Many fungal root symbionts of the genus Trichoderma are well-known for their beneficial effects on agronomic performance and protection against plant pathogens;moreover,they may enhance protection from insect pests,by triggering plant resistance mechanisms.Defense barriers against insects are induced by the activation of metabolic pathways involved in the production of defense-related plant compounds,either directly active against herbivore insects,or exerting an indirect effect,by increasing the attraction of herbivore natural enemies.In a model system composed of the tomato plant,the aphid Macrosiphum euphorbiae and the parasitoid Aphidius ervi,plant metabolic changes induced by Trichoderma harzianum and their effects on higher trophic levels have been assessed.T.harzianum T22 treatments induce a primed state that upon aphid attacks leads to an increased attraction of aphid parasitoids,mediated by the enhanced production of volatile organic compounds (VOCs) that are known to induce Aphidius ervi flight.Transcriptome sequencing of T22-treated plants infested by aphids showed a remarkable upregulation of genes involved in terpenoids biosynthesis and salicylic acid pathway,which are consistent with the observed flight response ofA.ervi and the VOC bouquet profile underlying this behavioral response.