11 - Membranes

BIOLOGICAL
MEMBRANES
AND
TRANSPORT
Chapter 11
MEMBRANES
Biological membranes allow life as we know it to exist.
They form cells and enable SEPARATION between the inside
and outside of an organism, controlling by means of their
SELECTIVE PERMEABILITY which substances enter and
leave.
INSIDE OUTSIDE
extracellular
cytoplasm
matrix
MEMBRANES
These BARRIERS prevent molecules generated inside the
cell from leaking out and unwanted molecules from
diffusing in.
TRANSPORT BARRIER
Dynamic
Steady
ComposiFon
Yet, they also contain TRANSPORT SYSTEMS that allow

SPECIFIC MOLECULES to be taken up and unwanted
compounds to be removed from the cell.
MEMBRANES
The main components of membranes are LIPIDS and
PROTEINS.
Membranes also contain CARBOHYDRATES that are linked

to lipids and proteins.
MEMBRANES
Lipids and proteins exist in the membrane as
separate but loosely-‐aKached molecules.
The membrane is not like a balloon that can expand and
contract; rather, it is fairly rigid and can burst if penetrated
or if a cell takes in too much water.
However, because of its mosaic nature,

a very fine needle can easily penetrate a
plasma membrane without causing it to
burst; the membrane will self-‐seal when
the needle is extracted.
MEMBRANES
The lipid components of the membrane form the permeability barrier,
LIPIDS PERMEABILITY BARRIER
and protein components act as a transport system of pumps and
channels that give the membrane selecEve permeability.
PROTEINS TRANSPORT SYSTEMS

MEMBRANES
1) Membrane lipids have both hydrophilic and hydrophobic
moieEes.
These lipids SPONTANEOUSLY form closed bimolecular
sheets in aqueous media.
Lipid
2) Membranes are very
bilayer
thin structures, only
two molecules thick,
(6-‐10 nm) that form
closed boundaries
between different
compartments.
MEMBRANES
3) Membranes are noncovalent assemblies.
The consEtuent protein and lipid molecules are held
together by many noncovalent interacEons.
protein
protein
phospholipids
MEMBRANES
4) Membranes consist mainly of LIPIDS and PROTEINS.
Membranes also contain CARBOHYDRATES that are linked

to lipids and proteins.
MEMBRANES
5) The lipid bilayers are barriers to the flow of polar
molecules.
MEMBRANES
6) Specific proteins mediate disFncFve funcFons of
membranes.
Transporters carry a molecule from one side of the plasma membrane
to the other. Receptors can bind an extracellular molecule, and this
acEvates an intracellular process. Enzymes in the membrane can do
the same thing they do in the cytoplasm of a cell: transform a
molecule into another form. Anchor proteins can physically link
intracellular structures with extracellular structures.
MEMBRANES
7) Membranes are asymmetric.
The two faces of biological membranes always differ from

each other.
MEMBRANES
8) Membranes are FLUID structures.
Lipid molecules diffuse rapidly in the plane of the

membrane, as do proteins, unless they are anchored by
specific interacEons.
MEMBRANES
9) Most cell membranes are electrically polarized, such
that the inside is negaEve.
+
-‐
Membrane potenEal plays a key role in transport, energy

conversion, and excitability.
CHEMICAL COMPOSITION OF MEMBRANES
Different membrane have different chemical composiEon.

