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The zoophytophagous mirid predator Nesidiocoris tenuis (Reuter) (Hemiptera: Miridae) is used in the augmentative biological control of Thrips palmi Karny (Thysanoptera: Thripidae), a fruit and vegetable pest in protected culture in Japan. N. tenuis can be maintained on certain plants without arthropod prey. Although such plant species may be useful as banker plants, their attractiveness to N. tenuis is underresearched. We investigated the behavioral responses of N. tenuis to (1) uninfested and conspecific-infested banker plants, Verbena × hybrida (verbena), Scaevola aemula (scaevola), Cleome hassleriana (cleome), and Lobularia maritima (sweet alyssum) and (2) conspecific-infested banker plants and T. palmi-infested eggplants, using a Y-tube olfactometer. Female N. tenuis preferred the volatiles emitted by (1) uninfested verbena over clean air but showed no preference for those emitted by the other uninfested banker plants, and (2) conspecific-infested verbena over clean air but showed no preference for those emitted by conspecific-infested scaevola over clean air. The attractiveness of the volatiles of T. palmi-infested eggplants for N. tenuis was not significantly different from those of N. tenuis-infested verbena and scaevola. As both uninfested and conspecific-infested verbena attracted female N. tenuis, this species is a promising banker plant. Chemical analyses and principal coordinate analyses of the volatiles from the headspaces of uninfested and N. tenuis-infested verbena and scaevola showed that 1-octen-3-ol, (E)-β-caryophyllene, and (E)-4,8-dimethyl-1,3,7-nonatriene distinguished these plant species and are candidate volatile N. tenuis attractants. [ABSTRACT FROM AUTHOR]

Zoophytophagous arthropods can elicit plant defense responses affecting potential prey beyond predation. Phytophagy prevails as the main trigger for these responses, as in the case of Euseius stipulatus (Athias-Henriot) (Mesostigmata: Phytoseiidae), a predator occurring in citrus. Because other triggers cannot be excluded, our aim was to examine whether other phytoseiids co-occurring with E. stipulatus but not engaged in plant feeding [Neoseiulus californicus (McGregor) and Phytoseiulus persimilis Athias-Henriot] could induce similar responses (in terms of herbivore induced plant volatiles, HIPVs, and main defensive pathways), and how these affected the behavior of conspecifics and the shared prey, Tetranychus urticae Koch (Prostigmata: Tetranychidae). N. californicus triggered plant genotype-specific defense responses, including the production of different HIPVs compared to clean plants. However, we could not observe these effects for P. persimilis. T. urticae avoided better protected plants, because of stronger direct or indirect defense. As plants with weaker direct defense levels should offer higher prey densities, and those harboring conspecific predators represent higher risk of cannibalism, predators were expected to behave similarly. However, they did not. Our results demonstrate that plant defense triggered by phytoseiids is species-specific, depend on plant genotype and can be triggered by non-feeding activities. As N. californicus is a highly efficient predator used worldwide, further studies with this species are needed. Likewise, cineol, one of the volatiles identified in the blends triggered by this phytoseiid, could be used to manipulate the prey. These studies could pave the way for a more efficient use of phytoseiids in agroecosystems. [ABSTRACT FROM AUTHOR]

Context: Tritrophic interactions may be affected by local factors and the broader landscape context. At small spatial scales, carnivorous enemies of herbivorous insects use herbivore-induced plant volatiles (HIPVs) to find herbivores, but it is unknown whether variation in plant attractiveness due to differential HIPV emission can enhance recruitment of carnivores from the wider landscape. Objectives: We studied whether parasitism of caterpillars of the butterfly Pieris brassicae on white cabbage was influenced by landscape composition and cover with brassicaceous species that provide hosts for parasitoids of P. brassicae in 19 landscapes in the Netherlands. We also investigated whether differential attraction of parasitoids to herbivore-infested plants affected parasitism of P. brassicae caterpillars by using different accessions as proxies for HIPV emission. Results: The cabbage accession that is highly attractive to parasitic wasps recruited more wasps than a less attractive accession, but only when parasitism rates were high. Parasitism rate as proxy of wasp recruitment correlated positively with the cover of brassicaceous plants and area of arable land, suggesting that these habitats support hosts for the wasps and their population growth. In contrast, forest area was negatively associated with parasitism rates. Conclusions: Our study shows that the degree of attractiveness of plants to parasitoids, which is probably mediated by HIPVs, can be a useful predictor of parasitism, but needs to be considered within the landscape context. To understand the strength of tritrophic interactions it is crucial to consider local-scale processes driven by plant-trait variation in combination with landscape-scale processes that determine carnivore abundance. [ABSTRACT FROM AUTHOR]

