Mechanisms of salt
tolerance in plants.
A review
Anton Mozqueira
Anton Mozqueira
Communication in Science Mechanisms of crop salt tolerance.
The problem of salinity tolerance in plants, especially those used as crops, is not a new one.
Dry-land salinity caused by irrigation, the removal of tress and the raising of the water table
affects around 2,223,948,433 acres of land worldwide(Colmer, Flowers & Munns 2006). This
area will continue to grow as the amount of irrigated land increases because there is a
strong link between irrigation and salinisation (Flowers 2004). Most of the important crops
that we rely on are salt intolerant, and growing them on saline soils reduces yields or
completely prevents growth. With the world population and the amount of irrigated land
increasing, the development of salt tolerant crops will become crucial if we are to continue
to produce sufficient amounts of food (Flowers 2004). Farmers, plant breeders and most
recently scientists continue to grapple with the problem of creating a plant that is capable of
being grown under saline conditions without the sacrifice of those traits that make it useful
as a crop plant (Cheeseman 1988). The reason for this is found in the complexity of a plants
response to salt stress, with mechanisms operating cooperatively at many levels of the
plants physiology, some of which remain to be fully investigated (Flowers 2005). This review
explores some of those natural mechanisms employed by plants with reference to their
presentation in the literature.
There are two main types of plants relative to salinity tolerance; they are halophytes and
glycophytes; those which can thrive in saline conditions and those which can’t (Parida, Das
2005). Halophytes are able to tolerate and in some cases require the presence of salt
whereas glycophytes don’t. Both types of plants use the same kinds of mechanisms to deal
with salt, but halophytes are more proficient at doing so (Yokoi, et al 2002).
All plants require some salts, especially potassium to function properly, (ASHRAF et al.
2008), but excessive concentrations of salts, particularly of sodium and chloride disrupt the
plants growth in several ways. Too much sodium or chloride is harmful to the plant because
it reduces its uptake and transport of water by changing the osmotic potential of the plants
cells (Parida, Das 2005). If there is too much salt outside the plants cells then the salt holds
all the water so that the plant cannot take any in. When the plant tries to adjust its osmotic
potential by taking in large amounts of salt, high concentrations can develop in sensitive
areas, such as the cytoplasm and chloroplast (Flowers 2005). High concentrations of sodium
or chloride will disrupt these cells function and inhibit the plants growth.
Plants which maintain high transpiration rates during salinity stress are less tolerant
(Flowers 2004). Transpiration, or the process of the plant exchanging oxygen for carbon
dioxide, requires that more water be taken into the plant, to replace what was lost to
evaporation. This process consumes more water during the day than when carried out at
night. Because the plant requires more water, greater amounts of salts will be absorbed
with the water by the plant. The ability to adjust transpiration rates and times will make a
plant more tolerant (Gorham, Jones & McDonnell 1985).
Anton Mozqueira
Communication in Science Mechanisms of crop salt tolerance.
Another problem caused by high salinity is the production of reactive oxygen species (ROS)
in the chloroplast, mitochondrion and peroxisome (Ashraf 2009). They include super oxide,
hydrogen peroxide, hydroxyl radical and singlet oxygen (Ashraf, Akram 2009). They are a
product of the plants metabolism and are a normal component of the plants chemistry
(Ashraf 2009). They are only harmful in higher concentrations, a result of the stress caused
by high salinity (Ashraf, Akram 2009, Apse 2002). Because less salt tolerant plants do not
normally produce enough anti-oxidants to scavenge all of the ROS produced under these
conditions, the reactive oxygen species will try to pair their single oxygen with other
elements to form a more stable compound. To complete this pairing, the ROS can break
down important molecules such as proteins, lipids, and nucleic acids within the cells (Ashraf
High levels of salinity also inhibit the plants growth by limiting or preventing membrane
transportation of nutrients from the ground to the plants cells (Ashraf et al. 2008). Because
there are high levels of sodium and chloride in the soil it is hard for the plant to select for
potassium and other useful ions over the more abundant sodium and chloride ions(Flowers
2004). This means that the plant will not receive sufficient nutrition to complete its
metabolic functions for growth, and that for any small amount of nutrition it receives, a
large amount of salt will have accumulated in its tissues.
Plant photosynthetic activity also decreases due to increased salinity (LI et al. 2008).
Because the plant tries to limit the loss of water through evaporation by closure of the
stomata, it is unable to exchange carbon dioxide for oxygen. This means that the plants
growth rate will be severely limited or stop altogether because it does not have any new
carbon to metabolise into the carbohydrates which it uses to grow (Parida, Das 2005). This
lack of oxygen exchange also leads to the development of excessive amounts of oxygen in
the chloroplast, leading to the formation of reactive oxygen species or ROS, which lead to a
further reduction of photosynthesis (Ashraf 2009). High salt content within the plants cells
also leads to the disruption of many complicated metabolic processes necessary for
photosynthesis (Flowers 2005).
Thus far the effect of high salinity on plants has been presented. Now the mechanisms used
to deal with high salinity will be examined. Plants employ many mechanisms at many levels
of their physiology in order to tolerate salinity. They try to employ several simpler
mechanisms cooperatively before altering the function of major processes such as
photosynthesis (Parida, Das 2005).
