Host resistance elicited by methyl jasmonate reduces emission of aggregation pheromones by the spruce bark beetle, Ips typographus

We treated Norway spruce (Picea abies) stems with methyl jasmonate (MeJA) to determine possible quantitative and qualitative effects of induced tree defenses on pheromone emission by the spruce bark beetle Ips typographus. We measured the amounts of 2-methyl-3-buten-2-ol and (S)-cis-verbenol, the two main components of the beetle's aggregation pheromone, released from beetle entrance holes, along with phloem terpene content and beetle performance in MeJA-treated and untreated Norway spruce logs. As expected, phloem terpene levels were higher and beetle tunnel length was shorter (an indication of poor performance) in MeJA-treated logs relative to untreated logs. Parallel to the higher phloem terpene content and poorer beetle performance, beetles in MeJA-treated logs released significantly less 2-methyl-3-buten-2-ol and (S)-cis-verbenol, and the ratio between the two pheromone components was significantly altered. These results suggest that host resistance elicited by MeJA application reduces pheromone emission by I. typographus and alters the critical ratio between the two main pheromone components needed to elicit aggregation. The results also provide a mechanistic explanation for the reduced performance and attractivity observed in earlier studies when bark beetles colonize trees with elicited host defenses, and extend our understanding of the ecological functions of conifer resistance against bark beetles.     

Host tree resistance influencing pheromone production in Ips typographus (Coleoptera: Scolytidae)

Analyses of volatiles in hindguts of Ips typgraphus males from different spruce trees and attack phases are reviewed. The composition of monoterpenes, and the chirality of α-pinene, have been determined in phloem samples. Relationships between compounds emanating from spruce trees and bark beetles, respectively, have been studied. Male beetles depend on their host tree for the production of pinene alcohols. The ratio between the pinene alcohols is almost constant in males boring in the same tree but can vary widely between males from different spruce trees. Very good correlations were found between some host tree monoterpene hydrocarbons and bark beetle produced pinene alcohols. The production of the essential pheromone component 2-methyl-3-buten-2-ol, was not correlated with the monoterpene content in the host trees, while the production of the other essential pheromone component, cis-verbenol, depends on the amount of the precursor, (−)-α-pinene in the phloem. Male beetles boring in a resistant spruce tree will continue to produce the pinene alcohols, including cis -verbenol, as long as the tree defends itself with resin.     

Influence of the proportion of (−) α-pinene in pheromone bait on Ips typographus (Col., Scolytidae) catch in pheromone trap barriers and in single traps

Abstract:  Further possibilities of increasing catch efficiency of Ips typographus L. in pheromone trap barriers with two different levels of release of (4S)- cis -verbenol (cV) and in single traps were tested. Pheromone mixtures containing cV, 2-methyl-3-buten-2-ol (MB) and 0, 1, 10, 35, 60 and 90% (−) α -pinene were prepared. Basically, MB was successively replaced by (−) α -pinene in the mixtures. Tests were performed on the edge of a spruce forest when outbreaks of I. typographus occurred. The substitution of MB with (−) α -pinene had different effects in different types of pheromone trap barriers and in single traps. It is not possible to increase the attractiveness of the bait having a higher percentage of cV by supplementing a certain percentage of MB with (−) α -pinene. The catch of I. typographus decreased with an increase in the percentage of (−) α -pinene in the bait. It is possible to increase the attractiveness of the bait having a lower percentage of cV by supplementing certain percentages of MB with (−) α -pinene. A slight synergistic effect of (−) α -pinene with MB/cV mixture is visible in this type of bait. In the case of single traps, supplementing MB with (−) α -pinene possibly causes a slight synergistic effect. An increase in the percentage of (−) α -pinene increases the percentage of males in the catch.     

