How plants give early herbivore alert: Volatile terpenoids attract parasitoids to egg-infested elms

Abstract

Plants can defend themselves against insect attack prior to larval feeding damage by responding to eggs laid on their leaves. Insect egg deposition can induce leaves to release a complex blend of volatiles attracting egg parasitoids which kill the eggs. Only a few studies have addressed the question which of these egg-induced plant volatiles are relevant for parasitoid attraction. Egg deposition by the elm leaf beetle Xanthogaleruca luteola on leaves of the European field elm Ulmus minor is known to induce the emission of a blend consisting mainly of terpenoids and some green leaf volatiles, which attracts a specialised egg parasitoid of X. luteola, the eulophid wasp Oomyzus gallerucae. Here, we investigated the role of oviposition-induced terpenoids from elm leaves for parasitoid attraction. Quantitative GC–MS analyses showed that inhibition of terpene biosynthesis in leaves by treatment with cerivastatin ® and fosmidomycin reduced emission of ( E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), ( E)- β-caryophyllene and a yet unidentified oxygenated sesquiterpene, but unexpectedly also that of green leaf volatiles (GLVs). Laboratory olfactometer assays revealed that inhibitor treatment rendered oviposition-induced elm leaves unattractive for the parasitoids. Further bioassays showed that single terpenoids per se attracted the parasitoids. Although the only tested GLV 1-hexanol was not attractive in olfactometer tests, we cannot rule out that other GLVs might play a role in parasitoid attraction. When attractiveness of DMNT was tested in the field, parasitoids were attracted to DMNT-baited traps in the presence of background odour emitted by a natural elm stand. We conclude that elms alert their egg parasitoid “helpers” after elm leaf beetle oviposition by means of one or more terpenoid volatiles.