What Is Reverse Osmosis?

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How Does It Work?

Diffusion is the movement of molecules from a region of
higher concentration to a region of lower concentration.
Osmosis is a special case of diffusion in which the molecules
are water and the concentration gradient occurs across a
semipermeable membrane. The semipermeable membrane
allows the passage of water, but not ions (e.g., Na+, Ca2+,
Cl-) or larger molecules (e.g., glucose, urea, bacteria).
Diffusion and osmosis are thermodynamically favorable and
will continue until equilibrium is reached. Osmosis can be
slowed, stopped, or even reversed if sufficient pressure is
applied to the membrane from the 'concentrated' side of the
membrane.

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Reverse osmosis occurs when the water is moved across the membrane against the
concentration gradient, from lower concentration to higher concentration. To
illustrate, imagine a semipermeable membrane with fresh water on one side and a
concentrated aqueous solution on the other side. If normal osmosis takes place, the
fresh water will cross the membrane to dilute the concentrated solution. In reverse
osmosis, pressure is exerted on the side with the concentrated solution to force the
water molecules across the membrane to the fresh water side.
Reverse osmosis is often used in commercial and residential water filtration. It is also
one of the methods used to desalinate seawater. Sometimes reverse osmosis is used
to purify liquids in which water is an undesirable impurity (e.g., ethanol).
Would you like to know more about diffusion, osmosis, and reverse osmosis? Here
are a couple of additional resources:

Osmosis, Reverse Osmosis, and Osmotic Pressure - This site provides the
basic equations used to describe osmosis. Several printed and internet
references are also provided.
What Is Reverse Osmosis? - Osmonics defines reverse osmosis and provides
several links to online technical papers.

last change: 07/31/2003 20:50:00

Active Transport

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SciLinks program, a
service of National

Science Teachers
Association. Copyright
2001.

Participation of a carrier molecule in the active transport of certain molecules

through the membrane. The transport process is specific and energy-consuming. The
conformation of the carrier changes during transport (ping-pong mechanism).

Active transport is the movement of a molecule across a membrane or another barrier that
is driven by energy other than stored in the concentration gradient or the electrochemical
gradient of the transported molecule. This type of transport requires usually the
expenditure of ATP and the help of specific transport proteins. In this way can even large
molecules can be channelled through the membrane. For the understanding of the details
of the mechanism are both the structure of the involved transport molecules and the
question how the energy for the transport is supplied and how it is used important. We
will meet the electrical membrane gradient again in this context.
Active transport can only occur at intact, closed membranes. Such membranes can
envelop very different compartments, like the whole cell, vesicles, the vacuole, the
mitochondrial matrix, the inner thylacoid space of the chloroplasts, etc. As a result of
active transport can ions and metabolites be concentrated within the respective
compartment or the cell and the steady state of the metabolism can be kept constant
despite of large fluctuations in the external medium's composition. Ions, especially
potassium, calcium, magnesium and phosphate have an important part in the regulation of
the metabolism.

The transport direction is thermodynamically determined by coupling the transport with a

gradient, usually an electron gradient. The direction can be reversed if the appropriate
substrate concentrations are chosen.
As a consequence of the respiratory chain and of photosynthesis are protons channelled
out of the mitochondrial matrix and the thylacoids of the chloroplasts, respectively. At the
same time causes the efflux of protons a translocation of electric charges. That way is
both a chemical proton gradient and an electric potential generated. The tendency of the
protons (H+) to return to the inside of the compartment is therefore quite large. It is called
the proton motive force. P. MITCHELL (1966, 1974) studied the process and his
observations pointed at a key position of the proton translocation. The energy set free
when the protons flow back is used for the synthesis of ATP, the involved enzyme is the
ATP synthethase. Depending on whether the reaction takes place in mitochondria or in
chloroplasts is it spoken of oxidative phosphorylation and photophosphorylation,
respectively. But protons can flow back under two further conditions:
1. By antiport, i.e. a proton is exchanged with another cation and fuels in
that way the transport of its counterion out of the compartment.
2. By symport. A proton is transported into a compartment and the energy
for the transport is supplied by the simultaneous transport of an anion
or a substrate molecule in the same direction. The process allows the
uptake and accumulation of anions or (small) molecules by the
compartment. In other words: an electrochemical gradient is used as a
source of energy for the active transport of molecules and ions through
the membrane. A second possible energy source for transport processes
is the breakdown of ATP.
ATP hydrolysis is actually the reversion of oxidative or photophosphorylation. The ATP
synthethase works the other way round: protons (and other cations like potassium) are
actively transported out of the compartment (proton pump) and build as a consequence a
pH gradient and a membrane potential up. These two forces again drive the translocation
of other molecules (or ions) via antiport or symport. As mentioned at the beginning is
active transport the name of two principally different but coupled processes:
1. The transport performed by a carrier. The direction is solely dependent
on the substrate gradient and serves to even out substrate
concentrations.
2. The energy-dependent process that leads to an asymmetry of the
transport so that the transported molecules are accumulated at one side
of the membrane against an (electro-) chemical gradient.

