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Co-infestations by herbivores, a common situation found in natural settings, can distinctly affect induced plant defenses compared to single infestations. Related tritrophic interactions might be affected through the emission of changed blends of herbivore-induced plant volatiles (HIPVs). In a previous study, we observed that the infestation by red spider mite (Oligonychus ilicis) on coffee plants facilitated the infestation by white mealybug (Planococcus minor), whereas the reverse sequence of infestation did not occur. Here, we examined the involvement of the jasmonate and salicylate pathways in the plant-mediated asymmetrical facilitation between red spider mites and white mealybugs as well as the effect of multiple herbivory on attractiveness to the predatory mite Euseius concordis and the ladybug Cryptolaemus montrouzieri. Both mite and mealybug herbivory led to the accumulation of JA-Ile, JA, and cis-OPDA in plants, although the catabolic reactions of JA-Ile were specifically regulated by each herbivore. Infestation by mites or mealybugs induced the release of novel volatiles by coffee plants, which selectively attracted their respective predators. Even though the co-infestation by mites and mealybugs resulted in a stronger accumulation of JA-Ile, JA and SA than the single infestation treatments, the volatile emission was similar to that of mite-infested or mealybug-infested plants. However, multiple infestation had a negative impact on the attractiveness of HIPVs to the predators, making them less attractive to the predatory mite and a repellent to the ladybug. We discuss the potential underlying mechanisms of the susceptibility induced by mites, and the effect of multiple infestation on each predator.

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BACKGROUND: Plant defenses activated by European zoophytophagous predators trigger behavioral responses in arthropods, benefiting pest management. However, repellence or attraction of pests and beneficial insects seems to be species-specific. In the neotropical region, the mirid predator Macrolophus basicornis has proved to be a promising biological control agent of important tomato pests; nevertheless, the benefits of its phytophagous behavior have never been explored. Therefore, we investigated if M. basicornis phytophagy activates tomato plant defenses and the consequences for herbivores and natural enemies. RESULTS: Regardless of the induction period of M. basicornis on tomato plants, Tuta absoluta females showed no preference for the odors emitted by induced or control plants. However, Tuta absoluta oviposited less on plants induced by M. basicornis for 72 h than on control plants. In contrast, induced plants repelled Bemisia tabaci females, and the number of eggs laid was reduced. Although females of Trichogramma pretiosum showed no preference between mirid-induced or control plants, we observed high attraction of the parasitoid Encarsia inaron and conspecifics to plants induced by M. basicornis. While the mirid-induced plants down-regulated the expression of genes involving the salicylic acid (SA) pathway over time, the genes related to the jasmonic acid (JA) pathway were up-regulated, increasing emissions of fatty-acid derivatives and terpenes, which might have influenced the arthropods' host/prey choices. CONCLUSION: Based on both the molecular and behavioral findings, our results indicated that in addition to predation, M. basicornis benefits tomato plant resistance indirectly through its phytophagy. This study is a starting point to pave the way for a novel and sustainable pest-management strategy in the neotropical region. © 2022 Society of Chemical Industry.

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The use of nectar-producing companion plants in crops is a well-known strategy of conserving natural enemies in biological control. However, the role of floral volatiles in attracting parasitoids and effects on host location via herbivore-induced plant volatiles is poorly known. Here, we examined the role of floral volatiles from marigold (Tagetes erecta), alone or in combination with volatiles from sweet pepper plant (Capsicum annuum), in recruiting Aphidius platensis, an important parasitoid of the green peach aphid Myzus persicae. We also investigated whether marigold floral volatiles are more attractive to the parasitoid than those emitted by sweet pepper plants infested by M. persicae. Olfactometry assays indicated that floral volatiles attracted A. platensis to the marigold plant and are more attractive than sweet pepper plant volatiles. However, volatiles emitted by aphid-infested sweet pepper were as attractive to the parasitoid as those of uninfested or aphid-infested blooming marigold. The composition of volatile blends released by uninfested and aphid-infested plants differed between both blooming marigold and sweet pepper, but the parasitoid did not discriminate aphid-infested from uninfested blooming marigold. Volatile released from blooming marigold and sweet pepper shared several compounds, but that of blooming marigold contained larger amounts of fatty-acid derivatives and a different composition of terpenes. We discuss the potential implications of the aphid parasitoid attraction in a diversified crop management strategy.

