For the transformation of a seed to a seedling complex physical and biochemical changes occur within a seed before germination can proceed. Germination is controlled by diverse seed dormancy mechanisms in plant species that delays germination until the conditions are most favourable for seed germination and seedling establishment (Thompson 1970). Baskin and Baskin (1998) identified four benefits for the evolution of seed dormancy in plants: (i) persistence in risky environments as seed banks, (ii) decreased intraspecific competition, (iii) improved chances of seedling establishment and (iv) increased fitness (seed production) of the individual and the species as a whole. They showed that seed dormancy may be caused by any one of physiological, morphological, physical, chemical and mechanical constraints or by a combination of more than one of these factors. For instance, seeds may possess an embryo with a physiological inhibiting mechanism, immature embryo, impermeable seed coat or may contain chemical inhibitors and hard woody fruit walls. In all of these cases seed dormancy is eventually broken by one or more of the following treatments: after ripening, heat treatment, cold temperature stratification, prolonged exposure to high temperatures, exposure to light, softening of seed coat by microbes or physical scarification, leaching of inhibiting chemicals, ageing of seeds and other subtle changes in the habitat. In temperate North America with snow cover during winter months the seeds of a large majority of sand dune species—Cakile edentula, Ammophila breviligulata, Calamovilfa longifolia, Iva imbricata, Croton punctatus, Uniola paniculata—and others require cold stratification at <4°C for 4–6 weeks to break their dormancy requirements. Seeds of some species such as A. breviligulata and U. paniculata that require cold stratification at the northern end of their range lose this requirement in the south (Seneca 1972). At southern locations exposure to high temperatures may be required to fulfil the dormancy requirements. Winter annuals, Vulpia ciliata, Cerastium atrovirens, Mibora minima and Saxifraga tridactylites, that grow and mature their seeds in early summer on sand dunes at Aberffraw, North Wales, require exposure to high soil temperatures to overcome a state of dormancy in a certain proportion of seeds at the time of dispersal (Carey and Watkinson 1993; Pemadasa and Lovell 1975).
Effects of cold stratification on the endogenous hormone, dormancy and germination of Cornus walteri Wanger seeds
