WRKY7, -11 and -17 Transcription Factors Are Modulators of the BZIP28 Branch of the Unfolded Protein Response During PAMP-triggered Immunity in Arabidopsis Thaliana

Plants must defend themselves against pathogens. The defense response requires greater protein synthesis, which generates endoplasmic reticulum (ER) stress, yet failure to attenuate this stress has detrimental effects. WRKY7/11/17 transcription factors (TFs) are negative regulators of immunity since mutants are more resistant to Pseudomonas syringae pv tomato (Pst) infection. Here, we reveal a connection between ER-stress and the molecular mechanisms underlying the wrky mutant phenotype. The bZIP28 TF upregulates ER-chaperone expression (BiP1/2, ERdj3B, and SDF2) upon exposure of Arabidopsis to a bacterial defense elicitor, flagellin 22 (Flg22). Also, the activation of ER-chaperones is more sustained in double and triple wrky mutants treated with Flg22, suggesting that WRKY7/11/17 TFs downregulate these genes. Moreover, wrky mutants accumulate more bZIP28 transcripts in response to Flg22, indicating that WRKY7/11/17 transcriptionally repress this TF. Using Arabidopsis protoplasts, we also demonstrate that WRKYs bind to the bZIP28 promoter via W-box elements. Additionally, triple wrky mutants are more resistant, whilst bzip28 mutants are more susceptible, to Pst infection. Finally, we postulate a model of PAMP-Triggered Immunity regulation, where Flg22 activates bZIP28-signaling inducing the expression of ER-stress genes, as well as WRKY7/11/17 expression, which in turn inhibits PTI by downregulating bZIP28, controlling physiological responses in the Arabidopsis-Pst interaction.