Non-Host Volatile Blend Optimization for Forest Protection against the European Spruce Bark Beetle,Ips typographus

Conifer feeding bark beetles (Coleoptera, Curculionidae, Scolytinae) pose a serious economic threat to forest production. Volatiles released by non-host angiosperm plants (so called non-host volatiles, NHV) have been shown to reduce the risk of attack by many bark beetle species, including the European spruce bark beetle, Ips typographus. However, the most active blend for I. typographus, containing three green leaf volatiles (GLVs) in addition to the key compounds trans-conophthorin (tC) and verbenone, has been considered too expensive for use in large-scale management. To lower the cost and improve the applicability of NHV, we aim to simplify the blend without compromising its anti-attractant potency. Since the key compound tC is expensive in pure form, we also tested a crude version: technical grade trans-conophthorin (T-tC). In another attempt to find a more cost effective substitute for tC, we evaluated a more readily synthesized analog: dehydro-conophthorin (DHC). Our results showed that 1-hexanol alone could replace the three-component GLV blend containing 1-hexanol, (3Z)-hexen-1-ol, and (2E)-hexen-1-ol. Furthermore, the release rate of tC could be reduced from 5 mg/day to 0.5 mg/day in a blend with 1-hexanol and (–)-verbenone without compromising the anti-attractant activity. We further show that T-tC was comparable with tC, whereas DHC was a less effective anti-attractant. DHC also elicited weaker physiological responses in the tC-responding olfactory receptor neuron class, providing a likely mechanistic explanation for its weaker anti-attractive effect. Our results suggest a blend consisting of (–)-verbenone, 1-hexanol and technical trans-conophthorin as a cost-efficient anti-attractant for forest protection against I. typographus.     

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 further sex pheromone synergists in the codling moth, Cydia pomonella

A number of angiosperm nonhost volatiles (NHVs) and green leaf volatiles (GLVs) were tested alone and as supplements to the antiaggregation pheromone, verbenone, for their ability to disrupt attack by the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae), on lodgepole pine, Pinus contorta Dougl. ex Loud. var. latifolia Engel. Preliminary experiments led to a refined NHV blend [benzyl alcohol, guaiacol, benzaldehyde, nonanal, salicylaldehyde, and conophthorin] and a refined GLV blend [(Z)-3-hexen-1-ol and (E)-2-hexen-1-ol]. In a 20-replicate experiment, NHV and GLV groups both singly, and verbenone alone, significantly reduced MPB mass attack on pheromone-baited trees and on trees within 5 m of the pheromone-baited trees. Both blends in combination with verbenone reduced the number of mass attacked, baited trees to three out of 20 compared to 20 out of 20 of the baited controls. Each binary combination was also effective at reducing mass attack. In these experiments, all tested repellents were released from devices stapled to trees at the same point as the pheromone bait, suggesting that the repellency could have been to a point source, rather than to the whole tree. Therefore, in two further experiments bands of release devices were wrapped around the treated trees and the pheromone bait was removed from the treated trees. In one experiment, when the aggregation pheromone bait was suspended between pairs of trees treated with the NHV blend plus GLV blend plus verbenone, only three out of 25 treated pairs had mass attack on at least one member of the pair. In the other 60-replicate experiment, with no pheromone baits present, attack occurred on 13 untreated and 11 banded trees, all in the path of a large advancing infestation. However, the mean attack density on the banded trees was significantly reduced to a level below the 40 attacks m^–2 of bark surface required to kill a healthy lodgepole pine. As a result of these experiments, operational trials are recommended.     

Inhibition of Predator Attraction to Kairomones by Non-Host Plant Volatiles for Herbivores: A Bypass-Trophic Signal

Background

Insect predators and parasitoids exploit attractive chemical signals from lower trophic levels as kairomones to locate their herbivore prey and hosts. We hypothesized that specific chemical cues from prey non-hosts and non-habitats, which are not part of the trophic chain, are also recognized by predators and would inhibit attraction to the host/prey kairomone signals. To test our hypothesis, we studied the olfactory physiology and behavior of a predaceous beetle, Thanasimus formicarius (L.) (Coleoptera: Cleridae), in relation to specific angiosperm plant volatiles, which are non-host volatiles (NHV) for its conifer-feeding bark beetle prey.

