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

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Salt <strong>Stress</strong><br />

51<br />

(Echeverria, 2000). The efficacy <strong>of</strong> salt gl<strong>and</strong>s has been established by the presence <strong>of</strong><br />

low concentrations <strong>of</strong> salts in leaves <strong>and</strong> a high K/Na ratio <strong>of</strong> species possessing such<br />

structures, even in conditions <strong>of</strong> rapidly increasing external salinity (Bradley <strong>and</strong> Morris,<br />

1991).<br />

It seems that in salt-secreting plants the mechanism <strong>of</strong> salt exclusion is operating<br />

as well, since it was demonstrated that in mangrove plants Aegiceras corniculatum<br />

<strong>and</strong> Avicennia marina Na + excretion does not keep pace with Na + uptake over a range<br />

<strong>of</strong> salinities, reducing plant growth at high external salinity (Ball, 1988).<br />

Salt bladders, probably best studied in several species <strong>of</strong> the genus Atriplex<br />

(Chenopodiaceae), are also <strong>of</strong> particular importance in the regulation <strong>of</strong> salt tolerance.<br />

They are composed <strong>of</strong> two cells: the small, basal <strong>and</strong> the upper, bladder cell (Osmond et<br />

al., 1980). The latter dies, once it has accumulated a sufficient amount <strong>of</strong> salt in its<br />

vacuole, leaving a salt crust at the leaf surface (Figure 3).<br />

Figure 3. Leaf surface <strong>of</strong> the salt-excreting halophyte Atriplex tatarica L. var. diffusa Ten.<br />

(x 1000, Dajic, 1996)<br />

Many Atriplex species are characterized by the presence <strong>of</strong> salt bladders at the<br />

surface <strong>of</strong> young leaves only (Breckle et al., 1990), <strong>and</strong> these, in same cases, may<br />

contain almost the whole leaf sodium (Jeschke <strong>and</strong> Stelter, 1983). It is considered that<br />

the salt bladders <strong>of</strong> Atriplex represent a key adaptation to salt tolerance for their natural<br />

habitats (Freitas <strong>and</strong> Breckle, 1992).<br />

Altogether, the prevention <strong>of</strong> excessive salt accumulation within the plant body is<br />

achieved through the following modes, such as: a) control <strong>of</strong> the salt uptake at the root<br />

level <strong>and</strong> regulation <strong>of</strong> Na + delivery to the shoot by loading <strong>of</strong> the xylem <strong>and</strong> retrieval <strong>of</strong><br />

ions from the xylem before reaching the shoot), b) K/Na selectivity, c) recirculation <strong>of</strong><br />

salts via the phloem, d) allocation <strong>of</strong> salts within particular parts <strong>of</strong> the plant, e) ion<br />

leakage <strong>and</strong> abscission <strong>of</strong> organs loaded with salts, f) control <strong>of</strong> transpiration, <strong>and</strong> g)<br />

secretion <strong>of</strong> ions by salt-excreting structures.

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