Herbivore-induced plant volatiles (HIPVs) are important cues for natural enemies to find their hosts. HIPVs are usually present as blends and the effects of combinations of individual components are less studied. Here, we investigated plant volatiles in a tritrophic system, comprising the parasitoid wasp Lytopylus rufipes Nees (Hymenoptera: Braconidae), the Oriental fruit moth Grapholita molesta (Busck) (Lepidoptera: Tortricidae), and Japanese pear, Pyrus pyrifolia 'Kosui', so as to elucidate the effects of single components and blends on wasp behaviors. Bioassays in a four-arm olfactometer, using either shoots or their isolated volatiles collected on adsorbent, revealed that female wasps preferred volatiles from host-infested shoots over those from intact shoots. Analyses identified (Z)-3-hexenyl acetate (H), linalool (L), (E)-β-ocimene (O), (E)-3,8-dimethyl-1,4,7-nonatriene (D), and (E,E)-α-farnesene (F). Among them, only F was induced by infestation with G. molesta. When tested singly, only O and D elicited positive responses by L. rufipes. Binary blends of HO and DF elicited a positive response, but that of HD elicited a negative one, even though D alone elicited a positive response. Remarkably, wasps did not prefer either the ODF or HL blends, but showed a highest positive response to a quinary blend (HLODF). These results show that synergism among volatiles released from host-infested plants is necessary for eliciting high behavioral responses in L. rufipes, enabling L. rufipes to find its host efficiently. [ABSTRACT FROM AUTHOR]

Sour orange, Citrus aurantium, displays higher constitutive and earlier inducible direct defenses against the two-spotted spider mite, Tetranychus urticae, than Cleopatra mandarin, Citrus reshni. Moreover, herbivore-induced plant volatiles (HIPVs) produced by sour orange upon infestation can induce resistance in Cleopatra mandarin but not vice versa. Because the role of these HIPVs in indirect resistance remains ignored, we have carried out a series of behavioral assays with three predatory mites with different levels of specialization on this herbivore, from strict entomophagy to omnivory. We have further characterized the volatile blend associated with T. urticae, which interestingly includes the HIPV methyl salicylate, as well as that produced by induced Cleopatra mandarin plants. Although a preference for less defended plants with presumably higher prey densities (i.e., C. reshni) was expected, this was not always the case. Because predators' responses changed with diet width, with omnivore predators responding to both HIPVs and prey-related odors and specialized ones mostly to prey, our results reveal that these responses depend on plant genotype, prey presence and predator diet specialization. As the different volatile blends produced by infested sour orange, induced Cleopatra mandarin and T. urticae itself are attractive to T. urticae natural enemies but not to the herbivore, they may provide clues to develop new more sustainable tools to manipulate these agriculturally relevant species. [ABSTRACT FROM AUTHOR]

In tri-trophic interactions among plants, herbivorous insects, and parasitoids, the food plant species of herbivores may affect parasitoid performance, such as host searching behavior and development and survival of immature parasitoids in the host body. Here, we tested the effects of food plant species of the host Mythimna separata (Lepidoptera: Noctuidae) on host searching behavior (i.e., flight responses to host-infested plant volatiles [HIPVs] and antennal responses to host feces or fecal extract) by the larval parasitoid Cotesia kariyai (Hymenoptera: Braconidae). In laboratory tests, female wasps responded differentially to HIPVs and feces produced from host caterpillars on different diets. Female wasps showed stronger responses to HIPVs from infested maize than to those from infested daikon. Similarly, wasps showed stronger contact responses to feces from caterpillars fed on maize or kidney bean than to those fed on daikon or an artificial diet. Previous studies demonstrated that maize-fed caterpillars are suitable hosts, whereas daikon-fed caterpillars are unsuitable hosts, for parasitism by C. kariyai. Therefore, our results indicate that female wasps show stronger responses to the chemicals related to suitable hosts than to those of unsuitable hosts. [ABSTRACT FROM AUTHOR]

