The SnRK2.2-ZmHsf28-JAZ14/17 module regulates drought tolerance in maize

Abscisic acid (ABA) and jasmonic acid (JA) are important plant hormones in response to drought stress. We have identified that ZmHsf28 elevated ABA and JA accumulation to confer drought tolerance in maize; however, the underlying mechanism still remains elusive. The knockout line zmhsf28 is generated to confirm the positive role of ZmHsf28 in drought response. Multiple approaches are combined to reveal protein interaction among ZmHsf28, ZmSnRK2.2 and ZmJAZ14/17, which form a regulatory module to mediate maize drought tolerance through regulating ABA and JA key biosynthetic genes ZmNCED3 and ZmLOX8. Upon drought stress, zmhsf28 plants exhibit weaker tolerance than the WT plants with slower stomatal closure and more reactive oxygen species accumulation. ZmHsf28 interacted with ZmSnRK2.2 physically, resulting in phosphorylation at Ser220, which enhances binding to the heat shock elements of ZmNECD3 and ZmLOX8 promoters and subsequent gene expression. Meanwhile, ZmMYC2 upregulates ZmHsf28 gene expression through acting on the G-box of its promoter. Besides, ZmJAZ14/17 competitively interact with ZmHsf28 to interfere with protein interaction between ZmHsf28 and ZmSnRK2.2, blocking ZmHsf28 phosphorylation and impairing downstream gene regulation. The ZmSnRK2.2-ZmHsf28-ZmJAZ14/17 module is identified to regulate drought tolerance through coordinating ABA and JA signaling, providing the insights for breeding to improve drought resistance in maize.