The plant sesquiterpene germacrene D specifically activates a major type of antennal receptor neuron of the tobacco budworm moth Heliothis virescens

Plants release hundreds of volatiles that are important in interactions with insects or other organisms. However, knowledge is scarce as to which of the compounds are detected by the organism's olfactory receptor neurons. In the present study, single receptor neurons on the antennae of the tobacco budworm moth, Heliothis virescens, were screened for their sensitivities to naturally produced plant volatiles by the use of gas chromatography linked to electrophysiological recordings from single cells (GC-SCR). Plant volatiles, collected by aeration of host and non-host plants, were tested on each receptor neuron via parallel GC-columns. Thus, simultaneous recordings of the gas chromatogram and the neuron responses to each component were obtained. One type of receptor neuron, appearing in 80% of all experiments, responded with high sensitivity and selectivity to one particular component, present in host as well as non-host mixtures. The component, identified as a sesquiterpene hydrocarbon by linked gas chromatography-mass spectrometry, was isolated from a sesquiterpene fraction of cubebe oil and identified by NMR as germacrene D. The purified compound was then re-tested via gas chromatography on the same receptor neuron type, verifying the identification. A weaker response to another sesquiterpene hydrocarbon was also recorded.     

Receptor neuron discrimination of the germacrene D enantiomers in the moth Helicoverpa armigera

Plants release complex mixtures of volatiles, including chiral constituents. In the search for the biologically relevant plant odorants, gas chromatography linked to electrophysiological recordings from single receptor neurons has been employed. In heliothine moths, including the females of the Eurasian cotton bollworm moth Helicoverpa armigera, a major type of receptor neurons is identified, showing high sensitivity and selectivity for the sesquiterpene germacrene D. In the present study, gas chromatography with a chiral column linked to single cell recordings were performed. It was found that all germacrene D neurons belonged to one type; all responded to both enantiomers, but (-)-germacrene D had approximately 10 times stronger effect than (+)-germacrene D. Parallel dose-response curves for the two enantiomers were obtained by direct stimulations. The enantiomeric composition of germacrene D, which differed in six plant species and in different individuals of one species, was determined on the basis of the neuron responses. The results, showing the presence of one neuron type for receiving the information about germacrene D in the various plants, suggests that the two enantiomers mediate the same kind of information to the moth, but with different intensity.     

Detection of fruit- and flower-emitted volatiles by olfactory receptor neurons in the polyphagous fruit chafer Pachnoda marginata (Coleoptera: Cetoniinae)

Olfactory receptor neurons on the antennae of the African fruit chafer species Pachnoda marginata (Coleoptera: Scarabaeidae) were examined through extensive use of gas chromatography linked with electrophysiological recordings from single olfactory receptor neurons. Contacted neurons were stimulated with a large number of extracted volatiles from 22 different fruits and with 64 synthetic plant compounds. Extracted fruit volatiles were identified using linked gas chromatography-mass spectrometry. In total, 48 different odor compounds were found to elicit responses. Analysis of the response spectra of the contacted neurons (n = 232) revealed the presence of 28 classes of receptor neurons. The neurons exhibited strong selectivity as well as high sensitivity. Eleven of the identified classes were selectively activated by single compounds, while the remaining were activated by 2-6 compounds. Several receptor neurons that were activated by more than one compound responded to compounds sharing basic structural similarities. The results support the growing hypothesis that a significant proportion of plant-odor receptor neurons in insects are highly sensitive and selective for single odors.     

