Root herbivory reduces growth and survival of the shoot feeding specialist Pieris rapae on Brassica nigra

Plants may respond to herbivore attacks by changing their chemical profile. Such induced responses occur both locally and systemically throughout the plant. In this paper we studied how Brassica nigra (L.) Koch (Brassicaceae) plants respond to two different root feeders, the endoparasitic nematode Pratylenchus penetrans Cobb (Tylenchida: Pratylenchidae) and the larvae of the cabbage root fly Delia radicum L. (Diptera: Anthomyiidae). We tested whether the activities of the root feeders affected the survival and development of the shoot feeding crucifer specialist Pieris rapae (L.) (Lepidoptera: Pieridae) via systemically induced changes in the shoots. Overall, P. rapae larvae grew slower and produced fewer pupae on plants that were infested with root feeders, especially on plants infested with P. penetrans. This effect could not be attributed to lower water or protein levels in these plants, as the percentage of water in the controls and root infested shoots was similar, and protein content was even higher in root infested plants. Both glucosinolate as well as phenolic levels were affected by root feeding. Initially, glucosinolate levels were the lowest in root infested plants, but on P. penetrans infested plants they increased more rapidly after P. rapae started feeding than in controls or D. radicum infested plants. Plants with D. radicum feeding on their roots had the highest phenolic levels at all harvest dates. Our results indicate that root feeding can significantly alter the nutritional quality of shoots by changes in secondary metabolite levels and hence the performance of a specialist shoot feede