Maize Membrane-Bound Transcription Factor Zmbzip17 is a Key Regulator in the Cross-Talk of ER Quality Control and ABA Signaling

Abiotic stresses disrupt protein folding and induce endoplasmic reticulum (ER) stress, which in turn activates the unfolded protein response (UPR) to aid in the refolding or degradation of misfolded proteins. The phytohormone ABA regulates many aspects of plant development and plays a central role in the stress response; however, the role of ABA in transducing stress signals to activate the UPR has not been recognized. In this study, a gene encoding the maize ortholog of AtbZIP17, a transmembrane transcription factor functioning as an ER stress transducer, was identified from the MaizeGDB database, and designated ZmbZIP17. ZmbZIP17 was induced by both ABA and ER stress-eliciting agents such as dithiotreitol (DTT) and tunicamycin (TM). Transiently expressed yellow fluorescent protein (YFP)-ZmbZIP17 co-localized with the ER marker HDEL-mCherry under control conditions, but partially translocated into the nucleus upon TM treatment or removal of the transmembrane domain. TM-induced processing of ZmbZIP17 was confirmed by Western blot analysis. When overexpressed in Arabidopsis, ZmbZIP17 triggered ER stress response gene expression and tolerance to DTT and TM, elevated ABA-responsive gene expression and ABA sensitivity both pre- and post-germination. Additionally, ABA was found to induce ER stress response gene expression, alone or synergistically with ZmbZIP17, in the absence of DTT or TM; while ZmbZIP17 was capable of interacting with ABA-responsive cis-elements (ABREs) that exist in promoters of known ABA-responsive genes. Together, our results reveal a direct link between ER stress and ABA signaling pathways involving the ZmbZIP17 transcription factor.