Behavioural responses of larvae of Colorado potato beetle,Leptinotarsa decemlineata (Coleoptera: Chrysomelidae), to host plant volatile blends attractive to adults

Abstract

• 1 Orientation of second‐ and fourth‐instar Colorado potato beetle (CPB) Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), to volatiles emitted from a solanaceous host, potato, and seven synthetic blends or three individual chemicals emitted by potato plants were investigated in laboratory bioassays.
• 2 Both second‐ and fourth‐instar CPB were attracted to intact and mechanically damaged (MD) potato foliage. When offered a choice between intact and MD foliage, no preference was observed.
• 3 Among seven synthetic blends tested (of which six are attractive to adult CPB), second‐ and fourth‐instar CPB were attracted only to a single three‐component blend comprising (±)‐linalool, methyl salicylate, and (Z)‐3‐hexenyl acetate. Individual compounds and two‐component blends were inactive. No significant difference was noted between larval responses to the attractive synthetic blend vs. MD potato foliage.
• 4 Second‐and fourth‐instar larvae had similar thresholds for behaviour for the three‐component blend (50 µg source load). Female CPB were attracted to source loads 10× below the larval threshold (5 µg). Male CPB were the most sensitive life form tested with a behavioural threshold at 0.5 µg source load which was 10× and 100× below female and larval thresholds, respectively
• 5 This is the first report of a synthetic chemical attractant for CPB larvae. As both larval and adult CPB are attracted to a single chemical blend, the usefulness of the attractant as a component of an attracticide or ‘push‐pull’ strategies for management of pestiferous populations is enhanced.

     

Plant volatiles in the sexual communication of Melolontha hippocastani: response towards time-dependent bouquets and novel function of (Z)-3-hexen-1-ol as a sexual kairomone

1. Swarming males of Melolontha hippocastani are known to locate females that stay feeding within the host trees by orienting towards damage‐induced plant volatiles (green leaf volatiles) and a sex pheromone. Thus, volatiles emitted by freshly damaged leaves might indicate to a male the presence of currently feeding females. 2. The hypothesis was studied that volatiles from freshly damaged leaves are more attractive to males than volatiles from old damaged leaves. The odour bouquets of damaged leaves from three plant species that have been shown to attract male M. hippocastani in the field were analysed 10 min (fresh damage) and 1.5 h (old damage) after damaging, using coupled gas chromatography–mass spectrometry. The results showed clear differences between the bouquets: the bouquet of freshly damaged leaves of all species consisted of typical leaf aldehydes, i.e. hexanal, (Z)‐3‐hexenal, (Z)‐2‐hexenal, (E)‐2‐hexenal, the leaf alcohol (Z)‐3‐hexen‐1‐ol, and the corresponding acetate. One and a half hours after damage, aldehydes had almost vanished and (Z)‐3‐hexen‐1‐ol and (Z)‐3‐hexenyl acetate predominated; however males of M. hippocastani were equally attracted to traps baited with volatiles from old and freshly damaged leaves in field experiments. When traps were baited with synthetic volatile mixtures mimicking the bouquets of old and freshly damaged leaves, M. hippocastani males even preferred the old damage mixture. 3. Experiments addressing the role of individual green leaf volatiles revealed that only (Z)‐3‐hexen‐1‐ol was highly attractive while the other compounds tested individually were behaviourally inactive, however all tested compounds elicited comparable electrophysiological responses on male M. hippocastani antennae. 4. In analogy to the term aggregation kairomone used for feeding‐induced plant volatiles that attract both sexes of an insect, the term sexual kairomone is suggested to describe the novel function of (Z)‐3‐hexen‐1‐ol in the sexual communication of M. hippocastani.     

