Unit 1 Introduction to Biomembranes

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UNIT 4

MEMBRANE
TRANSPORT-I

Structure
4.1 Introduction 4.4 Cytosis

Expected Learning Outcomes 4.5 Summary

4.2 Simple and Facilitated Diffusion 4.6 Terminal Questions

Simple Diffusion 4.7 Answers

Facilitated Diffusion 4.8 Suggested Readings
Osmosis

4.3 Active and Passive Transport

Active Transport
Passive Transport

The FRAP does not

distinguish the protein
that are immobilized or
can only diffuse over a
limited distance in a
4.1 INTRODUCTION given time. The
alternative technique,
In the previous block, you studied about the structural aspects of biological SPT (Single Particle
Tracking) can overcome
membrane. Now you know in detail about membrane composition and fluid such hurdles. In SPT,
mosaic model. You have also studied some of the characteristic features like individual protein
membrane asymmetry, membrane fluidity, membrane dynamics as well as molecule is labeled with
techniques like freeze fracture and FRAP (Fluorescence Recovery after antibody coated with
gold particles. The
Photobleaching) which are employed to study membrane and its properties. In movement of the labeled
this unit you will learn about one of the functional aspects of biomembrane. molecule is followed by
Apart from providing barrier and protection from the external environment, Computer Enhanced
biomembrane plays a crucial role in transport of substances, ions, molecules Video Microscopy.
and drugs across biological membranes.
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Membrane transport is essential for the life of a cell. At cellular level vast
amount of exchange is necessary to maintain its structure and function. The
transport may involve the addition of biological molecules and the discharge of
waste products. Membrane transport refers to the movement of particles
(solute and ions) across or through the phospholipid bilayer membrane. It
depends upon the permeability of the membrane, transmembrane
concentration difference and the size and charge of the solute. Transport
across membrane may be by passive, facilitated or active mechanisms. Some
of these mechanisms require the expenditure of energy and use of
transmembrane proteins. We will begin the unit with simple phenomenon like
diffusion and go into the details of active and passive transport. We will also
focus on some ATP powered pumps as an example to understand their mode
of functioning.

Expected Learning Outcomes

After studying this unit, you should be able to:
™ describe the process of diffusion;

™ differentiate between simple and facilitated diffusion;

™ describe active and passive transport;

™ differentiate between active and passive transport;

™ describe various types of transport mechanisms; and

™ explain various types of ion channel proteins.

4.2 SIMPLE AND FACILITATED DIFFUSION

Biological membranes are composed of lipids and proteins. The lipid bilayer
serves as a barrier to the movement of hydrophilic (water soluble) molecules
across the membrane. However, it selectively allows the passage of selected
ions and molecules which contribute to the steady state condition required for
The concentration
biological processes. Generally, the proteins embedded in the lipid bilayer
difference of a molecule
ion across the two sides function as transporter or receptor of extracellular stimuli.
of a membrane is
Let us recall the structural features of biological membranes which you have
termed as gradient.
studied in Unit 1. Biological membranes are composed of lipids and proteins.
Movement from a high The major lipid in membranes is phospholipid where two fatty acid chains are
concentration to a low
attached to OH of the 1st and 2nd carbon of glycerol. The OH on the 3rd carbon
concentration is known
as movement with or in is attached to a polar organic alcoholic molecule and phosphate group. These
the direction of side chains make one end as polar and the other end as non-polar in the
concentration phospholipid molecule. In the lipid bilayer, the polar part is outside and non-
gradient or downhill.
polar part faces inside. The non-polar interior prevents the passage of water
Movement from a low
concentration to a high soluble substances through the bilayer. The ability of the lipid bilayer to
concentration is known discriminate between low and high molecular weight hydrophilic molecules,
as movement against permitting the former to diffuse across but not the latter is due to the presence
concentration
of pores in the bilayer. These pores are formed randomly as a result of thermal
gradient or uphill.
movement of acyl phospholipid chains.

