The MKK3 MAPK cascade integrates temperature and after-ripening signals to modulate seed germination

Temperature is a major environmental cue for seed germination. The permissive temperature range for germination is narrow in dormant seeds and expands during after-ripening. Quantitative trait loci analyses of pre-harvest sprouting in cereals have revealed that MKK3, a mitogen-activated protein kinase (MAPK) cascade protein, is a negative regulator of grain dormancy. Here we show that the MAPKKK19/20-MKK3-MPK1/2/7/14 cascade modulates germination temperature range in Arabidopsis seeds by elevating germinability of the seeds at sub- and supra-optimal temperatures. The expression of MAPKKK19 and MAPKKK20 is regulated by an unidentified temperature sensing and signaling mechanism the sensitivity of which is modulated during after-ripening of the seeds, and MPK7 is activated at the permissive temperature for germination regulated by expression levels of MAPKKK19/20. Activation of the MKK3 cascade represses abscisic acid (ABA) biosynthesis enzyme gene expression, and induces expression of ABA catabolic enzyme and gibberellic acid biosynthesis enzyme genes, resulting in expansion of the germinable temperature range. Our data demonstrate that the MKK3 cascade integrates temperature and after-ripening signals to germination processes including phytohormone metabolism.