Behavioural and electrophysiological responses of the European corn borer Ostrinia nubilalis to host‐plant volatiles and related chemicals

The European corn borer Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae) is a worldwide pest of maize (Zea mays L.) and other crops. The semiochemicals released by maize plants and structurally‐related compounds can be used by adult female O. nubilalis for host‐plant location and oviposition. Headspace volatile compounds emitted by watered and water‐deprived maize plants are collected and identified by their retention indices and mass spectra. The most abundant compounds from watered plants are limonene, linalool, benzoic acid, indole, β‐caryophyllene and acetophenone, whereas, in water‐deprived plants, limonene, acetophenone, hexanoic acid, benzoic acid and indole are dominant. In addition, (E)‐4,8‐dimethyl‐1,3,7‐nonatriene, 6‐methyl‐5‐hepten‐2‐one, anisole and 1‐carvone are undetected in the water‐deprived plants. Some of the identified compounds show electrophysiological activity (electroantennogram) in the antennae of both sexes, with the responses elicited by tridecane, tetradecane, dodecane, nonanal, decanal and 2‐ethylhexanol on males being particularly noteworthy. In a dual‐choice olfactometer, adult females show a preference for 2‐hexanol, heptanal, methyl salicylate, hexyl acetate, nonanal, methyl dodecanoate, β‐pinene and (E)‐2‐hexenyl acetate over hexane controls. Tetradecane, linalool, methyl hexanoate, methyl nonanoate, (Z)‐3‐hexenyl benzoate, tridecane, 2‐cyclopentylcyclopentanone, 3‐methylbutyl acetate, β‐myrcene and (Z)‐3‐hexenyl butanoate result in fewer females in the test arm compared with the control arm. No single compound displays an activity similar to watered maize plants, supporting the hypothesis that blends of volatiles in specific ratios are more effective than single volatile chemicals. The results of the present study suggest that methyl salicylate, which elicits also one of the highest electrophysiological responses in female antennae, plays a role in host preference by O. nubilalis females.     

Behavioral responses for evaluating the attractiveness of specific tea shoot volatiles to the tea green leafhopper,Empoaca vitis

Abstract The tea green leafhopper, Empoasca vitis Göthe, is one of the most serious insect pests of tea plantations in mainland China. Over the past decades, this pest has been controlled mainly by spraying pesticides. Insecticide applications not only have become less effective in controlling damage, but even more seriously, have caused high levels of toxic residues in teas, which ultimately threatens human health. Therefore, we should seek a safer biological control approach. In the present study, key components of tea shoot volatiles were identified and behaviorally tested as potential leafhopper attractants. The following 13 volatile compounds were identified from aeration samples of tea shoots using gas chromatography‐mass spectrometry (GC‐MS): (E)‐2‐hexenal, (Z)‐3‐hexen‐1‐ol, (Z)‐3‐hexenyl acetate, 2‐ethyl‐1‐hexanol, (E)‐ocimene, linalool, nonanol, (Z)‐butanoic acid, 3‐hexenyl ester, decanal, tetradecane, β‐caryophyllene, geraniol and hexadecane. In Y‐tube olfactometer tests, the following individual compounds were identified: (E)‐2‐hexenal, (E)‐ocimene, (Z)‐3‐hexenyl acetate and linalool, as well as two synthetic mixtures (called blend 1 and blend 2) elicited significant taxis, with blend 2 being the most attractive. Blend 1 included linalool, (Z)‐3‐hexen‐1‐ol and (E)‐2‐hexenal at a 1 : 1 : 1 ratio, whereas blend 2 was a mixture of eight compounds at the same loading ratio: (E)‐2‐hexenal, (Z)‐3‐hexen‐1‐ol, (Z)‐3‐hexenyl acetate, 2‐penten‐1‐ol, (E)‐2‐pentenal, pentanol, hexanol and 1‐penten‐3‐ol. In tea fields, the bud‐green sticky board traps baited with blend 2, (E)‐2‐hexenal or hexane captured adults and nymphs of the leafhoppers, with blend 2 being the most attractive, followed by (E)‐2‐hexenal and hexane. Placing sticky traps baited with blend 2 or (E)‐2‐hexenal in the tea fields significantly reduced leafhopper populations. Our results indicate that the bud‐green sticky traps baited with tea shoot volatiles can provide a new tool for monitoring and managing the tea leafhopper.     

