The Layers of Plant Responses to Insect Herbivores

Plants collectively produce hundreds of thousands of specialized metabolites that are not required for growth or development. Each species has a qualitatively unique profile, with variation among individuals, growth stages, and tissues. By the 1950s, entomologists began to recognize the supreme importance of these metabolites in shaping insect herbivore communities. Plant defense theories arose to address observed patterns of variation, but provided few testable hypotheses because they did not distinguish clearly among proximate and ultimate causes. Molecular plant-insect interaction research has since revealed the sophistication of plant metabolic, developmental, and signaling networks. This understanding at the molecular level, rather than theoretical predictions, has driven the development of new hypotheses and tools and pushed the field forward. We reflect on the utility of the functional perspective provided by the optimal defense theory, and propose a conceptual model of plant defense as a series of layers each at a different level of analysis, illustrated by advances in the molecular ecology of plant-insect interactions.