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Summary: Herbivore‐induced plant volatiles (HIPVs) are widely recognized as an ecologically important defensive response of plants against herbivory. Although the induction of this 'cry for help' has been well documented, only a few studies have investigated the inhibition of HIPVs by herbivores and little is known about whether herbivores have evolved mechanisms to inhibit the release of HIPVs.To examine the role of herbivore effectors in modulating HIPVs and stomatal dynamics, we conducted series of experiments combining pharmacological, surgical, genetic (CRISPR‐Cas9) and chemical (GC‐MS analysis) approaches.We show that the salivary enzyme, glucose oxidase (GOX), secreted by the caterpillar Helicoverpa zea on leaves, causes stomatal closure in tomato (Solanum lycopersicum) within 5 min, and in both tomato and soybean (Glycine max) for at least 48 h. GOX also inhibits the emission of several HIPVs during feeding by H. zea, including (Z)‐3‐hexenol, (Z)‐jasmone and (Z)‐3‐hexenyl acetate, which are important airborne signals in plant defenses.Our findings highlight a potential adaptive strategy where an insect herbivore inhibits plant airborne defenses during feeding by exploiting the association between stomatal dynamics and HIPV emission. [ABSTRACT FROM AUTHOR]

Parasitoids need to find their plant-feeding hosts in complex environments that contain multiple other plant and insect species. They usually rely on herbivore-induced plant volatiles to locate herbivore-infested plants from a distance and their foraging efficiency may be reduced when volatile information is not available. Downwind foraging during times when high wind speeds prevent odour-guided upwind flights may create foraging situations with limited accessibiliy to volatile information. We hypothesized that parasitoids forage less efficiently by landing on nonhost-damaged or undamaged plants when they are forced to fly downwind and tested this in a wind tunnel experiment. We released the parasitoid Cotesia glomerata (Hymenoptera, Braconidae) either upwind or downwind of a plant stand and observed their foraging behaviour. During downwind foraging, parasitoids were less successful in host finding and needed more time until they managed to oviposit in a host caterpillar compared to the upwind foraging situation. The observed increase in foraging time was caused by prolonged foraging on nonhost-infested and undamaged plants in the downwind situation, indicating that parasitoids leave an unprofitable patch that does not contain host caterpillars earlier, when they perceive volatiles from other herbivore-infested plants located upwind. Volatile information on the availability of herbivore-infested plants within a plant stand seems to be crucial for efficient foraging in plant stands that contain a mixture of host-infested, nonhost-infested and undamaged plants. Parasitoid foraging efficiency may thus be strongly reduced when high wind speed prevents odour-guided upwind flight. • Downwind-flying parasitoids foraged less efficiently than upwind-flying parasitoids. • Upwind-flying parasitoids had access to HIPVs from all herbivore-infested plants. • Low HIPV availability reduced between-plant movement of downwind-flying parasitoids. • Downwind-flying parasitoids foraged more intensively on nonhost-infested plants. • Intensive foraging on nonhost-infested plants delayed host finding. [ABSTRACT FROM AUTHOR]

Nine herbivore-induced plant volatiles (HIPVs) and one methyl jasmonate were field-tested for their attractiveness to the main predators in cotton fields of North China in 2 yr. The main predators including ladybird beetles (Propylaea japonica (Thunberg), Harmonia axyridis (Pallas)), green lacewings (Chrysoplera sinica (Tjeder), Chrysopa spp.), predatory bugs (Geocoris pallidipennis (Costa), Orius spp., Nabis spp.) and spiders (Misumenops tricuspidatus (Fabricius), Erigonidium graminicolum (Sundevall)) were investigated. Two-way ANOVA indicated that the volatile compound, year, and the volatile compound x year interaction affected the behavioral responses of predators. It was found that indole significantly attracted the ladybird beetle P. japonica, H. axyridis. Linalool could attract P. japonica. Green lacewing C. sinica was significantly attracted by α-pinene and β-pinene, whereas indole significantly attracted Chrysopa spp. Methyl jasmonate and α-pinene showed significant attraction to small-flower bug Orius spp. In addition, the attraction of α-humulene to C. sinica, attractiveness of β-pinene to Orius spp. and Chrysopa spp., were observed only in one of the two years. However, the big-eyed bug G. pallidipennis, damsel bug Nabis spp., spiders M. tricuspiata and E. graminicolum did not respond to any of the tested HIPVs. These results are discussed with respect to possible applications of a synthetic attractant for main predators in cotton fields. [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]

Silicon (Si) is important in plant defenses that operate in a direct manner against herbivores, and work in rice (Oryza sativa) has established that this is mediated by the jasmonate signaling pathway. Plant defenses also operate indirectly, by the production of herbivore induced plant volatiles (HIPVs) that attract predators and parasitoids of herbivores. These indirect defenses too are mediated by the jasmonate pathway but no earlier work has demonstrated an effect of Si on HIPVs. In this study, we tested the effect of Si supplementation versus Si deprivation to rice plants on subsequent HIPV production following feeding by the important pest, rice leaffolder (Cnaphalocrocis medinalis). Gas chromatography-mass spectrometry analyses showed lower production of a-bergamotene, β-sesquiohellandrene, hexanal 2-ethyl, and cedrol from +Si herbivore-infested plants compared with -Si infested plants. These changes in plant chemistry were ecologically significant in altering the extent to which parasitoids were attracted to infested plants. Adult females of Trathala flavo-orbitalis and Microplitis mediator both exhibited greater attraction to the HIPV blend of +Si plants infested with their respective insect hosts compared to -Si infested plants. In equivalent studies using RNAi rice plants in which jasmonate perception was silenced there was no equivalent change to the HIPV blend associated with Si treatment; indicating that the effects of Si on HIPVs are modulated by the jasmonate pathway. Further, this work demonstrates that silicon alters the HIPV blend of herbivore-infested rice plants. The significance of this finding is that there are no earlier-published studies of this phenomenon in rice or any other plant species. Si treatment to crops offers scope for enhancing induced, indirect defenses and associated biological control of pests because parasitoids are more strongly attracted by the HIPVs produced by +Si plants. [ABSTRACT FROM AUTHOR]

