A novel cost of R gene resistance in the presence of disease

Resistance responses can impose fitness costs when pests are absent. Here, we test whether the induction of resistance can decrease fitness even in plants under attack; we call this potential outcome a net cost with attack. Using lines in which genetic background was controlled, we investigated whether susceptible Arabidopsis thaliana plants can outperform R gene resistant plants when infected with pathogens. For the R gene RPS2, there was a fitness benefit of resistance in the presence of intraspecific competition, but there was a net cost in the absence of competition: resistant plants produced less seed than susceptible plants even though infected with Pseudomonas syringae. This net cost was primarily due to overcompensation by susceptible plants, which occurred because of a developmental response to infection. For the R gene RPP5, there was no fitness effect of resistance without competition but a net cost when plants were infected with Peronospora parasitica in the presence of competition. This net cost was due to a reduction in the fitness of infected, resistant plants and complete compensation in susceptible plants. A spatially variable model suggests that a trade-off between net benefits and net costs with attack may help explain the persistence of individuals lacking R gene resistance to disease.