In the physiological sense, germination begins with seed water uptake
and ends with the initiation of elongation by the embryonic axis, usua
lly the radicle. The driving forces and constraints on expansion by th
e embryo are examined, particularly for seeds in which the embryo is s
urrounded by endosperm and testa tissues that restrict growth. Models
have been developed to predict germination based on thermal time, hydr
otime and combined hydrothermal time. These population-based models in
dicate that the timing of germination is closely tied to physiological
ly determined temperature and water potential thresholds for radicle e
mergence which vary among individual seeds in a population. The restra
int imposed by tissues surrounding the radicle is a major determinant
of the threshold water potential. Enzymatic weakening of these tissues
is a key event regulating the timing of radicle emergence. Considerab
le evidence suggests that endo-beta-mannanase is involved in this proc
ess in a number of species, although it is doubtful that it is the sol
e determinant of when radicle emergence occurs. Molecular and biochemi
cal studies are revealing the complexity of events occurring in endosp
erm and embryo cells associated with the completion of germination. Un
ique permeability properties and the presence of enzymes associated wi
th pathogen resistance suggest additional functional roles for the tis
sues enclosing the embryo. The insights gained from physiology and mod
elling are being extended by the application of molecular techniques t
o identify and determine the function of genes expressed in associatio
n with germination. Single-seed assay methods, in vivo reporters, spec
ific modification of gene expression and mutagenesis will be critical
technologies for advancing our understanding of germination.