| 1. |
- Axelsson, Karolin, et al.
(author)
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Chemical defence responses of Norway spruce to two fungal pathogens
- 2020
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In: Forest Pathology. - : Blackwell Verlag. - 1437-4781 .- 1439-0329. ; 50:6
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Journal article (peer-reviewed)abstract
- Constitutive and inducible terpene production is involved in conifer resistance against insects and fungal infestations. To gain knowledge about local defence responses of Norway spruce bark against pathogens and to find potential chemical markers for resistance breeding, we inoculated the stem of 8-year-old Norway spruce (Picea abies) clonal trees with bothEndoconidiophora polonica(Ep, a common fungal pathogen associated with the spruce bark beetleIps typographus) andHeterobasidion parviporum(Hp, a severe pathogen causing root and stem rot disease). Three weeks after inoculation, the fungal-inoculated and intact bark from each tree was sampled. The terpenes in tree bark were extracted with hexane and characterized by gas chromatography-mass spectrometry (GC-MS). The two fungi induced varied terpene responses in the four spruce clones used. Three of the clones showed a 2.3-fold to 5.7-fold stronger terpene response to Hp relative to Ep inoculation, while one clone responded similarly to inoculation with the two fungal pathogens. The amount of the diterpenes thunbergol and geranyllinalool varied between the clones. The level of thunbergol was higher in both intact and fungal-inoculated bark from the less susceptible clones compared with the more susceptible clones. Geranyllinalool was present in higher amounts in the susceptible clones and is thus a possible marker for susceptibility. Our observations show that Norway spruce employs a similar chemical mechanism against the two fungal pathogens. Based on the present and earlier published data, we suggest that certain Norway spruce genotypes have a strong defence reaction against these two pathogens. The diterpenes thunbergol and geranyllinalool might be useful markers of susceptibility in tree-breeding programmes and should be the focus of further detailed investigations.
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| 2. |
- Danielsson, Marie, et al.
(author)
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Arthropod infestation sites and induced defence can be traced by emission from single spruce needle
- 2019
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In: Arthropod-Plant Interactions. - : Springer. - 1872-8855 .- 1872-8847. ; 13:2, s. 253-259
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Journal article (peer-reviewed)abstract
- Emissions of defence chemicals from Norway spruce seedlings can be induced by feeding arthropods or by exogenous hormonal application. Some defence chemicals may attract or repel associated arthropods. The aim of this study was to show that it is possible to detect and collect stress-induced volatiles from micro sites, such as at the scale of a single needle, in vivo by using SPME. Methyl jasmonate application on the stem of Norway spruce seedlings induced emission of (E)-beta-farnesene only from the needles closest to the application site. Emissions of (E)-beta-farnesene, (E,E)-alpha-farnesene and (E)-alpha-bisabolene were only detected from needles infested by the spider mite Oligonychus ununguis. The total volatile amount detected by SPME-GC-MS reached a considerable mass of 14 ng/needle/24 h, suggesting that emission from damaged and stressed conifers might have a larger impact on the macro climate than previously estimated.
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| 3. |
- Kandasamy, Dineshkumar, et al.
(author)
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Conifer-killing bark beetles locate fungal symbionts by detecting volatile fungal metabolites of host tree resin monoterpenes
- 2023
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In: PLoS biology. - : Public Library of Science (PLoS). - 1544-9173 .- 1545-7885. ; 21:2
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Journal article (peer-reviewed)abstract
- Outbreaks of the Eurasian spruce bark beetle (Ips typographus) have decimated millions of hectares of conifer forests in Europe in recent years. The ability of these 4.0 to 5.5 mm long insects to kill mature trees over a short period has been sometimes ascribed to two main factors: (1) mass attacks on the host tree to overcome tree defenses and (2) the presence of fungal symbionts that support successful beetle development in the tree. While the role of pheromones in coordinating mass attacks has been well studied, the role of chemical communication in maintaining the fungal symbiosis is poorly understood. Previous evidence indicates that I. typographus can distinguish fungal symbionts of the genera Grosmannia, Endoconidiophora, and Ophiostoma by their de novo synthesized volatile compounds. Here, we hypothesize that the fungal symbionts of this bark beetle species metabolize spruce resin monoterpenes of the beetle's host tree, Norway spruce (Picea abies), and that the volatile products are used as cues by beetles for locating breeding sites with beneficial symbionts. We show that Grosmannia penicillata and other fungal symbionts alter the profile of spruce bark volatiles by converting the major monoterpenes into an attractive blend of oxygenated derivatives. Bornyl acetate was metabolized to camphor, and α- and β-pinene to trans-4-thujanol and other oxygenated products. Electrophysiological measurements showed that I. typographus possesses dedicated olfactory sensory neurons for oxygenated metabolites. Both camphor and trans-4-thujanol attracted beetles at specific doses in walking olfactometer experiments, and the presence of symbiotic fungi enhanced attraction of females to pheromones. Another co-occurring nonbeneficial fungus (Trichoderma sp.) also produced oxygenated monoterpenes, but these were not attractive to I. typographus. Finally, we show that colonization of fungal symbionts on spruce bark diet stimulated beetles to make tunnels into the diet. Collectively, our study suggests that the blends of oxygenated metabolites of conifer monoterpenes produced by fungal symbionts are used by walking bark beetles as attractive or repellent cues to locate breeding or feeding sites containing beneficial microbial symbionts. The oxygenated metabolites may aid beetles in assessing the presence of the fungus, the defense status of the host tree and the density of conspecifics at potential feeding and breeding sites.