The funcEon of a membrane depends on its chemical composiEon.
CHEMICAL
FUNCTION
COMPOSITION
So before describing membrane structure and funcEon we need to

review the molecular components of membranes.
CHEMICAL COMPOSITION
OF MEMBRANES
CHEMICAL COMPOSITION OF MEMBRANES
Molecular COMPONENTS of membranes:
1) lipids
2) proteins
3) carbohydrates (present as part of glycoproteins and
glycolipids)
LIPIDS
IN
MEMBRANES
MEMBRANES LIPIDS
The three common types of membrane LIPIDS are:
glycerophospholipids
1) phospholipids
phosphosfingolipids
2) Glycolipids (sphingolipids)
3) cholesterol
Each kingdom, each species, each Essue or cell type, and
the organelles of each cell type have a CHARACTERISTIC SET
OF MEMBRANE LIPIDS.
MEMBRANES LIPIDS
The MAJOR COMPONENT of membrane is PHOSPHOLIPIDS.
MEMBRANES LIPIDS
Their polar head groups favor contact with water, whereas
their hydrocarbon tails interact with one another, away from
water.
H2O H2O H2O H2O

H2O
H2O
PHOSPHOLIPID BILAYER
MEMBRANES LIPIDS
The formaEon of lipid bilayers is a
self-‐assembly process
In other words, the structure of a bimolecular sheet is

inherent in the structure of the consEtuent lipid molecules.
The growth of lipid bilayers from phospholipids is a rapid

and spontaneous process in water.
Hydrophobic interacFons
are the major driving force
for the formaFon of lipid bilayers
MEMBRANES LIPIDS
Dispersion of lipids in H2O Cluster of lipid molecules
Only lipid porEons at the edge of

the cluster force the ordering of
Each lipid molecule forces water. Fewer H2O molecules
surrounding H2O molecules to are ordered and entropy
become highly ordered. increases.
MEMBRANES LIPIDS
1) Lipid clustering reduces the amount of hydrophobic
surface exposed to water and thus minimizes the number of
molecules in the shell of ordered water at the lipid-‐water
interface resulEng in an increase in entropy.
Hydrophobic interacFons among lipid molecules provide
the thermodynamic driving force for the formaEon and
maintenance of these clusters.
2) Furthermore, van der Waals a[racEve forces between

the hydrocarbon tails favor close packing of the tails.
3) Finally, there are electrostaFc and hydrogen-‐bonding
aKracFons between the polar head groups and water
molecules.
MEMBRANES LIPIDS
These hydrophobic interacEons have three significant
biological consequences:
•  Lipid bilayers have an inherent tendency to be extensive
•  Lipid bilayers will tend to close on themselves so that
there are no edges with exposed hydrocarbon chains, and
so they form compartments.
•  Lipid bilayers are self-‐sealing because a hole in a bilayer is

energeEcally unfavorable.
MEMBRANE GLYCOLIPIDS
A glycolipid is a lipid that has an a[ached carbohydrate.
The sugar part may vary from one saccharide unit to several
saccharide units.
The lipid porEon of a glycolipid is a sphingolipid.
GLYCOLIPID
The carbohydrates
appear on the
EXTERIOR
SURFACE
of the cell membrane
for all eukaryoEc cells.
PROTEINS
IN MEMBRANES
MEMBRANE PROTEINS
We now turn to membrane proteins, which are responsible
for most of the DYNAMIC PROCESSES carried out by
membranes.
Membranes differ in their protein content.
Nerve fibers 18%
(Myelin, insulaEon) proteins
Most cells 50% proteins

(Pumps, channels,
receptors, enzymes)
Mitochondria 75%
(energy transducEon) proteins
MEMBRANE PROTEINS
Membrane proteins may be divided operaEonally into two
groups:
1) Integral proteins (intrinsic proteins)
2) Periferal proteins (extrinsic proteins)

INTEGRAL MEMBRANE PROTEINS
Are very FIRMLY ASSOCIATED with the membrane, they

oên interact EXTENSIVELY with the hydrocarbon chains of
membrane lipids, and are removable only by agents that
interfere with hydrophobic interacEons, such as detergents
or organic solvents.
Most integral membrane proteins SPAN the lipid bilayer.


