Bumblebee electric charge stimulates floral volatile emissions in Petunia integrifolia but not in Antirrhinum majus.

The timing of volatile organic compound (VOC) emission by flowering plants often coincides with pollinator foraging activity. Volatile emission is often considered to be paced by environmental variables, such as light intensity, and/or by circadian rhythmicity. The question arises as to what extent pollinators themselves provide information about their presence, in keeping with their long co-evolution with flowering plants. Bumblebees are electrically charged and provide electrical stimulation when visiting plants, as measured via the depolarisation of electric potential in the stem of flowers. Here we test the hypothesis that the electric charge of foraging bumblebees increases the floral volatile emissions of bee pollinated plants. We investigate the change in VOC emissions of two bee-pollinated plants (Petunia integrifolia and Antirrhinum majus) exposed to the electric charge typical of foraging bumblebees. P. integrifolia slightly increases its emissions of a behaviorally and physiologically active compound in response to visits by foraging bumblebees, presenting on average 121 pC of electric charge. We show that for P. integrifolia, strong electrical stimulation (600-700 pC) promotes increased volatile emissions, but this is not found when using weaker electrical charges more representative of flying pollinators (100 pC). Floral volatile emissions of A. majus were not affected by either strong (600-700 pC) or weak electric charges (100 pC). This study opens a new area of research whereby the electrical charge of flying insects may provide information to plants on the presence and phenology of their pollinators. As a form of electroreception, this sensory process would bear adaptive value, enabling plants to better ensure that their attractive chemical messages are released when a potential recipient is present.

Identification of Semiochemicals from Cowpea, Vigna unguiculata, for Low-input Management of the Legume Pod Borer, Maruca vitrata.

Cowpea, Vigna unguiculata L. Walp. (Fabaceae), is one of the most important food legumes grown on the African continent, as it provides an affordable source of dietary protein. Yields of cowpea are significantly reduced through damage by legume pod-borer, Maruca vitrata (Lepidoptera: Crambidae), caterpillars to flowers, tender leaves and pods. Semiochemical-based strategies are considered as environmentally benign and affordable for pest management, particularly on smallholder farms. In this study, we investigated the importance of cowpea flower volatiles as host location cues for egg-laying M. vitrata, and herbivore-induced plant volatiles (HIPVs) as M. vitrata repellents and natural enemy (Apanteles taragamae and Phanerotoma syleptae parasitoid) attractants. In oviposition choice assays, M. vitrata laid more eggs on flowering cowpea plants than non-flowering plants. Coupled gas chromatography-electrophysiology (GC-EAG) analysis using the antennae of female M. vitrata and an extract of flower volatiles collected by dynamic headspace collection revealed the presence of five EAG-active components that were identified by coupled GC-mass spectrometry (GC-MS) analysis as benzaldehyde, benzyl alcohol, acetophenone, a vinylbenzaldehyde isomer and (E)-cinnamaldehyde. A synthetic blend of the identified compounds, prepared using 3-vinylbenzaldehyde, induced M. vitrata to lay as many eggs on non-flowering cowpea as on flowering plants. The moths also preferred laying eggs on intact plants compared to M. vitrata-infested plants. As the emission of EAG-active floral compounds was determined to be lower in the headspace of infested cowpea flowers, the role of HIPVs emitted by M. vitrata-damaged leaves was also investigated. Of the compounds induced by larval damage, (E)-DMNT, indole, n-hexyl acetate, 1-octen-3-ol and linalool were shown by GC-EAG to possess electrophysiological activity. A synthetic blend of the EAG-active compounds, using racemic 1-octen-3-ol and linalool, significantly reduced egg numbers on flowering cowpea. Larval and egg parasitoids, i.e. A. taragamae and Ph. syleptae, respectively, of M. vitrata both preferred the Y-tube olfactometer arm treated with synthetic (E)-DMNT, whereas preference for racemic linalool and (E)-nerolidol was dose-dependent in A. taragamae. Our results provide the platform for the development of future semiochemical-based pest management strategies against M. vitrata on smallholder farms in West Africa.