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Plants are known to increase the emission of volatile organic compounds upon the damage of phytophagous insects. However, very little is known about the composition and temporal dynamics of volatiles released by wild plants of the genus Crotalaria (Fabaceae) attacked with the specialist lepidopteran caterpillar Utetheisa ornatrix (Linnaeus) (Erebidae). In this work, the herbivore-induced plant volatiles (HIPV) emitted by Crotalaria nitens Kunth plants were isolated with solid phase micro-extraction and the conventional purge and trap technique, and their identification was carried out by GC/MS. The poly-dimethylsiloxane/divinylbenzene fiber showed higher affinity for the extraction of apolar compounds (e.g., trans-ß-caryophyllene) compared to the Porapak™-Q adsorbent from the purge & trap method that extracted more polar compounds (e.g., trans-nerolidol and indole). The compounds emitted by C. nitens were mainly green leaf volatile substances, terpenoids, aromatics, and aldoximes (isobutyraldoxime and 2-methylbutyraldoxime), whose maximum emission was six hours after the attack. The attack by caterpillars significantly increased the volatile compounds emission in the C. nitens leaves compared to those subjected to mechanical damage. This result indicated that the U. ornatrix caterpillar is responsible for generating a specific response in C. nitens plants. It was demonstrated that HIPVs repelled conspecific moths from attacked plants and favored oviposition in those without damage. The results showed the importance of volatiles in plant-insect interactions, as well as the choice of appropriate extraction and analytical methods for their study.

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BACKGROUND: Plants in nature can be sequentially attacked by different arthropod herbivores. Feeding by one arthropod species may induce plant-defense responses that might affect the performance of a later-arriving herbivorous species. Understanding these interactions can help in developing pest-management strategies. In tomato, the sweet-potato whitefly Bemisia tabaci and the two-spotted spider mite Tetranychus urticae are key pests that frequently cohabit on the same plant. We studied whether colonization by one species can either facilitate or impede later colonization of tomato plants by conspecific or heterospecific individuals. RESULTS: B. tabaci females showed a strong preference for and increased oviposition on plants previously colonized by conspecifics. In contrast, plants infested with T. urticae repelled B. tabaci females and reduced their oviposition rate by 86%. Although females of T. urticae showed no preference between conspecific-infested or uninfested plants, we observed a 50% reduction in the number of eggs laid on conspecific-infested plants. Both herbivorous arthropods up-regulated the expression of genes involving the jasmonic acid and abscisic acid pathways, increasing emissions of fatty-acid derivatives, but only B. tabaci increased the expression of genes related to the salicylic acid pathway and the total amount of phenylpropanoids released. Terpenoids were the most abundant compounds in the volatile blends; many terpenoids were emitted at different rates, which might have influenced the arthropods' host selection. CONCLUSION: Our results indicate that B. tabaci infestation facilitated subsequent infestations by conspecifics and mites, while T. urticae infestation promoted herbivore-induced resistance. Based on both the molecular and behavioral findings, a novel sustainable pest-management strategy is discussed.

