic environment

Sea water is also hypertonic to the ECF of most marine vertebrates. To avoid fatal dehydration, these animals (e.g., bony
fishes like the cod) must continuously drink sea water and then desalt it by pumping ions out of their gills by active
transport. Marine birds and sea turtles use a similar desalting mechanism. They drink salt water to take care of their water
needs and use metabolic energy to desalt it. In the herring gull, the salt is extracted by two glands in the head and released
(in a very concentrated solution to the outside through the nostrils.

2. Water balance of cells with walls

The cells of plants, prokaryotes, fungi, and some protists are surrounded by walls. A plant cell in a hypotonic solution
takes up water by osmosis and swells until the elastic cell wall exerts turgor pressure on the cell content that opposes
further water uptake. A dynamic equilibrium is established- water enters and leaves the cell at the same rate and the cell
becomes turgid (very firm), and this is a healthy state for most plant cells. Turgid cells contribute to the mechanical
support of the plant (as water provides hydrostatic skeleton for soft plant organs such as leaves and young stem). Thus, it
requires cells to be hyperosmotic(hypertonic) to their environment.
Plants because of their cell walls and large central vacuoles can take advantage of steep water gradients to maintain the
shape and rigidity of leaves and stems.
Plant cells and bacterial cells avoid bursting in hypotonic surroundings by their strong cell walls. These allow the buildup
of turgor within the cell. When the turgor pressure equals the osmotic pressure, osmosis ceases.
If a plant cell is isotonic to its surrounding, there is no movement of water into the cell. The cell becomes flaccid (limp),
and loss of structural support from turgor pressure causes the plant to wilt. The cell wall provides no advantages when a
plant cell is immersed in a hypertonic solution. As the plant cell loses water, its volume shrinks. Eventually, the plasma
membrane pulls away from the cell wall as the cell loses water to a hypertonic environment. This plasmolysis is usually
lethal.

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Importance of turgor pressure for plant cells
 It keeps the cells and their organelles stretched which is essential for proper functioning of a cell.

 It provides support to non-woody tissues

 It is essential for cell enlargement, during growth.

 It keeps the leaves fully expanded, and properly oriented to light.

 The opening and closing of stomata are caused due to turgidity of guard cells

Reverse osmosis
Reverse osmosis is the expulsion of pure water from osmotically active solution, through a semipermeable membrane,
under the influence of pressure higher than the osmotic pressure of the solution. It is used in removing salts from saline
water as well as in greater purification of water.
Importance of osmosis
Plants very much rely on osmosis, and osmosis is responsible for:
(i) entry of soil water into root.
(ii) Cell to cell movement of water.
(iii) Living cells remain distended or turgid only by the osmotic entry of water into them.
(iv) Various cell organelles like mitochondria and chloroplasts will collapse, if they are not able to maintain a proper
osmotic concentration.
(v) 70% of cell water is held in vacuoles. It enters through endosmosis, due to solutes found dissolved in it.
(vi) The soft organs like leaves, flowers, fruits and young stems are able to keep themselves stretched and swollen, due to
turgidity of their cell, which is dependent on osmosis.
(vii) Osmosis plays a key role in the germination of seeds.
(viii) Many plant movements like the folding and drooping of leaves are brought about by osmosis.
(ix) The stomata open and close, only in response to increase or decrease of the osmotic pressure of the guard cells
(x) a high osmotic pressure has been found to protect the plants against drought and frost injury.
(xi) Seeds and spores are similarly able to pass through the unfavorable periods, due to high osmotic pressure.

