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In higher plants under low oxygen or hypoxic conditions, the phytohormone ethylene and hydrogen peroxide (H 2 O 2 ) are involved in complex regulatory mechanisms in hypoxia signaling pathways. The respiratory burst oxidase homolog D (RbohD), an NADPH oxidase, is involved in the primary stages of hypoxia signaling, modulating the expression of downstream hypoxia-inducible genes under hypoxic stress. In this study, our data revealed that under normoxic conditions, seed germination was delayed in the  rbohD / ein2-5  double mutant, whereas postgermination stage root growth was promoted. Under submergence, the  rbohD/ein2-5  double mutant line had an inhibited root growth phenotype. Furthermore, chlorophyll content and leaf survival were reduced in the  rbohD/ein2-5  double mutant compared with wild-type plants under submerged conditions. In quantitative RT-PCR analysis, the induction of  Ethylene-responsive factor 73 / hypoxia responsive 1  ( AtERF73/HRE1 ) and  alcohol dehydrogenase 1  ( AtADH1 ) transcripts was lower in the  rbohD/ein2-5  double mutant during hypoxic stress than in wild-type plants and in  rbohD  and  ein2-5  mutant lines. Taken together, our results indicate that an interplay of ethylene and RbohD is involved in regulating seed germination and post-germination stages under normoxic conditions. Moreover, ethylene and RbohD are involved in modulating seedling root growth, leaf chlorophyll content, and hypoxia-inducible gene expression under hypoxic conditions.

NADPH Oxidase RbohD and Ethylene Signaling are Involved in Modulating Seedling Growth and Survival Under Submergence Stress