Chemical signatures affecting host choice in the Eucalyptus herbivore,Gonipterus sp. (Curculionidae: Coleoptera)

It is well known that herbivorous insects respond to host plant volatiles. Yet details of how these insects perceive the complex profile of volatiles from different potential host plants have not been studied for most insects. Gonipterus spp. are important pests of Eucalyptus worldwide, but differ in their preference for different species of this host. In this study, we consider whether host volatiles affect the host choice for a Gonipterus sp. and we characterize the response of the female insect to the volatile profiles from these hosts in an electro-antennographic experiment. We sampled volatiles from freshly damaged leaves of three Eucalyptus species and analysed the profiles by gas chromatography coupled to electro-antennography (GC-EAD) and gas chromatography coupled to mass spectrometry. Female weevils gave a mixed range of electro-physiological responses to volatile puffs from leaves of different tree species. This suggests that differences in volatile profiles of different trees play a role in how these beetles discriminate between potential hosts. GC-EAD analysis showed that responses were as complex as the volatile chemical compositions of the leaves. A number of these chemicals were identified, and responses were mostly due to general green leaf volatiles. This was also evident from the fact that the insects showed a markedly greater response to the total volatile profile from freshly damaged leaves for all species. The females of the Gonipterus sp. can therefore detect damaged leaves, which may indicate host quality. Host specificity information is further expected to lie in the relative differences in emission ratios and synergism between different host chemical compounds, rather than specific individual compounds.     

Discrimination between enantiomers of linalool by olfactory receptor neurons in the cabbage moth Mamestra brassicae (L.)

Plants emit complex blends of volatiles, including chiral compounds that might be detected by vertebrates and invertebrates. Insects are ideal model organisms for studying the underlying receptor neuron mechanisms involved in olfactory discrimination of enantiomers. In the present study, we have employed two-column gas chromatography linked to recordings from single olfactory receptor neurons of Mamestra brassicae, in which separation of volatiles in a polar and a chiral column was performed. We here present the response properties of olfactory receptor neurons tuned to linalool. The narrow tuning of these receptor neurons was demonstrated by their strong responses to (R)-(-)-linalool, the weaker responses to the (+)-enantiomer as well as a few structurally related compounds, and no responses to the other numerous plant released volatiles. The enantioselectivity was verified by parallel dose-response curves, that of (R)-(-)-linalool shifted 1 log unit to the left of the (S)-(+)-linalool curve. A complete overlap of the temporal response pattern was found when comparing the responses of the same strength. Analysis of the spike amplitude and waveform indicated that the responses to the two enantiomers originated from the same neuron.     

Selective receptor neurone responses to E-β-ocimene, β-myrcene, E,E-α-farnesene and homo-farnesene in the moth Heliothis virescens, identified by gas chromatography linked to electrophysiology

An important question in olfaction is for which odorants receptor neurones have evolved. In the present study, olfactory receptor neurones on the antennae of the tobacco budworm moth Heliothis virescens were screened for sensitivity to naturally occurring plant-produced volatiles by the use of gas chromatography linked to electrophysiology. Volatiles of host as well as non-host plants collected by headspace techniques were used for stimulating the neurones, sequentially via two columns, one polar and one nonpolar installed in parallel in the gas chromatograph. Three types of neurones presented in this paper responded to one, two or three compounds for which the retention times were determined in both column types. The chemical structures of the active components were determined on the basis of mass spectrometry linked to gas chromatography, indicating E-beta-ocimene and beta-myrcene as stimulants for neurone type 1, E,E-alpha-farnesene for neurone type 2 and homo-farnesene for neurone type 3. Re-testing authentic materials verified the identifications for the type 1 neurones. The results demonstrate a high specificity for the three types of neurones by strong responses to one or two structurally similar compounds out of hundreds present in a large variety of plants. The study exemplifies plant odour detection by narrowly tuned receptor neurones in a polyphagous moth species.     

