Plant responses to variable timing of aboveground clipping and belowground herbivory depend on plant age

Abstract

Aims Plants use different types of responses such as tolerance and induced defense to mitigate the effects of herbivores. The direction and magnitude of both these plant responses can vary with plant age. However, most studies have focused on aboveground herbivory, whereas important feeding occurs belowground. Here, we tested the hypothesis that plant tolerance and defense following shoot damage or root herbivory depends on plant age. Methods In order to test our hypothesis, we exposed the perennial grass species Holcus lanatus to defoliation and root nematode inoculation at three growth stages (young, intermediate and old plants), and examined responses of plant traits related to tolerance (regrowth following defoliation) and defense (leaf and root nitrogen and phenolics). Important Findings Defoliation overall reduced plant shoot and root biomass as well as foliar concentrations of phenolics regardless of plant age at defoliation. In contrast, defoliation increased foliar N concentrations, but only when defoliation occurred at intermediate and old plant age. Inoculation with root-feeding nematodes reduced root N concentrations after a prolonged period of growth, but only when nematodes had been inoculated when plants were young. The relative shoot regrowth rate of plants increased immediately after defoliation but this was independent of the plant age at which defoliation occurred, i.e. was not stronger in plants that were defoliated at a more advanced age, as hypothesized. Similarly, relative root growth rates increased shortly after defoliation, but this was only observed for plants defoliated when they were young. We conclude that plant responses to aboveground and belowground herbivory in traits related to both defense and tolerance are affected by plant age, but do not generally change with plant age.