8 Abstract In the last decade, biological control programs 9 for greenhouse tomatoes and other crops have been suc-10 cessfully implemented using zoophytophagous plant bugs 11 (Miridae), which can feed on both plant tissues and insect 12 prey. It is well known that plants respond to herbivore 13 attacks by releasing volatile compounds through diverse 14 pathways triggered by phytohormones. These herbivore-15 induced plant volatiles can alert neighboring plants, repel 16 or attract herbivores, and attract natural enemies of these 17 herbivores. Nevertheless, the possible benefits of induced 18 plant responses by zoophytophagous predators that could 19 add to their usefulness as biocontrol agents have not been 20 studied until now. Here we show that the zoophytophagous 21 predator Nesidiocoris tenuis activated abscisic acid and 22 jasmonic acid (JA) signaling pathways in tomato plants, 23 which made them less attractive to the whitefly Bemisia 24 tabaci, a major tomato pest worldwide, and more attractive 25 to the whitefly parasitoid, Encarsia formosa. We also found 26 that intact tomato plants exposed to volatiles from N. ten-27 uis-punctured plants activated the JA pathway, and as a 28 consequence, E. formosa was also attracted to these intact 29 plants with activated defense systems. Thus, our results 30 demonstrate that N. tenuis not only benefits tomato plants 31 directly by entomophagy but also indirectly by phyto-32 phagy, which induces a physiological response in the 33 tomato plant.
38We have proved that the zoophytophagous predator Nesi-39 diocoris tenuis induces plant benefits directly by its ento-40 mophagy and also indirectly by its phytophagy, which 41 induces the attraction of a whitefly parasitoid (Encarsia 42 formosa) and antixenosis to the whitefly Bemisia tabaci. 43 Furthermore, N. tenuis-punctured plants induce plant 44 defenses in intact plants that result in attraction of E. for-45 mosa. Our results might be one reasonable explanation for 46 the great success achieved by N. tenuis as a key biocontrol 47 agent in tomatoes.