华北大黑鳃金龟对桃树挥发物的电生理反应

【目的】华北大黑鳃金龟Holotrichia oblita是一种分布范围广,寄主种类多的农业害虫。本研究鉴定了桃树顶空挥发物中对华北大黑鳃金龟雌虫具有触角电生理活性的化合物,并探索桃树挥发物活性成分的剂量对华北大黑鳃金龟雌、雄虫触角电生理反应的影响。【方法】采用顶空吸附法收集桃树枝叶挥发物,用气相色谱 触角电位联用技术(GC-EAD)和气相色谱 质谱联用技术(GC-MS)鉴定有电生理活性的桃树挥发物成分,最后用触角电位(electroantennograph, EAG)技术测试这些挥发物在0.01,0.1, 1, 10和100 μg剂量下引起的华北大黑鳃金龟雌、雄成虫EAG反应。【结果】桃树挥发物中能引起华北大黑鳃金龟雌成虫触角电生理反应的成分有7种,包括Z-β-罗勒烯、顺-3-己烯乙酸酯、顺-3-己烯醇、壬醛、水杨酸甲酯以及2种未知化合物。所测试的对华北大黑鳃金龟雌成虫触角具有电生理活性的5种桃树挥发物成分在各剂量下都能引起雌、雄成虫明显的EAG反应。雌、雄虫对同种物质的EAG反应随着剂量的增加而有增大的趋势,并且雌虫的EAG反应显著高于雄虫。【结论】桃树挥发物中,罗勒烯、顺-3-己烯乙酸酯、顺-3-己烯醇、壬醛、水杨酸甲酯均能引起华北大黑鳃金龟雌、雄成虫的剂量依赖性的触角电生理反应,而且存在显著的性别差异。可利用这些化合物开展行为反应和田间诱集试验,进一步验证其对华北大黑鳃金龟的引诱活性。     

Identification of olfactory receptor neurons in Uraba lugens (Lepidoptera: Nolidae) and its implications for host range

Phytophagous insects detect volatile compounds produced by host and non-host plants, using species-specific sets of olfactory receptor neurons (ORNs). To investigate the relationship between the range of host plants and the profile of ORNs, single sensillum recordings were carried out to identify ORNs and corresponding active compounds in female Uraba lugens (Lepidoptera: Nolidae), an oligophagous eucalypt feeder. Based on the response profiles to 39 plant volatile compounds, 13 classes of sensilla containing 40 classes of ORNs were identified in female U. lugens. More than 95% (163 out of 171) of these sensilla contained 16 classes of ORNs with narrow response spectra, and 62.6% (107 out of 171) 18 classes of ORNs with broad response spectra. Among the specialized ORNs, seven classes of ORNs exhibited high specificity to 1,8-cineole, (±)-citronellal, myrcene, (±)-linalool and (E)-β-caryophyllene, major volatiles produced by eucalypts, while nine other classes of ORNs showed highly specialized responses to green leaf volatiles, germacrene D, (E)-β-farnesene and geranyl acetate that are not produced by most eucalypts. We hypothesize that female U. lugens can recognize their host plants by detecting key host volatile compounds, using a set of ORNs tuned to host volatiles, and discriminate them from non-host plants using another set of ORNs specialized for non-host volatiles. The ORNs with broad response spectra may enhance the discrimination between host and non-host plants by adding moderately selective sensitivity. Based on our finding, it is suggested that phytophagous insects use the combinational input from both host-specific and non-host specific ORNs for locating their host plants, and the electrophysiological characterization of ORN profiles would be useful in predicting the range of host plants in phytophagous insects.     

Identification of Olfactory Volatiles using Gas Chromatography-Multi-unit Recordings (GCMR) in the Insect Antennal Lobe