Molasses Grass Induces Direct and Indirect Defense Responses in Neighbouring Maize Plants.

Plants have evolved intricate defence strategies against herbivore attack which can include activation of defence in response to stress-related volatile organic compounds (VOCs) emitted by neighbouring plants. VOCs released by intact molasses grass (Melinis minutiflora), have been shown to repel stemborer, Chilo partellus (Swinhoe), from maize and enhance parasitism by Cotesia sesamiae (Cameron). In this study, we tested whether the molasses grass VOCs have a role in plant-plant communication by exposing different maize cultivars to molasses grass for a 3-week induction period and then observing insect responses to the exposed plants. In bioassays, C. partellus preferred non-exposed maize landrace plants for egg deposition to those exposed to molasses grass. Conversely, C. sesamiae parasitoid wasps preferred volatiles from molasses grass exposed maize landraces compared to volatiles from unexposed control plants. Interestingly, the molasses grass induced defence responses were not observed on hybrid maize varieties tested, suggesting that the effect was not simply due to absorption and re-emission of VOCs. Chemical and electrophysiological analyses revealed strong induction of bioactive compounds such as (R)-linalool, (E)-4,8-dimethyl-1,3,7-nonatriene and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene from maize landraces exposed to molasses grass volatiles. Our results suggest that constitutively emitted molasses grass VOCs can induce direct and indirect defence responses in neighbouring maize landraces. Plants activating defences by VOC exposure alone could realize enhanced levels of resistance and fitness compared to those that launch defence responses upon herbivore attack. Opportunities for exploiting plant-plant signalling to develop ecologically sustainable crop protection strategies against devastating insect pests such as stemborer C. partellus are discussed.

(2R,5S)-Theaspirane Identified as the Kairomone for the Banana Weevil, Cosmopolites sordidus, from Attractive Senesced Leaves of the Host Banana, Musa spp.

The principal active component produced by highly attractive senesced host banana leaves, Musa spp., for the banana weevil, Cosmopolites sordidus, is shown by coupled gas chromatography-electroantennography (GC-EAG), coupled GC-mass spectrometry (GC-MS), chemical synthesis and coupled enantioselective (chiral) GC-EAG to be (2R,5S)-theaspirane. In laboratory behaviour tests, the synthetic compound is as attractive as natural host leaf material and presents a new opportunity for pest control.

cis-Jasmone Elicits Aphid-Induced Stress Signalling in Potatoes.

Elicitation of plant defense signaling that results in altered emission of volatile organic compounds (VOCs) offers opportunities for protecting plants against arthropod pests. In this study, we treated potato, Solanum tuberosum L., with the plant defense elicitor cis-jasmone (CJ), which induces the emission of defense VOCs and thus affects the behavior of herbivores. Using chemical analysis, electrophysiological and behavioral assays with the potato-feeding aphid Macrosiphum euphorbiae, we showed that CJ treatment substantially increased the emission of defense VOCs from potatoes compared to no treatment. Coupled GC-electroantennogram (GC-EAG) recordings from the antennae of M. euphorbiae showed robust responses to 14 compounds present in induced VOCs, suggesting their behavioral role in potato/aphid interactions. Plants treated with CJ and then challenged with M. euphorbiae were most repellent to alate M. euphorbiae. Principal component analysis (PCA) of VOC collections suggested that (E)-2-hexenal, (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene (TMTT), (E)-ß-farnesene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), methyl salicylate (MeSA), CJ, and methyl benzoate (MeBA) were the main VOCs contributing to aphid behavioral responses, and that production of TMTT, (E)-ß-farnesene, CJ, and DMNT correlated most strongly with aphid repellency. Our findings confirm that CJ can enhance potato defense against aphids by inducing production of VOCs involved in aphid-induced signalling.

Electrophysiological and behavioral responses of female African rice gall midge, Orseolia oryzivora Harris and Gagné, to host plant volatiles.