The polypepEde chain of some

integral membrane proteins
does not enter the bilayer but
rather is anchored in one leaflet
by a covalently aKached
hydrocarbon chain.
PERIFERAL MEMBRANE PROTEINS
2) Periferal proteins (extrinsic proteins)
Peripheral membrane proteins do not interact with the
hydrophobic core of the phospholipid bilayer.
They associate with the membrane through electrostaFc
interacFons and hydrogen bonding with the hydrophilic
domains of integral proteins and with the polar head
groups of membrane lipids.
These polar
interacEons can be
disrupted by adding
salts or by changing
the pH.
MEMBRANE PROTEINS
peripheral
integral integral
(transmembrane)
integral peripheral
transmembrane
In ALL the integral transmembrane proteins examined to
date, the membrane-‐spanning domains are
α helices
or
mulFple β strands
In a transmembrane protein with α helices, the α helices
are located in the hydrophobic of the bilayer.
Most of the amino acids in

these membrane-‐spanning
α helices are nonpolar and
only a very few are charged.
The porEon that projects out of the bilayer tends to have a
large amount of hydrophilic amino acids.
In a transmembrane protein with beta sheets, the β sheet
curls up to form a hollow cylinder.
Beta barrel structures are common in

porins, proteins that funcEon as
transporters for ions and small polar
molecules.
The outside surface of a beta barrel is
nonpolar, given that it interacts with the
hydrocarbon core of the membrane.
In contrast, the inside of of a beta barrel is

quite hydrophilic and is filled with water.
MEMBRANE GLYCOPROTEINS
Glycoproteins are proteins to which oligosaccharides
(usually two to ten sugar residues in length) are covalently
a[ached.
The glycoprotein
carbohydrate chains are
oên branched instead of
linear
Glycoproteins of the plasma membrane are always situated

with their sugar residues on the OUTER SURFACE of the
cell.
Membrane glycoproteins play important roles in cell recogniEon and
cell-‐cell interacEon and anEgenicity (anEbodies, immunoglobulins,
are glycoproteins).
MEMBRANE DYNAMICS
MEMBRANE DYNAMICS
One remarkable feature of all biological membranes is their
FLEXIBILITY
their ability to change shape without losing their integrity

and becoming leaky.
The basis for flexibility is the
NONCOVALENT INTERACTIONS
among lipids in the bilayer
Individual lipids are able to move because they are not
covalently anchored to one another.
MEMBRANE DYNAMICS
Individual phospholipid and proteins (and cholesterol) have
some freedom of moEon.
The more movement

we have among the lipids and proteins,
the more fluid and less rigid the membrane is.
The less movement,

we have among the lipids and proteins,
the less fluid and more rigid the membrane is.
If the fluidity is determined by the relaEve movement, what

determines the relaEve movement itself?
MEMBRANE DYNAMICS
The relaEve movement of the molecules in the membrane
is determined by
strength of the aKracFon

between the lipid molecules
strength of non covalent intermolecular forces that exist

between the phospholipids in the membrane.
MEMBRANE DYNAMICS
WEAK STRONG
ATTRACTIONS ATTRACTIONS
MORE LESS
MOVEMENT MOVEMENT
MORE FLUID MORE RIGID

MEMBRANE DYNAMICS
Let’s imagine what happens to a membrane when we go from a low
temperature to a high temperature (the temperature of the
environment in which the membrane is, increases).
What happens?
low temperature high temperature
Let’s imagine that we start with a relaEvely rigid membrane and we
start increasing the temperature.
We are transferring kineEc energy to the membrane lipids which start
to move with greater velocity.
Because they start to move with greater velocity, the intermolecular
bonds that are holding that rigid, well defined, ordered structure,
cannot maintain that rigidity any longer.
MEMBRANE DYNAMICS
As the
temperature
Tm
Fluidity
increases, there is is the
a sharp transiEon temperature
from the “solid” at which this
state to the TRANSITION
“liquid” state. takes place.
At a specific temperature called Tm, the melEng

temperature, a phase transiEon takes place.
The collapse of that well defined and ordered structure
leads into the fluid state.
MEMBRANE DYNAMICS
Membrane with Membrane with

