To protect themselves from infectious diseases,plants have de-ployed a two-tiered surveillance system to recognize invading pathogens(Fu and Dong,2013).In the first layer of the immune system,plants evolved extracellular pattern recognition receptors to detect conserved pathogen-associated molecular patterns(PAMPs)and activate PAMP-triggered immunity(PTI)(Boller and Felix,2009).The second layer of the immune system,which is termed effector-triggered immunity(ETI),relies on an array of nucleotide-binding leucine-rich repeat receptor(NLR)proteins to detect pathogen effectors,either directly or the activities of these effectors indirectly(Figure 1A)(Sun et al.,2020).NLR activation often triggers a rapid localized cell death called hypersensitive response(HR).