The social wasp Polybia fastidiosuscula Saussure (Hymenoptera: Vespidae) uses herbivore‐induced maize plant volatiles to locate its prey

Social wasps in the Polybia genus are important for use as pest‐control agents in agricultural systems. The objective of this study was to investigate the behavioural responses of Polybia fastidiosuscula Saussure (Hymenoptera: Vespidae) to volatiles from maize, both constitutive volatiles and those induced by the herbivory of Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae). To assess the behavioural response of P. fastidiosuscula to S. frugiperda larvae, undamaged plants, S. frugiperda‐damaged plants, mechanically damaged plants, mechanically damaged plants plus regurgitant from larvae and extracts from various treatments, bioassays were conducted in a Y‐olfactometer. In addition, the volatiles from plants subjected to different treatments were collected via aeration, and they were quantified and identified. The wasps showed a greater preference for plants with damage induced either by larval feeding or by being mechanically damaged plus regurgitant than for undamaged plants or either larvae alone or mechanically damaged plants. Wasps were more attracted to extracts from plants + S. frugiperda larvae and to an extract from mechanically damaged plants + the regurgitant of larvae compared to hexane. The primary compounds induced by herbivory for 5–6 h after the beginning of the damage or regurgitant treatment were identified as α‐pinene, β‐myrcene, (Z)‐3‐hexenyl acetate, limonene, (E)‐ocimene, linalool, DMNT, (E)‐β‐farnesene, TMTT and indole. The results presented here show that the social wasp P. fastidiosuscula uses herbivore‐induced plant volatiles from maize to locate its prey.     

Laboratory and greenhouse evaluation of a synthetic host volatile attractant for Colorado potato beetle, Leptinotarsa decemlineata (Say)

Abstract 1 The attractiveness of potato plants treated with a synthetic host volatile blend [(Z)‐3‐hexenyl acetate (+/–)‐linalool, and methyl salicylate] to newly emerged and 5‐day‐old adult Colorado potato beetle, Leptinotarsa decemlineata (Say), was compared at four doses against untreated control plants and plants treated with an azadirachtin‐based antifeedant in greenhouse cage arenas. 2 Attractant‐treated plants (derived release rates of 0, 5.7, 17.1 or 57 µg/h) were significantly more attractive than untreated control plants to newly emerged and 5‐day‐old adults only at 57 µg/h. 3 Attractant‐treated plants were significantly more attractive than antifeedant‐treated plants to newly emerged and 5‐day‐old adults at the 5.7 µg/h treatment level and higher. Mean insect density on attractant‐treated plants in the attractant/antifeedant study was significantly higher than in the attractant/control study. 4 Habituation to the synthetic attractant was evaluated by exposing adult beetles to the synthetic attractant for 0, 1, 2.5, 4, 8, 12 or 16 h, before release into a wind tunnel in which an attractant‐baited plant model was placed at the upwind end. Insects exposed to the synthetic host attractant for ≤ 8 h moved to the synthetic attractant‐baited plant model whereas insects exposed to the synthetic host attractant for 12 and 16 h did not. Furthermore, beetles exposed to the synthetic attractant for 0 and 1 h moved at rates greater than, or equal to, the median whereas beetles exposed for longer time periods moved at rates significantly less than the median. 5 These results demonstrate the potential for using the synthetic plant attractant and an antifeedant as components in a stimulo‐deterrent strategy for management of the Colorado potato beetle as shown by us in another study.     

Maize plants sprayed with either jasmonic acid or its precursor,methyl linolenate,attract armyworm parasitoids,but the composition of attractants differs

Treatment of both uninfested and armyworm‐infested maize plants with jasmonic acid (JA) is known to attract the parasitic wasp, Cotesia kariyai Watanabe (Hymenoptera: Braconidae). Here, we show that treatment with a methyl ester of a JA precursor, methyl linolenate (MeLin), also causes maize plants to attract this wasp, yet does not cause elevated levels of endogenous JA. The volatile chemicals emitted from either infested or uninfested maize plants treated with MeLin were qualitatively and quantitatively different from those emitted from JA‐treated plants. Among compounds emitted from MeLin‐treated plants, α‐pinene and menthol attracted wasps in pure form in a two‐choice test using a choice chamber. A mixture of methyl salicylate, α‐copaene, and β‐myrcene also attracted wasps. In contrast, (Z)‐3‐hexenyl acetate was among the main attractants for C. kariyai in JA‐treated plants. These data show that in addition to JA, MeLin also has the potential to increase the host‐finding ability of C. kariyai, but that the composition of attractants they induce differs.     

