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Ectopic expression of the Arabidopsis  RESISTANCE TO POWDERY MILDEW8.1  ( RPW8.1 ) boosts pattern-triggered immunity leading to enhanced resistance to different pathogens in Arabidopsis and rice. However, the underlying regulatory mechanism remains largely elusive. Here, we report that  XAP5 CIRCADIAN TIMEKEEPER  ( XCT, At2g21150 ) positively regulates  RPW8.1 -mediated cell death and disease resistance. Forward genetic screen identified the  b3-17  mutant that exhibited less cell death and susceptibility to powdery mildew and bacterial pathogens. Map-based cloning identified a G-to-A point mutation at the 3′ splice site of the 8th intron, which resulted in splice shift to 8-bp down-stream of the original splice site of  XCT  in  b3-17 , and introduced into a stop codon after two codons leading to a truncated XCT.  XCT  has previously been identified as a circadian clock gene required for small RNA biogenesis and acting down-stream of  ETHYLENE-INSENSITIVE3  ( EIN3 ) in the ethylene-signaling pathway. Here we further showed that mutation or down-regulation of  XCT  by artificial microRNA reduced  RPW8.1 -mediated immunity in R1Y4, a transgenic line expressing  RPW8.1-YFP  from the  RPW8.1  native promoter. On the contrary, overexpression of  XCT  in R1Y4 background enhanced  RPW8.1 -mediated cell death, H 2 O 2  production and resistance against powdery mildew. Consistently, the expression of  RPW8.1  was down- and up-regulated in  xct  mutant and  XCT  overexpression lines, respectively. Taken together, these results indicate that  XCT  positively regulates  RPW8.1 -mediated cell death and disease resistance, and provide new insight into the regulatory mechanism of  RPW8.1 -mediated immunity.

XAP5 CIRCADIAN TIMEKEEPER Positively Regulates RESISTANCE TO POWDERY MILDEW8.1–Mediated Immunity in Arabidopsis