Methodology/Principal Findings

Olfactory detection in the clerid was confirmed by gas chromatography coupled to electroantennographic detection (GC-EAD) for a subset of NHV components. Among NHV, we identified two strongly antennally active molecules, 3-octanol and 1-octen-3-ol. We tested the potential inhibition of the combination of these two NHV on the walking and flight responses of the clerid to known kairomonal attractants such as synthetic mixtures of bark beetle (Ips spp.) aggregation pheromone components (cis-verbenol, ipsdienol, and E-myrcenol) combined with conifer (Picea and Pinus spp.) monoterpenes (α-pinene, terpinolene, and Δ³-carene). There was a strong inhibitory effect, both in the laboratory (effect size d = −3.2, walking bioassay) and in the field (d = −1.0, flight trapping). This is the first report of combining antennal detection (GC-EAD) and behavioral responses to identify semiochemical molecules that bypass the trophic system, signaling habitat information rather than food related information.

Conclusions/Significance

Our results, along with recent reports on hymenopteran parasitoids and coleopteran predators, suggest that some NHV chemicals for herbivores are part of specific behavioral signals for the higher trophic level and not part of a background noise. Such bypass-trophic signals could be of general importance for third trophic level players in avoiding unsuitable habitats with non-host plants of their prey.     

Smoke,pheromone and kairomone olfactory receptor neurons in males and females of the pine sawyer Monochamus galloprovincialis (Olivier) (Coleoptera: Cerambycidae)

The response of antennal olfactory receptor neurons (ORNs) of Monochamus galloprovincialis to several odourants was tested using single sensillum electrophysiology. Behaviourally active pheromone, and kairomone (host and sympatric bark beetle pheromone) odours were tested alongside smoke compounds released by burnt wood that are potentially attractive to the insect. The antennae bore several types of sensilla. Two plate areas in the proximal and distal ends of each antennal segment were covered with basiconic sensilla that responded to the odour stimuli. Sensilla basiconica contained one or two cells of different spike amplitude. The 32 male and 38 female ORNs tested responded with excitations or inhibitions to the different plant odours. In general the response of male and female receptors was very similar so they were pooled to perform a cluster analysis on ORN responses. Six ORNs were clearly specialised for pheromone reception. Responses to kairomone and smoke odours were less specific than those of pheromone, but a group of 9 cells was clearly excited by smoke compounds (mainly eugenol and 4-methyl 2-methoxyphenol), a group of 8 cells was very responsive to α-pinene, β-pinene and cis-verbenol, and a group of 14 cells responded to a wider range of compounds. The rest of the cells (47%) were either non-responsive or slightly inhibited by smoke compounds. Dose–response curves were obtained for several compounds. Different compounds induced significantly different latencies and these appeared to be unrelated to their boiling point.     

Representation of a mixture of pheromone and host plant odor by antennal lobe projection neurons of the silkmoth Bombyx mori

Pheromone-source orientation behavior can be modified by coexisting plant volatiles. Some host plant volatiles enhance the pheromonal responses of olfactory receptor neurons and increase the sensitivity of orientation behavior in the Lepidoptera species. Although many electrophysiological studies have focused on the pheromonal response of olfactory interneurons, the response to the mixture of pheromone and plant odor is not yet known. Using the silkmoth, Bombyx mori, we investigated the physiology of interneurons in the antennal lobe (AL), the primary olfactory center in the insect brain, in response to a mixture of the primary pheromone component bombykol and cis-3-hexen-1-ol, a mulberry leaf volatile. Application of the mixture enhanced the pheromonal responses of projection neurons innervating the macroglomerular complex in the AL. In contrast, the mixture of pheromone and cis-3-hexen-1-ol had little influence on the responses of projection neurons innervating the ordinary glomeruli whereas other plant odors dynamically modified the response. Together this suggests moths can process plant odor information under conditions of simultaneous exposure to sex pheromone.     

