In exo70B1-mediated autoimmunity, TIR-NBS2 associates with CPK5, likely stabilizing the enzyme to maintain kinase activity beyond the initiating calcium-mediate

Obligate biotrophs, such as the powdery mildew pathogens, deliver effectors to the host cell and obtain nutrients from the infection site. The interface between the plant host and the biotrophic pathogen thus represents a major battleground for plant-pathogen interactions. Increasing evidence shows that cellular trafficking plays an important role in plant immunity. Here, we report that Arabidopsis thaliana ENHANCED DISEASE RESISTANCE4 (EDR4) plays a negative role in resistance to powdery mildew and that the enhanced disease resistance in edr4 mutants requires salicylic acid signaling. EDR4 mainly localizes to the plasma membrane and endosomal compartments. Genetic analyses show that EDR4 and EDR1 function in the same genetic pathway. EDR1 and EDR4 accumulate at the penetration site of powdery mildew infection, and EDR4 physically interacts with EDR1, recruiting EDR1 to the fungal penetration site. In addition, EDR4 interacts with CLATHRIN HEAVY CHAIN2 (CHC2), and edr4 mutants show reduced endocytosis rates. Taken together, our data indicate that EDR4 associates with CHC2 and modulates plant immunity by regulating the relocation of EDR1 in Arabidopsis.

Many bacterial pathogens secret effectors into host cells to disable host defenses and thus promote infection. The exocyst complex functions in the transport and secretion of defense molecules, and loss of function of the EXO70B1 subunit leads to autoimmunity by activation of a truncated Toll/interleukin-1 receptor–nucleotide-binding sequence protein (TIR-NBS2; herein referred to as TN2). Here, we show that EXO70B1 is required for pathogen-associated molecular pattern-triggered immune responses in Arabidopsis thaliana. The effector AvrPtoB, an E3 ligase from Pseudomonas syringae pv. tomato (Pto) strain DC3000, associates with EXO70B1. AvrPtoB ubiquitinates EXO70B1 and mediates EXO70B1 degradation via the host’s 26S proteasome in a manner requiring E3 ligase activity. AvrPtoB enhances Pto DC3000 virulence by overcoming EXO70B1-mediated resistance. Moreover, overexpression of AvrPtoB in Arabidopsis leads to autoimmunity, which is partially dependent on TN2. Expression of TN2 in tobacco (Nicotiana tabacum and Nicotiana benthamiana) triggers strong and rapid cell death, which is suppressed by co-expression with EXO70B1 but reoccurs when co-expressed with AvrPtoB. Taken together, our data highlight that AvrPtoB targets the Arabidopsis thaliana EXO70 protein family member EXO70B1 to manipulate the defense molecule secretion machinery or immunity.