INVITED REVIEW Functions and regulation of -1,3-glucanases during seed germination, dormancy release and after-ripening

-1,3-Glucanase (Glu) expression in seeds plays important roles in the regulation of seed germination, dormancy and in the defence against seed pathogens. A thick -1,3-glucan layer is typical for the seed envelope of cucurbitaceous species, confers seed semipermeability and is degraded during germination. In many species with coat-imposed dormancy, the seed envelope confers a physical constraint to radicle emergence. In the solanaceous species, the micropylar endosperm and testa have this function, and endosperm weakening appears to be a prerequisite for germination. Class I Glu is transcriptionally induced in the micropylar endosperm of tobacco, tomato and other solanaceous seeds just prior to radicle emergence. Glu induction and germination are tightly linked in response to plant hormones and environmental factors, e.g. they are both promoted by gibberellins and inhibited by abscisic acid (ABA). Sense and antisense transformation of tobacco reveals two sites of Glu action: after-ripening-mediated release of testa-imposed dormancy and endosperm rupture during germination. The use of an ABA-inducible chimeric sense-transgene resulted in overexpression of class I Glu in seeds and provided direct evidence that Glu contributes to endosperm rupture. A model integrating Glu, seed dormancy, after-ripening and germination is presented, and possible mechanisms for Glu action are discussed. It is proposed that Glu not only helps defend seeds against pathogens, but is also a key factor in regulating coat-imposed dormancy and germination of seeds in response to environmental and hormonal cues