Specialist and generalist herbivores exert opposing selection on a chemical defense

Plant defense traits often show high levels of genetic variation, despite clear impacts on plant fitness. This variation may be partly maintained by trade-offs in the defense against multiple herbivore species, for example between generalists and coevolved specialists. Despite a long-standing discussion in the literature on the subject, no study to date has specifically manipulated specialist and generalist herbivores independently of one another to determine whether the two guilds exert opposing selection pressures on specific defensive traits. In two separate experiments, the dominant specialist and generalist herbivores of Brassica nigra were independently manipulated to test whether the composition of the herbivore community altered the direction of selection on a major defensive trait of the plant, sinigrin concentration. It was found that generalist damage was negatively correlated but specialist loads were positively correlated with increasing sinigrin concentrations; and sinigrin concentration was favored when specialists were removed, disfavored (past an intermediate point) when generalists were removed and selectively neutral when plants faced both generalists and specialists. These results suggest that specialist and generalist herbivores can exert opposing selection pressures on chemical defenses, and thus that changes in herbivore community composition can alter the net selective value of defensive traits.