Predator–prey interactions: olfactory adaptations of generalist and specialist predators

1 Olfactory responses of the Colorado potato beetle (CPB), Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), a generalist predator, Podisus maculiventris (Say) (Hemiptera, Heteroptera: Pentatomidae) (Pm), and a specialist predator, Perillus bioculatus (F.) (Hemiptera, Heteroptera: Pentatomidae) (Pb) were investigated. Volatiles tested included 20 compounds emitted by undamaged potato plants (Solanum tuberosum), plants that had been artificially damaged, or plants damaged by feeding by CPB larvae. 2 Coupled gas chromatography/electroantennogram detector (GC/EAD) recordings revealed five compounds for which reliable responses were recorded from CPB antennae: (E)-2-hexen-1-ol, (Z)-3-hexen-1-ol, (±)-linalool, nonanal, methyl salicylate, and indole. Both Pm and Pb responded selectively to the same compounds as the CPB with exceptions: (1) (Z)-3-hexenyl butyrate elicited reliable responses for both Pm and Pb, and (2) (E)-2-hexen-1-ol and (Z)-3-hexen-1-ol were inactive for Pm and Pb under these conditions. Dose–response curves showed that CPB was at least 100 times more sensitive to (E)-2-hexen-1-ol than were the predators. Both predators were more sensitive to each of the other compounds than were CPB. Both CPB and Pm were attracted to a five component blend comprising (E)-2-hexen-1-ol, (Z)-3-hexen-1-ol, (±)-linalool, nonanal and methyl salicylate. However, attraction of CPB to the blend occurred only with lower doses of (E)-2-hexen-1-ol and (Z)-3-hexen-1-ol. 3 These results show that the herbivore (CPB) has olfactory receptors which are more sensitive to constitutive host plant volatiles, e.g. green leaf volatiles, while both generalist (Pm) and specialist (Pb) predators are more sensitive to systemic volatiles produced in response to prey feeding. Keywords Colorado potato beetle, constitutive compounds, host plant, induced compounds, olfaction, Perillus bioculatus, Podisus maculiventris, predator, prey, tritrophic.     

Synergism between aromatic compounds and green leaf volatiles derived from the host plant underlies female attraction in the oriental fruit moth

Blends of volatile compounds emitted by host plants are known to mediate the attraction of gravid female herbivores to oviposition sites, but the role of individual odor components is still little understood. We characterized the olfactory response of mated female Cydia (Grapholita) molesta (Busck) (Lepidoptera: Tortricidae) to synthetic mixtures of compounds emitted by peach shoot, a key host plant of this herbivore, and investigated the role of important constituents of bioactive mixtures in moth attraction. Relative ratios of constituents of the mixtures corresponded to the natural ratio of volatile compounds collected in the plant's headspace. A significant attractant effect was found for a comparatively complex 10‐compound mixture that included four green leaf volatiles [(Z)‐3‐hexen‐1‐ol, 1‐hexanol, (E)‐2‐hexenal, and (Z)‐3‐hexen‐1‐yl acetate], five aromatics (benzaldehyde, methyl salicylate, methyl benzoate, benzonitrile, and phenylacetonitrile), and a carboxylic acid (valeric acid). Using a subtraction approach, the number of compounds was progressively decreased, resulting in a bioactive 5‐compound mixture composed of two constituents, green leaf volatiles and aromatic compounds. Further evaluations revealed that benzaldehyde and benzonitrile must be present in association with three distinct green leaf volatiles to produce an attractant effect on the female moths. This 5‐compound mixture was as attractive as natural peach shoot volatiles, which are known to comprise over 20 compounds. Results are discussed in light of the documented synergistic effect between the three general green leaf volatiles and the two specific aromatic compounds.     

