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

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Water <strong>Stress</strong><br />

29<br />

include water <strong>and</strong> ion channels, enzymes necessary for the formation <strong>of</strong> compatible<br />

solutes, molecular chaperons, late embryogenesis abundant (LEA) proteins, proteins<br />

involved in protein degradation, <strong>and</strong> reactive oxygen-scavenging enzymes.<br />

Figure 3. Transcriptional factors <strong>and</strong> cis-acting elements involved in water stressresponsive<br />

gene expression. Transcriptional factors <strong>and</strong> cis-acting elements are shown in<br />

oval <strong>and</strong> square boxes, respectively. Abbreviations: ABA, abscisic acid; AREB, ABREbinding<br />

proteins; ABRE, ABA-responsive element; CE, coupling element; MYBR, MYB<br />

recognition site; MYC, MYC recognition site; DREB, DRE-binding protein; CBF, C-<br />

repeat-binding factor; DRE/CRT, dehydration-responsive element/C-repeat; LEA, late<br />

embryogenesis abundant protein<br />

Expression <strong>of</strong> many genes for transcription factors is up-regulated under<br />

drought stress conditions. These transcriptional factors include the dehydration-responsible<br />

element (DRE)/C-repeat (CRT)-binding (DREB/CBF) protein family, ethyleneresponsible<br />

element binding factor (ERF), zinc-finger family, WRKY family, basic helixloop-helix<br />

(bHLH) family, basic-domain leucine zipper (bZIP) family, NAC family, <strong>and</strong><br />

homeodomain transcription family. Drought stress also induces expression <strong>of</strong> proteins<br />

functioning in phosphorylation signaling cascades such as MAP kinase, calcium-dependent<br />

kinase, receptor kinase <strong>and</strong> histidine kinase, <strong>and</strong> in phospholipid metabolism<br />

(Zhu, 2002).<br />

Drought-responsive genes can be divided into two groups, ABA-dependent<br />

<strong>and</strong> ABA-independent genes, according to their dependency on ABA for induction<br />

(Yamaguchi-Shinozaki <strong>and</strong> Shinozaki, 2005). The ABA-responsive element (ABRE) is a<br />

major cis-sequence in the promoter <strong>of</strong> many ABA-responsive genes. The cis-element<br />

was first identified in the Em gene <strong>of</strong> wheat <strong>and</strong> has since been found in various plants<br />

such as maize, oat, rice, tobacco <strong>and</strong> Arabidopsis where it functions as an ABRE.<br />

Expression <strong>of</strong> ABA-responsive genes is activated through binding <strong>of</strong> AREB/ABF, a<br />

bZIP-type transcription factor, to the ABRE sequence (Uno et al., 2000; Choi et al.,<br />

2000).

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