A systematic review of chemical dialogues mediated by volatile compounds in plants associated with entomopathogenic–endophytic fungi: from herbivores to natural enemies

Rice production faces significant losses from Spodoptera frugiperda, with current control strategies relying heavily on chemical insecticides and Bt cultivars, leading to resistance development and environmental concerns. In search of alternative control methods this systematic review examined multitrophic interactions mediated by volatile organic compounds (VOCs) in Oryza sativa inoculated with the entomopathogenic fungus Beauveria bassiana, evaluating impacts on army worms Spodoptera spp. and parasitoids of the genus Trichogramma. Following PRISMA-ScR guidelines, 20 studies from 2000-2025 were analyzed. The results indicated that B. bassiana endophytic colonization modulates VOC profiles, including emission of green leaf volatiles, monoterpenes (e.g., α-pinene), and sesquiterpenes (e.g., β-caryophyllene), enhancing parasitoid attraction and reducing pest oviposition and larval survival. The review also identified fungal VOCs, such as 3-methylbutanol, with direct behavioral effects on pests. Bibliometric analysis revealed increasing research interest but thematic fragmentation between applied entomopathogen studies and chemical ecology. Despite evidence supporting VOC-mediated indirect plant defenses, methodological limitations persist, including insufficient compound characterization and limited field validation. This synthesis underscored the potential of integrating endophytic entomopathogenic fungi to manipulate plant chemical signaling, offering a promising strategy for sustainable and ecologically based pest management in rice cultivation, particularly in tropical agroecosystems.