STRONG WEAK intermolecular
intermolecular interacFons
interacFons
HIGH Tm LOW Tm

MEMBRANE DYNAMICS
We know that membrane fluidity is determined by the relaEve
movement of the lipids, and the relaEve movement is determined by
the strength of the intermolecular interacEons between lipid
molecules.
But what determines the strength of the intermolecular

interacEons within the membrane?
(1) Length of the faKy acids
(2) Degree of unsaturaFon of the faKy acids
(3) Cholesterol
MEMBRANE DYNAMICS
(1) Length of the faKy acids
Long faKy acids Short faKy acids
More hydrophobic Less hydrophobic

interacFons between interacFons between
hydrocarbon chains hydrocarbon chains
LOW FLUIDITY HIGH FLUIDIY
HIGH Tm LOW Tm

MEMBRANE DYNAMICS
(2) Degree of unsaturaFon
Saturated faKy acids Unsaturated faKy acids
More hydrophobic Less hydrophobic

interacFons between interacFons between
hydrocarbon chains hydrocarbon chains
LOW FLUIDITY HIGH FLUIDIY
HIGH Tm LOW Tm

MEMBRANE DYNAMICS
Trans fats, which result from the parEal hydrogenaEon of some
unsaturated fa[y acids, have been banned from certain stores and
ciEes because of their health risks.
Part of the health concern is due to their ability to lower membrane
fluidity, in addiEon to the tendency of trans fats to accumulate and
form plaques in blood vessels.
Foods with trans fats:
Cakes, pies and cookies

(especially with frosEng),
biscuits, margarine, crackers,
microwave popcorn, candies,
doughnuts, fried fast foods, Manufacturers can list 0
frozen pizza. grams if the trans fat
content is under 0.5 grams.
MEMBRANE DYNAMICS
To control the fluidity of their
membrane, bacteria synthesize
more unsaturated fa[y acids and
fewer saturated ones when
cultured at low temperatures than
when cultured at higher
temperatures.
In ANIMALS, the factor

that keeps the
membrane fluid is
cholesterol.
MEMBRANE DYNAMICS
(3) Cholesterol
Cholesterol interacts with phospholipids by orienEng its
polar hydroxyl head group close to the polar lipid head
group.
The hydrophobic four ring structure lies alongside the

phospholipids in the membrane.
MEMBRANE DYNAMICS
(3) Cholesterol
Cholesterol tends to dampen the effects of temperature on the
membrane. Thus, cholesterol funcFons as a buffer,
prevenEng lower temperatures from inhibiEng fluidity and prevenEng
higher temperatures from increasing fluidity too much.
•  At T below Tm, cholesterol inserEon

prevents the phospholipid tails to pack too
Eghtly, so it increases fluidity.
•  At T above Tm, rigid ring system of sterol

reduces freedom of neighboring tails to
rotate about their C-‐C bonds, so reduces
fluidity in core of bilayer.
•  Result: thermal transiEon becomes broader/less sharp.

MEMBRANE DYNAMICS
Although lipids are mobile in the plane of the bilayer, lipid
movement from one face of the bilayer and the other is a
much slower process.
FAST SLOW
This allows the generaEon and maintenance of
LIPID
ASYMMETRY
MEMBRANE DYNAMICS
At physiological temperature, transbilayer—or “flipflop”—
diffusion of a lipid molecule from one leaflet of the bilayer to the
other occurs very slowly if at all in most membranes.
Transbilayer movement requires that a polar or charged head group

leaves its aqueous environment and move into the hydrophobic
interior of the bilayer, a process with a large, posiEve free-‐energy
change.
MEMBRANE DYNAMICS
There are, however, situaEons, like during membrane synthesis, in
which such movement is essenEal.
A family of proteins, the flippases, facilitates flipflop

diffusion, providing a transmembrane path that is
energeEcally more favorable and much faster than the
uncatalyzed movement.
TRANSPORT ACROSS
MEMBRANES
TRANSPORT ACROSS MEMBRANE
Cell membranes are selecFvely permeable.
  Some solutes cross the membrane freely

  Some solutes cross with assistance
  Some solutes do not cross at all.
The ability of small molecules to freely cross the

phospolipid bilayer is a funcEon of their hydrophobicity.
The permeability of small molecules is correlated with their solubility
in a nonpolar solvent relaEve to their solubility in water.
Lipid bilayer membranes have a VERY LOW PERMEABILITY
for IONS and MOST POLAR MOLECULES.
Water is an excepEon to this generalizaEon.

