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No.42 - 農業生物資源研究所

No.42 - 農業生物資源研究所

No.42 - 農業生物資源研究所

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2<br />

Yuji KAMIYA<br />

Fig. 1. Gibberellin biosynthesis in Arabidopsis.<br />

It has been known that GA promotes seed germination in many plant species. In Arabidopsis,<br />

severe GA deficient-mutants, such as ga1-3 and ga2-1, are defective in seed germination<br />

(KOORNNEEF and VAN DER VEEN 1980), and chemical inhibitors of GA biosynthesis inhibit<br />

germination (NAMBARA et al. 1991). These observations indicate that de novo GA biosynthesis is<br />

necessary for seed germination in Arabidopsis (HEDDEN and KAMIYA 1997).<br />

Light is a critical environmental factor for seed germination in some small-seeded plants such<br />

as lettuce, tomato and Arabidopsis (SHINOMURA, 1997). The effect of light on seed germination is<br />

primarily mediated by phytochromes (BORTHWICK et al. 1952; BUTLER et al. 1959). Genes<br />

encoding GA 3-oxidases, which convert inactive precursor to active GAs, are regulated by<br />

phytochromes in germinating lettuce and Arabidopsis seeds (TOYOMASU et al. 1998; YAMAGUCHI<br />

et al. 1998).<br />

Temperature is another crucial external cue that controls seed germination (BEWLEY and<br />

BLACK 1982). In many plant species, exposure of seeds to low temperature (typically 2-5)<br />

immediately after imbibition is effective to promote germination. (SHROPSHIRE et al. 1961; CONE<br />

and SPRUIT 1983). This treatment is called “stratification”. Although this method is widely used to<br />

improve the frequency and synchronization of germination, the mechanism for the thermoregulation<br />

of seed germination has been unclear.<br />

The effect of cold treatment on GA content has been reported in the 1970s, based on semiquantitative<br />

analysis of endogenous GAs using -amylase bioassay and/or gas chromatographymass<br />

spectrometry (GC-MS) (ROSS and BRADBEER 1971; SINSKA et al. 1973; WILLIAMS et al.<br />

1974). DERKX et al. (1994) analyzed the effect of pre-chilling of Arabidopsis seeds and reported<br />

that bioactive GA4 was detectable only in pre-chilled seeds, but not in dark-imbibed seeds. It has<br />

not been clear whether the effect of pre-chilling is a direct response to low temperature because pre-

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