Impacts of plant symbiotic fungi on insect herbivores: mutualism in a multitrophic context.

We consider how fungi that form symbiotic associations with plants interact with insect herbivores attacking the same plants. Both endophytes and mycorrhizae have significant impacts on herbivores with which they are in relatively intimate contact, but weaker effects on those from which they are spatially separated. Generalist insects are usually adversely affected by the presence of endophytes and mycorrhizae, whereas specialist insects may often benefit. Effects on feeding guilds vary according to type of fungi; for example, aphids are often negatively affected by endophytes but respond positively to mycorrhizae, and leaf-chewers are usually negatively affected by both types of fungi. There is a strong taxonomic bias in the literature and many interactions remain little studied; laboratory studies predominate over field studies. Although some patterns emerge, there is a large amount of specificity and context dependency in the outcome of interactions, reflecting the influence of fungal and host genotype, fungal, host, and insect species, and environmental factors. Whereas some of the mechanisms underpinning these interactions are relatively well characterized, others remain unclear and await elucidation by molecular and metabolomic techniques.