Plants synthesize variable mixtures of herbivore-induced plant volatiles (HIPVs) as part of their evolutionary conserved defense. To elucidate the impact of chewing herbivores with different level of adaptation on HIPV profiles in rice, we measured HIPVs released from rice seedlings challenged by either the generalist herbivore Mythimna loreyi (MYL) or the specialist Parnara guttata (PAG). Both herbivores markedly elicited the emission of HIPVs, mainly on the second and third days after attack compared to control plants. In addition, side-by-side HIPV comparisons using MYL and PAG caterpillars revealed that generalist feeding induced comparably more HIPVs relative to specialist, particularly on day two as highlighted by multivariate analysis (PLS-DA) of emitted HIPVs, and further confirmed in mimicked herbivory experiments. Here, mechanically wounded plants treated with water (WW) released more VOCs than untreated controls, and on top of this, oral secretions (OS) from both herbivores showed differential effects on volatile emissions from the wounded plants. Similar to actual herbivory, MYL OS promoted higher amounts of HIPVs relative to PAG OS, thus supporting disparate induction of rice indirect defenses in response to generalist and specialist herbivores, which could be due to the differential composition of their OS. (196 words). [ABSTRACT FROM AUTHOR]

We compared the attractiveness of a mixture of cis-3-hexen-1-ol, cis-3-hexenyl acetate, and methyl salicylate (MeSA), and of each of these compounds singly, to a predacious ladybird beetle, Stethorus punctum picipes (Casey), to make it clear whether mixture of several volatiles may attract more natural enemies at different spatial scales. In an olfactometer experiment, S. p. picipes females were attracted to the volatile mixture, but not to single volatiles. Male S. p. picipes were not attracted to the mixture. In a small-scale open-field test using sticky cards, S. p. picipes were attracted to the volatile mixture and to MeSA alone. In a vineyard trapping test, significantly more S. p. picipes were trapped on sticky cards baited with a mixture of volatiles than on single-volatile-baited cards. [ABSTRACT FROM AUTHOR]

Using herbivore-induced plant volatiles (HIPVs) to attract specific natural enemies in the field has proven challenging, partly because of a poor understanding of: (i) which compound(s) to manipulate to attract specific taxa, and (ii) the ecological conditions over which HIPVs are effective. To address these issues, we quantified the response of a complex arthropod community to three common HIPVs (methyl salicylate, cis-3-hexen-1-ol, and phenylethyl alcohol) as individual compounds and equal part blends in corn and soybean fields. Of 119 arthropod taxa surveyed, we found significant responses by four species in corn fields (2 parasitoids, 1 herbivore, and 1 detritivore) and 16 in soybean fields (8 parasitoids, 3 predators, 4 herbivores, and 1 detritivore), with both attractive and repellent effects of the HIPVs observed. For example, tachinid flies were highly attracted to cis-3-hexen-1-ol (ca. 3-fold increase), but repelled by methyl salicylate (ca. 60 % decrease). Surprisingly, we found very few cases in which HIPVs acted synergistically; only two arthropod groups (ichneumonid wasps and phorid flies) were more attracted by a blend of the HIPVs than by the individual compounds composing the blend. Crop type, however, had a strong impact on the strength of arthropod responses to HIPVs. A few arthropod species were broadly affected across both crops (i.e., the herbivore Halticus bractatus was repelled by most of our treatments, regardless of crop background), but overall more arthropod groups responded to HIPVs released in soybean fields compared with corn. This was true despite the fact that taxa responding to HIPVs were present and abundant in both systems, suggesting that crop-based outcomes were likely driven by the plant matrix rather than mere differences in taxonomic composition of the arthropod community in corn vs. soybean fields. As a whole, these results suggest that: (i) repellent effects of HIPVs on natural enemies of herbivorous insects can be observed as frequently as attractive effects; (ii) odor blends may be no more effective than single-compound lures for some taxa; and (iii) crop background alters the magnitude of attraction to HIPVs, depending on the species being targeted. [ABSTRACT FROM AUTHOR]