African rice gall midge, Orseolia oryzivora Harris and Gagné, is a major pest of rice in Africa. Depsite its economic importance, its chemical ecology is not well understood. Here, we assessed behavioral and electrophysiological responses of O. oryzivora to host plant volatiles. In olfactometer bioassays, mated female O. oryzivora were attracted to volatiles emitted from intact rice plants but were repelled by volatiles collected from plants infested by conspecifics. In a choice test, there was a preference for volatiles from uninfested plants over those from infested plants. Coupled gas chromatography-electroantennography analyses of panicle volatiles isolated four electrophysiologically active components: (S)-linalool, 4,8-dimethyl-1,3,7-nonatriene, (E)-caryophyllene, and (R/S)-(E)-nerolidol. A synthetic blend of volatiles at the same concentration and ratio as that from an intact plant was attractive to mated females, whereas a blend based on the ratio of volatiles from an infested plant was repellent. This suggests that O. oryzivora uses olfaction for host plant recognition. The identification of blends of volatiles emitted by plants that can both attract and repel O. oryzivora may aid the development of sustainable control measures.

Sharing a Host Plant (Wheat [Triticum aestivum]) Increases the Fitness of Fusarium graminearum and the Severity of Fusarium Head Blight but Reduces the Fitness of Grain Aphids (Sitobion avenae).

We hypothesized that interactions between fusarium head blight-causing pathogens and herbivores are likely to occur because they share wheat as a host plant. Our aim was to investigate the interactions between the grain aphid, Sitobion avenae, and Fusarium graminearum on wheat ears and the role that host volatile chemicals play in mediating interactions. Wheat ears were treated with aphids and F. graminearum inoculum, together or separately, and disease progress was monitored by visual assessment and by quantification of pathogen DNA and mycotoxins. Plants exposed to both aphids and F. graminearum inoculum showed accelerated disease progression, with a 2-fold increase in disease severity and 5-fold increase in mycotoxin accumulation over those of plants treated only with F. graminearum. Furthermore, the longer the period of aphid colonization of the host prior to inoculation with F. graminearum, the greater the amount of pathogen DNA that accumulated. Headspace samples of plant volatiles were collected for use in aphid olfactometer assays and were analyzed by gas chromatography-mass spectrometry (GC-MS) and GC-coupled electroantennography. Disease-induced plant volatiles were repellent to aphids, and 2-pentadecanone was the key semiochemical underpinning the repellent effect. We measured aphid survival and fecundity on infected wheat ears and found that both were markedly reduced on infected ears. Thus, interactions between F. graminearum and grain aphids on wheat ears benefit the pathogen at the expense of the pest. Our findings have important consequences for disease epidemiology, because we show increased spread and development of host disease, together with greater disease severity and greater accumulation of pathogen DNA and mycotoxin, when aphids are present.

Responses of parasitoids to volatiles induced by Chilo partellus oviposition on teosinte, a wild ancestor of maize.

Maize, a genetically diverse crop, is the domesticated descendent of its wild ancestor, teosinte. Recently, we have shown that certain maize landraces possess a valuable indirect defense trait not present in commercial hybrids. Plants of these landraces release herbivore-induced plant volatiles (HIPVs) that attract both egg [Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)] and larval [Cotesia sesamiae Cameron (Hymenoptera: Braconidae)] parasitoids in response to stemborer egg deposition. In this study, we tested whether this trait also exists in the germplasm of wild Zea species. Headspace samples were collected from plants exposed to egg deposition by Chilo partellus Swinhoe (Lepidoptera: Crambidae) moths and unexposed control plants. Four-arm olfactometer bioassays with parasitic wasps, T. bournieri and C. sesamiae, indicated that both egg and larval parasitoids preferred HIPVs from plants with eggs in four of the five teosinte species sampled. Headspace samples from oviposited plants released higher amounts of EAG-active compounds such as (E)-4,8-dimethyl-1,3,7-nonatriene. In oviposition choice bioassays, plants without eggs were significantly preferred for subsequent oviposition by moths compared to plants with prior oviposition. These results suggest that this induced indirect defence trait is not limited to landraces but occurs in wild Zea species and appears to be an ancestral trait. Hence, these species possess a valuable trait that could be introgressed into domesticated maize lines to provide indirect defense mechanisms against stemborers.