Plants respond to herbivory with the emission of plant volatiles, which can be used by the herbivores' natural enemies to locate their hosts or prey. In nature, plants are often simultaneously confronted with insect herbivores and phytopathogens, potentially interfering with the attraction of the herbivores' enemies as a result of modifications of the induced volatile blend. Here, we investigated parasitoid ( C otesia glomerata) attraction to volatiles of plants challenged by different attackers, either alone or in combination with P ieris brassicae caterpillars, hosts of C . glomerata. We used a natural system consisting of B rassica nigra plants, eggs and larvae of P . brassicae, B revicoryne brassicae aphids and the bacterial phytopathogen X anthomonas campestris pv. campestris. In all cases, parasitoids successfully located host-infested plants, and wasp foraging behaviour was unaffected by the simultaneous presence of a non-host attacker or host eggs. Analysis of the volatile emissions show that the volatile blends of caterpillar-infested treatments were different from those without caterpillars. Furthermore, dually attacked plants could not be separated from those with only caterpillars, regardless of non-host identity, supporting the behavioural data. Our results suggest that, in this system, indirect plant defences may be more resistant to interference than is generally assumed, with volatiles induced during dual attack remaining reliable indicators of host presence for parasitoids. [ABSTRACT FROM AUTHOR]

Parasitoid foraging decisions are often affected by community characteristics such as community diversity and complexity. As part of a complex habitat, the presence of unsuitable hosts may affect foraging behaviour of parasitoids. First, unsuitable herbivores may affect the localization of patches where hosts are present. Second, encounters with unsuitable herbivores in the food plant patch may affect parasitoid decisions during their searching behaviour in the patch. In this review, we outline the importance of the presence of unsuitable herbivores on the behavioural responses of parasitoids during both these foraging phases. Nonhosts feeding on a neighbouring plant or on the same plant individual the host is feeding from may affect odour-based searching by parasitoids in a way specific for the species combination studied. Feeding by specific host and nonhost–herbivore combinations may induce volatiles that are more, less or equally attractive compared to those from plants infested by the host only. Within the food patch, mixed presence of host and nonhost may reduce the number of hosts parasitized per time unit and reduce parasitoid foraging efficiency. Importantly, we show that a single nonhost species may have contrasting effects in terms of its effects on odour-based searching and patch residence decisions. We conclude that studying host searching behaviour at both phases of foraging is essential for our understanding of parasitoid foraging behaviour in natural and agricultural settings. We further speculate on the ecological context in which unsuitable herbivores affect either of the two phases of parasitoid foraging. [ABSTRACT FROM AUTHOR]

Herbivore-induced plant volatiles (HIPV) provide important information that influences host location behaviour for insect natural enemies, such as parasitoid wasps, that develop in the bodies of herbivorous insects. The dietary breadth of both the parasitoid and its host may affect the extent to which a searching parasitoid relies on HIPV. Specialist species are expected to rely on specific volatile cues to which they respond innately, whereas generalists are expected to show a higher degree of phenotypic plasticity that depends on foraging experience in the parasitoid. We compared the response to HIPV emitted by different plant species damaged by host and nonhost caterpillars for two congeneric parasitoid species, the specialist Diadegma semiclausum and the generalist Diadegma fenestrale, attacking caterpillars of the diamondback moth, Plutella xylostella. For the three tested plant species, Brassica oleracea, a feral Brassica population and Sinapis alba, both parasitoid species preferred volatiles from host-infested plants over those produced by undamaged plants. However, both parasitoid species only distinguished between volatiles induced by host and nonhosts when the caterpillars had been feeding on B. oleracea, the plant on which they had been reared. Chemical analysis of the volatile blends could not explain volatile preferences of the parasitoids. Despite the difference in their dietary breadth, the two parasitoids responded similarly to HIPV and experience treatments. A flexible response to a wide array of volatile blends by parasitoids is probably important in nature, given that different generations of the host and the parasitoid probably develop on different food plants. [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]

Abstract: We evaluated five herbivore-induced plant volatiles plus a male-produced pheromone as attractants for adult green lacewings in Washington apple orchards in 2008. We found at least five attractants or combinations of attractants were attractive to the three most abundant green lacewing species in our trials. Chrysopa nigricornis and Chrysopa oculata were attracted to the combination of methyl salicylate and iridodial with iridodial alone being the second best attractant. Chrysoperla plorabunda was found in lower numbers than C. nigricornis and C. oculata, but did exhibit a significant attraction to benzaldehyde. In mid-summer, we added the herbivore-induced plant volatile squalene to the study and found it to be exceedingly attractive, but only to male C. nigricornis. Whether alone or in combination, squalene attracted 4–5-fold more C. nigricornis than any other compound tested. Our data have revealed C. nigricornis to be an abundant orchard predator that can be readily monitored with squalene-baited traps. Despite the obvious promise of HIPVs in biological control programs, we urge caution in their deployment as large-scale attractants, at least until further studies have investigated potential disruption of natural enemy population dynamics. [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]