All organisms inhabit a world full of sensory stimuli that determine their behavioral and physiological response to their environment. Olfaction is especially important in insects, which use their olfactory systems to respond to, and discriminate amongst, complex odor stimuli. These odors elicit behaviors that mediate processes such as reproduction and habitat selection^1-3. Additionally, chemical sensing by insects mediates behaviors that are highly significant for agriculture and human health, including pollination^4-6, herbivory of food crops⁷, and transmission of disease^8,9. Identification of olfactory signals and their role in insect behavior is thus important for understanding both ecological processes and human food resources and well-being.To date, the identification of volatiles that drive insect behavior has been difficult and often tedious. Current techniques include gas chromatography-coupled electroantennogram recording (GC-EAG), and gas chromatography-coupled single sensillum recordings (GC-SSR)^10-12. These techniques proved to be vital in the identification of bioactive compounds. We have developed a method that uses gas chromatography coupled to multi-channel electrophysiological recordings (termed ''GCMR'') from neurons in the antennal lobe (AL; the insect''s primary olfactory center)^13,14. This state-of-the-art technique allows us to probe how odor information is represented in the insect brain. Moreover, because neural responses to odors at this level of olfactory processing are highly sensitive owing to the degree of convergence of the antenna''s receptor neurons into AL neurons, AL recordings will allow the detection of active constituents of natural odors efficiently and with high sensitivity. Here we describe GCMR and give an example of its use.Several general steps are involved in the detection of bioactive volatiles and insect response. Volatiles first need to be collected from sources of interest (in this example we use flowers from the genus Mimulus (Phyrmaceae)) and characterized as needed using standard GC-MS techniques^14-16. Insects are prepared for study using minimal dissection, after which a recording electrode is inserted into the antennal lobe and multi-channel neural recording begins. Post-processing of the neural data then reveals which particular odorants cause significant neural responses by the insect nervous system.Although the example we present here is specific to pollination studies, GCMR can be expanded to a wide range of study organisms and volatile sources. For instance, this method can be used in the identification of odorants attracting or repelling vector insects and crop pests. Moreover, GCMR can also be used to identify attractants for beneficial insects, such as pollinators. The technique may be expanded to non-insect subjects as well.     

Discrimination of oilseed rape volatiles by the honeybee: combined chemical and biological approaches

Honeybees (Apis mellifera L.) were individually subjected to a classical conditioning procedure in order to obtain an olfactory conditioned proboscis extension response. To relate the behavioural response directly to antennal detection abilities, a technique was developped for coupling proboscis extension responses and electroantennogram recordings, with the stimulation being provided by the effluent of a gas chromatograph (GC). Bees were conditioned with a six-component mixture being part of oilseed rape (Brassica napus L.) floral volatiles, and tested with the individual components separated by GC. Responses of the conditioned bees were compared to those of unconditioned bees. No behavioural response was obtained in the control group, neither to the individual components nor to the mixture. Conditioning induced behavioural responses for three components, and an increase of electroantennogram responses for all components. A second experiment was conducted with an air entrainment extract of oilseed rape flower volatiles. Behavioural responses of conditioned and unconditioned bees were recorded. Responses obtained from conditioned bees tested with the air entrainment extract showed six groups of behaviourally active GC peaks. Unconditioned bees showed the same pattern of responses but at a lower level. The coupled technique described here appears to be a reliable tool for locating active components in a synthetic as well as in a natural mixture of floral volatiles. The effects of conditioning on odour discrimination and on its sensory correlates are discussed.     

Plant volatiles as method of communication

Plants emit volatile compounds that can act as a communication method to insects, neighboring plants and pathogens. Plants respond to leaf and root damage by herbivores and pathogens by emitting these compounds. The volatile compounds can deter the herbivores or pathogens directly or indirectly by attracting their natural enemies to kill them. The simultaneous damage of plants by herbivores and pathogens can influence plant defense. The induced plant volatiles can also make neighboring plants ready for defense or induce defense in parts distant from the damaged area of the same plant. Belowground root herbivory can alter the defense response to aboveground leaf herbivory. In addition, most plants normally emit volatile compounds from their flowers that directly attract foraging mutualistic insects for nectar, which in turn perform the very important function of pollination for subsequent reproduction. The volatile compounds emitted from the floral and vegetative parts of plants belong to three main classes of compounds: terpenoids, phenylpropanoids/benzenoids, and C6-aldehydes (green-leaf volatiles). The volatile phytohormones methyl salicylate and methyl jasmonate serve as important signaling molecules for communication purposes, and interact with each other to optimize the plant defense response. Here we discuss and integrate the current knowledge on all types of communication between plants and insects, neighboring plants and pathogens that are mediated through plant volatiles.     

