Summary• Environmental conditions during seed maturation influence germination, but the genetic basis of maternal environmental effects on germination is virtually unknown.• Using single and multiple mutants of phytochromes, it is shown here that different phytochromes contributed to germination differently, depending on seed-maturation conditions.• Arabidopsis thaliana wild-type seeds that were matured under cool temperatures were intensely dormant compared with seeds matured at warmer temperature, and this dormancy was broken only after warm seed-stratification followed by cold seedstratification. The warm-cold stratification broke dormancy in fresh seeds but not in dry after-ripened seeds. Functional PHYB and PHYD were necessary to break cool-induced dormancy, which indicates a previously unknown and ecologically important function for PHYD. Disruption of PHYA in combination with PHYD (but not PHYB) restored germination to near wild-type levels, indicating that PHYA contributes to the maintenance of cool-induced dormancy on a phyD background. Effects of seed-maturation temperature were much stronger than effects of seed-maturation photoperiod. PHYB contributed to germination somewhat more strongly in seeds matured under short days, whereas PHYD contributed to germination somewhat more strongly in seeds matured under long days.• The variable contributions of different phytochromes to germination as a function of seed-maturation conditions reveal further functional diversification of the phytochromes during the process of germination. This study identifies among the first genes to be associated with maternal environmental effects on germination.

Summary• Germination timing is a fundamental life-history trait, as seedling establishment predicates realized fitness in the wild. Light and temperature are two important cues by which seeds sense the proper season of germination. Using Arabidopsis thaliana , we provide evidence that phytochrome-mediated germination pathways simultaneously respond to light and temperature cues in ways that affect germination.• Phytochrome mutant seeds were sown on agar plates and allowed to germinate in lit, growth chambers across a range of temperatures (7 ° C to 28 ° C).• phyA had an important role in promoting germination at warmer temperatures, phyE was important to germination at colder temperatures and phyB was important to germination across a range of temperatures.• Different phytochromes were required for germination at different temperatures, indicating a restriction or even a potential specialization of individual phytochrome activity as a function of temperature. This temperature-dependent activity of particular phytochromes reveals a potentially novel role for phytochrome pathways in regulating the seasonal timing of germination.

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We identified a new role of phytochrome in mediating germination responses to seasonal cues and thereby identified for the first time a gene involved in maternal environmental effects on germination. We examined the germination responses of a mutant, hy2-1, which is deficient in the phytochrome chromophore. The background genotype, Landsberg erecta (Ler), lacked dormancy in most treatments, while hy2-1 required cold stratification for germination in a manner that resembled a more dormant ecotype, Columbia (Col). Unlike Col, hy2-1 was not induced into dormancy by warm stratification. Therefore, the down-regulation of phytochrome-mediated germination pathways results in sensitivity to cold, but we found no evidence that reduced phytochrome activity enables the warm-induction of dormancy. Cool temperatures during seed maturation induced dormancy. The hy2-1 mutants did not overcome this dormancy, indicating that phytochrome-mediated pathways are required to break cold-induced dormancy. Ler did not respond to post-stratification temperature, but hy2-1 did respond, suggesting phytochrome pathways are involved in germination responses to temperature. In summary, phytochromes mediate dormancy and germination responses to seasonal cues experienced both during seed maturation and after dispersal. Phytochromes therefore appear to be involved in mediating seasonal germination timing, a trait of great ecological importance and one that is under strong natural selection.

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Among the most important environmental sensors in plants are the phytochromes, members of a duplicatedgene family of photoreceptors. We characterized the functional diversification of this gene family with respect to a single ecologically important and highly environmentally sensitive process-germination-and thereby addressed the consequence of gene duplication in the evolution of environmental sensitivity. We exposed single and multiple mutants of different phytochromes of Arabidopsis thaliana to dormancy-breaking or dormancyinducing treatments that resemble different seasonal environmental conditions, and we recorded their germination responses. Much redundancy of function occurred, but an impressive degree of functional diversification was also apparent. Specifically, PHYB was an important contributor to germination when seeds did not receive lowtemperature imbibition. It interacted nonadditively with PHYA, whose contribution to germination was redundant to that of PHYB and most apparent when seeds experienced high followed by low temperatures. We also determined that PHYD was necessary for full germination when seeds were exposed to a high temperature during imbibition. PHYE appears to contribute redundantly to germination, except when seeds experienced high followed by low temperatures. Finally, we found new evidence that PHYA and possibly PHYD redundantly contribute to the suppression of germination. The diversification of activities of all these phytochromes during this one process of germination suggests that they all have a combined function in the regulation of germination responses to complex seasonal conditions and that gene duplication can contribute to the evolution of precise responses to variable environments.