Mostly, the globular proteins in the membrane are inserted in the bilayer with
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Unit 4
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Transport-I
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their non-polar portion in contact with lipid and polar portion emerging out from
the membrane surface. These proteins serve as channels for specific ions or
molecules to cross this hydrophobic barrier, since many molecules required
by the cell are polar and cannot enter freely into the cell. These polar
molecules enter the cell through specific channel proteins found in the plasma
membrane. The channels are specific transmembrane proteins having polar
amino acids in the interior area. On the other hand hydrophobic non polar
molecules can enter or leave freely the cell through lipid bilayer. This
spontaneous process is termed diffusion. This process allows the small and
hydrophobic (or lipophilic) molecule to move inside and outside the cell
membrane. Diffusion is movement of substances (atoms, ions, molecules)
from a region of higher concentration to a region of lower concentration. This
results in uniform distribution of the substance. The driving force for diffusion
is concentration gradient. Diffusion obeys Fick’s law, which states that the
rate of movement of a substance is directly proportional to the concentration
gradient. Diffusion can be of three types:

x Simple diffusion
x Facilitated diffusion

x Osmosis (in aqueous medium)

4.2.1 Simple diffusion

Smaller molecules and nonpolar substances move freely through lipid bilayer
from the side of higher concentration to lower concentration. This type of
movement is shown by oxygen, nitrogen, benzene, water, urea, glycerol,
carbon dioxide etc. The rate of diffusion is directly proportional to the
concentration difference, surface area, distance, and temperature and
inversely proportional to the size of molecule. Simple diffusion thus shows a
linear relationship between the concentration of solute and its rate of transport
across the membrane.
All water-soluble molecules or ions that enter or leave the cell are either
transported by carriers or pass through channels. Ions and some other polar
molecules diffuse across the cell membrane through channels. Ion channels
are proteins containing a hydrated interior portion that spans throughout the
bilayer. Ions diffuse through the channel in either direction without meeting the
lipid bilayer. Furthermore, these ions do not interact or bind to the channel
proteins. The diffusion through ion channels is an essential component of
signaling in the nervous system. Some of the molecules and their permeability
across the plasma membrane are given in Table 4.1.

4.2.2 Facilitated diffusion

The diffusion of some ions and other solutes like sugars and amino acids
across the membrane is facilitated by carriers. Carriers bind their substrates
with high stereospecificity and undergo cyclic conformational changes while
channels do not do so. Channels are less specific to substrates but facilitate
faster transport. In this type of transport the molecule binds physically to the
carrier protein on one side of the membrane and is released on the other side
(Fig. 4.1). This movement is from a higher concentration side to the lower
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concentration side. The cell uptakes the required essential molecules like
sugars and prevent the accumulation of unwanted molecules by facilitated
diffusion. However, the rate of movement through channel proteins is much
higher than transport through carrier proteins. Membrane proteins that speed
the movement of a solute across the membrane by facilitating diffusion are
called transporters or permeases.

Table 4.1: Examples of some molecules and their permeability across

the plasma membran

S. Category Molecules Permeability

No.

1. Gases CO2, O2, N2 Permeable

2. Small uncharged polar Ethanol Permeable

molecules Urea, Water Slightly Permeable

3. Large uncharged polar Glucose, Fructose Impermeable

molecules

4. Ions K+, Mg2+,Ca2+, Cl-, Impermeable

HCO3-, HPO42-

5. Charged polar Amino acids, ATP, Impermeable

molecules Glucose-6-phosphate,
Proteins, Nucleic acids

In hypertonic
solutions, there are
more solute molecules
outside the cell which
causes the water
molecules to be sucked
out of the cell and the
cell shrinks.
In hypotonic solutions,
there is less solute
molecules outside the Fig. 4.1: Overview of membrane transport proteins. Passive transport across ion
cell, water will move channels or carriers is driven by a chemical gradient. Active transport by carrier
inside the cell. The cell proteins/ transporters allows a transport against a chemical gradient, thanks to
will grow larger and the consumption of energy, e.g. by ATP hydrolysis.
finally burst.
In isotonic solutions,
4.2.3 Osmosis
there is equal Osmosis is a special example of diffusion. It is diffusion of water molecule
concentration of solute
on both side of the through semipermeable membranes from a dilute solution to a concentrated
membrane; the solution. This process does not require any energy. Absorption of water by
movement rate of water plant roots is an example of osmosis in plant. Plant cells in a hypotonic
will be equal in both the solution gain water by osmosis and the plant tissue becomes stiffer (turgid)
directions.
while in a hypertonic solution plants loose water by osmosis and tissue
58 becomes flaccid (shrunken).
Unit 4
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Let us understand some of the terms which you will come across frequently in
this block.