There are four molecules that pass the lipid bilayer readily:
•  H2O
•  O2
•  CO2
•  CO
Membrane transport is essenEal for cellular life.
Membrane transport is dependent:
1) SIZE and POLARITY of solute

Large molecule would not pass due to their size, and polar
molecules would not pass because they have high affinity
for the extracellular water molecules in which they are
dissolved and because they cannot easily pass through the
non-‐polar membrane.
2) transmembrane solute CONCENTRATION

There are TWO means for the movement of substances through a
membrane:
Simple diffusion through the lipid bilayer

1) PASSIVELY
by diffusion through an aqueous,
protein-‐lined channel
Facilitated diffusion
by a protein transporter
(carrier)
or
2) ACTIVELY which requires an energy-‐driven protein

by an energy-‐ “pump” capable of moving substances against
coupled transport a concentraEon gradient
process
PASSIVE
TRANSPORT
Simple diffusion
In simple diffusion, small noncharged molecules or lipid soluble
molecules pass between the phospholipids, moving from areas of
high concentraEon to areas of low concentraEon (“they MOVE DOWN
their concentraFon gradient”).
high concentraEon
O2, CO, CO2, H2O and

most lipids enter and
leave cells by simple
diffusion.
low
concentraEon
Simple diffusion
Simple diffusion
In accord with the Second Law of Thermodynamics,

molecules spontaneously move from a region of higher
concentraEon to one of lower concentraEon
An entropy increase powers transport across the
membrane.
Ma[ers become more complicated when the molecule is highly polar
or charged.
Facilitated diffusion (passive transport)
Why is it so difficult for a polar or charged solute to cross the
phopsholipid bilayer?
To pass through a lipid bilayer, a polar or charged solute must first
give up its interacEons with the water molecules in its hydraEon shell,
then diffuse through a solvent (lipid) in which it is poorly soluble.
The energy of acEvaEon for translocaEon of a polar solute across the
bilayer is so large that pure lipid bilayers are virtually IMPERMEABLE
TO POLAR AND CHARGED SPECIES.
Membrane proteins lower the acEvaEon energy for transport of polar
compounds and ions by providing an alternaEve path through the
bilayer for specific solutes.
Facilitated diffusion (passive transport)
There are two categories of transporters:
CHANNELS CARRIERS
The diffusion across the membrane is facilitated by the channel or a
carrier.
It is also called passive transport, because the energy driving the ion
movement originates from the ion GRADIENT itself, WITHOUT any
ENERGY contribuEon by the transport system.
-‐ CHANNELS
Channels are proteins that allow transmembrane
movement at vey high rates approaching the limit of
unhindered diffusion.
High
concentraEon
Low
concentraEon
-‐ CHANNELS
Ion channels can be highly selecFve for parEcular ions.
For example, some channels
allow the flow of K+ very
effecEvely but do not allow
appreciable levels of Na+ to
cross the membrane.
Other channels transport

posiEvely charged ions
(caEons), but block the flow of
negaEvely charged ions
(anions).
Channels are usually not saturable.