Encoding of plant odours by receptor neurons in the pine weevil (Hylobius abietis) studied by linked gas chromatography-electrophysiology

Receptor neuron responses to plant volatiles, trapped by head-space procedures, were examined in the pine weevil Hylobius abietis, using gas chromatography linked with electrophysiological recordings from single neurons. Seventy-two receptor neurons were tested 173 times for various plant volatile mixtures, either via a polar or a non-polar column.

1. All responses appeared as increased firing rates which followed the concentration profiles of the GC-eluted compounds.
2. The neurons were classified separately for the two column types in 17 and 19 groups respectively, according to the compounds they responded to. It suggests that the plant odour information is encoded by a large, but limited number of receptor neuron types.
3. Most neurons responded to a limited number of compounds (1–5) and showed a marked best response to one of them, whereas additional responses to several other components which seems to be structurally similar, was recorded for some neurons. It suggests that the plant odour receptor neurons are rather narrowly than broadly tuned, and that each neuron is specialized for receiving information about one or a few related compounds.
4. Most neurons responded to monoterpenes, whereas the other neurons responded to compounds of other categories.
5. Both major and minor plant volatile components activated specifically receptor neurons.

     

Perception of plant volatile blends by herbivorous insects--finding the right mix

Volatile plant secondary metabolites are detected by the highly sensitive olfactory system employed by insects to locate suitable plants as hosts and to avoid unsuitable hosts. Perception of these compounds depends on olfactory receptor neurones (ORNs) in sensillae, mostly on the insect antennae, which can recognise individual molecular structures. Perception of blends of plant volatiles plays a pivotal role in host recognition, non-host avoidance and ensuing behavioural responses as different responses can occur to a whole blend compared to individual components. There are emergent properties of blend perception because components of the host blend may not be recognised as host when perceived outside the context of that blend. Often there is redundancy in the composition of blends recognised as host because certain compounds can be substituted by others. Fine spatio-temporal resolution of the synchronous firing of ORNs tuned to specific compounds enables insects to pick out relevant host odour cues against high background noise and with ephemeral exposure to the volatiles at varying concentrations. This task is challenging as they usually rely on ubiquitous plant volatiles and not those taxonomically characteristic of host plants. However, such an odour coding system has the advantage of providing flexibility; it allows for adaptation to changing environments by alterations in signal processing while maintaining the same peripheral olfactory receptors.     

Five types of olfactory receptor neurons in the strawberry blossom weevil Anthonomus rubi: selective responses to inducible host-plant volatiles

Plants release hundreds of volatiles that are important in the interaction with herbivorous animals, but which odorants are detected by which species? In this study, single receptor neurons on the antenna of the oligophagous strawberry blossom weevil Anthonomus rubi were screened for sensitivity to naturally produced plant compounds by the use of gas chromatography linked to electrophysiological recordings from single cells. The narrow tuning of the neurons was demonstrated by responses solely to a few structurally related sesquiterpenes, aromatics or monoterpene hydrocarbons out of hundreds of plant constituents tested. We present five olfactory receptor neuron types, identified according to one primary odorant i.e. the compound to which the neurons are most sensitive. These odorants, (-)-germacrene D, (-)-beta-caryophyllene, methyl salicylate, E-beta-ocimene and (3E)-4,8-dimethyl-1,3,7-nonatriene, present in the intact strawberry plant, are induced in higher amounts by weevil feeding. This suggests that these compounds can provide information about the presence of conspecifics. We used protocols especially designed to allow comparison with previously investigated species. Striking similarities, but also differences, are demonstrated between receptor neuron specificity in the strawberry weevil and moths.     