Chemiosmotic coupling of the transport of

molecules through a membrane according to
the principles of antiport (1) and symport (2,
3). S = substrate
(according to C. L. SLAYMAN und D.
GRADMANN, 1975).

The resulting asymmetric transport of the carrier is based on an energy-consuming

conformational change of the molecule (for example a phosphorylation of the carrier).
The two states of conformation (phosphorylated and non-phosphorylated) are
characterized by different affinities for their substrates. The velocity constants of the
permeation can vary by scales. A number of toxic substances like ammonium ions or
dinitrophenyl (DNP) destroy the proton gradient so that no more energy is available for
active transport. Such substances are called uncoupling agents.
The membrane-bound proteins needed for active transport are without exception
oligomerous complexes, i.e. they consist of several protein subunits. They are often
termed pumps, too.
A reversible ATP synthethase (able to hydrolyze ATP) that gets its energy from the
electrochemical potential difference between inside and outside is among the most
important components of the whole complex. The transport kinetics of the pumps can similar to those of facilitated diffusion - be compared with enzyme kinetics.
The proton complexes work analogous to enzymes and are therefore often also classified
as permeases. Proteins of animal membranes (like erythrocytes) and micro-organisms
have been studied especially extensively.
The number of polypeptide chains that make up a complex, the molecular weight and the
number of ions or molecules transported per time unit is often known. It is also known,
that many pumps work only when in their specific surrounding, in other words when in
close vicinity to the right phospholipids. And finally is it known that pumps are integrated
into the membrane in always the same orientation because transport is always directed.
But in no case is the exact tertiary or the quaternary structure of the complex known and
only these structures would explain, why a specific molecule or ion and no other, similar
ones are transported.

The best-known and best-studied ion pumps are

the sodium-potassium- and the calcium pump.
The sodium / potassium pump was isolated from
the most different types of membranes.
It is generally known that cells contain more
potassium than sodium ions. This is also true for
plants living in saline (sodium containing) soils.
Such plants are also said to be halophilic.
Especially in the cells of halophilic plants was a
very large activity of sodium / potassium
exchanges detected. The cells invest a
considerable amount of energy in order to
maintain within the cytosol as low a concentration
as possible. Part of the sodium is pumped into the
vacuole where it does not interfere but helps to
maintain the osmotic pressure. A number of
secondary plant products like the glycosides
oubain (strophanthin) and digitonin inhibit the
sodium / potassium pump selectively.
The calcium pump located especially in intracellular and organelle-membranes is of no
lesser importance. The calcium concentrations within the different compartments differ
by scales. On the other hand is calcium known for its regulating effects on the
metabolism. The calcium pump has accordingly a direct influence on the throughput
ratios of many biosynthetic pathways.

Proton pumps: 1. electrogenic pump, 2. electro-neutral pump, 3. electro-neutral pump

with calcium as its counterion, 4. electrogenic proton transport, 5. electro-neutral anion /
OH antiport (according to R. E. CLELAND, 1982).

Our knowledge about the exact distribution of ions within the single compartments is
rather sketchy, especially in plants.
A large advantage of micro-organisms is that mutants with defects in the transport system
can be isolated. The defects can be traced back to the losses or the changes of particular

subunits of the protein complex. Two principally different sugar transport mechanisms
were detected in micro-organisms:
1. Active sugar transport via indirect coupling to an energy source. The
substrate is transported through the membrane without changes. The
carrier changes its affinity for the substrate during its translocation
through the membrane. The affinity is high at the outside of the
membrane and low at its inside. In many cell types is the transport of
metabolites (and substrates, respectively) coupled to a functioning
sodium / potassium pump.
2. Active sugar transport by means of substrate modification. The
substrate is chemically modified during the transport. Examples are the
phosphorylation of sugars and the glycosylation of adenine. The
process is also called vectorial phosphorylation. The transport is
initiated by a chemical reaction during which the substrate is
'phosphorylated into the cell'.
Peter v. Sengbusch - Impressum

Comparison of
passive and active
transport.
1998 by Alberts, Bray, Johnson, Lewis, Raff, Roberts, Walter.
Published by Garland Publishing, a member of the Taylor & Francis Group.

http://www.garlandscience.com

Legend:
If uncharged solutes are small enough, they can move down their concentration gradients
directly across the lipid bilayer itself by simple diffusion. Examples of such solutes are ethanol,
carbon dioxide, and oxygen. Most solutes, however, can cross the membrane only if there is a
membrane transport protein (a carrier protein or a channel protein) to transfer them. As
indicated, passive transport, in the same direction as a concentration gradient, occurs
spontaneously, whereas transport against a concentration gradient (active transport) requires an
input of energy. Only carrier proteins can carry out active transport, but both carrier proteins and
channel proteins can carry out passive transport.
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