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    Antagonists and defense mechanisms of entomopathogenic nematodes and their mutualistic bacteria
    (Academic Press Inc Elsevier Science, 2021) Raja, Ramalingam Karthik; Arun, Alagarsamy; Touray, Mustapha; Gulsen, Sebnem Hazal; Cimen, Harun; Gulcu, Baris; Hazir, Selcuk
    Entomopathogenic nematodes (EPNs) in the families Steinernematidae and Heterorhabditidae are effective biocontrol agents against a number of important soil insect pests and are safe to vertebrates, plants and other non-target organisms with no negative effects on the environment. They are associated with mutualistic bacteria in the genus Xenorhabdus for Steinernematidae and Photorhabdus for Heterorhabditidae and are often referred to as the nematode/bacterium complex. The natural habitat of EPNs is the soil where the third-stage infective juveniles (IJs), the only free-living stage, can infect an array of insect hosts. However, the survival of the IJs is affected by various abiotic (i.e., temperature, moisture, soil texture, soil salinity, UV light, oxygen, and pH) and biotic (i.e., natural enemies, omnivores, scavengers, competitors, and plants) factors in the soil. In this review, we summarize the impact of the biotic factors, especially natural enemies, omnivores, scavengers, and competitors against EPNs and conclude that the major predators of the IJs appear to be nematophagous fungi, mites, and collembolans. Omnivores and scavengers, particularly, mites, collembolans, ants and birds, feeding on EPNkilled insects appear to be the major mortality factors on the developing nematodes especially of insects that are 2 days or less post infection. In addition, we discuss the defense mechanisms of EPN IJs against nematophagous fungi, and the protection of the nematode-killed insects by production of the scavenger deterrent factor produced by the mutualistic bacteria against omnivores and scavengers. By understanding the interactions between EPNs and their antagonists we can improve and increase the successful use of EPNs in biological control programs.