Semiochemistry of the Scarabaeoidea.

The superfamily Scarabaeoidea comprises a large and diverse monophyletic group. Members share ancestral characteristics, but often exhibit considerable differences in their ecology, physiology, or mating strategies. A large number of species are regarded as pests of crop or amenity plants, while others are beneficial to humans and even may be extremely rare as a result of anthropogenic activities. A significant number of chemical ecology-based studies have been conducted with the Scarabaeoidea in order to characterize semiochemicals influencing their behavior, such as pheromones and plant-derived allelochemicals. These may be used either to control or preserve populations of the beetles, depending upon pest or beneficial status. This paper is a review of the role and identity of the semiochemicals of the Scarabaeoidea, with comments on possible future research and applied opportunities in the field of chemical ecology.

Novel pheromone-mediated reproductive behaviour in the stag beetle, Lucanus cervus.

The iconic European stag beetle (Lucanus cervus) (Coleoptera: Lucanidae) is one of the largest terrestrial beetles in Europe. Due to decreasing population numbers, thought to be a consequence of habitat loss, this beetle has become a near-threatened species across much of Europe, and a reliable monitoring system is required to measure its future population trends. As part of a programme aimed at conserving UK populations, we have investigated the chemical ecology of the beetle, with a view to developing an efficient semiochemical-based monitoring system. Such a scheme will be beneficial not only in the UK but across the European range of the species, where the beetle is of conservation concern. Here, we report on a surprising discovery of a male-produced pheromone, which provokes initial sexual receptivity in females, and which has not been previously identified in the animal kingdom. Furthermore, we assign sex pheromone function to a previously described female-specific compound.

Underground signals carried through common mycelial networks warn neighbouring plants of aphid attack.

The roots of most land plants are colonised by mycorrhizal fungi that provide mineral nutrients in exchange for carbon. Here, we show that mycorrhizal mycelia can also act as a conduit for signalling between plants, acting as an early warning system for herbivore attack. Insect herbivory causes systemic changes in the production of plant volatiles, particularly methyl salicylate, making bean plants, Vicia faba, repellent to aphids but attractive to aphid enemies such as parasitoids. We demonstrate that these effects can also occur in aphid-free plants but only when they are connected to aphid-infested plants via a common mycorrhizal mycelial network. This underground messaging system allows neighbouring plants to invoke herbivore defences before attack. Our findings demonstrate that common mycorrhizal mycelial networks can determine the outcome of multitrophic interactions by communicating information on herbivore attack between plants, thereby influencing the behaviour of both herbivores and their natural enemies.

Do asexual morphs of the peach-potato aphid, Myzus persicae, utilise the aphid sex pheromone? Behavioural and electrophysiological responses of M. persicae virginoparae to (4aS,7S,7aR)-nepetalactone and its effect on aphid performance.

The aphid sex pheromone component (4aS,7S,7aR)-nepetalactone is considered to be a potential tool for enhancing biological control of aphids. Studies have confirmed its potential to attract parasitoids, increase parasitism rates in the field and also alter the spatial distribution of parasitoids. An important aspect that has been overlooked is the impact that the introduction of nepetalactone may have on aphid populations already present in field or glasshouse environments. The most prevalent pest aphid populations in glasshouse and field environments are the asexual morphs, which are capable of exponential growth if populations are not controlled. The short-term implications of the sex pheromone on asexual aphids were observed through their behavioural response. Using Y-tube olfactometry, it is shown that virginoparae of the peach-potato aphid, Myzus persicae, are repelled by high concentrations of nepetalactone. Long-term effects of the pheromone which may span the aphid's life, or even generations, were assessed via mean relative growth rate (MRGR) and the intrinsic rate of natural increase (rm). Electroantennography also demonstrated that asexual female aphids are able to detect aphid sex pheromone components. To our knowledge, this is the first time it has been reported that M. persicae virginoparae are able to detect aphid sex pheromone components or that their behavioural response and/or performance has been studied. The implications of these results and their significance in understanding semiochemical communication are discussed.