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Plants produce species-specific herbivore-induced plant volatiles (HIPVs) after damage. We tested the hypothesis that herbivore-specific HIPVs prime neighboring plants to induce defenses specific to the priming herbivore. Since Manduca sexta (specialist) and Heliothis virescens (generalist) herbivory induced unique HIPV profiles in Nicotiana benthamiana, we used these HIPVs to prime receiver plants for defense responses to simulated herbivory (mechanical wounding and herbivore regurgitant application). Jasmonic acid (JA) accumulations and emitted volatile profiles were monitored as representative defense responses since JA is the major plant hormone involved in wound and defense signaling and HIPVs have been implicated as signals in tritrophic interactions. Herbivore species-specific HIPVs primed neighboring plants, which produced 2 to 4 times more volatiles and JA after simulated herbivory when compared to similarly treated constitutive volatile-exposed plants. However, HIPV-exposed plants accumulated similar amounts of volatiles and JA independent of the combination of priming or challenging herbivore. Furthermore, volatile profiles emitted by primed plants depended only on the challenging herbivore species but not on the species-specific HIPV profile of damaged emitter plants. This suggests that feeding by either herbivore species primed neighboring plants for increased HIPV emissions specific to the subsequently attacking herbivore and is probably controlled by JA.

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BACKGROUND: Whether tactics to manipulate natural enemies in agro-ecosystems enhance their ecosystem function and services remains debatable. We conducted field experiments in 2015-2016 to test the hypothesis that attraction of natural enemies to herbivore-induced plant volatiles (HIPVs), alone or in combination with companion plants, increases crop productivity. Our treatments consisted of bean plants alone or baited with methyl salicylate (MeSA; an HIPV), or combined with coriander (a companion plant), or with both MeSA and coriander. Numbers of arthropods were visually sampled in each treatment. Sentinel aphids were used to measure ecosystem function (i.e. predation). Plant damage and biomass, and the number and weight of pods and seeds, were measured as a proxy for ecosystem services. RESULTS: MeSA and coriander, when alone or combined, increased the abundance of insect predators from six families, reduced herbivore (e.g. spider mite and thrips) populations, and increased aphid predation. MeSA and coriander also reduced damage by spider mites. MeSA with or without coriander did not, however, increase crop biomass or any yield parameters. CONCLUSIONS: MeSA alone or combined with coriander attracted different predator communities, altered pest communities, and reduced damage; however, these results did not cascade down to improve crop productivity. © 2018 Society of Chemical Industry.

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Herbivory can induce several structural and functional alterations in the plant secretory system. Glandular trichomes are the main sites of production of volatile organic compounds (VOCs) with several chemical properties in Lamiaceae species. Ocimum species usually have three morphotypes of glandular trichomes (morphotype I is peltate and has a wide four-celled head; morphotype II is capitate and has a unicellular head; and morphotype III is capitate with a bicellular head) which produce a great amount of terpenes, although other chemical categories of substances are also produced. Despite the abundance of trichomes producing important anti-herbivory components in their leaves, the association between Ocimum species and leaf-cutter ants has been commonly registered in Brazil. We investigated the effect of leaf-cutter ant attack on the density of the glandular trichomes and on the chemistry of the VOCs released from leaves of O. gratissimum. Plants were subjected to Acromyrmex rugosus attack until 90 % of leaves were removed. After 40 days from the leaf-cutter attack, both treatments were sampled. The glandular trichome density was analysed by scanning electron microscopy. The VOCs were extracted utilizing headspace solid-phase microextraction (HS-SPME) technique and analysed by gas chromatography. Generally, the density of glandular trichomes increased in the adaxial leaf surface of the attacked plants. However, we bring novelties on this topic since we analysed the density of each morphotype separately. The morphotype I decreased in the abaxial leaf surface, and increased in the adaxial leaf surface; the morphotype II increased in both leaf surfaces; and the morphotype III decreased in the abaxial leaf surface and remained constant in the adaxial leaf surface of attacked plants. In leaves of attacked plants, the (Z)-ß-ocimene increased by 50 %, the α-selinene by 13 % and the germacrene D by 126 %, whereas the eugenol decreased by 70 %. Our data point to a differential response of each glandular morphotype in O. gratissimum and are consistent with the idea of a compartmentalization of functions among the different glandular morphotypes in the plant defence against environmental factors.