Channels: Channels are proteins having several helical segments that are
spread back and forth through the membrane, forming a channel. For
example, the bacteriorhodopsin contains seven nonpolar helical segments
that traverse the membrane, forming a circular pore through which protons
pass during the light-driven pumping of protons.
Pores: Some transmembrane proteins have extensive nonpolar regions with
secondary configurations of E-pleated sheets instead of D helices. The E
sheets form a characteristic motif, folding back and forth in a circle so the
sheets come to be arranged like the staves/poles of a barrel. This E- barrel,
open on both the ends, is a common feature of the porin class of proteins that
are found within the outer membrane of some bacteria.

Transmembrane proteins: Cells contain a variety of different

transmembrane proteins, which differ in the way they traverse the bilayer,
depending on their functions. They are anchored into the bilayer by their
nonpolar segments. While anchor proteins may pass through the bilayer only
once, many channels and pores are created by proteins that pass back and
forth through the bilayer repeatedly, creating a circular hole in the bilayer.

Tick [9] mark the correct option:

a) Biological membrane provide barrier to hydrophobic molecules.
[True/False]

b) Glycerol is a three carbon molecule. [True/False]

c) A phospholipid contains three fatty acid chains. [True/False]

d) Active transport does not require energy. [True/False]

e) Active transport is required for concentrating metabolites in a particular

organelle of the cell. [True/False]

Fill in the blanks with appropriate words:

a) Polar molecule enters the cell through specific …………. (channels/

anchors) in the plasma membrane.
b) Osmosis is the phenomenon of diffusion of water molecules through
…………... (permeable/semipermeable) membrane from a dilute solution
to a more …………… (concentrated/diluted) solution.

c) The interior side of plasma membrane is ……………. (positively/

negatively) charged relative to the extracellular side of the membrane.

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4.3 ACTIVE AND PASSIVE TRANSPORT
Transport of substances against their concentration gradient (from lower
concentration to higher concentration) is energetically unfavorable; therefore
energy is required for this process. However, this process is essential for:

i) Maintaining the balance of ions across the membrane

ii) Concentrating metabolites in a particular organ of cellular compartment

iii) Exporting foreign/ toxic substances from the cell

As mentioned in the previous section, carrier proteins and channels are two
major classes of membrane transport proteins. The transport through these
proteins can be further classified into two types depending on the requirement
of energy for the transport process:
i) Active transport

ii) Passive transport

4.3.1 Active Transport

It is further classified into two types: primary active transport (PAT) and
secondary active transport (SAT). Primary active transport uses the energy
stored in ATP, photons and electrochemical gradients directly for the transport
of molecules from low concentration areas to high concentration area across
the cellular membrane. The PAT is facilitated by P-class pump, V-class pump,
F-class pump and ABC transporters. Let us discuss briefly, the main energy
sources driving PAT.
i) ATP hydrolysis: The enzyme catalyzed hydrolysis of ATP forming ADP
induces a conformational change in the transport protein facilitating the
influx or efflux of particles. The enzyme catalyzing this process is known
as ATPase. The most common example of ATP hydrolysis during primary
active transport in cells is Na+/ K+ pump.
ii) Electrochemical gradient energy: An electrochemical gradient has two
components: (1). An electric component resulting from difference in
charge on either side of the membrane and (2). A chemical component
resulting from the difference in concentration of ions across the cellular
membrane. The electrochemical gradient is generated in terms of the
presence of proton (H+) gradient.
iii) Photon energy: The energy stored in a photon, is used to generate a
proton gradient. The stepwise movement of electrons in an electron
transport chain reduces NADPH and subsequently generates proton
gradient.
In secondary active transport the movement of molecules is from lower
concentration to higher concentration utilizing energy. However, in this case the
direct hydrolysis of ATP does not occur. Instead, the electrochemical gradient
generated from pumping of ions out of the cell is used. Secondary active
transport is further classified as uniport, symport or antiport (Fig. 4.2).