-‐ CHANNELS
Normally K+ float around surrounded by 8 water molecules.
In order to pass through the selecEvity filter, each
potassium ion has to shed these water molecules.
This is how the selecEvity filter works:

the dimensions of the channel are
designed to mimic this shell of water.
Protein’s O atoms that line the pore
are oriented toward the center of the
channel. 8 of these O atoms surround
each K+, and act as a perfect
replacement for the normal layer of
water molecules.
-‐ CARRIERS
The binding of the solute to the carrier on one side of the
membrane triggers a conformaFonal change in the
protein, exposing the solute to the other surface of the
membrane, from where it can diffuse down its
concentraFon gradient.
-‐ CARRIERS
These specific protein carriers move molecules, such are

sugars and amino acids one at a Eme down a concentraEon
gradient.
-‐ CARRIERS
Facilitated diffusion by carriers, is similar in many ways to an enzyme-‐
catalyzed reacEon.
Their “substrates” are moved from one compartment to another, but
are not chemically altered.
They also called

TRANSPORTERS
or
PERMEASES
In addiEon, both enzymes

and transporters exhibit
saturaEon-‐type kineEcs.
-‐ CARRIERS
Like enzymes, transporters bind their substrates with stereochemical
specificity through mulEple weak, noncovalent interacEons.
Unlike ion channels, which can conduct millions of ions per
second, most carriers can move only hundreds to thousands of solute
molecules per second across the membrane.
Another important feature of facilitaEve transporters is that, like

enzymes and ion channels, their acEvity can be regulated.
Facilitated diffusion is parEcularly important in mediaEng the entry

and exit of polar solutes, such as sugars and amino acids, that do not
penetrate the lipid bilayer.
-‐ CARRIERS
Glucose is the body’s primary source of direct energy, and most
mammalian cells contain a membrane protein that facilitates the
diffusion of glucose from the bloodstream into the cell.
A gradient favoring the

conEnued diffusion of
glucose into the cell is
maintained by
phosphorylaEng the
sugar aêr it enters the
cytoplasm, thus lowering
the intracellular glucose
concentraEon.
CARRIERS
CHANNELS
Summary of passive transport
ACTIVE
TRANSPORT
AcFve Transport
In acFve transport, substances are moved AGAINST their
concentraFon gradient.
AcFve transport:
Transport AGAINST the
concentraEon gradient
AcEve transport results in the accumulaFon of a solute on one side of
the membrane.
AcFve Transport
AcFve transport is thermodynamically unfavorable (endergonic)
AcEve transport takes place only when coupled (directly or

indirectly) to an exergonic process such as
•  the absorpEon of sunlight
•  an oxidaEon reacEon
•  the breakdown of ATP
•  or the concomitant flow of some other chemical species
down its electrochemical gradient.
AcFve Transport
For examples, typically, the K+ concentraEon inside a mammalian cell
is about 100 mM, whereas that outside the cell is only about 5 mM.
[K+] = 100 mM

[K+] = 5 mM
The ability of a cell to generate such steep CONCENTRATION

GRADIENTS across its plasma membrane cannot occur by either
simple or facilitated diffusion.
AcFve Transport
Like facilitated diffusion, acEve transport depends on
integral membrane proteins that selecEvely bind a
parEcular solute and move it across the membrane in a
process driven by changes in the protein’s conformaFon.
Unlike facilitated diffusion, however, movement of a solute
against a gradient requires the coupled input of
ENERGY
Proteins that carry out acFve transport are oên referred

to as “PUMPS.”
AcFve transport mechanisms can be divided into two major
categories:
1) PRIMARY acFve transport

Directly uses a source of chemical energy (e.g., ATP) to move ions
across a membrane.
2) SECONDARY acFve transport

