Physiology and Molecular Biology of Stress ... - KHAM PHA MOI
Physiology and Molecular Biology of Stress ... - KHAM PHA MOI
Physiology and Molecular Biology of Stress ... - KHAM PHA MOI
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Salt <strong>Stress</strong><br />
47<br />
tolerance research is to determine those transporters which function in the control <strong>of</strong><br />
Na + entry, to find a way to limit Na + influx <strong>and</strong> thus to improve salinity tolerance (Zhu,<br />
2001).<br />
4.2. Significance <strong>of</strong> K/Na Selectivity<br />
Uptake <strong>and</strong> distribution <strong>of</strong> sodium ions within the root is to a large extent connected<br />
with the effects <strong>of</strong> potassium, since Na + efflux in root cortex cells is stimulated by K +<br />
influx, which is related to the K/Na root selectivity (Jeschke, 1972). The presence <strong>of</strong><br />
potassium (<strong>and</strong> calcium) ions has been shown to decrease Na + influx into plant cells<br />
(e.g. Laz<strong>of</strong> <strong>and</strong> Bernstein, 1999).<br />
In wheat, salt tolerance is related to the enhanced K/Na selectivity (Gorham,<br />
1990) <strong>and</strong> it has recently been reported that the genus Triticum expresses a range <strong>of</strong><br />
genetic variation related to K/Na discrimination. The trait is highly heritable (Munns et<br />
al., 2002). In contrast to dicotyledonous plants, in monocots the maintenance <strong>of</strong> a lower<br />
Na/K ratio in shoots is <strong>of</strong> greater significance, because <strong>of</strong> their lower capacity for<br />
sodium storage <strong>and</strong> higher requirement for K + <strong>and</strong> compatible organic solutes (Glenn et<br />
al., 1999).<br />
Time-course measurements <strong>of</strong> root ion content have indicated the possibility<br />
<strong>of</strong> a sodium-excluding mechanism in roots <strong>of</strong> Kosteletzkya virginica through low Na + /<br />
K + ratios brought about not only by exclusion <strong>of</strong> Na + , but also by a strong K + affinity<br />
which was ascribed to a change <strong>of</strong> the sterol/phospholipid ratio in salinized roots (Blits<br />
<strong>and</strong> Gallagher, 1990). A wide range <strong>of</strong> K + /Na + selectivity <strong>and</strong> sodium exclusion were<br />
demonstrated for halophytic <strong>and</strong> non-halophytic Chenopods (Reimann, 1992), with<br />
Na + accumulation being most pronounced in the shoots.<br />
To confer salt stress tolerance, in many species, the achievement <strong>of</strong> a high K + :<br />
Na + ratio is more important than simply maintaining low concentrations <strong>of</strong> sodium ions<br />
(Maathuis <strong>and</strong> Amtmann, 1999). A highly significant correlation between K + <strong>and</strong> Na + in<br />
leaves <strong>and</strong> salinity-induced yield reduction was reported for a number <strong>of</strong> rice genotypes<br />
irrigated with moderately saline water (Asch et al., 2000). The significance <strong>of</strong> a<br />
high K/Na ratio for maintenance <strong>of</strong> yield components <strong>and</strong> fiber characteristic has also<br />
been evaluated in salt-tolerant cotton lines (Ashraf <strong>and</strong> Ahmad, 2000). The maintenance<br />
<strong>of</strong> higher K + <strong>and</strong> Ca 2+ to Na + ratios, especially in young growing <strong>and</strong> recently<br />
exp<strong>and</strong>ed tissues, appeared to be important characteristics <strong>of</strong> salt tolerance in barley<br />
cultivars (Wei et al., 2003). In a study with four barley cultivars exposed to a range <strong>of</strong><br />
salinity concentrations (Figure 1) it was noticed that the highest differences in Na + : K +<br />
ratio, were expressed at 200 mol m -3 NaCl (Dajic, unpublished data). The Na + : K + ratio<br />
seems to be valuable criterion for screening <strong>of</strong> salt tolerance within salt-sensitive species,<br />
such as maize. It was found that among four maize cultivars differing in drought<br />
tolerance, <strong>and</strong> tested for their responses to salt stress (Dajic, unpublished data), the<br />
Polj 17, previously characterized for high capacity <strong>of</strong> ABA accumulation under drought<br />
stress <strong>and</strong> related tolerance to water deficit (Pekic <strong>and</strong> Quarrie, 1988) has maintained the<br />
lowest values <strong>of</strong> Na/K under salinity conditions (Figure 2).