60 i) Uniport: Uniporters transport a single type of molecules/ions down its

Unit 4
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concentration gradient via facilitated diffusion. For example, glucose and
amino acids cross the plasma membrane in most mammalian cells with
the help of uniporters.

ii) Symport: When the carrier protein transports two solutes in the same
direction, it is called symport. It uses a downhill movement of one
molecule/ion to transport other molecule against its concentration
gradient. Symporters move both the molecules or ions in the same
direction through a transmembrane transport protein. They usually have
opposite charge.

iii) Antiport: When an ion traverses in one direction and simultaneously

another metabolite is transported in the other direction, then it is called
antiport. It moves two or more different molecules/ions across the cellular
membrane in opposite directions. Secondary active transporter antiport
protein moves one molecule/ion down its concentration gradient and uses
the energy produced from this process for the movement of another
molecule/ion up its concentration gradient. This is similar to symport
mechanism. The only difference being instead of moving in same
direction molecules move in opposite direction.

Fig. 4.2: Overview of transporters.

Now, let us focus our attention on a major class of proteins, the ATP-powered
pumps, that use the energy released by hydrolysis of the terminal
phosphoanhydride bond of ATP to transport ions and various small molecules
across membranes against their concentration gradients. Although these
proteins are commonly called ATPases, they normally do not hydrolyze
ATP into ADP and Pi unless ions or other molecules are simultaneously
transported. For example Na+- K+ pump (also known as Na+- K+ ATPase),
Ca++- ATPase, H+- K+ATPase etc. These ATP driven ion pumps and transporters
utilize ATP for uphill transport whereas in case of coupled transport, the energy
stored in ion gradients is used. Coupled transporters are also known as
secondary active transporters, such as symporter & antiporters.

ATP-Powered Pumps
All ATP-powered pumps are transmembrane proteins with one or more
binding sites for ATP located on subunits or segments of the protein that face
the cytosol. These pumps are grouped into four classes. Three of the classes
P-type, F-type, and V-type only transport ions, whereas members of the ABC
family primarily transport small molecules such as amino acids and sugars.
Here you must note that all of these ATP powered pumps are involved in
primary active transport as mentioned in the beginning of this section. 61
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Na+-K+ ATPase in the plasma membrane of animal cells and Ca2+ ATPase
pumps are examples of P-type of ATP powered pumps. During transport, at
least one of the subunits is phosphorylated (hence the name “P” class), and
the transported ions move through the phosphorylated subunit of the pump. All
known V and F pumps transport only protons. The H+ pump that generates
and maintains the plasma membrane electric potential in plant, fungal, and
bacterial cells, also belong to V-type. F-class pumps are found in bacterial
plasma membranes and in mitochondria and chloroplasts. In contrast to V-
class pumps, F-class pump generally function as a kind of reverse proton
pump. The energy released by the movement of protons form the exoplasmic
to the cytosolic face of the membrane down the proton electrochemical
gradient is used to power the synthesis of ATP from ADP and P i. Because of
their importance in ATP synthesis in chloroplasts and mitochondria, F-class
proton pumps are commonly called ATP synthases. ABC (ATP-binding
cassette) is a large family of multiple members that are more diverse in
function, for example multidrug-resistance proteins. You will study about ABC
family of transporters in detail in next unit.

Have you ever wondered why do we need so many diverse transporters? Just
think about the ionic composition.

The ion composition inside the cell is different from that in the extracellular fluid,
and this difference has to be maintained for cell survival. For example the
gradients of sodium (Na+), calcium (Ca2+), potassium (K+) and chloride
(Cl-), must be maintained within proper limit to ensure normal functioning of the
cell and maintenance of the membrane potential. In fact, all plasma membranes
have electric potentials (transmembrane potential) across them with inside of
the cell being more negative compared to the outside. This is due to active
transport of ions particularly H+ ions out of the cell. This potential difference
allows for the entry of positively charged ions into the cell but opposes the entry
of negatively charged ions. For example, the sodium ion (Na+) has higher
concentration outside of the cell than inside. This is maintained by a constant
active transport of Na+ out of the cell, which is also helpful in maintaining the
osmotic pressure on both the sides of the membrane. Ca2+ and Cl-
concentrations are also higher outside the cell than cell interior. On the other
hand, K+ is higher inside than the extracellular concentrations. This difference in
the concentration gradient leads to a change in electrochemical gradient across
the cell membrane. Table 4.2 gives the concentration of some of the commonly
occurring ions across the biological membrane.
Table 4.2: Concentration of some important physiological ions across
cell membrane.
S. No. Ion Concentration in Concentration in
cytosol (mM) extracellular space (mM)

