Salt tolerance and avoidance mechanisms at germination of annual pasture legumes: importance for adaptation to saline environments

Mechanisms for tolerance and avoidance of salinity at germination were examined in five self-regenerating annual pasture legumes of Mediterranean origin (Medicago polymorpha, Melilotus siculus, Trifolium subterraneum, T. michelianum and T. tomentosum). The maximum NaCl concentrations, for which no reduction in germination percentage occurred, were 300 mM for M. siculus, 240 mM for M. polymorpha and 120 mM for T. michelianum, T. subterraneum and T. tomentosum. Following 21 days exposure to 300 mM NaCl, imbibed seeds of T. subterraneum showed reduced germinability upon transfer to non-saline solution, whereas those of the four other species recovered full germinability. Following exposure to 600 mM NaCl, however, only T. michelianum and Melilotus siculus had some degree of recovery, with 38% and 31% germinability, respectively. Na⁺ in imbibed seed tissues of all species increased markedly with increasing NaCl concentration, while K⁺ decreased in all but Melilotus siculus. Seed coat impermeability ('hard seeds') acted as protection against the toxic effects of salinity. Melilotus siculus, Medicago polymorpha and T. tomentosum showed a delay in hard-seed breakdown ('seed softening') over the summer-autumn period, compared to T. subterraneum and T. michelianum, which acts to defer germination until soil surface salinity levels are likely to be lower. These three species also had higher levels of residual hard seeds. The results from this study support field observations that Melilotus siculus and Medicago polymorpha are the best adapted annual pasture legumes for highly saline soils in southern Australia, T. tomentosum and T. michelianum have some adaptation to moderately and mildly saline soils, respectively, whereas T. subterraneum has no adaptive traits for even mildly saline environments. A model for annual legume germination on saline land in southern Australia is also presented.