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Plants under pathogen attack produce high levels of ethylene, which plays important roles in plant immunity. Previously, we reported the involvement of ACS2 and ACS6, two Type I ACS isoforms, in  Botrytis cinerea –induced ethylene biosynthesis and their regulation at the protein stability level by MPK3 and MPK6, two Arabidopsis pathogen-responsive mitogen-activated protein kinases (MAPKs). The residual ethylene induction in the  acs2/acs6  double mutant suggests the involvement of additional ACS isoforms. It is also known that a subset of  ACS  genes, including  ACS6 , is transcriptionally induced in plants under stress or pathogen attack. However, the importance of  ACS  gene activation and the regulatory mechanism(s) are not clear. In this report, we demonstrate using genetic analysis that ACS7 and ACS11, two Type III ACS isoforms, and ACS8, a Type II ACS isoform, also contribute to the  B. cinerea –induced ethylene production. In addition to post-translational regulation, transcriptional activation of the  ACS  genes also plays a critical role in sustaining high levels of ethylene induction. Interestingly, MPK3 and MPK6 not only control the stability of ACS2 and ACS6 proteins via direct protein phosphorylation but also regulate the expression of  ACS2  and  ACS6  genes. WRKY33, another MPK3/MPK6 substrate, is involved in the MPK3/MPK6-induced  ACS2/ACS6  gene expression based on genetic analyses. Furthermore, chromatin-immunoprecipitation assay reveals the direct binding of WRKY33 to the W-boxes in the promoters of  ACS2  and  ACS6  genes  in vivo , suggesting that WRKY33 is directly involved in the activation of  ACS2  and  ACS6  expression downstream of MPK3/MPK6 cascade in response to pathogen invasion. Regulation of ACS activity by MPK3/MPK6 at both transcriptional and protein stability levels plays a key role in determining the kinetics and magnitude of ethylene induction.

plos genetics

Dual-Level Regulation of ACC Synthase Activity by MPK3/MPK6 Cascade and Its Downstream WRKY Transcription Factor during Ethylene Induction in Arabidopsis