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The effects of multiple insect attacks on herbivore-induced plant volatiles and carnivorous arthropods are increasingly studied. Phytopathogens also represent an important threat to plants, and plant defence strategies against pathogens and insects are strongly interconnected, yet the potential impact of pathogens on insect-induced volatiles has been largely overlooked, and degree of pathogenicity is rarely considered. We investigated how pathogen challenge, with virulent and avirulent strains of Xanthomonas campestris either alone or with simultaneous Pieris brassicae caterpillar herbivory, affected the volatile emissions of Brassica nigra plants. The impact of these volatiles on the foraging behaviour of Cotesia glomerata parasitoids wasps was then assessed. Pathogens themselves induced volatiles that were highly attractive to parasitoids, and enhanced the attractiveness of host-infested plant volatiles. Chemical analyses revealed that virulent and avirulent strains differentially induced plant volatiles, with primarily sesquiterpene, homoterpene and green leaf volatile compounds contributing to the differences. Strong similarities were found in the blends induced by the virulent strain and caterpillar herbivory. Challenge by either virulent or avirulent pathogens has a significant impact on plant chemistry and its interactions with other community members, demonstrating the importance of integrating pathogen- and insect-based research to broaden our knowledge of plant defences under conditions of increasing complexity.

1. Plant traits that enhance the attraction of the natural enemies of their herbivores have been postulated to function as an ‘indirect defence’. An important underlying assumption is that this enhanced attraction results in increased plant fitness due to reduced herbivory. This assumption has been rarely tested. 2. We investigated whether there are fitness consequences for the charlock mustard Sinapis arvensis, a short-lived outcrossing annual weedy plant, when exposed to groups of large cabbage white (Pieris brassicae) caterpillars parasitized by either one of two wasp species, Hyposoter ebeninus and Cotesia glomerata, that allow the host to grow during parasitism. Hyposoter ebeninus is solitary and greatly reduces host growth compared with healthy caterpillars, whereas C. glomerata is gregarious and allows the host to grow approximately as large as unparasitized caterpillars. Both healthy and parasitized P. brassicae caterpillars initially feed on the foliage, but later stages preferentially consume the flowers. 3. In a garden experiment, plants damaged by parasitized caterpillars produced more seeds than conspecific plants damaged by unparasitized caterpillars. Reproductive potential (germination success multiplied by total seed number) was similar for plants that were not exposed to herbivory and those that were damaged by parasitized caterpillars and lower for plants that were damaged by healthy unparasitized caterpillars. However, these quantitative seed traits negatively correlated with the qualitative seed traits, individual seed size and germination success, suggesting a trade-off between these two types of traits. 4. We show that parasitism of insect herbivores that feed on reproductive plant tissues may have positive fitness consequences for S. arvensis. The extent to which plant fitness may benefit depends on parasitoid lifestyle (solitary or gregarious), which is correlated with the amount of damage inflicted on these tissues by the parasitized host.

1.Plants respond to insect herbivores with changes in physical and chemical traits, both locally and systemically, in leaves and flowers. Such phenotypic changes may influence the behaviour of every community member that interacts with the plant. Here, we address effects of plant responses to eggs and subsequent herbivory by caterpillars on plant-mediated interactions with pollinators and consequences for plant fitness. 2.Using a common garden set-up, we have investigated responses of Brassica nigra plants to herbivore exposure from egg deposition onwards throughout larval development. We quantified effects of infestation by the specialist Pieris brassicae on: 1. behaviour of pollinators; 2. volatile emission and 3. timing and number of seeds produced. 3.Egg deposition and folivory did not influence visitation by pollinators to plots of infested or control plants. Effects of herbivore infestation on both pollinator visitation and volatile emission were observed only at a later stage, when caterpillars were feeding on the flowers. 4.Remarkably, before eggs had hatched, infested plants accelerated seed production. The caterpillars that developed from the eggs fed on flowers but not on seeds and thus seed production prior to herbivory on flowers safeguarded reproductive output. 5.The results of this study show that early plant investments in reproduction can successfully prevent consumption of expensive reproductive tissues. By accelerating seed production, plants prevented consumption of flowers and effectively defended themselves against the herbivores