多种植物挥发物及马铃薯甲虫聚集素对马铃薯甲虫的引诱作用

马铃薯甲虫Leptinotarsa decemlineata (Say)是马铃薯上一种最主要的毁灭性害虫。由于其严重的抗药性, 急需寻找一种替代杀虫剂的防治方法。本实验从开发马铃薯甲虫引诱剂的目的出发,用“Y”型嗅觉仪测定了马铃薯甲虫对来自植物的7种挥发物单体、8个挥发物混合物配方以及马铃薯甲虫聚集素的行为反应,并进一步进行了田间诱集试验。室内生测结果表明,2-苯乙醇对马铃薯甲虫雌雄虫都有明显的引诱作用。芳樟醇+水杨酸甲酯+顺乙酸-3-己烯酯的混合物对马铃薯甲虫雄虫有很强的引诱作用(81.67%), 但是对雌虫引诱作用不明显(63.33%)。在其中加入马铃薯甲虫聚集素不但提高了对雄虫的引诱率（88.33%）,还消除了之前的雌雄性别间反应差异现象,使对雌虫的引诱率达到了83.33%。田间诱集结果显示,芳樟醇+水杨酸甲酯+顺乙酸-3-己烯酯+马铃薯甲虫聚集素在所研究的引诱剂配方中引诱效果最好。从而为马铃薯甲虫发生的预测预报和诱杀技术提供了研究基础。     

Antennal and behavioural responses of Heortia vitessoides females to host plant volatiles of Aquilaria sinensis

This study investigated volatiles from Aquilaria sinensis (Lour.) Gilg (Thymelaeaceae) leaves that attracted Heortia vitessoides Moore (Lepidoptera: Crambidae). Volatiles from young and old A. sinensis leaves were identified and quantified by gas chromatography–mass spectrometry and gas chromatography‐electroantennogram detection. Both wind tunnel bioassays and field tests were conducted to measure the attraction of adults to synthetic blends of volatiles from leaves of different maturations. Consistent electroantennographic activity was obtained for nine and three compounds from headspace collections of young and old A. sinensis leaves, respectively. Qualitative and quantitative differences were found among two odour profiles. In wind tunnel experiments, the fresh young leaves proved to be more attractive to females than old leaves. A nine‐component mixture, including hexanal, limonene, 2‐hexanol, octanal, (Z)‐3‐hexenyl acetate, (Z)‐3‐hexen‐1‐ol, nonanal, decanal, and 2,6,10‐trimethyl‐dodecane (with a ratio of 2:16:9:4:63:100:13:10:5) from young leaves attracted moths significantly more than the three‐component mixture of nonanal, decanal, and 2,6,10‐trimethyl‐dodecane (with a ratio of 11:14:26) from old leaves. Further subtractive bioassays conducted in the wind tunnel showed that both the complete nine‐component mixture and a subtracted four‐component mixture of hexanal, (Z)‐3‐hexenyl acetate, nonanal, and decanal (with a ratio of 2:63:13:10) elicited equivalent responses in females. All components in the four‐component blend were essential for optimal attraction. In a field trial using the nine‐ and four‐component blends, more moths were captured using both blends than in traps baited with hexane only. Our study indicates that the odour blends of young leaves play an important role in H. vitessoides host plant recognition. The mechanisms behind host recognition and age‐dependent changes in leaf chemistry are discussed.     

Volatiles released from bean plants in response to agromyzid flies

Liriomyza sativae Blanchard and Liriomyza huidobrensis (Blanchard) (Diptera: Agromyzidae) are two invasive flies in China that have caused economical damage on vegetables and ornamental plants. In this article, we report the profiles of emitted volatiles from healthy, mechanically damaged, and leafminer-damaged bean, Phaseolus vulgaris L., plants. Among 25 emitted volatiles identified, (E)-2-hexen-1-al, (3E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), (Z)-3-hexenyl acetate, (Z)-3-hexen-1-ol, (syn)- and (anti)-2-methylpropanal oxime, (syn)-2-methylbutanal oxime, linalool, and (E,E)-α-farnesene were consistently released from damaged bean plants. Combined amounts of these nine compounds made up more than 70% of the total volatiles emitted from each treatment. No qualitative differences in volatile emission were found between bean plants damaged by the two fly species; however, amounts of several major compounds induced by L. huidobrensis damage were significantly higher than those from plants damaged by L. sativae. The mechanically damaged plants released a higher proportion of green leaf volatiles than plants in the other treatments, whereas leafminer-damaged plants produced more terpenoids and oximes. Furthermore, the volatile profiles emitted from plants, damaged by adult leafminers, by second instar larvae, and even the plants with empty mines left by leafminer larvae (the pupal stage) were significantly different. The identification of volatile oximes released from damaged plants was confirmed and is discussed in a behavioral and biological control context.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Effects of single and multiple herbivory by host and non‐host caterpillars on the attractiveness of herbivore‐induced volatiles of sugarcane to the generalist parasitoid Cotesia flavipes