Attractive responses by Monochamus galloprovincialis (Col., Cerambycidae) to host and bark beetle semiochemicals

Abstract:  The pine sawyer Monochamus galloprovincialis is the European vector of the recently introduced pine wood nematode. This nematode is the causal organism of pine wilt disease, a serious tree killer in East Asia. Efficacious baits and traps to monitor and control this beetle are now required. The effect of bark beetle ( Ips spp.) pheromone components, released individually (ipsenol) or in blends (ipsenol, ipsdienol, cis -verbenol and methyl-butenol), together with host volatiles (turpentine or α -pinene and ethanol) on M. galloprovincialis trap catches has been studied in Spain. A kairomonal response by male and female of M. galloprovincialis to Ips semiochemicals was found. Beetles were more attracted to host blends supplemented with bark beetle pheromones than to host volatiles alone. Ipsenol alone was attractive to pine sawyers, and was synergistic with α -pinene and ethanol. The full blend of the four Ips semiochemicals and the host compounds was highly attractive. Multiple-funnel traps were as effective as black cross-vane traps in capturing this insect when the escape of trapped beetles was prevented. Trapping of non-target bark beetle predators was also evaluated. The trogossitid Temnochila coerulea and clerid Thanasimus formicarius were kairomonally attracted to and killed in traps baited with bark beetle pheromones. These results suggest that effective monitoring of M. galloprovincialis would be possible by baiting any of these traps with host volatiles and Ips semiochemicals, but reduction of the lure components and trap modification to minimize impact on predators should be considered.     

Behavioural and electrophysiological responses of the bark beetle, Ips typographus, to potential pheromone components

ABSTRACT. Field tests showed ( S )-(-)- cis -verbenol and ( R )-(+)- trans -verbenol in combination with a second pheromonal component, 2-methyl-3-buten-2-ol, more attractive than the combination with their optical antipodes. Inhibition of response to the attractant component, ( S )-(-)- cis -verbenol, by high concentrations of its optical antipode did not occur. No significant differences were noted for response to the attractant, ( S )-(-)- cis -verbenol and 2-methyl-3-buten-2-ol, with the addition of either ipsdienol enantiomer or a racemic mixture of ipsdienol enantiomers. Electroantennogram (EAG) studies correlated well with the behavioural studies. EAGs recorded from male and female beetles revealed both sexes to have a lower threshold for the pheromone, ( S )-(-)- cis -verbenol, than its host terpene precursor, (-)- alpha -pinene. EAGs showed a greater number of acceptors for (-)- alpha-pinene in males than in females. EAGs at acceptor saturation to the enantiomers of alpha -pinene and the verbenol isomers showed males more responsive to (-)- alpha -pinene, (±)- cis -verbenol, and ( R )-(+)- trans -verbenol. Significantly greater EAGs were elicited in females than in males to (-)- alpha -pinene, and (±)- and ( S )-(-)- cis -verbenol. No significant differences in EAGs of females to the enantiomers of trans -verbenol were noted. EAGs showed similar thresholds in males and females to the pheromone component, 2-methyl-3-buten-2-ol; however, female response at threshold was significantly greater than male response. The results are discussed with regard to olfactory acceptor evolution.     

Aggregation response of European engraver beetles of the genus Ips mediated by terpenoid pheromones

Upon exposure to vapours of the host tree's monoterpene (–)--pinene, Ips acuminatus produces the terpene alcohol cis-verbenol which, besides ipsdienol and ipsenol, is a component of its aggregation pheromone. I. cembrae transforms the same monoterpene predominantly into myrtenol and trans-verbenol but little into cis-verbenol, which interrupts pheromone response under field conditions.On the other hand, I. cembrae releases, upon feeding in the bark, the terpene alcohol amitinol in major amounts. Amitinol significantly enhances field response to the aggregation pheromone that also includes ipsdienol, ipsenol, and 3-methyl-3-buten-1-ol. However, amitinol reduces pheromone response in I. acuminatus and I. erosus. Also, males of I. sexdentatus release amitinol which appears to increase response to its attractive principle pheromone component, racemic ipsdienol. There is some evidence that present knowledge of the chemical communication systems among European Ips spp. still lacks satisfactory explanation of the naturally occuring aggregation en masse, perhaps with the exception of I. erosus and I. typographus.     