2. Active transport mechanisms

If a molecule is to be transported from an area of low concentration to an area of high concentration, work must be done
to overcome the influences of diffusion and osmosis. Since in the normal state of a cell, large concentration differences in
K+, Na+ and Ca2+ are maintained, it is evident that active transport mechanisms are at work.
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Active transport is the movement of a substance across a membrane against its concentration gradient and involves the use
of cellular energy to power special movable carrier transmembrane proteins known as pumps or solute pumps, to bring
material into the cell or take them out against their concentration gradients. This is usually to accumulate high
concentrations of molecules that a cell needs, such as ions, glucose or amino acids. The energy for active transport
mechanisms may come from the hydrolysis of ATP, the absorbance of light, the transport of electrons, or coupling with
other processes that are moving particles down their concentration gradients.
Generally, active transport is
ῼ uphill transport, i.e., moves substances against concentration gradient and thus it is energetically uphill and
requires the cell to expend energy. Energy-requiring process during which a transport protein pumps a molecule
across a membrane, against its concentration gradient.
ῼ is faster than passive transport.
ῼ The rate of active transport reaches the maximum, when all the protein pumps are used in transport, which is
called saturation effect. Amino acids, some sugars, and ions are transported by protein carriers called pumps by
utilizing energy. This process is considered to be responsible for the accumulation of an excess amount of solute
within the cells.
ῼ Helps cells maintain steep ionic gradients across the cell membrane (e.g., Na +, K+, Mg++, Ca++and Cl-).

In Active transport (aka solute pumping), the energy of ATP may be used directly ( in which case it is known as Primary
Active transport) or indirectly (in which case it is known as Secondary Active transport).
1. Primary Active transport
2. Secondary Active transport

1. Primary Active transport (direct Active Transport)

Some transporters bind ATP directly and use the energy of its hydrolysis to drive active transport. Such active transport
mechanisms are primary active transports. The proteins involved with active transport are all carrier proteins and also
known as ion pumps. and enable a cell to maintain internal concentrations of ions and small molecules that would
otherwise diffuse across the membrane. ATP energizes the pumps which act as catalysts in the splitting of ATP → ADP +
phosphate, so the pumps are also called ATPases.
Some common Primary Active transporters
1. The Na+/K+ ATPase or The Na+/K+ pump
2. The H+/K+ ATPase or the H+/K+ pump
3. The Ca2+ ATPase or The Ca2+ pump
4. ABC Transporters
Example: The Na+/K+ ATPase or the Na+/K+ pump.
Compared with its surroundings, an animal cell has a much higher concentration of potassium ions and a much lower
concentration of sodium ions. The cytosol of animal cells contains a concentration of potassium ions (K +) as much as 20
times higher than that in the extracellular fluid. Conversely, the extracellular fluid contains a concentration of sodium ions
(Na+) as much as 10 times greater than that within the cell.
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  K+ / inside a cell is 100mM and outside a cell is 5mM
 Na+ inside a cell is 15mM outside a cell is 150mM
The plasma membrane helps maintain these steep gradients by pumping sodium out of the cell and potassium into the cell.
The sodium-potassium pump (also called the Na+/K+-ATPase enzyme) uses the energy from the hydrolysis of ATP to
actively transport 3 Na+ ions out of the cell for each 2 K+ ions pumped into the cell.
These pumps are found in the membrane of virtually every cell, and are essential in transmission of nerve impulses and in
muscular contractions.
The hydrolysis of a single ATP molecule by the Na+/K+-ATPase is required to transport five ions 3Na+ ions out of the
cell and 2K+ ions into the cell. Thus it is an antiport that transports the two ions simultaneously in opposite directions
across the membrane.
Functions of the pump
The crucial roles of the Na+/K+ ATPase are reflected in the fact that almost one-third of all the energy generated by the
mitochondria in animal cells is used just to run this pump.
 The primary function is to maintain constant cell volume by decreasing the intracellular ion concentration (and
thus the osmotic pressure) and increasing the extracellular ion concentration, thus decreasing the flow of water
into the cell. In other words, the accumulation of sodium ions outside of the cell draws water out of the cell and
thus enables it to maintain osmotic balance (otherwise it would swell and burst from the inward diffusion of
water).
 The Na+ -K+ pump also plays a minor role in the maintenance of a potential difference across the plasma
membrane. It pumps 3 Na ions out in exchange for 2 K ions pumped in. It is an electrogenic pump, and thus helps
establish a net charge across the plasma membrane with the interior of the cell being negatively charged with
respect to the exterior. This resting potential prepares nerve and muscle cells for the propagation of nerve
impulses. The gradient of sodium ions is also harnessed to provide the energy to run several types of indirect
pumps.

