Seed dormancy can prevent germination under unfavourable conditions that reduce the chances of seedling survival. Freshly harvested seeds often have strong primary dormancy that depends on the temperature experienced by the maternal plant and which is gradually released through afterripening. However, seeds can be induced into secondary dormancy if they experience conditions or cues of future unfavourable conditions. Whether this secondary dormancy induction is influenced by seed-maturation conditions and primary dormancy has not been explored in depth. In this study, we examined secondary dormancy induction in seeds of Arabidopsis thaliana matured under different temperatures and with different levels of afterripening. We found that low water potential and a range of temperatures, from 8°C to 35°C, induced secondary dormancy. Secondary dormancy induction was affected by the state of primary dormancy of the seeds. Specifically, afterripening had a non-monotonic effect on the ability to be induced into secondary dormancy by stratification; first increasing in sensitivity as afterripening proceeded, then declining in sensitivity after 5 months of afterripening, finally increasing again by 18 months of afterripening. Seed-maturation temperature sometimes had effects that were independent of expressed primary dormancy, such that seeds that had matured at low temperature, but which had comparable germination proportions as seeds matured at warmer temperatures, were more easily induced into secondary dormancy. Because seed-maturation temperature is a cue of when seeds were matured and dispersed, these results suggest that the interaction of seed-maturation temperature, afterripening and post-dispersal conditions all combine to regulate the time of year of seed germination.

Different life stages frequently respond to the same environmental cue to regulate development so that each life stage is matched to its appropriate season. We investigated how independently each life stage can respond to shared environmental cues, focusing on vernalization, in Arabidopsis thaliana plants. We first tested whether effects of rosette vernalization persisted to influence seed germination. To test whether genes in the vernalization flowering pathway also influence germination, we assessed germination of functional and nonfunctional alleles of these genes and measured their level of expression at different life stages in response to rosette vernalization. Rosette vernalization increased seed germination in diverse ecotypes. Genes in the vernalization flowering pathway also influenced seed germination. In the Columbia accession, functional alleles of most of these genes opposed the germination response observed in the ecotypes. Some genes influenced germination in a manner consistent with their known effects on FLOWERING LOCUS C gene regulation during the transition to flowering. Others did not, suggesting functional divergence across life stages. Despite persistent effects of environmental conditions across life stages, and despite pleiotropy of genes that affect both flowering and germination, the function of these genes can differ across life stages, potentially mitigating pleiotropic constraints and enabling independent environmental regulation of different life stages.

FLOWERING LOCUS C (FLC) has a major regulatory role in the timing of flowering in Arabidopsis thaliana (L.) Heynh. and has more recently been shown to influence germination. Here, we investigated the conditions under which FLC influences germination, and demonstrated that its effect depends on the level of primary and secondary dormancy and the temperature of seed imbibition. We tested the germination response of genotypes with different degrees of FLC activity over the course of after-ripening and after secondary dormancy induction by hot stratification. Genotypes with high FLC-activity showed higher germination; this response was greatest when seeds exhibited primary dormancy or were induced into secondary dormancy by hot stratification. In this study, which used less dormant seeds, the effect of FLC was more evident at 22°C, the less permissive germination temperature, than at 10°C, in contrast to prior published results that used more dormant seeds. Thus, because effects of FLC variation depend on dormancy, and because the range of temperature that permits germination also depends on dormancy, the temperature at which FLC affects germination can also vary with dormancy. Finally, we document that the effect of FLC can depend on FRIGIDA and that FRIGIDA itself appears to influence germination. Thus, pleiotropy between germination and flowering pathways in A. thaliana extends beyond FLC and involves other genes in the FLC genetic pathway.

Despite some degree of functional divergence between the regulation of flowering and germination by autonomous-pathway genes, the autonomous pathway is highly functionally conserved across life stages. Therefore, genes in the autonomous flowering-time pathway are likely to contribute to genetic correlations between flowering and seed germination, possibly contributing to the winter-annual life history.

Acquisition of secondary dormancy enables spatio-temporal control of germination. Evidence suggests that PHYD influences secondary dormancy induction by high temperature in Arabidopsis and correlates with removal of the germination repressor PIL5.