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Physiology and Molecular Biology of Stress ... - KHAM PHA MOI

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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).

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