Melatonin antagonizes ABA action to promote seed germination by regulating Ca2+ efflux and H2O2 accumulation

Seed germination is a vital stage in the plant life-cycle that greatly contributes to plant establishment. Melatonin has been shown to promote seed germination under various environmental stresses; however, the mechanism remains largely underexplored. Here, we reported that melatonin antagonized abscisic acid (ABA) to promote seed germination by regulating ABA and gibberellic acid (GA3) balance. Transcriptomic analysis revealed that such a role of melatonin was associated with Ca2+ and redox signaling. Melatonin pretreatment induced Ca2+ efflux accompanied by an up-regulation of vacuolar H+/Ca2+ antiporter 3 (CAX3). AtCAX3 deletion in Arabidopsis exhibited reduced Ca2+ efflux. Inhibition of Ca2+ efflux in the seeds of melon and Arabidopsis mutant AtCAX3 compromised melatonin-induced germination under ABA stress. Melatonin increased H2O2 accumulation, and H2O2 pretreatment decreased ABA/GA3 ratio and promoted seed germination under ABA stress. However, complete inhibition of H2O2 accumulation abolished melatonin-induced ABA and GA3 balance and seed germination. Our study reveals a novel regulatory mechanism in which melatonin counteracts ABA to induce seed germination that essentially involves CAX3-mediated Ca2+ efflux and H2O2 accumulation, which, in turn, regulate ABA and GA3 balance by promoting ABA catabolism and/or GA3 biosynthesis.