It is well known that parasitoids are attracted to volatiles emitted by host‐damaged plants; however, this tritrophic interaction may change if plants are attacked by more than one herbivore species. The larval parasitoid Cotesia flavipesCameron (Hymenoptera: Braconidae) has been used intensively in Brazil to control the sugarcane borer, Diatraea saccharalisFabricius (Lepidoptera: Pyralidae) in sugarcane crops, where Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae), a non‐stemborer lepidopteran, is also a pest. Here, we investigated the ability of C. flavipes to discriminate between an unsuitable host (S. frugiperda) and a suitable host (D. saccharalis) based on herbivore‐induced plant volatiles (HIPVs) emitted by sugarcane, and whether multiple herbivory (D. saccharalis feeding on stalk + S. frugiperda feeding on leaves) in sugarcane affected the attractiveness of HIPVs to C. flavipes. Olfactometer assays indicated that volatiles of host and non‐host‐damaged plants were attractive to C. flavipes. Even though host‐ and non‐host‐damaged plants emitted considerably different volatile blends, neither naïve nor experienced wasps discriminated suitable and unsuitable hosts by means of HIPVs emitted by sugarcane. With regard to multiple herbivory, wasps innately preferred the odor blend emitted by sugarcane upon non‐host + host herbivory over host‐only damaged plants. Multiple herbivory caused a suppression of some volatiles relative to non‐host‐damaged sugarcane that may have resulted from the unaltered levels of jasmonic acid in host‐damaged plants, or from reduced palatability of host‐damaged plants to S. frugiperda. In conclusion, our study showed that C. flavipes responds to a wide range of plant volatile blends, and does not discriminate host from non‐host and non‐stemborer caterpillars based on HIPVs emitted from sugarcane. Moreover, we showed that multiple herbivory by the sugarcane borer and fall armyworm increases the attractiveness of sugarcane plants to the parasitoids.     

Antennal and behavioural response of codling moth Cydia pomonella to plant volatiles

Identification of host volatile compounds attractive to codling moth Cydia pomonella, a most important insect of apple, will contribute to the development of safe control techniques. Synthetic apple volatiles in two doses were tested for antennal and behavioural activity in codling moth. Female antennae strongly responded to (Z)3‐hexenol, (Z)3‐hexenyl benzoate, (Z)3‐hexenyl hexanoate, (±)‐linalool and E,E‐α‐farnesene. Two other compounds eliciting a strong antennal response were the pear ester, ethyl (E,Z)‐2,4‐decadienoate, and its corresponding aldehyde, E,E‐2,4‐decadienal, which is a component of the larval defence secretion of the European apple sawfly. Attraction of codling moth to compounds eliciting a strong antennal response was tested in a wind tunnel. Male moths were best attracted to a blend of (E,E)‐α‐farnesene, (E)‐beta‐farnesene and ethyl (E,Z)‐2,4‐decadienoate. The aldehyde E,E‐2,4‐decadienal had an antagonistic effect when added to the above mixture.     

Similar attractiveness of maize volatiles induced by Helicoverpa armigera and Pseudaletia separata to the generalist parasitoid Campoletis chlorideae

Campoletis chlorideae Uchida (Hymenoptera: Ichneumonidae), a major larval endoparasitoid of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), also attacks many other noctuid caterpillars. We investigated the attractiveness of H. armigera‐ and Pseudaletia separata (Lepidoptera: Noctuidae)‐infested maize [Zea mays L. (Poaceae)] plants to C. chlorideae, and analyzed the volatiles emitted from infested plants and undamaged plants. Considering the reported specific induction of plant volatiles by elicitors in the caterpillar regurgitant, we also tested the response of the parasitoid to mechanically damaged plants treated with caterpillar regurgitant or water and measured the volatiles released by these plants. In wind‐tunnel bioassays, C. chlorideae was strongly attracted to herbivore‐induced maize volatiles. Mechanically damaged plants, whether they were treated with caterpillar regurgitant or water, were more attractive to the parasitoid than undamaged plants. The parasitoid did not distinguish between maize seedlings infested by the two noctuid insects, nor did they show a difference in attraction to mechanically damaged plants treated with caterpillar regurgitant or water. Coupled gas chromatography–mass spectrometer (GC‐MS) analysis revealed that 15 compounds were commonly emitted by herbivore‐infested and mechanically damaged maize plants, whereas only two compounds were released in minor amounts from undamaged plants. Infestation by H. armigera specifically induced four terpenoids, β‐pinene, β‐myrcene, D‐limonene, and (E)‐nerolidol, which were not induced by infestation of P. separata and mechanical damage, plus caterpillar regurgitant or water. Two compounds, geranyl acetate and β‐sesquiphellandrene, were also induced by the infestation of H. armigera, but not by the infestation of P. separata. All treated maize plants released volatiles in significantly larger total amounts than did undamaged plants. Maize plants infested by H. armigera emitted greater amounts of volatiles than plants infested by P. separata. The treatment with caterpillar regurgitant resulted in larger amounts of volatile emission than the treatment with water did in mechanically damaged plants. The amounts of emissions of individual compounds were also different between differently treated plants.     

