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Secreted papain-like Cys proteases are important players in plant immunity. We previously reported that the C14 protease of tomato is targeted by cystatin-like EPIC proteins that are secreted by the oomycete pathogen  Phytophthora infestans  ( Pinf ) during infection. C14 has been under diversifying selection in wild potato species coevolving with  Pinf  and reduced  C14  levels result in enhanced susceptibility for  Pinf.  Here, we investigated the role C14-EPIC-like interactions in the natural pathosystem of Arabidopsis with the oomycete pathogen  Hyaloperonospora arabidopsidis  ( Hpa ). In contrast to the  Pinf -solanaceae pathosystem, the C14 orthologous protease of Arabidopsis, RD21, does not evolve under diversifying selection in Arabidopsis, and  rd21  null mutants do not show phenotypes upon compatible and incompatible  Hpa  interactions, despite the evident lack of a major leaf protease.  Hpa  isolates express highly conserved EPIC-like proteins during infections, but it is unknown if these  Hpa EPICs can inhibit RD21 and one of these  Hpa EPICs even lacks the canonical cystatin motifs. The  rd21  mutants are unaffected in compatible and incompatible interactions with  Pseudomonas syringae  pv.  tomato , but are significantly more susceptible for the necrotrophic fungal pathogen  Botrytis cinerea , demonstrating that RD21 provides immunity to a necrotrophic pathogen.

A Role in Immunity for Arabidopsis Cysteine Protease RD21, the Ortholog of the Tomato Immune Protease C14