Can tolerance traits impose selection on herbivores?

Plant tolerance reduces the fitness consequences of herbivore and natural enemy damage, while resistance reduces the amount of damage suffered. In contrast to resistance, tolerance is often assumed to not affect herbivore performance and evolution. Evidence from the literature, however, suggests that it is possible for plant tolerance to affect herbivore performance and evolution, and potentially plant-herbivore coevolution. First, for cases when genetic correlations between resistance and tolerance are due to pleiotropy, the genes and loci for tolerance and resistance are the same, and as such both traits will affect herbivore performance directly. Second, it is possible that the physiological basis and mechanisms of plant tolerance-for example, changes in plant physiology and resource allocation-directly alter herbivore fitness characters. In this paper, I review the evidence for these potential effects of plant tolerance on herbivore performance, and suggest straightforward experiments to evaluate these possibilities. More generally, I propose that this untested assumption is constraining our view of plant-herbivore coevolution.