Phytochrome-induced softening of the micropylar endosperm of Datura ferox seeds is followed by radicle protrusion and preceded by a significant increase in the extractable activity of β-mannanase and β-mannosidase (6 and 3 times over the far-red light controls, respectively) and a decrease in the mannan content of the micropylar endosperm cell walls. This relationship between phytochrome-induced germination, endosperm softening and mannan degradation was studied in Datura ferox seeds subjected to a range of water potentials during the presence of Pfr. Low water potential inhibited the decrease in mannose-rich cell-wall insoluble polysaccharides, the increase in activity of β-mannosidase and endosperm softening. A good correlation was found between activity of β-mannosidase measured at 45 h after exposure to a red light pulse and germination counted 24 h later when, with different external water potentials, germination varied between 0 and 90%. In contrast, no inhibitory effect of low water potential on in vitro β-mannanase activity was detected. We suggest that the in vivo action of β-mannanase might be limited by accumulation of manno-oligosaccharides caused by inhibition of β-mannosidase. These data are consistent with the concept of a link between mannan mobilization and endosperm softening. Part of the effect of low water potential on Pfr-induced germination may be mediated by inhibition of β-mannosidase activity, with consequences for mannan hydrolysis and endosperm softening.