Selective manipulation of predators using pheromones: responses to frontalin and ipsdienol pheromone components of bark beetles in the Great Lakes region

Abstract 1 One proposed approach to improving biological control of bark beetles (Coleoptera: Scolytidae; alt. Curculionidae: Scolytinae) is to manipulate predator movement using semiochemicals. However, selective manipulation is impeded by attraction of both predators and pests to bark beetle pheromones. 2 The primary bark beetle affecting pine plantations in Wisconsin, U.S.A., is the pine engraver, Ips pini (Say). Other herbivores include Ips grandicollis (Eichhoff) and Dryophthorus americanus Bedel (Curculionidae). The predominant predators are the beetles Thanasimus dubius (Cleridae) and Platysoma cylindrica (Histeridae). 3 We conducted field assays using two enantiomeric ratios of ipsdienol, and frontalin plus α‐pinene. Ipsdienol is the principal pheromone component of I. pini, and frontalin is produced by a number of Dendroctonus species. α‐Pinene is a host monoterpene commonly incorporated into commercial frontalin lures. 4 Thanasimus dubius was attracted to frontalin plus α‐pinene, and also to racemic ipsdienol. By contrast, I. pini was attracted to racemic ipsdienol, but showed no attraction to frontalin plus α‐pinene. Platysoma cylindrica was attracted to 97%‐(–)‐ipsdienol and, to a lesser extent, racemic ipsdienol, but not to frontalin plus α‐pinene. Ips grandicollis was attracted to frontalin plus α‐pinene but not to ipsdienol. Dryophthorus americanus was attracted to both ipsdienol and frontalin plus α‐pinene. 5 This ability to selectively attract the predator T. dubius without attracting the principal bark beetle in the system, I. pini, provides new opportunities for research into augmentative biological control and basic population dynamics. Moreover, the attraction of T. dubius, but not P. cylindrica, to frontalin plus α‐pinene creates opportunities for selective manipulation of just one predator. 6 Patterns of attraction by predators and bark beetles to these compounds appear to reflect various degrees of geographical and host tree overlap with several pheromone‐producing species.     

Geographic variation in prey preference in bark beetle predators

Abstract 1. Bark beetles and their predators are useful systems for addressing questions concerning diet breadth and prey preference in arthropod natural enemies. These predators use bark beetle pheromones to locate their prey, and the response to different pheromones is a measure of prey preference. 2. Trapping experiments were conducted to examine geographic variation in the response to prey pheromones by two bark beetle predators, Thanasimus dubius and Temnochila virescens. The experiments used pheromones for several Dendroctonus and Ips prey species (frontalin, ipsdienol, and ipsenol) and manipulated visual cues involved in prey location (black vs. white traps). The study sites included regions where the frontalin‐emitter Dendroctonus frontalis was in outbreak vs. endemic or absent. 3. There was significant geographic variation in pheromone preference for T. dubius. This predator strongly preferred a pheromone (frontalin) associated with D. frontalis at outbreak sites, while preference was more even at endemic and absent sites. No geographic variation was found in the response by T. virescens. White traps caught fewer insects than black traps for both predators, suggesting that visual cues are also important in prey location. 4. The overall pattern for T. dubius is consistent with switching or optimal foraging theory, assuming D. frontalis is a higher quality prey than Ips. The two predator species partition the prey pheromones in areas where D. frontalis is abundant, possibly to minimise competition and intraguild predation.     

