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Literature review<br />
oligosaccharides (RAGHAVAN, 1976; BEWLEY & BLACK, 1994). Most seeds store<br />
their major food reserves within the embryo; usually the cotyledons (BEWLEY &<br />
BLACK, 1994). Some plants also have their seed storage reserves within extra-<br />
embryonic tissues. These extra-embryonic tissues used for storage include the<br />
endosperm (Gymnosperms) or the perisperm (C<strong>of</strong>fea arabica). Both embryonic and<br />
extra-embryonic tissues can also be used for storage; such as in maize (BEWLEY &<br />
BLACK, 1994).<br />
2.8.2 Seed germination<br />
Germination starts with the uptake <strong>of</strong> water by a seed and ends with the onset <strong>of</strong><br />
elongation <strong>of</strong> the embryonic axis, usually the radicle (BEWLEY & BLACK, 1994). It<br />
also includes the steps <strong>of</strong> protein hydration, sub cellular changes, respiration,<br />
macromolecular syntheses and cell elongation (RAGHAVAN, 1976; BEWLEY &<br />
BLACK, 1994). The combined effect <strong>of</strong> these steps is to transform a dehydrated,<br />
dormant embryo into an embryo which grows actively and accumulates water<br />
(MAYER, 1977; BEWLEY & BLACK, 1994). A seed in which none <strong>of</strong> these processes<br />
have taken place is said to be quiescent. They characteristically have low moisture<br />
content (5-15%) and an extremely slow metabolic rate (BEWLEY & BLACK, 1994).<br />
Seeds are able to survive in this state for a number <strong>of</strong> years. Quiescent seeds<br />
require an environment <strong>of</strong> suitable temperature, hydration and the presence <strong>of</strong><br />
oxygen in order to germinate (BEWLEY & BLACK, 1994). The major cellular<br />
processes involved in the initiating and facilitating <strong>of</strong> radicle emergence include<br />
respiration, RNA and protein synthesis and enzyme and organelle activity<br />
(RAGHAVAN, 1976; BEWLEY & BLACK, 1994).<br />
During imbibition various structural and physical changes occur (BEWLEY & BLACK,<br />
1994). The completion <strong>of</strong> imbibition requires a small amount <strong>of</strong> water (not more than<br />
three times the seed’s dry weight). For successful subsequent root and shoot growth<br />
a larger and more constant supply <strong>of</strong> water is essential (BEWLEY & BLACK, 1994).<br />
The uptake <strong>of</strong> water by the seed can be seen as triphasic (Figure 2.31) (BEWLEY &<br />
BLACK, 1994). Phase 1 involves imbibition, where a strong osmotic gradient results<br />
in the uptake <strong>of</strong> water from the soil (BEWLEY & BLACK, 1994). Different areas <strong>of</strong> the<br />
testa and different organs within the seed <strong>of</strong>ten absorb variable amounts <strong>of</strong> water.<br />
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