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
Create successful ePaper yourself
Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.
Salt <strong>Stress</strong><br />
61<br />
A significant number <strong>of</strong> halophytes are C 4<br />
species, which are characterized by<br />
their higher requirements for sodium ions compared with C 3<br />
species (Brownell <strong>and</strong><br />
Crossl<strong>and</strong>, 1972). In conditions <strong>of</strong> osmotic stress <strong>and</strong> high temperatures, C 4<br />
plants have<br />
an advantage in comparison with C 3<br />
plants, because <strong>of</strong> their ability to carry on photosynthesis<br />
when stomata are to a large extent closed, coupled with the absence <strong>of</strong><br />
photorespiration in the mesophyll cells (Larcher, 1995). Photosynthetic responses to<br />
salinity in the halophytic tribe Salsoleae (family Chenopodiaceae) have been reviewed,<br />
with particular attention paid to relations between the C4 NAD-ME (malic enzyme)<br />
Salsoloid type <strong>of</strong> carboxylation <strong>and</strong> the chloroplast structure (Voznesenskaya et al.,<br />
1999).<br />
Abscisic acid is well recognized as an important stress hormone. The concentration<br />
<strong>of</strong> ABA increases when water deficits occur, with its de novo synthesis beginning<br />
in the roots, in response to sensing an insufficient supply <strong>of</strong> water (Zhang et al.,<br />
1989). In halophytes, which grow in conditions <strong>of</strong> “physiological drought”, due to low<br />
water potential in the root medium, the lowest concentrations <strong>of</strong> ABA were found<br />
under salinity concentrations optimum for growth (Clipson et al., 1988). Such case was<br />
reported for the highly tolerant halophytic species Suaeda maritima, which exhibited<br />
the lowest seasonal range <strong>of</strong> ABA contents (from 649.4 ng g -1 to 835.6 ng g -1 dry<br />
weight) in comparison with several other species, where higher ABA concentrations<br />
were correlated with increased sodium content <strong>of</strong> the shoot (Dajic et al., 1997a).<br />
In glycophytes, salinity leads to the accumulation <strong>of</strong> ABA (Asch et al., 1995),<br />
as in tomato (Chen <strong>and</strong> Plant, 1999; Yurekli et al., 2001) <strong>and</strong> wheat (Aldesuquy <strong>and</strong><br />
Ibrahim, 2002). In bush bean plants exposed to 75 mM NaCl, inhibition <strong>of</strong> leaf expansion<br />
was mediated by ABA rather than by Na + or Cl - toxicity, <strong>and</strong> the increase <strong>of</strong> ABA<br />
induced by a salt-pretreatment limited the accumulation <strong>of</strong> Na + <strong>and</strong> Cl - in the leaves,<br />
resulting in adaptation to salinity stress (Montero et al., 1997). Besides the significant<br />
role <strong>of</strong> ABA, favorable effects <strong>of</strong> other hormones in plant responses to salinity, such as<br />
cytokinins (Kuiper <strong>and</strong> Steingrover, 1991) <strong>and</strong> gibberellins (Kaur et al., 1998; Ashraf et<br />
al., 2002) have been documented.<br />
8. MOLECULAR BASIS OF SALT TOLERANCE<br />
According to Hasegawa et al. (2000) determinants <strong>of</strong> salt stress tolerance include effector<br />
molecules that enable adaptive reactions <strong>and</strong> mechanisms <strong>of</strong> plants in saline environments<br />
<strong>and</strong> regulatory molecules that control these pathways. Effectors are proteins<br />
<strong>and</strong> metabolites involved in ion homeostasis (membrane proteins involved in regulation<br />
<strong>of</strong> ionic transport), osmotic adjustment <strong>and</strong> water regime regulation (osmolytes)<br />
<strong>and</strong> toxic radical scavenging (mainly enzymes), while regulatory molecules are cellular<br />
signal pathway components <strong>and</strong> transducers <strong>of</strong> long-distance response coordination<br />
(hormones, mediators, transcription factors <strong>and</strong> regulatory genes).