Isolation and identification of host cues from mango, Mangifera indica, that attract gravid female oriental fruit fly, Bactrocera dorsalis.

The oriental fruit fly, Bactrocera dorsalis, is an economically damaging, polyphagous pest of fruit crops in South-East Asia and Hawaii, and a quarantine pest in other parts of the world. The objective of our study was to identify new attractants for B. dorsalis from overripe mango fruits. Headspace samples of volatiles were collected from two cultivars of mango, 'Alphonso' and 'Chausa', and a strong positive behavioral response was observed when female B. dorsalis were exposed to these volatiles in olfactometer bioassays. Coupled GC-EAG with female B. dorsalis revealed 7 compounds from 'Alphonso' headspace and 15 compounds from 'Chausa' headspace that elicited an EAG response. The EAG-active compounds, from 'Alphonso', were identified, using GC-MS, as heptane, myrcene, (Z)-ocimene, (E)-ocimene, allo-ocimene, (Z)-myroxide, and γ-octalactone, with the two ocimene isomers being the dominant compounds. The EAG-active compounds from 'Chausa' were 3-hydroxy-2-butanone, 3-methyl-1-butanol, ethyl butanoate, ethyl methacrylate, ethyl crotonate, ethyl tiglate, 1-octen-3-ol, ethyl hexanoate, 3-carene, p-cymene, ethyl sorbate, α-terpinolene, phenyl ethyl alcohol, ethyl octanoate, and benzothiazole. Individual compounds were significantly attractive when a standard dose (1 µg on filter paper) was tested in the olfactometer. Furthermore, synthetic blends with the same concentration and ratio of compounds as in the natural headspace samples were highly attractive (P < 0.001), and in a choice test, fruit flies did not show any preference for the natural samples over the synthetic blends. Results are discussed in relation to developing a lure for female B. dorsalis to bait traps with.

Identification of host kairomones from maize, Zea mays, for the maize weevil, Sitophilus zeamais.

The maize weevil, Sitophilus zeamais, is an economically important pest of stored grains in tropical and subtropical regions of the world. The behavioral responses of adult S. zeamais to volatile organic compounds (VOCs) from maize seeds, Zea mays, were studied to identify semiochemicals used in host location and provide potential tools for managing this pest. VOCs released directly from crude seed extracts, vacuum distilled extracts, hexane and diethyl ether fractions from silica gel chromatography of the vacuum distillates, air entrainment samples, and identified volatile compounds were assayed using a Perspex four-arm olfactometer. Weevils spent significantly more time, and made a higher number of visits, to the region of the olfactometer where Z. mays volatiles were present than in control regions comprising solvent only. When white and yellow Z. mays VOCs were compared in a choice test, the mean time spent in the two olfactometer treatment arms was significantly greater than the mean time spent in the control arms. However, weevils did not show any preference for either of the two treatments, thus demonstrating that both varieties of maize have similar activity. Gas chromatography (GC), coupled gas chromatography-mass spectrometry (GC-MS), GC peak enhancement and electroantennography (EAG) identified hexanal, (E)-2-heptenal, and octanal as biologically active compounds in air entrainment samples and diethyl ether fractions of vacuum distillates. A 3-component synthetic blend of the identified compounds was significantly attractive to both sexes of the weevil. These host kairomones could be deployed in semiochemical based monitoring and management of S. zeamais in the tropics.

Oviposition induced volatile emissions from African smallholder farmers' maize varieties.