Interruption of aggregation pheromone in Ips typographus (L.) (Col. Scolytidae) by non-host bark volatiles

Abstract 1 The antennally active nonhost bark volatiles (NHVs): trans‐conophthorin (tC), C₆‐alcohols (green leaf volatiles; GLVs) and C₈‐alcohols, were tested for their ability to reduce attraction of the spruce bark beetle Ips typographus (L) (Col. Scolytidae) to its pheromone sources in both laboratory walking bioassy and field trapping experiments. 2 In the walking bioassay with I. typographus females, individual NHVs such as tC, 3‐octanol and 1‐octen‐3‐ol, and the unsuitable host signal, verbenone (Vn), were inactive at the doses tested. However, the blend of C₆‐alcohols (3GLVs) and all the binary, ternary, or quarternary blends significantly reduced the female attraction to the pheromone sources. 3 In the field trapping experiments, individual NHV signals (tC, C6‐alcohols and C₈‐alcohols) all reduced catch of I. typographus in pheromone‐baited traps, with their inhibitory effects similar to that of the known inhibitor, Vn. The binary, ternary or quarternary combinations of these NHV signals or Vn, all caused significantly stronger reductions in trap catches than the individual signals. The blends showed similar levels of interruption, except the binary blend of C₈‐alcohols (2C8OH) and Vn. 4 Difference in trapping mechanism between pipe traps (attraction and landing) and Lindgren funnel traps (attraction) did not affect the pattern of inhibition of these active NHV signals and Vn. 5 These behaviourally active nonhost volatiles and Vn might be used effectively to protect spruce trees or stands against attacks by I. typographus.     

Specialization of the olfactory receptor cells in the bark beetleIps typographus and its predatorThanasimus formicarius to bark beetle pheromones and host tree volatiles

Summary Olfactory receptor cells of the spruce bark beetle,Ips typographus, and its predator, the clerid beetleThanasimus formicarius, were studied using electrophysiological techniques. Recordings were made of nerve impulses from single cells and of the summated receptor potential (electroantennogram).Information from bark beetle pheromones and host volatiles is detected by separate olfactory receptor cells inI. typographus. Those which detected bark beetle pheromones responded to only one key substance. Some receptor cells which responded to spruce bark volatiles were strongly activated by one of the synthetic host compounds tested. However, too few host compounds were tested to reach definite conclusions about the specialization of host odour cells. T. formicarius has evolved olfactory receptor cells for bark beetle pheromones. These have similar specificities (specialist types) to those of the bark beetles. Furthermore, the predator has olfactory receptor cells for many bark beetle pheromones. This indicates thatT. formicarius is able to detect and discriminate between many bark beetle species. No significant differences were found between prey and predator cells which responded to host volatiles.     

Olfactory recognition and behavioural avoidance of angiosperm nonhost volatiles by conifer-inhabiting bark beetles

Abstract 1 When searching for suitable hosts in flight, especially in mixed forests, conifer‐inhabiting bark beetles will encounter not only suitable host trees and their odours, but also unsuitable hosts and nonhost trees. Rejection of these trees could be based on an imbalance of certain host characteristics and/or a negative response to some nonhost stimuli, such as nonhost volatiles (NHV). 2 Recent electrophysiological and behavioural studies clearly indicate that conifer‐inhabiting bark beetles are not only able to recognize, but also to avoid, nonhost habitats or trees by olfactory means. Green leaf volatiles (GLV), especially C₆‐alcohols, from the leaves (and partly from bark) of nonhost angiosperm trees, may represent nonhost odour signals at the habitat level. Specific bark volatiles such as trans‐conophthorin, C₈‐alcohols, and some aromatic compounds, may indicate nonhosts at the tree species level. Flying bark beetles are also capable of determining whether a possible host is unsuitable by reacting to signals from conspecifics or sympatric heterospecifics that indicate old or colonized host tree individuals. 3 Combined NHV signals in blends showed both redundancy and synergism in their inhibitory effects. The coexistence of redundancy and synergism in negative NHV signals may indicate different functional levels (nonhost habitats, species, and unsuitable hosts) in the host selection process. Combinations of NHV and verbenone significantly reduced the number of mass attacked host trees or logs on several economically important species (e.g. Dendroctonus ponderosae, Ips typographus, and I. sexdentatus). 4 We suggest a semiochemical‐diversity hypothesis, based on the inhibition by NHV of bark beetle host‐location, which might partly explain the lower outbreak rates of forest insects in mixed forests. This ‘semiochemical‐diversity hypothesis’ would provide new support to the general ‘stability‐diversity hypothesis’. 5 Natural selection appears to have caused conifer‐inhabiting bark beetles to evolve several olfactory mechanisms for finding their hosts and avoiding unsuitable hosts and nonhost species. NHV and unsuitable host signals have potential for use in protecting trees from attack. The use of these signals may be facilitated by the fact that their combination has an active inhibition radius of several metres in trap test, and by the observation of area effects for several trees near inhibitor soruces in tree protection experiments. Furthermore, incorporation of negative signals (such as NHV and verbenone) and pheromone‐based mass‐trapping in a ‘push–pull’ fashion may significantly increase the options for control against outbreaks of conifer‐inhabiting bark beetles, especially in high risk areas.     