Pine monoterpenes and pine bark beetles: a marriage of convenience for defense and chemical communication

Pine-feeding bark beetles (Coleoptera: Scolytidae) interact chemically with their host pines (Coniferales: Pinaceae) via the behavioral, physiological, and biochemical effects of one class of isoprenoids, the monoterpenes and their derivatives. Pine monoterpenes occur in the oleoresin and function as behaviorally active kairomones for pine bark beetles and their predators, presenting a classic example of tri-trophic chemical communication. The monoterpenes are also essential co-attractants for pine bark beetle aggregation pheromones. Ironically, pine monoterpenes are also toxic physiologically to bark beetles at high vapor concentrations and are considered an important component of the defense of pines. Research over the last 30 years has demonstrated that some bark beetle aggregation pheromones arise through oxygenation of monoterpenes, linking pheromone biosynthesis to the host pines. Over the last 10 years, however, several frequently occurring oxygenated monoterpene pheromone components (e.g., ipsenol, ipsdienol and frontalin) have also been shown to arise through highly regulated de novo pathways in the beetles (reviewed in Seybold and Tittiger, 2003). The most interesting nexus between these insects and their plant hosts involves the late-stage reactions in the monoterpenoid biosynthetic pathway, during which isomeric dimethylallyl diphosphate and isopentenyl diphosphate are ultimately elaborated to stereospecific monoterpenes in the trees and to hydroxylated monoterpenes or bicyclic acetals in the insects. There is signal stereospecificity in both production of and response to the monoterpenoid aggregation pheromones of bark beetles and in response to␣the monoterpenes of the pines. In the California fivespined ips, Ips paraconfusus, we have discovered a number of cytochome P450 genes that have expression patterns indicating that they may be involved in detoxifying monoterpene secondary metabolites and/or biosynthesizing pheromone components. Both processes result in the production of oxygenated monoterpenes, likely with varying degrees of stereospecificity. A behavioral analysis of the stereospecific response of I. paraconfusus to its pheromone is providing new insights into the development of an efficacious bait for the detection of this polyphagous insect in areas outside the western United States. In contrast, a Eurasian species that has arrived in California, the Mediterranean pine engraver, Orthotomicus (Ips) erosus, utilizes both a monoterpenoid (ipsdienol) and a hemiterpenoid (2-methyl-3-buten-2-ol) in its pheromone blend. The stereospecificity of the response of O. erosus to the monoterpenoid appears to be the key factor to the improved potency of the attractant bait for this invasive species.Dedicated to Professor David L. Wood on the occasion of his 75th birthday, January 8, 2006     

Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: implications for pest management and ecology of bark beetles

Abstract 1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of α‐pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of α‐pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of α‐pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of α‐pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.     

Inducibility of chemical defenses in Norway spruce bark is correlated with unsuccessful mass attacks by the spruce bark beetle

Secondary attraction to aggregation pheromones plays a central role in the host colonization behavior of the European spruce bark beetle Ips typographus. However, it is largely unknown how the beetles pioneering an attack locate suitable host trees, and eventually accept or reject them. To find possible biomarkers for host choice by I. typographus, we analyzed the chemistry of 58 Norway spruce (Picea abies) trees that were subsequently either (1) successfully attacked and killed, (2) unsuccessfully attacked, or (3) left unattacked. The trees were sampled before the main beetle flight in a natural Norway spruce-dominated forest. No pheromones were used to attract beetles to the experimental trees. To test the trees' defense potential, each tree was treated in a local area with the defense hormone methyl jasmonate (MeJ), and treated and untreated bark were analyzed for 66 different compounds, including terpenes, phenolics and alkaloids. The chemistry of MeJ-treated bark correlated strongly with the success of I. typographus attack, revealing major chemical differences between killed trees and unsuccessfully attacked trees. Surviving trees produced significantly higher amounts of most of the 39 analyzed mono-, sesqui-, and diterpenes and of 4 of 20 phenolics. Alkaloids showed no clear pattern. Differences in untreated bark were less pronounced, where only 1,8-cineole and (-)-limonene were significantly higher in unsuccessfully attacked trees. Our results show that the potential of individual P. abies trees for inducing defense compounds upon I. typographus attack may partly determine tree resistance to this bark beetle by inhibiting its mass attack.     