1 Na+ 15 145

2 K+ 120 4.5

3 Cl- 20 116

62 4 Ca2+ 10-4 1.2

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The combined effect of electrical and concentration gradients are termed as
electrochemical gradient. In case of K+ and Cl-, the electrical and chemical
gradients of molecules work against each other. K+ is attracted towards the
inside of the cell because of the interior negative membrane potential, but its
concentration gradient works in opposite direction. Thus K+ reaches almost to
equilibrium across the plasma membrane and tries to move out of the cell. Cl -
also tends to move outside of the cell because of the negative interior
membrane potential although its concentration outside is much higher. In case
of Na+ and Ca2+, both tend to come inside the cell down the concentration
gradient and the negative membrane interior pulls them inside the cell. So, let
us try to understand how these pumps work in order to maintain the required
gradient.

Sodium- Potassium Pump

More than one third of all of the energy exhausted by a non-dividing animal cell
is utilized for the active transport of sodium (Na+) and potassium (K+) ions.
Most animal cells have a low internal concentration of Na+, relative to their
surroundings, and a high internal concentration of K+. They maintain these
concentration differences by actively pumping Na+ out of the cell and K+ in by
the sodium-potassium pump. The cell obtains the energy it needs to operate
the pump from hydrolysis of adenosine triphosphate (ATP).

The sodium-potassium pump is an active transport process, transporting Na +

and K+ from areas of low concentration to the areas of high concentration. The
sodium-potassium pump works through a series of conformational changes in
the trans-membrane protein. The main steps of the process are as follows:
1. Three sodium ions bind to the cytoplasmic side of the pump leading to
change in its conformation.
2. In this new conformation, the protein binds a molecule of ATP and cleaves
it into adenosine diphosphate and phosphate (ADP + Pi). ADP is
released; but the phosphate group remains bound to the protein. The
protein is now in the phosphorylated state.

3. The phosphorylation of the protein induces a second conformational

change in the protein. This change translocates three Na+ across the
membrane, so they now face the exterior. In this new conformation, the
protein has a low affinity for Na+, and the bound Na+ dissociate from the
protein and diffuse into the extracellular fluid.
4. However, the new conformation has a high affinity for K+, two of which
bind to the extracellular side of the protein as soon as the Na+ is released.
5. The binding of the K+ causes another conformational change in the
protein, and the bound phosphate group is released.

6. De-phosphorylated protein reverts back to its original conformation,

exposing the two K+ to the cytoplasm and releasing them into the interior
of the cell. The original conformation has a high affinity for Na+; when
these ions bind, they initiate another cycle.
Three Na+ ions leave the cell and two K+ ions enter inside every cycle. The
changes in protein conformation that occur during the cycle are rapid, 63
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enabling each carrier to operate about 100 cycles and transporting as many
as 300 Na+ per second.

Proton Pump

The proton pump pumps protons (H+ ions) across a membrane using energy
derived from energy-rich molecules or from photosynthesis, for example
NADH dehydrogenase, cytochromes c oxidase etc. This generates a proton
gradient across the membrane. Generally, membranes are impermeable to
protons; therefore protons can diffuse back down their concentration gradient
by another co-transport protein like ATP synthase. You will learn about the
functions of these transport proteins in next block. The movement of protons
through their co-transport protein is coupled to the production of ATP, the
energy-storing molecule.

4.3.2 Passive Transport

Through the channels, the substances simply diffuse down the concentration
gradient and usually do not require energy. This is called as passive transport.
Small hydrophobic molecules are able to cross membrane by simple diffusion.

Na+- Glucose Symporter

Many amino acids and sugars are accumulated in animal cells against their
concentration gradient i.e. they are transported into the cell from the extracellular
fluid, even though their concentrations are higher inside the cell. The transport of
these molecules is coupled with sodium ions to enter the cell down the Na+
concentration gradient established by the sodium-potassium pump. Thus, Na+
and a specific sugar or amino acid simultaneously binds to the same
transmembrane protein, on the outside of the cell and is an example of symport.
Both are then translocated to the inside of the cell, but in the process Na+ moves
down its concentration gradient while the sugar or amino acid moves against its
concentration gradient. The cell uses some of the energy stored in the Na+
concentration gradient, to accumulate sugars and amino acids.