Response of garden chafer, Phyllopertha horticola, to plant volatiles: from screening to application

Field tests were performed on a golf course and in an apple orchard to screen synthetic plant volatiles with respect to their attractiveness for the garden chafer, Phyllopertha horticola L. (Coleoptera: Scarabaeidae), and to investigate the possible application of plant volatiles for garden chafer control. The chemicals tested were green leaf volatiles (GLV), terpenoids, and phenylpropanoids. Funnel traps baited with the GLV (Z)‐3‐hexen‐1‐ol, 1‐hexanol (Z)‐3‐hexenal, and hexanal captured more P. horticola than unbaited controls. Furthermore, traps baited with all tested floral terpenoids (i.e., geraniol, geranyl acetate, citronellol, linalool, and nerol) and phenylpropanoids (i.e., eugenol, anethol, isoeugenol, eugenyl acetate, and isoeugenyl acetate) captured more garden chafers than controls. Different dispenser types loaded once with a mixture of (Z)‐3‐hexen‐1‐ol (50%), geraniol (11.5%), eugenol (27%), and 2‐phenylethyl propionate (11.5%) attracted P. horticola over a whole flight season. A commercially available membrane dispenser had the best properties, combining the highest number of captured beetles with a low release rate. A simple modification of the trap design, i.e., a reduction of the funnel outlet diameter, significantly reduced the capture of beneficial non‐target insects (Apoidea), without influencing the number of captured garden chafers. A mass trapping experiment in the apple orchard revealed that the use of attractant traps significantly reduced the percentage of apples disfigured by feeding holes of adult garden chafers (control area: 18.9%, test area: 11.6%). The possible application of synthetic plant volatiles in mass trapping and monitoring approaches for garden chafer control is discussed.     

Response of mirid predators to synthetic herbivore‐induced plant volatiles

Zoophytophagous plant bugs feed on plant tissue as a source of water and nutrients, besides feeding on prey. By phytophagy, mirid predators activate plant defense responses through different pathways, resulting, among others, in the release of herbivore‐induced plant volatiles (HIPVs). These compounds could repel herbivores and attract parasitoids and predators, and synthetic versions could potentially be used in biological control. Nevertheless, little is known about the influence of synthetic volatiles on mirid attraction. Using Y‐tube olfactometer trials, we evaluated the responses of Nesidiocoris tenuis (Reuter), Macrolophus pygmaeus (Rambur), and Dicyphus bolivari Lindberg (Hemiptera: Miridae), important natural enemies used to control various greenhouse pests, to 10 synthetic versions of HIPVs released from tomato (Solanum lycopersicum L., Solanaceae) plants induced by N. tenuis and M. pygmaeus. Nesidiocoris tenuis responded to five of the 10 HIPVs, whereas M. pygmaeus and D. bolivari responded to four of the 10 HIPVs. Two green leaf volatiles, (Z)‐3‐hexenyl propanoate and (Z)‐3‐hexenyl acetate, and the ester methyl salicylate (MeSA) were attractive to all three mirid predator species. Our results demonstrate that the volatiles released by tomato plants activated by N. tenuis and M. pygmaeus phytophagy are attractive to their conspecifics and also to D. bolivari. Further studies should evaluate the potential of these compounds to attract predatory mirids in the field.     

