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Engineering resistance genes to gain effector recognition is emerging as an important step in attaining broad, durable resistance. We engineered potato resistance gene  R3a  to gain recognition of the virulent AVR3a EM  effector form of  Phytophthora infestans . Random mutagenesis, gene shuffling and site-directed mutagenesis of  R3a  were conducted to produce R3a* variants with gain of recognition towards AVR3a EM . Programmed cell death following gain of recognition was enhanced in iterative rounds of artificial evolution and neared levels observed for recognition of AVR3a KI  by R3a. We demonstrated that R3a*-mediated recognition responses, like for R3a, are dependent on SGT1 and HSP90. In addition, this gain of response is associated with re-localisation of R3a* variants from the cytoplasm to late endosomes when co-expressed with either AVR3a KI  or AVR3a EM  a mechanism that was previously only seen for R3a upon co-infiltration with AVR3a KI . Similarly, AVR3a EM  specifically re-localised to the same vesicles upon recognition by R3a* variants, but not with R3a. R3a and R3a* provide resistance to  P. infestans  isolates expressing AVR3a KI  but not those homozygous for AVR3a EM .

Detection of the Virulent Form of AVR3a from Phytophthora infestans following Artificial Evolution of Potato Resistance Gene R3a