The Na+ - driven glucose pump is an example of symporter. It is found on the

apical side of intestinal epithelial cells. After a meal when the concentration of
sugar is higher inside the cell than outside, this pump transports glucose into
gut epithelial cells. This co-transporter is dependent on Na+ gradient for its
energy which flows into the cell down its gradient to allow the transport of
glucose into the cell.

Na+-Ca2+ Antiporter

In a related however different process, called counter-transport, the inward

movement of Na+ is coupled with the outward movement of another
substance, such as Ca++ or H+. Similar to co-transport, both Na+ and the other
substance bind to the same transport protein, in this case called an antiport,
but during antiport they bind on opposite sides of the membrane and are
moved in opposite directions. In counter-transport, the cell uses the energy
released from Na+ movement down its concentration gradient into the cell to
extrude a substance up its concentration gradient.
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The Na+/Ca2+ exchanger also uses the electrochemical gradient of Na+ to allow
the transport of Ca2+. In this case, Na+ flows into the cell down its
concentration gradient, which allows the export of Ca2+ from the cell against its
gradient. Three Na+ molecules enter the cell for exit of each Ca2+.

Anion Transporter
Bicarbonate (HCO3-) transport mechanisms are the principal regulators of pH
in animal cells. Such transport also plays a vital role in acid-base maintenance
in the stomach, pancreas, intestine, kidney, reproductive organs and the
central nervous system. For example, exchange of HCO 3- for Cl- in a
reversible, electroneutral manner or Na+/HCO3- co-transport where proteins
mediate the coupled movement of Na+ and HCO3- across plasma
membranes, often in an electrogenic manner.

Glucose Transporter

Several glucose transporters mediate the thermodynamically downhill

movement of glucose across the plasma membranes of animal cells. These
transporters are named as GLUT1 to GLUT5 and have distinct roles to
perform which is given briefly in Table 4.3. These transport proteins are made
up of about 500 residues long amino acids and possess a common 12
transmembrane helix structure.
Table 4.3: Glucose transporters and their functions
Name Location Function

GLUT1 All mammalian tissues Basal glucose uptake

GLUT2 Liver and pancreatic cells In liver, GLUT2 removes excess

glucose from the blood while in
pancreas, it adjusts the rate of
insulin secretion which accordingly
signals the need for removal of
glucose from the blood for storage
as glycogen or conversion into fat

GLUT3 All mammalian tissues Basal glucose uptake

GLUT4 Muscle and fat cells Amount in muscle plasma Mostly, the inner
membrane increases with membranes in a double
membrane system are
endurance training
intrinsically
impermeable to nearly
GLUT5 Small intestine Primarily a fructose transporter
all ions and polar
Porins molecules. For
instance, a large
Porins are proteins found in the outer membranes of many Gram-negative number of transporters
bacteria. They function to form a water-filled pore through the membrane, from are required to shuttle
metabolites such as
the exterior to the periplasm, which is a region located between the outer and ATP, pyruvate, and
inner membranes. The porin channel allows the diffusion of small hydrophilic citrate across the inner
molecules. mitochondrial
membrane.
Similarly, mitochondria also have two membrane systems, an outer and inner
membrane. The outer membrane is quite permeable to most small molecules 65
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and ions because of the presence of mitochondrial porin, a 30-35 kd pore
forming protein also known as voltage-dependent anion channel (VDAC). This
regulates the transport of usually anionic metabolites such as phosphate,
chloride, organic anions, and the adenine nucleotides across the outer
membrane. VDAC appears to form an open E-barrel structure similar to that of
the bacterial porins. The outer membrane of bacteria, like that of mitochondria, is
permeable to most small metabolites because of the presence of porins.
Beta-based membrane
potein is found in the
outer membranes of
4.4 CYTOSIS
bacteria, mitochondria Mechanism for transport of large quantities of molecules in and out of the cell
and chloroplasts. They
resemble barrels used to
is termed as cytosis. It is broadly divided into two types: endocytosis and
contain liquids. These E- exocytosis.
barrel proteins serve
essential functions in Endocytosis
cargo transport,
Most of the substances used as fuel molecules are polar and cannot move
signaling and re also
vital for membrane freely inside the lipid bilayer. The process used by single celled eukaryotes to
biogenesis. internalize these substances is known as endocytosis, in which the plasma
membrane extends outward and envelops the food particles. Endocytosis is
further divided into three types:
i) Phagocytosis
ii) Pinocytosis
iii) Receptor-mediated endocytosis

