The hemolymph microbiome of insects.

Hemolymph has long been recognized as a key mediator of nutritional and immunological homeostasis in insects, with the tacit understanding that hemolymph is a hostile environment for microorganisms, and microbiologically sterile in healthy insects. Recent research is overturning the conventional wisdom, and there is now overwhelming evidence that various non-pathogenic microorganisms can stably or transiently inhabit hemolymph in a diversity of insects. Most is known about Spiroplasma, especially in Drosophila species, and secondary symbionts of the Enterobacteriaceae, notably Hamiltonella defensa, in aphids. These bacteria require many nutrients, representing a likely drain on host nutritional resources, and they persist in the hemolymph by a combination of evasion and tolerance of insect immune effectors. These traits can be costly to the insect host. For some hemolymph microorganisms, these costs are balanced by other traits beneficial to the insect, notably protection against natural enemies mediated by specific toxins or competition for key nutrients. Three key priorities for future research are: to investigate the prevalence and taxonomic diversity of hemolymph microorganisms in insects; to establish the role of host nutritional and immune factors as determinants of the abundance and proliferation rates of hemolymph microorganisms; and to integrate the developing understanding of these microorganisms and their impacts (both costs and benefits) on insect nutrition and immune function into the wider study of insect physiology.