Uses an electrochemical gradient – generated by acEve transport – as
an energy source to move molecules, and thus does not directly
require a chemical source of energy such as ATP. The energy-‐releasing
movement of one molecule (down its gradient) is coupled with the
energy-‐requiring movement of another molecule (up its gradient).
PRIMARY AcFve Transport
Solute accumulaEon is coupled directly to an exergonic chemical
reacEon, such as conversion of ATP to ADP + Pi.
One of the most important pumps in animal cells is the sodium-‐
potassium pump, which moves Na+ out of the cells and and K+ into
them .
This process is responsible for maintaining the large excess of Na+
outside the cell and the large excess of K+ ions on the inside.
Na+ +
K+ Na Na +
Na
Na +
+
+ K+ Na+
Na Na+
Na+ K+
K+ K+ K+ +
Na + Na
Na+ K
+
K + K+
K+ K+
The sodium potassium pump, or Na+/K+-‐ATPase is an enzyme that
couples the breakdown of ATP to the simultaneous movement of
both Na+ and K+ against their electrochemical gradients.
+ + + +
+ + +
+ +
-‐ -‐ -‐ -‐
-‐ -‐ -‐
-‐ -‐
The sodium-‐potassium pump maintain correct concentraEons of Na+

and K+ in living cells, and generates the voltage across the cell
membrane.
Pumps involved in creaEng and maintaining membrane voltages are

known as electrogenic pumps, and the sodium-‐potassium pump is
generally considered the main electrogenic pump in animals.
This pump is called a P-‐type ion pump because the ATP interacEon
phosphorylates the pump and causes a change in its conformaFon.
SODIUM POTASSIUM ATPase
PhosphorylaEon makes the

To begin, the pump is When Na+ bind, they pump change shape, re-‐
open to the inside of the trigger the pump to orienEng itself so it opens
cell. hydrolyze ATP. towards the extracellular
In this form, the pump One phosphate group space.
really likes to bind (has a from ATP is a[ached to In this conformaEon, the
high affinity for) sodium the pump, which is then pump no longer likes to
ions, and will take up said to be bind to sodium ions (has a
three of them. phosphorylated. low affinity for them), so
ADP is released as a by-‐ the three sodium ions are
product. released outside the cell.
SODIUM POTASSIUM ATPase
In its outward-‐facing With the phosphate

form, the pump switches group gone, the pump In its inward-‐facing shape,
allegiances and now will change back to its the pump loses its interest
really likes to bind to (has original form, opening in (has a low affinity for)
a high affinity for) towards the interior of potassium ions, so the two
potassium ions. the cell. potassium ions will be
It will bind two of them, released into the
and this triggers removal cytoplasm. The pump is
of the phosphate group now back to where it was
a[ached to the pump in in step 1, and the cycle can
step 2. begin again.
For each molecule of ATP converted to ADP and Pi, the transporter
moves two K+ inward and three Na+ outward across the plasma
membrane.
2 K+
3 Na+
The process is electrogenic—it creates a net separaEon of charge

across the membrane.
The ATP-‐driven Na+-‐ K+ pump, STORES ENERGY by creaFng a steep
CONCENTRATION GRADIENT.
SECONDARY AcFve Transport
The electrochemical gradients created by acEve transport are like the
cellular equivalent of baKeries: they store energy, which can be
released as ions move back down their concentraEon gradients.
Secondary acFve transport takes advantage of the

gradients established by primary acFve transport,
using their energy to transport other substances
against their own gradients
Unlike in primary acEve transport, in secondary acEve

transport, ATP is not directly coupled to the molecule of
interest.
Secondary transport is accomplished by coupling the

movement of one molecule down its gradient (energy-‐
releasing process) to the transport of some other molecule
up its gradient (energy-‐requiring process).
movement of one molecule movement of one molecule

down its gradient up its gradient
(energy-‐releasing process) (energy-‐requiring process)
coupling
In other words, secondary acEve transport occurs when

endergonic (uphill) transport of one solute is coupled to
the exergonic (downhill) flow of a different solute that was
originally pumped uphill by primary acEve transport.
Secondary acEve transport is also known as COTRANSPORT,

and it's mediated by carrier proteins called cotransporters.
As an example of secondary acEve transport, let's suppose we have a
high concentraEon of sodium ions in the extracellular space (thanks
to the hard work of the sodium-‐potassium pump).
Na+ Na+ Na+ Na+ Na+ Na