Phagocytosis is the process by which a cell takes in/ingests or engulfs large

particles, microorganisms, dead cells or some fragment of big organic
molecule. For example, amoeba surrounds the target objects with pseudopods
for phagocytosis while white blood cells and macrophages engulf and kill
foreign invading pathogens.
Pinocytosis is common among eukaryotes. It is a process by which cell ingests
the fluid by the inward folding of the plasma membrane and the formation of
membrane-bound, fluid-filled vesicles (pinocytic/pinocytotic vesicles) either
clatherin coated or caveolae derived. This is also known as cell-drinking. Look at
(Fig. 4.3) and note the difference between the two processes.

(a) (b)

66 Fig. 4.3: Diagrammatic representation of phagocytosis and pinocytosis.

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Receptor-mediated endocytosis is a specialized form of endocytosis
involving specific receptor molecules. The molecule to be transported binds to
the receptors on the plasma membrane. The specificity of this type of transport
is provided by the structure of receptor in which only specific molecules can fit
into. The interior portion of the receptor molecules resembles a hook that is
trapped in an indented pit coated with the protein clathrin. The binding of
molecules to the receptor initiates the endocytosis process (Fig. 4.4). The
process is highly specific and very fast. Low density lipoprotein (LDL) is taken
up by receptor mediated endocytosis. The LDL molecules bring cholesterol into
the cell which is required for synthesis of membranes.

Fig. 4.4: Diagrammatic representation of receptor-mediated endocytosis.

Exocytosis

The discharge of larger material from vesicles at the cell surface is termed as
exocytosis. In plant cells, exocytosis helps in exporting materials required for
cell wall synthesis through the plasma membrane. In animal cells, exocytosis
provides a mechanism for secreting many hormones, neurotransmitters,
digestive enzymes and other substances.

Fill in the blanks:

a) When a carrier protein transport two solute in same direction, it is called

…………. .

b) The Na+ driven glucose pump is an example of ………….. .

c) The discharge of large material from vesicles at the cell surface is termed
as ………….. .
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Tick [9] mark the correct option:

a) One molecule of Na+ enters for the exit of each molecule of Ca2+.
[True/False]
b) Water is secreted out by the process of pinocytosis. [True/False]

c) Low density lipoproteins (LDL) is taken up by receptor mediated

endocytosis. [True/False]
d) Na+/ Ca2+exchanger is an example of antiport. [True/False]

4.5 SUMMARY
Let us recapitulate what we have learnt so far:

x The biological membranes allow the passage of selected ions and

molecules which contribute to the steady state condition required for
biological processes.
x Diffusion is the movement of substances (atoms, ions, molecules) from a
region of higher concentration to a region of lower concentration.

x Osmosis is diffusion of water molecule through semipermeable

membranes from a more dilute solution to a more concentrated solution.

x Transport of substances against its concentration gradient (from lower

concentration to higher concentration) is energetically unfavorable,
therefore requires energy and is known as active transport.

x When the carrier protein transports two solutes in the same direction, it is
called symport. When ion traverses in one direction and simultaneously
another metabolite is transported in the other direction, then it is called
antiport.

x Endocytosis is the process to internalize the substances, in which the

plasma membrane extends outward and envelops the food particles.

x The discharge of larger material from vesicles at the cell surface is termed
as exocytosis.

4.6 TERMINAL QUESTIONS

1. What is passive transport?
2. Explain the process of simple diffusion.

3. How facilitated diffusion is different from active transport?

4. Describe various types of ATP powered pumps?

5. Write a note on Na+-K+ pump.

6. Differentiate between phagocytosis and pinocytosis.

7. What is endocytosis? Explain.

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Unit 4
1 Introduction
Membrane
to Biomembranes
Transport-I
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4.7 ANSWERS
Self-Assessment Questions
1. a) false b) true c) false d) false e) true

2. a) channels b) semi permeable, concentrated c) negatively

3. a) symport b) symporter c) exocytosis

4. a) false b) false c) true d) false

Terminal Questions
1. Passive transport is the simplest method of transport and is dependent
upon the concentration gradient and size and charge of the solute. In
passive transport, small uncharged particles move across the membrane
until the concentration becomes same on both the sides of the
membrane. Passive transport is independent of membrane proteins and
does not require expenditure of energy.
2. In simple diffusion, small uncharged molecules of non-polar hydrophobic
molecules pass through the lipid bilayer to leave or enter the cell, moving
from areas of higher concentration towards areas of low concentration.
For example, oxygen, carbon dioxide and most lipids enter and leave cells
by simple diffusion.