Maize (corn), Zea mays, is a genetically diverse crop, and we have recently shown that certain open pollinated varieties (OPVs) of Latin American origin possess a trait not present in mainstream commercial varieties: they produce volatiles in response to stemborer oviposition that are attractive to stemborer parasitoids. Here, we tested whether a similar tritrophic effect occurs in the African OPVs 'Nyamula' and 'Jowi'. Herbivore induced plant volatiles (HIPVs) were collected from plants exposed to egg deposition by the stemborer Chilo partellus. In a four-arm olfactometer bioassay, the parasitic wasp Cotesia sesamiae preferred samples containing HIPVs from plants with eggs to samples collected from plants without eggs. EAG-active compounds, including (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), were released in higher amounts from the egg induced headspace samples. Our results suggest that this oviposition trait is not limited to S. American Z. mays germplasm, and that it could be used to increase indirect defense against attack by stemborers.

Maize landraces recruit egg and larval parasitoids in response to egg deposition by a herbivore.

Natural enemies respond to herbivore-induced plant volatiles (HIPVs), but an often overlooked aspect is that there may be genotypic variation in these 'indirect' plant defence traits within plant species. We found that egg deposition by stemborer moths (Chilo partellus) on maize landrace varieties caused emission of HIPVs that attract parasitic wasps. Notably, however, the oviposition-induced release of parasitoid attractants was completely absent in commercial hybrid maize varieties. In the landraces, not only were egg parasitoids (Trichogramma bournieri) attracted but also larval parasitoids (Cotesia sesamiae). This implies a sophisticated defence strategy whereby parasitoids are recruited in anticipation of egg hatching. The effect was systemic and caused by an elicitor, which could be extracted from egg materials associated with attachment to leaves. Our findings suggest that indirect plant defence traits may have become lost during crop breeding and could be valuable in new resistance breeding for sustainable agriculture.

Identification of semiochemicals released by cotton, Gossypium hirsutum, upon infestation by the cotton aphid, Aphis gossypii.

The cotton aphid, Aphis gossypii (Homoptera: Aphididae), is increasing in importance as a pest worldwide since the introduction of Bt-cotton, which controls lepidopteran but not homopteran pests. The chemical ecology of interactions between cotton, Gossypium hirsutum (Malvaceae), A. gossypii, and the predatory lacewing Chrysoperla lucasina (Neuroptera: Chrysopidae), was investigated with a view to providing new pest management strategies. Behavioral tests using a four-arm (Pettersson) olfactometer showed that alate A. gossypii spent significantly more time in the presence of odor from uninfested cotton seedlings compared to clean air, but significantly less time in the presence of odor from A. gossypii infested plants. A. gossypii also spent significantly more time in the presence of headspace samples of volatile organic compounds (VOCs) obtained from uninfested cotton seedlings, but significantly less time with those from A. gossypii infested plants. VOCs from uninfested and A. gossypii infested cotton seedlings were analyzed by gas chromatography (GC) and coupled GC-mass spectrometry (GC-MS), leading to the identification of (Z)-3-hexenyl acetate, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), methyl salicylate, and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene (TMTT), which were produced in larger amounts from A. gossypii infested plants compared to uninfested plants. In behavioral tests, A. gossypii spent significantly more time in the control (solvent) arms when presented with a synthetic blend of these four compounds, with and without the presence of VOCs from uninfested cotton. Coupled GC-electroantennogram (EAG) recordings with the lacewing C. lucasina showed significant antennal responses to VOCs from A. gossypii infested cotton, suggesting they have a role in indirect defense and indicating a likely behavioral role for these compounds for the predator as well as the aphid.

cis-Jasmone induces Arabidopsis genes that affect the chemical ecology of multitrophic interactions with aphids and their parasitoids.

