Genetic architecture and pleiotropy shape costs of Rps2-mediated resistance in Arabidopsis thaliana

The mounting evidence that R-genes incur large fitness costs raises a question: how can there be a 5-10% fitness reduction for all 149 R-genes in the Arabidopsis thaliana genome? The R-genes tested to date segregate for insertion-deletion (indel) polymorphisms where susceptible alleles are complete deletions. Since costs of resistance are measured as the differential fitness of isolines carrying resistant and susceptible alleles, indels reveal costs that may be masked when susceptible alleles are expressed. Rps2 segregates for two expressed clades of alleles, one resistant and one susceptible. Plants with resistant Rps2 are not less fit than those with a susceptible Rps2 allele in the absence of disease. Instead, all alleles provide a fitness benefit relative to an artificial deletion, due to the role of RPS2 as a negative regulator of defense. Our results highlight the interplay between genomic architecture and the magnitude of costs of resistance.