三种绿叶挥发性物质对云南切梢小蠹寄主定位行为的干扰作用

为了研究环境中非寄主阔叶植物释放出的绿叶挥发性物质（GLVs）对针叶树蛀干害虫云南切梢小蠹Tomicus yunnanesis的影响, 选取了(E)-2-己烯醛、 (E)-2-己烯醇和(Z)-3-己烯醇3种释放量较大的绿叶挥发性物质, 通过室内松梢取食试验测试了单组分及两两混合后对云南切梢小蠹寄主定位行为的干扰作用。结果表明： 源于阔叶植物的3种绿叶挥发性物质及其混合物能够不同程度干扰云南切梢小蠹的寄主定位行为。当虫放入广口瓶12 h后, 3个单组分绿叶挥发性物质处理组［A： (E)-2-己烯醛, P＜0.01; B： (E)-2-己烯醇, P＜0.01; C： (Z)-3-己烯醇, P＜0.01］及2个混合组分［D： (E)-2-己烯醛+(E)-2-己烯醇, P＜0.01）; E： (E)-2-己烯醛+(Z)-3-己烯醇, P＜0.01］中滞留在松梢外部的虫数与对照组相比都有显著性差异, 绿叶挥发性物质的存在显著降低了云南切梢小蠹侵害云南松松梢的概率。但是, 24 h后只有D组（P＜0.01）和E组（P＜0.01）滞留在松梢外部的虫数与对照组相比具有显著性差异, 在48 h后只有D组（P＜0.01）与对照相比仍具有显著性差异。本研究为利用非寄主植物的次生代谢产物防治云南切梢小蠹进行了有益的探索。     

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.     

Characterization of olfactory receptor neurons for pheromone candidate and plant volatile compounds in the clover root weevil,Sitona lepidus

Antennal olfactory receptor neurons (ORNs) for pheromone and plant volatile compounds were identified and characterized in male and female clover root weevil, Sitona lepidus (Gyllenhal), using the single sensillum recording technique with five pheromone-related compounds, and 40 host and non-host plant volatile compounds. Overall, seven different types of olfactory sensilla containing specialized ORNs were identified in each sex of S. lepidus. Among them, three different types of sensilla in the males and two types in the females housed ORNs specialized for pheromone-related compounds. The ORNs in males were specialized for 4-methyl-3,5-heptanedione or one or more of four stereoisomers of 5-hydroxy-4-methyl-3-heptanone. In contrast, female sensilla did not contain ORNs sensitive to 4-methyl-3,5-heptanedione while they contained ORNs sensitive to and specialized for the stereoisomers of (4S,5S)-5-hydroxy-4-methyl-3-heptanone. In addition to the pheromone-related ORNs, four types of olfactory sensilla contained ORNs responsive to plant volatile compounds in male S. lepidus, and five types in females. Most of the ORNs identified in S. lepidus showed a high degree of specificity to specific volatile compounds although some of the active compounds showed overlapping response spectra in the ORNs across different types of sensilla. The most active plant volatile compounds were the four green leaf volatile compounds, (E)-2-hexenol, (Z)-2-hexenol, (Z)-3-hexenol and (E)-2-hexenal, and isomers of two monoterpenols, (±)-linalool and (±)-α-terpineol, all eliciting strong responses from relatively large numbers of ORNs in male and female S. lepidus. Our study indicates that S. lepidus has a set of highly sensitive and selective ORNs for pheromone and plant volatile compounds. Further work is needed to elucidate the behavioral implications of these findings.     

