Identification of novel pathogen-responsive cis-elements and their binding proteins in the promoter of OsWRKY13, a gene regulating rice disease resistance

The WRKY transcription factor superfamily controls diverse developmental and physiological processes in plants. However, little is known about the factors that directly regulate the function of WRKY genes. In this study, we identified cis-acting elements and their binding proteins of rice OsWRKY13, a gene that plays a pivotal role in disease resistance against bacterial and fungal pathogens. Two novel pathogen-responsive cis-elements, PRE2 and PRE4, were characterized from the promoter region of OsWRKY13. The two cis-elements negatively regulate gene expression without pathogen challenge, and positively regulate gene expression after pathogen-induced protein binding. OsWRKY13 binds to PRE4, which harbours a novel W-like box. Another five proteins (Rad51-like; tubby-like; SWIM zinc finger and nucleotide-binding adaptor shared by APAF-1, certain R proteins and CED-4 (NB-ARC) domain containing proteins; and an unknown protein) also bind to one of the two cis-elements. Different proteins interacting with the same cis-element appear to have different DNA-binding core sequences. These proteins localize in the nucleus and show differential expression upon pathogen challenge. These results suggest that OsWRKY13 expression is regulated by multiple factors to achieve disease resistance.