It is of adaptive value for a plant to prepare its defenses when a threat is detected, and certain plant volatiles associated with insect damage, such as cis-jasmone (CJ), are known to switch-on defense metabolism. We used aphid and aphid parasitoid responses to Arabidopsis thaliana as a model system for studying gene expression and defense chemistry and its impact at different trophic levels. Differential responses to volatiles of induced Arabidopsis occurred for specialist and generalist insects: the generalist aphid, Myzus persicae, was repelled, whereas the specialist, Lipaphis erysimi, was attracted; the generalist aphid parasitoid Aphidius ervi was attracted, but the specialist parasitoid Diaeretiella rapae was not affected. A. ervi also spent longer foraging on induced plants than on untreated ones. Transcriptomic analyses of CJ-induced Arabidopsis plants revealed that a limited number of genes, including a gene for a cytochrome P450, CYP81D11, were strongly up-regulated in the treated plants. We examined transgenic Arabidopsis lines constitutively overexpressing this gene in bioassays and found insect responses similar to those obtained for wild-type plants induced with CJ, indicating the importance of this gene in the CJ-activated defense response. Genes involved in glucosinolate biosynthesis and catabolism are unaffected by CJ and, because these genes relate to interactions with herbivores and parasitoids specific to this family of plants (Brassicaceae), this finding may explain the differences in behavioral response of specialist and generalist insects.

Chemical Identity and Functional Characterization of Semiochemicals That Promote the Interactions between Rice Plant and Rice Major Pest Nilaparvata lugens.

The interaction between food crops and insect pests is mediated by semiochemicals emitted from host plants. These semiochemicals are natural behavioral modifiers and act on the insect olfactory system to locate hosts and preys. In this study, eight rice neuroactive semiochemicals were identified from rice varieties by GC-EAG and GC-MS. Their ability to modify rice pest behaviors was further studied as individual chemicals and physiologically relevant blend. The total amount of each semiochemical and the expression of their biosynthesis genes were significantly higher in pest susceptible variety than in pest-resistant variety and upregulated by the infestation of the pest Nilaparvata lugens (BPH). The semiochemicals emitted by uninfested plants (UIRVs) were more attractive to BPHs. Interestingly, the attractiveness of UIRVs was significantly reduced by the addition of the blend that mimics the natural composition of these semiochemicals emitted by infested plants (IRVs). Our study suggests a mechanism for the spread of pest infestation from infested plants to uninfested plants nearby. UIRVs initially serve as attractive signals to rice insect pests. The pest infestation changes the rice semiochemical profile to be less attractive or even repellent, which pushes further colonization to uninfested plants nearby. The identified semiochemicals can be used for crop protection based on a push-pull strategy.

Electrophysiological and behavioral responses of sorghum shoot fly, Atherigona soccata, to sorghum volatiles.

The sorghum shoot fly, Atherigona soccata, is an economically important pest of sorghum in Asia, Mediterranean Europe and Africa. Field observations have suggested that shoot fly susceptible sorghum varieties emit attractive volatiles, but the compounds involved were unknown. The objective of the present study was to identify plant-derived attractants for A. soccata. Headspace samples were collected from the susceptible cultivar 'Swarna,' and when female A. soccata were exposed to the volatiles in an olfactometer bioassay, a strong positive behavioral response was observed. Coupled GC-EAG with female A. soccata revealed eight compounds that elicited an EAG response, which were identified by coupled GC-MS and GC peak enhancement on two GC columns of different polarity as (Z)-3-hexen-1-yl acetate, (-)-α-pinene, (-)-(E)-caryophyllene, methyl salicylate, octanal, decanal, 6-methyl-5-hepten-2-one and nonanal. When an eight-component synthetic blend of the EAG active compounds, at the same concentration (2.64 µg) and ratio as in the natural headspace sample, was tested, A. soccata spent more time in the treated region of the olfactometer than controls (P = 0.001). Furthermore, when this synthetic blend and the natural headspace sample were tested in a choice test, the shoot flies did not show any preference for either of the two treatments, demonstrating that the synthetic blend had similar activity to the natural sample. Results are discussed in relation to breeding sorghum varieties less attractive to this pest.