植食性昆虫的嗅觉选食过程及其机制研究进展

植食性昆虫的嗅觉在其选择食物的过程中发挥了重要的作用,它能通过对植物挥发物的感受来定向和定位食物源并产生趋近行为,进而根据特殊的化合物或者多种化合物的特异浓度组合来区分寄主和非寄主植物.在这个过程中,昆虫嗅觉器官上相关的嗅觉感受蛋白被植物挥发物激活,形成特异的嗅觉感受通路,在行为上调控昆虫嗅觉选食的能力.本文主要从植食...     

Identification of plant odours activating receptor neurones in the weevil<Emphasis Type="Italic"> Pissodes notatus</Emphasis> F. (Coleoptera,Curculionidae)

Plants release complex mixtures of volatiles important in the interaction with insects and other organisms. In the search for compounds that contribute to the perception of odour quality in the weevil Pissodes notatus, single olfactory receptor neurones on the antennae were screened for sensitivity to naturally produced plant volatiles by the use of gas chromatography linked to single cell recordings. We here present 60 olfactory neurones responding to 25 of the numerous compounds released by host and non-host plants. All the neurones show high selectivity and are classified into 12 distinct types. The two most abundant types respond to alpha-pinene, beta-pinene, and 3-carene ( n=17), and to isopinocamphone and pinocamphone ( n=17), respectively. Other neurone types respond to limonene ( n=9), beta-phellandrene ( n=3), and fenchone ( n=4). Responses to beta-caryophyllene ( n=1) and to ethanol ( n=4) are also shown. Except for two pairs, the neurone types do not show overlap of the molecular receptive range. The active compounds are present in the host, Pinus pinaster, as well as in non-hosts, supporting the idea that plant odour quality is mediated by the ratio of the compounds rather than specific odorants.     

Induction of plant responses to oviposition and feeding by herbivorous arthropods: a comparison

Plants may respond both to feeding and oviposition by herbivorous insects. While responses of plants to feeding damage by herbivores have been studied intensively during the past decades, only a few, but growing number of studies consider the reactions of plants towards egg deposition by herbivorous insects. Plants showing defensive response to oviposition by herbivores do not `wait' until being damaged by feeding, but may instead react towards one of the initial steps of herbivore attack, the egg deposition. Direct plant defensive responses to feeding act directly against the feeding stages of the herbivores. However, a plant may also show direct defensive responses to egg deposition by (a) formation of neoplasms, (b) formation of necrotic tissue (= hypersensitive response), and (c) production of oviposition deterrents. All these plant reactions have directly negative effects on the eggs, hatching larvae, or on the ovipositing females. Indirect plant defensive responses to feeding result in the emission of volatiles (= synomones) that attract predators or parasitoids of the feeding stages. A few recent studies have shown that plants are able to emit volatiles also in response to egg deposition and that these volatiles attract egg parasitoids. Studies on the mechanisms of induction of synomones by egg deposition show several parallels to the mechanisms of induction of plant responses by feeding damage. When considering induced plant defence against herbivores from an evolutionary point of view, the question arises whether herbivores evolved the ability to circumvent or even to exploit the plant's defensive responses. The reactions of herbivores to oviposition induced plant responses are compared with their reactions to feeding induced plant responses.     

Identification of plant volatiles activating single receptor neurons in the pine weevil (Hylobius abietis)

Naturally produced plant volatiles, eliciting responses of single olfactory receptor neurons in the pine weevil, have been identified by gas chromatography linked with mass spectrometry. The receptor neurons (n = 72) were classified in 30 types, according to the compound which elicited the strongest response in each neuron, 20 of which compounds were identified. Most potent for 14 types of neurons (n = 50) were monoterpenes, including bicyclic (e.g. α-pinene, camphor and myrtenal) for 8 types (n = 32), monocyclic (limonene, carvone, α-terpinene) for 3 types (n = 12) and acyclic (e.g. β-myrcene and linalool) for 3 types (n = 6). Other compounds eliciting strongest responses of a neuron were five sesquiterpenes, including α-copaene and a farnesene-isomer, and an anethole type which has no biosynthetic relationship with terpenes. Within one type, receptor neurons with quite selective responses to the most potent compound as well as neurons with additional responses to several, structurally similar compounds were found, indicating that the neurons may have the same functional types of membrane receptors, but different sensitivities. Response spectra of neurons within the bicyclic-, mono-cyclic and acyclic types showed more overlapping than across the neuron types. Minimal overlapping response spectra was found between monoterpene and sesquiterpene neurons. The results suggest that this structure-activity relationship is significant for encoding plant odour information in the pipe weevil. Accepted: 6 January 1997     