+
Na +
Na+
Na + Na+
Na+
If a route such as a channel or carrier protein is open, sodium ions will
move down their concentraEon gradient and return to the interior of
the cell.
This downhill movement is coupled to the transport of other
substances by use of a shared carrier protein.
For instance, a carrier protein allows sodium ions to move down their
concentraEon gradient, while simultaneously transporEng a glucose
molecule up its gradient and into the cell.
The sodium ions moving down their concentraEon gradient have the
capacity to do work, and the carrier protein harnesses the energy of
the gradient to transport the glucose molecule.
Two molecules being transported may move either in the
same direcEon (i.e., both into the cell), or in opposite
direcEons (i.e., one into and one out of the cell)
When they move in the same direcEon, the protein that
transports them is called a symporter, while if they move in
opposite direcEons, the protein is called an anFporter.

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11 - Membranes

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BIOLOGICAL

Yet, they also contain TRANSPORT SYSTEMS that allow

Membranes also contain CARBOHYDRATES that are linked

However, because of its mosaic nature,

LIPIDS PERMEABILITY BARRIER

PROTEINS TRANSPORT SYSTEMS

Membranes also contain CARBOHYDRATES that are linked

7) Membranes are asymmetric.

The two faces of biological membranes always diﬀer from

Lipid molecules diﬀuse rapidly in the plane of the

Membrane potenEal plays a key role in transport, energy

Diﬀerent membrane have diﬀerent chemical composiEon.

So before describing membrane structure and funcEon we need to

H2O H2O H2O H2O

In other words, the structure of a bimolecular sheet is

The growth of lipid bilayers from phospholipids is a rapid

Only lipid porEons at the edge of

2) Furthermore, van der Waals a[racEve forces between

• Lipid bilayers are self-­‐sealing because a hole in a bilayer is

Most cells 50% proteins

1) Integral proteins (intrinsic proteins)

2) Periferal proteins (extrinsic proteins)

1) Integral proteins (intrinsic proteins)

Are very FIRMLY ASSOCIATED with the membrane, they

Most integral membrane proteins SPAN the lipid bilayer.

1) Integral proteins (intrinsic proteins)

1) Integral proteins (intrinsic proteins)

The polypepEde chain of some

Most of the amino acids in

Beta barrel structures are common in

In contrast, the inside of of a beta barrel is

Glycoproteins of the plasma membrane are always situated

their ability to change shape without losing their integrity

The more movement

The less movement,

If the ﬂuidity is determined by the relaEve movement, what

strength of the aKracFon

strength of non covalent intermolecular forces that exist

MORE FLUID MORE RIGID

At a speciﬁc temperature called Tm, the melEng

Membrane with Membrane with

HIGH Tm LOW Tm

But what determines the strength of the intermolecular

(2) Degree of unsaturaFon of the faKy acids

Long faKy acids Short faKy acids

More hydrophobic Less hydrophobic

LOW FLUIDITY HIGH FLUIDIY

HIGH Tm LOW Tm

Saturated faKy acids Unsaturated faKy acids

More hydrophobic Less hydrophobic

LOW FLUIDITY HIGH FLUIDIY

HIGH Tm LOW Tm

Foods with trans fats:

Cakes, pies and cookies

In ANIMALS, the factor

The hydrophobic four ring structure lies alongside the

• At T below Tm, cholesterol inserEon

• At T above Tm, rigid ring system of sterol

• Result: thermal transiEon becomes broader/less sharp.

•  Lipid bilayers are self-‐sealing because a hole in a bilayer is

•  At T below Tm, cholesterol inserEon

•  At T above Tm, rigid ring system of sterol

•  Result: thermal transiEon becomes broader/less sharp.

  Some solutes cross the membrane freely

2) ACTIVELY which requires an energy-‐driven protein

The sodium-‐potassium pump maintain correct concentraEons of Na+

In its outward-‐facing With the phosphate