Many parasitoids use volatiles produced by plants as important cues during their food and host search process. We investigated the attraction of the parasitic wasp Trybliographa rapae Westwood (Hymenoptera: Figitidae) to volatiles emitted from plants infested by the cabbage root fly Delia radicum L. (Diptera: Anthomyiidae), as well as to volatiles from a nectar food plant. Behavioural choice tests showed that male parasitoids were not attracted to any volatiles from plants infested by D. radicum or from nectar plants, while females showed clear attraction to both volatile sources. Young females were more attracted to combined volatiles of host and food plants over those from only the host plant, whereas older females showed no differences in attraction to the two odour sources. This suggests that intercropping attractive flowers with host plants could potentially be used to recruit newly emerged parasitoids from surrounding fields while older parasitoids invest more energy in host location than in additional food search. Volatiles from a whole infested plant were chosen over those emitted from separated above- and below-ground parts from infested plants. It is important to consider the availability of both energy and host resources for parasitoids when designing an eco-compatible management of a vegetable crop system. [ABSTRACT FROM AUTHOR]

Despite a large number of studies on herbivore-induced plant volatiles (HIPVs), little is known about which specific compounds are used by natural enemies to locate prey- or host- infested plants. In addition, the role of HIPVs in attracting natural enemies has been restricted largely to agricultural systems. Differences in volatile blends emitted by cultivars and plants that originate from wild populations may be attributed to potentially contrasting selection regimes: natural selection among the wild types and artificial selection among cultivars. A more realistic understanding of these interactions in a broader ecological and evolutionary framework should include studies that involve insect herbivores, parasitoids, and wild plants on which they naturally interact in the field. We compared the attractiveness of HIPVs emitted by wild and cultivated cabbage to the parasitoid Cotesia rubecula, and determined the chemical composition of the HIPV blends to elucidate which compounds are involved in parasitoid attraction. Wild and cultivated cabbage differed significantly in their volatile emissions. Cotesia rubecula was differentially attracted to the wild cabbage populations and preferred wild over cultivated cabbage. Isothiocyanates, which were only emitted by the wild cabbages, may be the key components that explain the preference for wild over cultivated cabbage, whereas terpenes may be important for the differential attraction among the wild populations. Volatile analysis revealed that parasitoid attraction cannot be explained by simple linear relationships. Our results suggest that unraveling which compound(s) are innately attractive to parasitoids of cabbage pests should include wild Brassicaceae. [ABSTRACT FROM AUTHOR]

Herbivory by arthropods often induces the emission of plant volatiles, which attract natural enemies of the herbivores. This induced emission of volatiles is considered to be a strategy of plants to effectively defend against herbivores by employing bodyguards. Recent empirical research has revealed that these volatiles can also affect neighboring undamaged plants and cause them to emit volatiles secondarily. Provided that signal emission imposes some cost on plants, the evolutionary advantage to undamaged plants in the emission of such secondary signals is unclear. We hypothesized that plants have evolved to emit a secondary signal to help nearby relatives by promoting the recruitment of natural enemies, whereby they increase inclusive fitness. We constructed a simulation model to evaluate this hypothesis. Our simulations suggest that a secondary signal evolves if the following five conditions are met: the cost of the signal is low; the potential risk of infestation is high; the attractiveness of the signal to natural enemies is highly positively correlated with the local density of the signal chemical; dispersal of offspring is spatially restricted, causing population viscosity, and; sufficient vacant space is available, allowing the population to be elastic. [ABSTRACT FROM AUTHOR]