Plants that can regulate their intake of sodium and chloride are able to continue functioning
and grow unimpeded in highly saline environments (Flowers 2004). Sodium and potassium
share the same transporter protein, called the Na +/K+-ATPase, so it is easy for sodium to be
over selected for when it is in higher concentrations (Apse 2002). Sodium/potassium
transporters in halophytic plants are more selective than those in glycophytes (Apse 2002).
Another method of entry is the slow vacuolar (SV) ion channels. These do not select for
Anton Mozqueira
Communication in Science Mechanisms of crop salt tolerance.
specific ions and so allow a large amount of ions in when there are high external
concentrations (Ashraf, Akram 2009). It has been found that in halophytic plants, slow
vacuolar ion channels are open much less than glycophytic ion channels and so prevent
indiscriminate admission of unwanted ions, leading to increased tolerance.
Plants that can actively select for ions are able to transport them to the vacuole, away from
the sensitive metabolic functions of the cytoplasm (Apse 2002). Here they help the plant
maintain its osmotic potential in highly saline soils, allowing it continue to access water and
nutrients in the ground (Flowers 2005). This function allows the plant to continue to grow.
Some plants also specifically fill developed and older tissues and leaves with salt before
allowing them to fall off the plant, helping to maintain ionic balance within the plant if
growth rates alone are not sufficient (Parida, Das 2005).
Plants exposed to many different types of stresses produce compatible solutes (Ashraf et al.
2008). These solutes are produced by the plant and help to preserve the function of
enzymes within the cytoplasm when there are high concentrations of sodium in its tissues (
Yokoi et al. 2002). They can also help prevent ROS from damaging plant cells by reacting
with them (Ashraf et al. 2008). The most common compatible solutes made by plants are
glycinebetaine, mannitol and proline (Flowers 2005).
When the plants photosynthetic processes are disrupted and reactive oxygen species begin
to damage the plant, anti oxidants and detoxifying enzymes are produced (Ashraf 2009).
These help to scavenge and reduce the impact of the reactive oxygen species. Their
increased production is an indicator of salt tolerance in a plant. Some of the anti oxidants
produced are ascorbate, glutathione, a-tocopherol, and carotenoids, and the detoxifying
enzymes include catalase and peroxidase(Apse 2002).
Salt tolerance in plants can also be attributed to their ability to alter transpiration rates and
times (Flowers 2004). A shorter transpiration period, or opening of the stomata only at night
allows the plant to continue growing without losing too much precious water to higher
evaporation rates during the day. This type of transpiration is called crassulacean acid
metabolism or (CAM) and is only displayed naturally in halophytic plants, such as
Mesembryanthemum crystallinum(Parida, Das 2005).
Some halophytes have developed specialized organelles, such as salt excreting glands, to
help regulate the amount of sodium in their tissue, when it cannot be compensated for by
growth rates alone(Flowers 2005). Salt excretion glands show strong preference for specific
ions such as sodium, over useful ions like potassium, so that the plant does not waste this
important nutrient (Parida, Das 2005). Other plants allow salt to accumulate in old or
developed tissues, away from important new growth areas. Once the tissue is full of salt it is
allowed to drop off, thereby removing the salt and maintaining ionic balance within the
plant (Parida, Das 2005).
Anton Mozqueira
Communication in Science Mechanisms of crop salt tolerance.
Flowers (1997, 2004, 2005) identifies in several of his reviews of the literature that many
researchers claim increased salt tolerance in their plants after the transferral of one or two
genes in order to over express a single trait. He dismisses their claims and points to lack of
commercially viable salt tolerant crops as his evidence and proposes that yield is a better
indicator of tolerance than the ability to simply survive(Flowers 2004). Although it would be
more convenient for a single trait to confer tolerance, all of the mechanisms above can be
found working together in halophytic plants to help them tolerate saline conditions, with
many, many genes controlling them (Flowers 2005). In order for a commercially viable, salt
tolerant cultivar to be produced, most or all of these traits would have to be transferred to
confer tolerance and the maintenance of high yields. Flowers proposes that the
development of halophytic plants into crops, particularly those halophytic grasses in the
same family as wheat, the Triticea, might be a new direction that could lead to success in
this area(Colmer, Flowers & Munns 2006, Flowers 2004). Other researchers suggest that the
development of a more complete understanding of the complex genetics responsible for all
the traits, and use of more advance gene identification and transferral techniques will result
in increased success(Ashraf, Akram 2009). As the pressures of food demand and salinisation
of farmable land mount, increased resources and energy will be devoted to this problem
and a re-evaluation of the potential of existing halophytic species as crop plants might be
necessary. In the meantime, the methods of exogenous application of compatible solutes,
as well as increasing the amounts of calcium and potassium in saline soils as identified by
(Ashraf et al. 2008), might be a temporary solution for traditional crop plants.
Anton Mozqueira
Communication in Science Mechanisms of crop salt tolerance.
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Mechanisms of salt tolerance in plants. - Plant-salt