Nectar in Plant-Insect Mutualistic Relationships: From Food Reward to Partner Manipulation

It has been known for centuries that floral and extra-floral nectar secreted by plants attracts and rewards animals. Extra-floral nectar is involved in so-called indirect defense by attracting animals (generally ants) that prey on herbivores, or by discouraging herbivores from feeding on the plant. Floral nectar is presented inside the flower close to the reproductive organs and rewards animals that perform pollination while visiting the flower. In both cases nectar is a source of carbon and nitrogen compounds that feed animals, the most abundant solutes being sugars and amino acids. Plant-animal relationships involving the two types of nectar have therefore been used for a long time as text-book examples of symmetric mutualism: services provided by animals to plants in exchange for food provided by plants to animals. Cheating (or deception or exploitation), namely obtaining the reward/service without returning any counterpart, is however, well-known in mutualistic relationships, since the interacting partners have conflicting interests and selection may favor cheating strategies. A more subtle way of exploiting mutualism was recently highlighted. It implies the evolution of strategies to maximize the benefits obtained by one partner while still providing the reward/service to the other partner. Several substances other than sugars and amino acids have been found in nectar and some affect the foraging behavior of insects and potentially increase the benefits to the plant. Such substances can be considered plant cues to exploit mutualism. Recent evidence motivated some authors to use the term "manipulation" of animals by plants in nectar-mediated mutualistic relationships. This review highlights the recent background of the "manipulation" hypothesis, discussing it in the framework of new ecological and evolutionary scenarios in plant-animal interactions, as a stimulus for future research.