2. Secondary Active Transport (Indirect Active Transport)

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Other transporters use the energy already stored in the gradient of a directly-pumped ion. Direct active transport of the ion
establishes a concentration gradient. When this is relieved by facilitated diffusion, the energy released can be harnessed to
the pumping of some other ion or molecule. In other words, Indirect Active Transport uses the downhill flow of an ion to
pump some other molecule or ion against its gradient. The driving ion is usually sodium (Na +) with its gradient
established by the Na+/K+ ATPase
Examples:
The Na+/glucose transporter.
This transmembrane protein allows sodium ions and glucose to enter the cell together. The sodium ions flow down their
concentration gradient while the glucose molecules are pumped up theirs. Later the sodium is pumped back out of the cell
by the Na+/K+ ATPase.
The Na+/glucose transporter is used to actively transport glucose out of the intestine and also out of the kidney tubules and
back into the blood.

VESICULAR TRANSPORT/ BULK TRANSPORT

Small solutes and water enter or leave the cell through the lipid bilayer or by transport proteins (simple diffusion,
facilitated diffusion and active transport). Particles and large molecules, such as polysaccharides and proteins, cross the
membrane via packaging in vesicles or vacuoles, generally known as vesicle-mediated transport or bulk transport.
Vesicles and vacuoles that fuse with the cell membrane may be utilized to release or transport chemicals out of the cell or
to allow them to enter a cell. Like active transport, these processes require energy.

Types of vesicular transport include:

a. Exocytosis b. Endocytosis

Vesicles and vacuoles that fuse with the cell membrane may be utilized to release or transport chemicals out of
the cell or to allow them to enter a cell. Exocytosis is the term applied when transport is out of the cell.

a. Exocytosis
Exocytosis describes the process of vesicles fusing with the plasma membrane and releasing their contents to the outside
of the cell. Exocytosis is especially common in secretary cells- in cells that produce substances for export out of the cells.
The transport vesicles usually budded from the ER or Golgi (the two major organelles involved in producing and

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preparing substances for secretion out of the cell), and moved by the cytoskeleton to the plasma membrane. When the two
membranes come in contact, the bilayers fuse and spill the contents to the outside. Many secretory cells use exocytosis to
export products (e.g., insulin in pancreas, or neuro-transmitter from neuron). Pancreatic cells called β-cells secrete insulin
into the blood by exocytosis. Neurons use exocytosis to release neurotransmitters that signal other neurons or muscle
cells. When plant cells are making walls, exocytosis delivers proteins and certain carbohydrates from Golgi vesicles to the
outside of the cell.

Exocytosis:
i. Moves materials out of the cell
ii. Material is carried in a membranous vesicle
iii. Vesicle migrates to plasma membrane
iv. Vesicle combines with plasma membrane
v. Material is emptied to the outside

b. Endocytosis
Endocytosis is a reversal of exocytosis, and during endocytosis, a cell brings in biological molecules and particulate
matter by forming new vesicles from the plasma membrane. A small area of the plasma membrane sinks inward to form a
pocket. As the pocket deepens, pinched off into the cytoplasm forming a vesicle called an endosome containing the
material that had been outside the cell. The substance subsequently enters the cytoplasm enclosed in a vesicle.
Used by cells to incorporate extracellular substances.