Functional Evolution of a Bark Beetle Odorant Receptor Clade Detecting Monoterpenoids of Different Ecological Origins

Insects detect odors using an array of odorant receptors (ORs), which may expand through gene duplication. How and which new functions may evolve among related ORs within a species remain poorly investigated. We addressed this question by functionally characterizing ORs from the Eurasian spruce bark beetle Ips typographus, in which physiological and behavioral responses to pheromones, volatiles from host and nonhost trees, and fungal symbionts are well described. In contrast, knowledge of OR function is restricted to two receptors detecting the pheromone compounds (S)-(–)-ipsenol (ItypOR46) and (R)-(–)-ipsdienol (ItypOR49). These receptors belong to an Ips-specific OR-lineage comprising seven ItypORs. To gain insight into the functional evolution of related ORs, we characterized the five remaining ORs in this clade using Xenopus oocytes. Two receptors responded primarily to the host tree monoterpenes (+)-3-carene (ItypOR25) and p-cymene (ItypOR27). Two receptors responded to oxygenated monoterpenoids produced in larger relative amounts by the beetle-associated fungi, with ItypOR23 specific for (+)-trans-(1R, 4S)-4-thujanol, and ItypOR29 responding to (+)-isopinocamphone and similar ketones. ItypOR28 responded to the pheromone E-myrcenol from the competitor Ips duplicatus. Overall, the OR responses match well with those of previously characterized olfactory sensory neuron classes except that neurons detecting E-myrcenol have not been identified. The characterized ORs are under strong purifying selection and demonstrate a shared functional property in that they all primarily respond to monoterpenoids. The variation in functional groups among OR ligands and their diverse ecological origins suggest that neofunctionalization has occurred early in the evolution of this OR-lineage following gene duplication.     

Fungal,host and non-host volatiles modify attraction of the walnut twig beetle,Pityophthorus juglandis,to pheromone lures

1. Thousand Cankers Disease (TCD) of walnut trees is caused by the pathogenic fungus Geosmithia morbida vectored by the walnut twig beetle (WTB) Pityophthorus juglandis. Monitoring efforts for WTB rely on pheromone-baited traps, but lures are likely effective at attracting beetles only over short distances. Fungal-derived kairomones may increase the efficacy of current lures, while additional volatiles may repel beetles from valuable trees.
2. The objective of this study was to determine the extent to which fungal, host and non-host volatiles modify the attraction of WTB to pheromone-baited traps. A trapping study that combined fungal, host-associated and non-host compounds with WTB-pheromone lures was conducted over three years in black walnut plantations experiencing a TCD outbreak in Walla Walla, WA.
3. Traps baited with pheromone and G. morbida volatiles (i.e., isoamyl and isobutyl alcohol) consistently attracted more WTB, while other fungal volatiles inconsistently increased attraction compared to those baited with pheromone lure alone. This is the first field study that demonstrates fungal volatiles can increase the attraction of a bark beetle to its pheromone in a hardwood system.
4. One fungal (benzyl alcohol) and two additional volatiles (limonene, piperitone) repelled WTB from pheromone-baited traps. Although limonene is known to repel WTB, this is the first demonstration that benzyl alcohol and piperitone repel a bark beetle.
5. Fungal volatiles may increase the efficacy of monitoring efforts and may play an important role in management tactics for WTB, especially in detecting the introduction and establishment of nascent populations and protecting trees from colonizing beetles.

