Systemically-induced response of cabbage plants against a specialist herbivore, Pieris brassicae

Plant responses to herbivory might directly affect the herbivore (direct defences) or might benefit the plant by promoting the effectiveness of natural antagonists of the herbivores (indirect defences). Brussels sprouts attacked by Pieris brassicae larvae release volatiles that attract a natural antagonist of the herbivores, the parasitoid Cotesia glomerata, to the damaged plant. In a previous study, we observed that feeding by caterpillars on the lower leaves of the plant triggers the systemic release of volatiles detectable by the parasitoids from upper leaves of the same plant. The role of these systemically induced volatiles as indirect defence and the dynamics of their emission were investigated in wind-tunnel dual choice tests with C. glomerata. The systemically induced emission of volatiles varied depending on leaf age and on plant age. Systemic induction affected parasitoid effectiveness, as induced plants could be more easily located by parasitoids than non-induced ones. The role of the systemic induction as a direct defence was investigated through behavioural and feeding tests with P. brassicae. In dual choice assays, Ist instar larvae preferred to feed and fed more on systemically induced than on non-induced leaves. In single choice assays, the leaf area consumed by caterpillars was larger on systemically induced leaves than on non-induced control leaves. However, caterpillars fed on systemically induced leaves attained the same weight as those feeding on non-induced controls. In addition, P. brassicae pupae whose larvae were fed on systemically induced leaves had longer developmental times than those of larvae fed on non-induced leaves. Adult oviposition behavior was not influenced by systemic induction. We conclude that systemically induced responses in cabbage might reduce P. brassicae fitness both directly, by affecting their development and feeding behavior and indirectly by making caterpillars and pupae more vulnerable to attack by carnivores. The occurrence of a possible relationship between direct and indirect defence is discussed.