巴氏新小绥螨对柑桔全爪螨处理的枳橙叶片挥发物的行为反应

【目的】研究巴氏新小绥螨Neoseiulus barkeri对柑桔全爪螨Panonychus citri及刺吸式昆虫为害柑桔叶片释放的挥发物的行为反应,揭示巴氏新小绥螨的嗅觉反应特点。【方法】采用GC-MS顶空进样法对枳橙叶片常见挥发性化合物、针刺枳橙叶片及其柑桔全爪螨雌成螨取食枳橙叶片挥发物进行定性分析,确定每类化合物的相对保留指数,构建枳橙叶片常见及受害挥发性化合物特征谱库。利用嗅觉测定技术分析巴氏新小绥螨对枳橙叶片挥发物的行为反应。【结果】针刺处理和柑桔全爪螨取食影响枳橙叶片挥发物的组成和含量。两种方法处理时枳橙叶片释放的主要物质为α-蒎烯、水芹烯、4-异丙基甲苯。随着处理加重,增量释放的物质为:cis-π-罗勒烯、月桂烯、柠檬烯、异松油烯,减量表达的物质为2-乙基-1-乙醇和十一烷。在10~(-2)、10~(-4)、10~(-6)和10~(-8)g/m L浓度下,正己醛、正壬醛、乙酸辛酯和正庚醛对巴氏新小绥螨有强烈的引诱作用(P>0.05);月桂烯在10~(-2)、10~(-4)和10~(-6)g/m L浓度下对巴氏新小绥螨有强烈的引诱作用(P>0.05);正壬醇和正辛醇随着浓度增加,对巴氏新小绥螨的引诱作用降低;苯甲醛对巴氏新小绥螨的引诱作用较弱。【结论】巴氏新小绥螨对柑桔全爪螨及刺吸式口器昆虫为害柑桔叶片释放出的挥发物各组分具有不同的行为反应,柑桔及其刺吸式害虫生境中的嗅觉线索在巴氏新小绥螨的寄主定位和生境选择中起着重要作用。     

Pea plant volatiles guide host location behaviour in the pea moth

Identification of plant volatiles that attract mated insect females for oviposition can provide important information about plant–insect relationships that can be used to develop pest control strategies involving manipulation of the female host search. Our study represents a first step towards identifying volatiles that affect the host location behaviour of the pea moth Cydia nigricana. The behaviours of virgin and mated males and females were analysed in cage experiments testing a two-choice situation at close range and in wind tunnel experiments evaluating upwind orientation over a distance. In both experimental setups, flowering pea plants constituted the most attractive phenological stage for mated females, with 58 % landing on such plants in the wind tunnel. Testing headspace extracts of different phenological stages of pea and of detached pea buds and flowers in the wind tunnel, mated females showed the highest landing responses to volatiles during flower development (budding 42 % and flowering 56 %) and from detached buds (46 %) and flowers (66 %). Volatile compounds collected from the various phenological stages of pea were analysed by gas chromatography–mass spectrometry, and the antennal responses to these headspace collections were evaluated by gas chromatography–electroantennography. Ten antennally active compounds were identified, nine of which were present in the headspace extracts of the whole pea plants at all tested phenological stages and in detached buds and flowers. Overall, our results demonstrate a clear link between host plant phenology, the corresponding plant odour, and the behaviour of mated C. nigricana females.