     

Role of plant volatiles in host plant location of the leafminer, Liriomyza sativae (Diptera: Agromyzidae)

Abstract The role of plant volatiles in host plant location of the leafminer Liriomyza sativae Blanchard was studied. Four types of antennal sensilla were identified on the funiculus by scanning electron microscopy: trichoid, basiconic, clavate and grooved sensilla. An olfactory pit, containing groups of sensilla, was present on the ventral side of the funiculus. No sexual difference was detected in sensilla diversity and distribution. In behavioural assays, both males and females were attracted by the odour of the bean Phaseolus vulgaris L. They had distinct EAG responses to the bean odour. No significant sexual difference was found in behaviour or EAG responses.
Electroantennograms were recorded from female L. sativae to 14 plant volatile compounds. The most distinct EAG responses were obtained for: (1) the general green leaf volatiles 1-hexanol (E)-2-hexen-1-ol, (E)-3-hexen-1-ol and its isomers, (Z)-3-hexen-1-ol, the acetate (E)-3-hexenylacetate and the aldehyde hexanal; and (2) limonene, a compound associated with tomato, which is a key host plant of this insect. Other volatile compounds associated with host plants, such as α-pinene, myrcene, β-caryophyllene, and eugenol did not elicit responses. The ability of this insect to locate a host plant appears to be augmented by the perception of a combination of host-specific and general green leaf volatiles. A modification of the EAG recording method of Dipteran species was provided.     

Pine shoot beetle, Tomicus piniperda (Col., Scolytidae), responses to common green leaf volatiles

We tested the hypothesis that green leaf volatiles (GLVs) disrupt the response of overwintered pine shoot beetles, Tomicus piniperda (L.) to multiple-funnel traps baited with the attractive host volatile α-pinene. A combination of four GLV alcohols, 1-hexanol ( E )-2-hexen-1-ol ( Z )-2-hexen-1-ol, and ( Z )-3-hexen-1-ol, caused 54 and 36% reduction in the number of pine shoot beetles captured in two separate trapping experiments. Similarly, a combination of the four alcohols plus two GLV aldehydes, hexanal and ( E )-2-hexenal, caused 38% reduction in the number of pine shoot beetles captured compared with α-pinene alone. A blend of the two GLV aldehydes was not disruptive. None of the four GLV alcohols nor the two GLV aldehydes were disruptive when tested individually. The finding that the blend of four GLV alcohols reduced attraction of T. piniperda supports the general hypothesis that GLVs common to nonhost angiosperms are disruptive to conifer-attacking bark beetles (Scolytidae).     

Host and non-host odour signals governing host selection by the pine shoot beetle, Tomicus piniperda and the spruce bark beetle, Hylurgops palliatus (Col., Scolytidae)

Abstract:  Tomicus piniperda and Hylurgops palliatus colonize susceptible host trees by responding to host-specific odour signals as well as by avoiding volatiles emanating from non-host conifers. In the field, the pine shoot beetle, T. piniperda , responded in high numbers to natural odour sources provided by their host tree, Pinus sylvestris , while the non-host conifers Larix decidua , Picea abies , or Pseudotsuga menziesii were significantly less attractive. In contrast, the spruce bark beetle, Hylurgops palliatus , preferentially responded to its main host, P. abies . Furthermore, T. piniperda attacks on P. sylvestris bolts decreased in presence of bark and wood particles from the non-host P. abies , whereas particles from P. menziesii appeared not to affect T. piniperda attacks. Apparently, tree-specific volatiles act at close range as specific signals that lead to the successful discrimination and colonization of the respective host tree species.     

Electroantennogram responses of the Colorado beetle, Leptinotarsa decemlineata, to plant volatiles

Electroantennogram responses of Colorado beetles were recorded to 53 plant volatiles including isomers. The system of antennal olfactory receptors is selective, even at high doses several compounds cause fairly small responses. Diminishing the concentration to a moderate stimulus strength reduces the number of perceptible chemicals. Distinct electroantennogram responses are obtained to a group of closely related components, namely the general green leaf volatiles trans-2-hexen-1-ol, cis-3-hexen-1-ol, hexanol-1, trans-2-hexenal, hexanal and cis-3-hexenyl-acetate, and to isomers such as trans-3-hexen-1-ol and cis-2-hexen-1-ol. The threshold concentration of the most effective compound, trans-2-hexen-1-ol is 1.2×10⁸ molecules per ml of air. The antennal olfactory receptors of the Colorado beetle are sensitively tuned to the perception of these general green leaf volatiles. As olfactory receptors of a number of phytophagous insects have been reported to respond to these components, this volatile complex probably plays a part in the host selection behaviour of various phytophagous insects.

