WRKY70 and its homolog WRKY54 negatively modulate the cell wall-associated defenses to necrotrophic pathogens in Arabidopsis

Previous studies have identified the Arabidopsis thaliana transcription factor WRKY70 as a node of convergence for salicylic acid (SA) and jasmonic acid (JA)-mediated defense signal pathways and, together with its closest homolog WRKY54, as a negative regulator of SA biosynthesis. Here, we demonstrate that WRKY70 together with WRKY54 negatively affect the response of Arabidopsis to the necrotrophic pathogens Pectobacterium carotovorum and Botrytis cinerea, but not to the hemibiotroph Pseudomonas syringae pv tomato (Pst) DC3000, as revealed by mutants studies. Unstressed wrky54wrky70 double mutants exhibited increased levels of SA, accumulation of hydrogen peroxide (H2O2) and up-regulated expression of both SA and JA/ethylene (ET) responsive defense related genes. Additionally, protein cross-linking in cell wall was promoted by endogenous SA, suggesting involvement of wall-associated defenses against necrotrophs. This response to necrotrophs was compromised by introducing the sid2-1 allele impairing SA biosynthesis and leading to reduction of H2O2 content and of defense gene expression. The data suggest that the elevated SA level in the wrky54wrky70 double mutant results in moderate accumulation of H2O2, in promoting cell wall fortification and consequently enhanced resistance to necrotrophs but is not sufficient to trigger hypersensitive reaction (HR)-like cell death and resistance to biotrophs/hemibiotrophs like Pst DC3000.