3. Facilitated diffusion is a form of passive transport mediated by transport

proteins situated within the plasma membrane. Facilitated diffusion allows
the movement of hydrophilic molecule through the lipid bilayer. The
movement is from high concentration towards lower concentration to
achieve the equilibrium. Facilitated diffusion utilizes channel proteins to
facilitate solute movement while active transport is the movement of
particles through a protein from low concentration area to a high
concentration area utilizing the energy of ATP hydrolysis. ATP hydrolysis
induces a conformational change in the transport protein which facilitates
the mechanical movement of the molecule. Active transport systems are
energy coupled devices as chemical and mechanical processes are
involved in the movement of the molecules and require energy.

4. ATP-powered pumps are transmembrane proteins with one or more

binding sites for ATP located on subunits or segments of the protein that
face the cytosol. These pumps are grouped into four classes. Three of
the classes P-type, F-type, and V-type only transport ions, whereas
members of the ABC family primarily transport small molecules such as
amino acids and sugars. Na+/K+ ATPase in the plasma membrane of
animal cells and Ca2+ ATPase pump are examples of P-type of ATP
powered pumps. During transport, at least one of the subunits is
phosphorylated (hence the name “P” class), and the transported ions
move through the phsophorylated subunit of the pump. All known V and F
pumps transport only protons. The H+ pump that generates and maintains
the plasma membrane electric potential in plant, fungal, and bacterial
cells also belongs to V-type. F-class pumps are found in bacterial plasma
membranes and in mitochondria and chloroplasts. F-type pump, use the 69
Block 2
1 Membrane
Biomembranes
Transport
....................................................................................................................................................................................................................................
energy released by the movement of protons to power the synthesis of
ATP from ADP and Pi (Refer section 4.3.1).

5. One of the most important pumps in animals cells is the sodium-

potassium pump (Na+-K+ ATPase), which maintains the electrochemical
gradient (and the correct concentrations of Na+ and K+) in living cells. The
sodium-potassium pump moves two K+ into the cell while moving three
Na+ out of the cell (Refer section 4.3.1).
6. Phagocytosis is the process by which a cell ingests or engulfs another
organism or some fragment of big organic molecule while pinocytosis is a
process by which cell ingests the fluid by the inward folding of the plasma
membrane and the formation of membrane-bound, fluid-filled vesicles.
This is also known as cell-drinking.

7. Endocytosis is the process in which large molecules enter into a cell. In

this a small piece of cell membrane covers the particles and in entrapped
and brought into the cell. If the particle is solid, it is called phagocytosis
and if it is liquid droplet, the process is termed as pinocytosis. Sometimes
specific receptors are involved for the process of endocytosis, then the
process is known as receptor mediated endocytosis. For more details
refer section 4.4.

4.8 SUGGESTED READINGS

x Garret, R.H., Grisham, C.M. (2016). Biochemistry (6th ed.). Boston,
Cengage Learning. ISBN-10: 1133106293, ISBN-13: 978-1133106296

x Berg, J.M., Tymoczko, J.L. and Stryer L., (2012) W.H. Biochemistry (7th
ed.), Freeman and Company (New York), ISBN:10: 1-4292-2936-5,
ISBN:13:978-1-4292-2936-4.
x Nelson, D.L., Cox, M.M. (2017). Lehninger: Principles of Biochemistry (7th
ed.). New York, WH: Freeman and Company. ISBN: 13: 978-1-4641-2611-
6 / ISBN:10:1-6412611-9.
x Lodish, H., Berk, A., Kaiser, C.A., Krieger, M., Bretscher, A., Ploegh, H.,
Amon, A., Scott, M.P. (2016). Molecular Cell Biology (8th ed.). New York,
WH: Freeman & Company. ISBN-13: 978-1-4641-0981-2.
Voet, D.J., Voet, J.G., Pratt, C.W. (2008). Principles of Biochemistry (3rd
ed.). New York, John Wiley & Sons, Inc. ISBN:13: 978-0470-

70