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Résumé Les électroantennogrammes de Leptinotarsa decemlineata en présence de 53 substances volatiles (y compris des isomères) ont été enregistrés. Le système des récepteurs olfactifs antennaires est sélectif, même à des doses élevées plusieurs composés ne provoquent que d'assez faibles réponses. La diminution de la concentration jusqu'à une puissance stimulante modérée réduit le nombre de substances perçues. Des électroantennogrammes différents ont été obtenus pour un groupe de composés très voisins, à savoir les substances volatiles de feuilles vertes: trans-2-hexen-1-ol, cis-3-hexen-1-ol, hexanol-1, trans-2-hexenal et cis-3-hexenyl-acetate, et aux isomères tels que trans-3-hexen-1-ol et cis-2-hexen-l-ol. La concentration seuil du composé le plus efficace, trans-2-hexen-1-ol, est de 1,2×10⁸ molécules par ml d'air. Les récepteurs olfactifs antennaires du Doryphore sont sensoriellement ajustés à la perception de ces substances volatiles générales des feuilles vertes. Comme les récepteurs olfactifs de nombreux insectes phytophages ont été signalés comme répondant à ces composés, ce complexe volatile joue probablement un rôle dans le comportement de sélection de l'hôte par différents insectes phytophages.

     

Additive and synergistic integration of multimodal cues of both hosts and non-hosts during host selection by woodboring insects

Host selection by herbivorous insects is primarily thought to depend on attraction to olfactory cues emitted from the host species. However, the discrimination of these hosts from non-host species may also arise from the adaptive detection and avoidance of non-host cues, including visual characteristics. Many generalist, conifer-colonising beetles, for example, use characteristic volatiles to identify and discriminate against non-host angiosperm trees such as aspens and birches while flying. These trees also differ in bark reflectance characteristics, which could also mediate host/non-host discrimination by interacting with semiochemicals. We tested this hypothesis by evaluating the responses of eight species of polyphagous woodboring beetles to traps which simulated the visual appearance of coniferous hosts (black) and angiosperm non-hosts (white), and which were baited with host or non-host volatiles. As predicted, three species of large woodboring beetle and a woodboring wasp all avoided white, non-host-simulating traps that were baited with attractive kairomones, and preferred black, host-simulating traps. Conversely, three ambrosia beetle species demonstrated weaker visual preferences, possibly because they preferentially colonise fallen hosts that would transmit less accurate visual information. However, the ambrosia beetle Trypodendron lineatum did show a greater preference for host-coloured traps when these released host-associated kairomones in addition to their pheromone, and also avoided white non-host traps, but only when these released non-host volatiles. To our knowledge, this is the first evidence that multiple non-host cues could synergistically mediate the adaptive discrimination of hosts and non-hosts. Our results suggest that successful host location by generalists arises from the complex integration of cues in multiple sensory modes, and that foraging herbivores evaluate both hosts and non-hosts during their search.     

Enhanced attraction of Plutella xylostella (Lepidoptera: Plutellidae) to pheromone-baited traps with the addition of green leaf volatiles

Diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), is one of the most serious pests of Brassicaceae crops worldwide. Electrophysiological and behavioral responses of P. xylostella to green leaf volatiles (GLVs) alone or together with its female sex pheromone were investigated in laboratory and field. GLVs 1-hexanol and (Z)-3-hexen-1-ol elicited strong electroantennographic responses from unmated male and female P. xylostella, whereas (Z)-3-hexenyl acetate only produced a relatively weak response. The behavioral responses of unmated moths to GLVs were further tested in Y-tube olfactometer experiments. (E)-2-Hexenal, (Z)-3-hexen-1-ol, and (Z)-3-hexenyl acetate induced attraction of males, reaching up to 50%, significantly higher than the response to the unbaited control arm. In field experiments conducted in 2008 and 2009, significantly more moths were captured in traps baited with synthetic sex pheromone with either (Z)-3-hexenyl acetate alone or a blend of (Z)-3-hexenyl acetate, (Z)-3-hexen-1-ol, and (E)-2-hexenal compared with sex pheromone alone and other blend mixtures. These results demonstrated that GLVs could be used to enhance the attraction of P. xylostella males to sex pheromone-baited traps.