There are three types of endocytosis:

(1) phagocytosis (“cellular eating”)
(2) pinocytosis (“cellular drinking”), and
(3) receptor-mediated endocytosis

1. Phagocytosis or ("cell eating")- endocytosis of solid particles

The plasma membrane engulfs solid material or particulate matter (e.g., bacteria engulfed by WBCs) from the ECF. That
is cells internalize particles or other cells by forming a phagocytic vesicle. The endosome is so large that it is called a
phagosome or vacuole. Phagosomes fuse their membrane with the lysosomal membrane, and then deliver their contents to
lysosomes. Once inside the lysosome, the contents of the phagosome are digested by the degradative enzymes of the
lysosome.
Phagocytosis occurs only in certain specialized cells (e.g. WBCs such as neutrophils and macrophages, and in
microorganisms like the amoeba)

2. Pinocytosis or (cell drinking)- endocytosis of fluid droplets

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occurs when the plasma membrane folds inward to form a pinocytic vesicle to take in droplets of extracellular fluid in
small vesicles. The process is not discriminating.
The cells of the kidney tubules reabsorb small proteins by pinocytosis, so that the protein is not lost in urine.

This electron micrograph shows a section of the wall of a

capillary. The small in-pocketings of the plasma membrane
are clearly seen (arrows). Most of these are open to the tissue
space but some can also be seen on the other side of the cell
apparently engulfing fluid from within the capillary. Perhaps
most of the vesicles facing the tissue space are not taking up
material by endocytosis but are instead discharging material
by exocytosis. If so, the pinocytic vesicles formed at one
surface of the cell may, after being detached, move through
the cell to the opposite surface and there discharge their
contents. In this way materials can be moved efficiently through the capillary wall. The pinocytosis vesicles in this image
represent a subtype called caveolae. In addition to their function in endo- and exocytosis, they also can serve as a reservoir
of plasma membrane. When a cell expands or is stretched, the caveolae flatten out providing more plasma membrane.
3. Receptor-mediated endocytosis
Receptor Mediated Endocytosis, as its name implies, it depends on the interaction of that molecule with a specific
binding protein in the cell membrane called a receptor. In receptor-mediated endocytosis, first the substance to be
transported binds to specific receptor proteins- membrane-embedded proteins with specific receptor sites exposed to the
cell's exterior, cluster in regions called coated pits. A molecule that binds to a specific receptor site of another molecule is
called a ligand. When the ligand binds to its specific receptor, the ligand-receptor complex accumulates in the coated pits
A layer of clathrin, a fibrous protein, lines and reinforces the coated pit on the cytoplasmic side and probably helps
deepen the pit to form a vesicle.
Progressive stages of receptor-mediated endocytosis:
Extracellular ligand binds to receptors in a coated pit. -----> Causes inward budding of the coated pit. -------> Forms a
coated vesicle inside a clathrin cage. -------> Ingested material is liberated from the vesicle. -------> Protein receptors can
be recycled to the plasma membrane.
Endosomes are formed by receptor-mediated endocytosis, and more discriminating process than pinocytosis.

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Receptor-mediated endocytosis enables cells to acquire large quantities of specific substances even if they are in low
concentration in extracellular fluid. For example, human cells use this process to take in cholesterol for use in the
synthesis of membranes and as a precursor for the synthesis of steroids.
Cholesterol travels in the blood in low-density lipoproteins (LDL), complexes of protein and lipid. These LDLs bind to
LDL receptors on cell membranes, initiating endocytosis, and entering the cell by endocytosis.
In an inherited disease called familial hypercholesterolemia, the LDL receptors are defective, leading to an accumulation
of LDL and cholesterol in the blood. This condition contributes to early atherosclerosis-deposition of LDL on the artery
walls and narrowing it.
Vesicles not only transport substances between the cell and its surroundings but also provide a mechanism for
rejuvenating or remodeling the plasma membrane. Endocytosis and exocytosis occur continually in most eukaryotic cells,
yet the amount of plasma membrane in a non-growing cell remains fairly constant. Apparently, the addition of membrane
by one process offsets the loss of membrane by the other. That is vesicle fusion with the plasma membrane offsets
membrane loss through endocytosis providing a mechanism to rejuvenate or remodel the plasma membrane.

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