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

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74<br />

Z . Dajic<br />

The small molecules designated as transcription factors bind to promoter regulatory<br />

elements in salt-inducible genes, whose role is associated with the transcriptional<br />

activation <strong>of</strong> genes regulated by salt <strong>and</strong> drought stress (Shinozaki <strong>and</strong> Yamaguchi-<br />

Shinozaki, 1997). An altered gene expression in salt-tolerant alfalfa seems to be connected<br />

with the involvement <strong>of</strong> a putative transcriptional factor Alfin1, while<br />

overexpression <strong>of</strong> the Alfin1 in transgenic alfalfa led to the activation <strong>of</strong> an additional<br />

salt-inducible gene, the MsPRP2 (Winicov, 1998), suggesting a role <strong>of</strong> transcription<br />

factors in coordinate gene regulation under salinity conditions.<br />

Salinity tolerance has been considered to be a quantitative trait (Foolad <strong>and</strong><br />

Jones, 1993). Hence, the identification <strong>of</strong> quantitative trait loci (QTL) that contribute to<br />

natural variation in responses to salt stress will be helpful in underst<strong>and</strong>ing the complexity<br />

<strong>of</strong> the genetic control <strong>of</strong> salt tolerance <strong>and</strong> provide opportunities for more efficient<br />

breeding for salt tolerance in crops. In the study <strong>of</strong> quantitative trait loci controlling<br />

various traits <strong>of</strong> rice seedlings exposed to salinity stress, out <strong>of</strong> a total <strong>of</strong> seven<br />

QTLs for traits associated with improved tolerance, four were located on chromosome<br />

6 (Prasad et al., 2000a). It was reported that QTLs for sodium <strong>and</strong> potassium uptake in<br />

rice were found on different chromosomes (Koyama et al., 2001), which is consistent<br />

with the independent inheritance <strong>of</strong> Na + <strong>and</strong> K + uptake in the mapping populations, <strong>and</strong><br />

with different uptake pathways (apoplastic leakage <strong>and</strong> membrane transport, respectively)<br />

for these cations under saline conditions. In rice, the putative locus for osmotic<br />

adjustment <strong>and</strong> two <strong>of</strong> five QTL associated with dehydration tolerance were close to<br />

chromosomal regions responsible for the root morphology (Lilley et al., 1996).<br />

Identification <strong>of</strong> QTLs for salt tolerance during vegetative growth in tomato<br />

led to detection <strong>of</strong> five genomic regions on chromosomes 1, 3, 5, 6, <strong>and</strong> 11 (Foolad et al.,<br />

2001). In Arabidopsis, 11 QTLs were detected, for variation in salt tolerance, <strong>of</strong> which 6<br />

were present during germination <strong>and</strong> 5 at the vegetative growth stage (Quesada et al.,<br />

2002). A survey <strong>of</strong> QTLs for salt tolerance in barley seedlings revealed 12 quantitative<br />

trait loci for seven traits (Ellis et al., 2002). A composite genomic map for the Triticeae <strong>of</strong><br />

QTLs affecting stress tolerance showed the highest concentration <strong>of</strong> QTLs <strong>and</strong> major<br />

loci controlling the plant’s adaptation to environmental stresses located on the chromosome<br />

group 5 (Cattivelli et al., 2002). The QTL analysis performed with 172 recombinant<br />

hybrids between Helianthus annuus <strong>and</strong> H. petiolaris, revealed a total <strong>of</strong> 14 QTLs<br />

for mineral uptake <strong>and</strong> three for survival, suggesting that salt tolerance in Helianthus is<br />

achieved through increased calcium uptake, coupled with greater sodium exclusion<br />

(Lexer et al., 2003). It was reported for several species that QTLs linked to salt tolerance<br />

vary with the developmental stage <strong>of</strong> the plant (Flowers, 2004).<br />

Significant progress in highlighting the physiological <strong>and</strong> biochemical responses<br />

<strong>of</strong> plants to salinity at different levels, as well as the molecular cloning <strong>of</strong><br />

genes involved in the different metabolic pathways altered by salts, has enabled the<br />

introduction <strong>of</strong> genetic engineering technologies in order to improve salt tolerance in<br />

crops.

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