Effects of plant volatiles on electrophysiological and behavioral responses of Cryptorrhynchus lapathi

Electroantennogram (EAG), Y‐tube olfactometer, and wind tunnel bioassays were conducted to test the electrophysiological and behavioral responses of Cryptorrhynchus lapathi L. (Coleoptera: Curculionidae) to six individual volatiles and seven essential oils (compounded volatiles). The aim of this study was to select effective plant compounds that can be used in the development of semiochemical‐based push–pull methods for the control of this harmful insect. Male and female C. lapathi displayed strong EAG responses to linoleic acid, α‐pinene, (Z)‐3‐hexen‐1‐ol, geraniol, turpentine oil, and salicylaldehyde. Y‐tube olfactometer assays indicated that salicylaldehyde and α‐pinene elicited strong repellent effects on female C. lapathi. Linoleic acid and (Z)‐3‐hexen‐1‐ol showed greater luring effects on male and female C. lapathi compared with the control. Turpentine oil was stronger repellent, and geraniol showed stronger luring effects on male C. lapathi than the control. Wind tunnel assays with both male and female C. lapathi indicated that salicylaldehyde, α‐pinene, and turpentine oil elicited repellent effects compared with the control. Linoleic acid, (Z)‐3‐hexen‐1‐ol, and geraniol were stronger lures of both male and female C. lapathi than the control. These results provide a basis for the further development of C. lapathi luring and repellent agents.     

Host plant choice experiments of Colorado potato beetle (Coleoptera: Chrysomelidae) in Virginia

Field and laboratory-choice experiments were conducted to understand aspects of host plant orientation by the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae), in Virginia. In laboratory bioassays, L. decemlineata oriented to volatiles emitted by potato, Solanum tuberosum L., foliage over both tomato, Lycopersicon esculentum L., and eggplant, Solanum melongena L., foliage, and eggplant over tomato foliage, all of which had been mechanically damaged. Field choice tests revealed more L. decemlineata adults, larvae, and egg masses on eggplant than on tomato. In other experiments, counts of live L. decemlineata on untreated paired plants and counts of dead beetles on imidacloprid-treated plants did not differ between potato and eggplant. L. decemlineata was significantly attracted to eggplant over both tomato and pepper. To determine whether feeding adults affected orientation to host plants, an imidacloprid-treated eggplant or potato plant was paired with an untreated eggplant or potato plant covered in a mesh bag containing two adult male beetles. Significantly more adults were attracted to eggplant with feeding male beetles paired with another eggplant than any other treatment combination. These results indicate that the presence of male L. decemlineata on plants affects host plant orientation and suggests that the male-produced aggregation pheromone may be involved.     

Repellent activity of Cymbopogon citratus and Tagetes minuta and their specific volatiles against Megalurothrips sjostedti

Cowpea is an important source of protein for people in Africa. However, the crop suffers major damage and yield losses due to bean flower thrips, Megalurothrips sjostedti Trybom (Thysanoptera: Thripidae). Although companion plants are known to reduce the damage caused by insect pests, the role of their volatiles in repelling pests from target plants has been the subject of few investigations. Here, we used the Y‐tube olfactometer experiments and chemical analyses to investigate the effect of volatiles from cowpea flowers and two companion plants; lemongrass, Cymbopogon citratus and Mexican marigold, Tagetes minuta on the olfactory responses of M. sjostedti. The results revealed that M. sjostedti males and females were repelled by the volatiles from freshly cut leaves of C. citratus. The combination of freshly cut leaves of C. citratus and cowpea flower was repellent to females but not to males. The female thrips, but not males, were repelled by the volatiles from the vegetative stage of T. minuta. Fifty‐four compounds were identified in the volatiles from two herbal plants. Among the major compounds, citral and a 4‐component blend comprised of dihydrotagetone, (Z)‐3‐hexenyl acetate, limonene and (Z)‐β‐ocimene repelled females but dihydrotagetone alone attracted females. While myrcene combined with cowpea flower volatiles enhanced the attraction of females M. sjostedti, when tested alone was not attractive. These results highlight the potential of volatiles from C. citratus and T. minuta to repel M. sjostedti females. The use of these plants as companion plants in a cowpea cropping system could reduce M. sjostedti infestation.