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Ashwani Kumar, Charu Lata, Pooja* and Anita Mann

ICAR-Central Soil Salinity Research Institute, Karnal 132001, India
*ICAR- Sugarcane Breeding Institute, Regional Center, Karnal 132001, India
Among abiotic stresses, drought and salinity stress are the major causes of historic
and modern agricultural productivity losses throughout the world. Both drought and salinity
result in osmotic stress that may lead to inhibition of growth. Consequently, most
angiosperms, including all major crop species, have a diminished capacity for Na+ transport
and tolerance to high salinity. New insights into the molecular mechanisms of Na+/K+
discrimination, Na+ extrusion and compartmentation, water transport, osmolyte biosynthesis
and function have led to genetically engineered plants with improved salt, drought, and cold
tolerance. Osmoprotectants are small molecules that act as osmolytes and help organisms
survive under extreme osmotic stress; examples include betaines, amino acids, and the sugar
trehalose. These molecules accumulate in cells and balance the osmotic difference between
the cell's surroundings and the cytosol. In extreme stress conditions, the cytosol and
osmoprotectants become a glass-like solid that helps stabilize proteins and cell membranes
from the damaging effects of desiccation. Compatible solutes have also been shown to play a
protective role by maintaining enzyme activity through freeze-thaw cycles and at higher
temperatures. Their specific action is unknown but is thought that they are preferentially
excluded from the proteins interface due to their propensity to form water structures. The
synthesis of glycine betaine, one of the most powerful osmoprotectants found in nature, also
plays an important role in the adaptation process of plants to a high-osmolarity environment.
Abiotic factors like drought, temperature, salt stress, etc. results in depletion of large number
food productions in todays world and these considerations arouse strong interest in plant
abiotic stress responses.
What is Osmoregulation?
The active regulation of the osmotic pressure of an organism's fluids to maintain the
homeostasis of the organism's water content is known as Osmoregulation; this means it keeps
the organism's fluids from becoming too diluted or too concentrated. Osmotic pressure is a
measure of the tendency of water to move into one solution from another by osmosis.
Osmosis is the movement of solvent molecules through a selectively-permeable membrane

into a region of higher solute concentration, aiming to equalize the solute concentrations on
the two sides. Higher the osmotic pressure of a solution, more water moves into the solution.
Pressure must be exerted on the hypertonic side of a selectively-permeable membrane to
prevent diffusion of water by osmosis from the side containing pure water. Various
metabolism involved in osmoregulation include polyoles, sugars, amino amino acids, proteins
and various other organic solutes. Act these metabolites play are important role in
maintaining osmotic potential of cytoplasm to maintain cell turgor. The concentration of these
compatible solutes/osmoprotectants within cell is maintained by either synthesis of these
compounds or combination of synthesis and degradation. Thus these osmolytes protect one
structure and maintain osmotic balance within the cell via continuous water influx.
Mechanism of Osmoregulation in Plants
While there are no specific osmo-regulatory organ in higher plants, the stomata are
important in regulating water loss through evapo-transpiration and on the cellular level,
vacuole plays an important role in regulatory the concentration of solvents in the cytoplasm.
The chemiosmotic regulatory systems of plant and fungal cells differ fundamentally from
those found in animal cells. In plants lack of plasma membrane Na +/K+-ATPases reported in
mechanism of osmoregulation. Thus, plants utilize H+-ATPases for primary extrusion or
sequestration of protons to generate H+ electrochemical gradients, which drive secondary ion
and nutrient transport processes via H+-symport/antiport systems. These H+-ATPase pumps
also modulate both intracellular and extracellular pH.
Except in the case of extreme halophytic archaebacteria, viable cellular processes in
animals, fungi, and plants depend upon the maintenance of low cytoplasmic Na+ and Cl
concentrations and a high K+/Na+ ratio, because K+ counteracts the inhibitory effects of Na+
(and Li+). Like animal cells, most plant cells maintain cytosolic K+ concentrations in the
range of 100200 mM and Na+ values in the low mM range (110 mM) up to a maximum of
100 mM (Maathuis and Amtmann, 1999). In contrast to K+, an essential cation for
maintaining biochemical interactions of the cytoplasm, Na+ is not essential for, but does
facilitate, volume regulation and growth in most plants. However, at high concentrations Na +
limits growth (Blumwald, 2000). Ironically, the productivity of irrigated agricultural regions
is generally many times greater than non-irrigated areas, yet irrigated crops are most
susceptible to detrimental salinity effects. Therefore, genetic engineering of crop plants to

improve their capacity for Na+ transport and sequestration is an important goal for meeting
the future food and fiber demands of a rapidly growing human population.
Many plants, such as extreme halophytes, display Na + dependence for optimal growth
and development and have developed specialized structures such as salt glands and bladders
to accommodate high salt concentrations in tissues (Glenn et al., 1999). Others have
developed whole plant strategies for avoiding stress such as accelerated completion of
ontogeny. However, these specialized adaptations are lacking in most major crop species.
Furthermore, the precise impact of osmotic and ionic effects on cell growth, division,
phytohormone balance, and death in the context of the whole plant are complex and require
further investigation (Munns, 2001). Therefore, emphasis is placed on the molecular genetic
mechanisms controlling osmotic regulation at the cellular level, mainly because the action
and regulation of most osmoregulatory components has not been fully explored in the context
of the whole plant. Some channels associated for osmoregulation process are cation and anion
selective channels. Plant genomes encode multiple classes of ion channels that mediate K +
and Na+ transport (see Amtmann and Sanders, 1999; Schachtman and Liu, 1999;
Zimmermann and Sentenac, 1999; Czempinski et al., 1999;). Ion channels transport ions
from the soil solution, secrete ions into xylem sap, and participate in signaling (Zimmermann
and Sentenac, 1999). In contrast to carriers, ion channels conduct rapid, downhill
dissipation of transmembrane electrochemical gradients, often under the control of membrane
potentials that dictate the gating properties of the channel. Monovalent cation-selective
plasma membrane channels are divided into three classes depending on their
electrophysiological behavior and ion selectivity.
In plants generally betA and betB genes synthesized against drought stress and
produce BetB and BetA proteins. The deduced BetB and BetA proteins showed significant
similarity to soluble glycine betaine aldehyde dehydrogenases and membrane-bound choline
dehydrogenases, respectively, from a variety of organisms. Evidence is presented that BetA is
able to oxidize both choline and glycine betaine aldehyde and therefore can mediate both
steps in the synthesis of glycine betaine. Finally, glycine betaine synthesis in higher plants
involves a BADH in combination with a choline monooxygenase (Munns, R. 2001.)The
challenge for the coming decade will be to integrate information gathered at the molecular
genetic and cellular levels with the complexity of whole plant physiology. But to understand
genetic and molecular mechanism of osmoregulation is very difficult due to different types

of stress, any type of stress is converted ultimately in to drought stress and the genes
produced for any particular stress are different and encode different proteins to encounter
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