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Literature review<br />

endogenous gibberellins increase during low temperature treatments (JONES &<br />

STODDART, 1977).<br />

Gibberellins (GA) promote the induction <strong>of</strong> cell wall hydrolases and thereby promote<br />

endosperm weakening and endosperm rupture (MARION-POLL, 1997). Abscisic acid<br />

(ABA) inhibits the induction <strong>of</strong> cell wall hydrolases and thereby inhibits this<br />

weakening and rupture (MARION-POLL, 1997).<br />

GA promotes and ABA inhibits the embryo growth potential. ABA, however, also<br />

plays an important role in seed development and germination, acquisition <strong>of</strong><br />

desiccation tolerance, accumulation <strong>of</strong> proteins and lipid reserves, and induction and<br />

maintenance <strong>of</strong> seed dormancy (MARION-POLL, 1997).<br />

The expression <strong>of</strong> genes encoding enzymes that mobilise food reserves is induced<br />

by several known GA signalling factors. These food reserves include the starches,<br />

proteins and lipids that are stored in the endosperm (PENG & HARBERD, 2002).<br />

During protein rehydration GA biosynthesis is induced by a phytochrome-mediated<br />

light signal and the newly synthesised GA down-regulates the expression <strong>of</strong> protein<br />

repressors <strong>of</strong> germination. These are also activated by rehydration, by protein<br />

degradation, suppression <strong>of</strong> transcription or mRNA degradation (MARION-POLL,<br />

1997). This newly synthesized GA also initiates signals to induce the expression <strong>of</strong><br />

hydrolytic enzymes that modify the cell wall and weaken the endosperm cap, thus<br />

facilitating germination (PENG & HARBERD, 2002).<br />

It has also been proposed that GA could promote the formation <strong>of</strong> low molecular<br />

weight mono- and disaccharides, which assist the intracellular generation <strong>of</strong> negative<br />

water potentials, thus aiding radicle emergence (TIAN et al., 2003).<br />

2.8.4.3 Cytokinins and auxins<br />

Cytokinins are involved in cell division and thus both radicle and cotyledon expansion<br />

(EMERY & ATKINS, 2006). Cytokinins accumulates mainly in the endosperm <strong>of</strong><br />

seeds (LEUBNER-METZGER, 2006). Auxins are involved in the coordination <strong>of</strong><br />

correct cellular patterning after the globular stage in embryogenesis (LEUBNER-<br />

METZGER, 2006).<br />

48

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