Biosynthesis of 2-methyl-3-buten-2-ol,a pheromone component of Ips typographus (Coleoptera: Scolytidae)

Exposure of beetles to ¹⁴C-labelled mevalonate by injection, resulted in significant incorporation of radioactivity in 2-methyl-3-buten-2-ol. Radiolabelled 2-methyl-3-buten-2-ol was obtained both by organic solvent extraction of beetle hindguts and by entrainment of volatiles in air surrounding logs with boring beetles. It is suggested that 2-methyl-3-buten-2-ol, an essential component of the aggregation pheromone of Ips typographus, can be synthesized from mevalonate.     

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.     

Pheromone production in bark beetles

The first aggregation pheromone components from bark beetles were identified in 1966 as a mixture of ipsdienol, ipsenol and verbenol. Since then, a number of additional components have been identified as both aggregation and anti-aggregation pheromones, with many of them being monoterpenoids or derived from monoterpenoids. The structural similarity between the major pheromone components of bark beetles and the monoterpenes found in the host trees, along with the association of monoterpenoid production with plant tissue, led to the paradigm that most if not all bark beetle pheromone components were derived from host tree precursors, often with a simple hydroxylation producing the pheromone. In the 1990s there was a paradigm shift as evidence for de novo biosynthesis of pheromone components began to accumulate, and it is now recognized that most bark beetle monoterpenoid aggregation pheromone components are biosynthesized de novo. The bark beetle aggregation pheromones are released from the frass, which is consistent with the isoprenoid aggregation pheromones, including ipsdienol, ipsenol and frontalin, being produced in midgut tissue. It appears that exo-brevocomin is produced de novo in fat body tissue, and that verbenol, verbenone and verbenene are produced from dietary α-pinene in fat body tissue. Combined biochemical, molecular and functional genomics studies in Ips pini yielded the discovery and characterization of the enzymes that convert mevalonate pathway intermediates to pheromone components, including a novel bifunctional geranyl diphosphate synthase/myrcene synthase, a cytochrome P450 that hydroxylates myrcene to ipsdienol, and an oxidoreductase that interconverts ipsdienol and ipsdienone to achieve the appropriate stereochemistry of ipsdienol for pheromonal activity. Furthermore, the regulation of these genes and their corresponding enzymes proved complex and diverse in different species. Mevalonate pathway genes in pheromone producing male I. pini have much higher basal levels than in females, and feeding induces their expression. In I. duplicatus and I. pini, juvenile hormone III (JH III) induces pheromone production in the absence of feeding, whereas in I. paraconfusus and I. confusus, topically applied JH III does not induce pheromone production. In all four species, feeding induces pheromone production. While many of the details of pheromone production, including the site of synthesis, pathways and knowledge of the enzymes involved are known for Ips, less is known about pheromone production in Dendroctonus. Functional genomics studies are under way in D. ponderosae, which should rapidly increase our understanding of pheromone production in this genus. This chapter presents a historical development of what is known about pheromone production in bark beetles, emphasizes the genomic and post-genomic work in I. pini and points out areas where research is needed to obtain a more complete understanding of pheromone production.     

不同小蠹类与天牛类引诱剂联用对蛀干甲虫的林间诱捕效果

松墨天牛Monochamus alternatus Hope是传播松材线虫病的主要媒介,引诱剂是抑制松墨天牛种群数量的无公害调控有效方式之一。本研究通过在中国浙江富阳的林间试验测定了6种小蠹类引诱剂与1种天牛引诱剂不同组合联用对松墨天牛成虫以及其他蛀干害虫的诱捕效果。结果表明： 横坑切梢小蠹Tomicus minor引诱剂3-carene-10-ol、云杉八齿小蠹Ips typographus引诱剂2-methyl-3-buten-2-ol、欧洲榆小蠹Scolytus multistriatus引诱剂4-methyl-3-heptanol和纵坑切梢小蠹T. piniperda引诱剂verbenol分别与松墨天牛引诱剂联用后,对松墨天牛的引诱效果无显著性差异(P<0.05)。西部松大小蠹Dendroctonus brevicomis 引诱剂exo-brevicomin或黄杉大小蠹D. pseudotsugae引诱剂1-methylcyclohex-2-en-l-ol与松墨天牛引诱剂联用后对松墨天牛的诱捕效果有减弱的趋势。另外,与单独使用中国天牛引诱剂相比,小蠹类引诱剂与其联用后在诱捕其他蛀干害虫的种类和个体数量上均有所减少。     

