Inducible plant responses linking above- and below-ground herbivory: ecological significance and underlying mechanisms

Above- and below-ground organisms can indirectly affect one another via several recently-described mechanisms, one such mechanism being herbivore-induced plant responses. Because plants leaves occur above-ground and roots below-ground, systemic plant responses to foliar- and root-feeding consumers can result in reciprocal interactions between above- and below-ground herbivory. To first address the broader theoretical context underlying plant-mediated herbivore interactions I conducted a meta-analytical review of interspecific interactions in phytophagous insects (Chapter 1). Using a data-set of 333 observations of interspecific herbivore interactions compiled from 145 independently published studies, I quantitatively assessed: (a) the overall importance of competition in the ecology of insect herbivores, and (b) whether plant-feeding insects conform to traditional competition paradigms. Despite finding frequent evidence for competition, I found very little evidence that phytophagous insects conform to theoretical predictions for interspecific competition. Notably, the strength of interactions between herbivores was largely unaffected by feeding guild, and occurred among distantly-related species that were spatially- and temporally-separated from one another. Moreover, in most cases plants mediated these indirect interactions. Next, I used the tobacco (Nicotiana tabacum) system to explore plant responses to foliar-feeding insects and root-feeding nematodes. I found that aboveground insect herbivores had limited impact on the secondary chemistry of roots, but belowground nematode herbivores strongly affected leaf chemistry (Chapter 2). However, the magnitude of leaf-root induction was also affected by vascular connectivity, with stronger induced responses among plant tissues that were more closely aligned (Chapter 3). Last, I assessed the impact of induced responses on the performance and abundance of foliar and root herbivores using manipulative greenhouse (Chapter 4) and field studies (Chapter 5). Overall, I documented that root-feeding nematodes positively affect leaf-chewing insects by interfering with aboveground nicotine dynamics, whereas aboveground insects benefit root-feeding nematodes via alteration of source-sink dynamics