Functional characterization of myrcene hydroxylases from two geographically distinct Ips pini populations

Ips pini bark beetles use myrcene hydroxylases to produce the aggregation pheromone component, ipsdienol, from myrcene. The enantiomeric ratio of pheromonal ipsdienol is an important prezygotic mating isolation mechanism of I. pini and differs among geographically distinct populations. We explored the substrate and product ranges of myrcene hydroxylases (CYP9T2 and CYP9T3) from reproductively-isolated western and eastern I. pini. The two cytochromes P450 share 94% amino acid identity. CYP9T2 mRNA levels were not induced in adults exposed to myrcene-saturated atmosphere. Functional assays of recombinant enzymes showed both hydroxylated myrcene, (+)- and (?)-α-pinene, 3-carene, and R-(+)-limonene, but not α-phellandrene, (?)-β-pinene, γ-terpinene, or terpinolene, with evidence that CYP9T2 strongly preferred myrcene over other substrates. They differed in the enantiomeric ratios of ipsdienol produced from myrcene, and in the products resulting from different α-pinene enantiomers. These data provide new information regarding bark beetle pheromone evolution and factors affecting cytochrome P450 structure–function relationships.     

A New Outbreak of the European Spruce Bark Beetle,Ips typographus (L.) (Coleoptera,Curculionidae) in Leningrad Province

Entomological Review - A new and potentially severe outbreak of the European spruce bark beetle Ips typographus began in Leningrad Province in 2021. The permanent presence of the species in...     

Successful reproduction and pheromone production by the spruce bark beetle in evolutionary naïve spruce hosts with familiar terpenoid defences

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Functional investigation of monoterpenes for improved understanding of the relationship between hosts and bark beetles

Spruce bark beetle (Ips typographus L.) is the most destructive insect pest of spruce forests in Eurasia. However, contact toxicity, in vivo metabolism, and ecological functions of host monoterpenes are poorly understood at the spruce tree–bark beetle–predator tritrophic level. Spruce monoterpenes including S-(–)-α-pinene, R-(+)-α-pinene, and myrcene showed contact toxicity to I. typographus, with LD₅₀ values ranging from 22–32 μg/mg. When topically treated with S-(–)-α-pinene or R-(+)-α-pinene, the amounts of volatile metabolites, including 4S-(–)-cis-verbenol, 4S-(+)-/4R-(–)-trans-verbenol, R-(+)-/S-(–)-verbenone and 1R-(–)-/1S-(+)-myrtenol, in the hindgut extracts of I. typographus varied significantly between sexes, and their quality (enantiomeric composition) varied significantly with the chirality of α-pinene. More importantly, S-(–)-α-pinene induced male adults to produce large amounts of 4S-(–)-cis-verbenol and S-(–)-verbenone. When topically treated with myrcene, the expected semiochemicals such as E-myrcenol, ipsenol and ipsdienol were not detected in the beetle hindguts, indicating that the pheromone biosynthetic system of I. typographus does not participate in the metabolism of host myrcene. In trap tests, S-(–)-α-pinene and R-(+)-α-pinene increased the catches of I. typographus and its predator Thanasimus substriatus in pheromone-baited traps, whereas myrcene exhibited a strong repellent (or inhibitory) effect on I. typographus but not on its predator. I. typographus seems to adopt different ecological strategies (e.g. avoidance to myrcene and preference for α-pinene) to adapt to and tolerate different host monoterpenes. Extensive investigation of these monoterpenes will help us understand their roles in manipulating the arms race between host trees and bark beetles, and potentially improve the efficacy of controlling I. typographus via the push-pull strategy using host kairomones.