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| 4. |
- Kännaste, Astrid, et al.
(author)
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Odors of Norway spruce (Picea abies L.) seedlings : differences due to age and chemotype
- 2013
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In: Trees. - : Springer. - 0931-1890 .- 1432-2285. ; 27, s. 149-159
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Journal article (peer-reviewed)abstract
- Small conifer seedlings (mini-seedlings) are less damaged by the large pine weevil Hylobius abietis (L.) (Coleoptera: Curculionidae) compared to conventional seedlings. Chemical difference between the seedling types is one possible explanation for this phenomenon. In the present paper, the emissions of volatile organic compounds (VOC) of 7- to 43-week-old Norway spruce [Picea abies (L.) Karst.] seedlings were analyzed. Collection and identification of the volatiles was made by solid phase micro-extraction and gas chromatography mass spectrometry (SPME–GC–MS). The enantiomers of α-pinene and limonene were separated in a two-dimensional GC (2D-GC). Most of the seedlings represented either a limonene- or a bornyl acetate-chemotype. Only minor changes in the volatile composition of the two types of seedlings were found during the first year. Age-related changes, however, were found in the volatiles released by wounded phloem where green leaf volatiles (GLVs) and borneol were the dominated VOC for young seedling. The attractive compound for the pine weevil, α-pinene, was first detected in the phloem emissions of 18- to 22-week-old seedlings. Different storage conditions of the seedlings during the winter/early spring-phase influenced the volatile composition in the phloem. High amount of GLVs was characteristic for the 43-week-old seedlings stored in naturally changing outdoor temperature, but not present in the seedlings winter-stored at a constant temperature of −4 °C. The possible role of these observed differences in odor emissions between seedlings of different age and physiological status for the feeding preferences of the large pine weevil is discussed.
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| 5. |
- Lu, Jun, et al.
(author)
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The shoot-feeding ecology of three Tomicus species in Yunnan Province, southwestern China
- 2014
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In: Journal of Insect Science. - : Oxford University Press (OUP). - 1536-2442. ; 14:37
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Journal article (peer-reviewed)abstract
- Three Tomicus pine shoot beetles, T. yunnanensis (Kirkendall and Faccoli) (Coleoptera: Scolytidae), T. minor (Hartig), and T. brevipilosus (Wood and Bright), have been causing serious damage to Yunnan pine (Pinus yunnanensis (Franchet) (Pinales: Pinaceae)) stands in Yunnan, southwestern China. However, their ability to coexist in the crowns of the same trees during the shoot-feeding phase has not been elucidated. In our study, we investigated and compared the shoot-feeding ecology of the three species of pine shoot beetle in P. yunnanensis in Anning County, Yunnan Province. Shoot-feeding by T. yunnanensis, T. minor, and T. brevipilosus initiated in April, May, and June, and ended in February, April, and May, respectively. Individual T. yunnanensis and T. minor adults fed in shoots for about seven months, and T. brevipilosus for nine months, before initiating reproduction. All three Tomicus species fed in the current-year shoots close to the apical bud. No specific overwintering behavior was observed prior to reproduction. The entrance hole of T. yunnanensis was furthest away from the apical bud, and T. minor was the closest to the apical bud. Differences in the spatial distribution of these shoot-feeding sites might reduce competition among the three beetle species. The long-lasting and overlapping shoot-feeding by the three Tomicus species may reduce the resistance of P. yunnanensis and facilitate the reproduction of these beetles in the trunks of living trees, and thus help explain the severe damage by Tomicus in P. yunnanensis.
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| 6. |
- Mageroy, Melissa H., et al.
(author)
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Molecular underpinnings of methyl jasmonate-induced resistance in Norway spruce
- 2020
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In: Plant, Cell and Environment. - : Blackwell Publishing. - 0140-7791 .- 1365-3040. ; 43:8, s. 1827-1843
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Journal article (peer-reviewed)abstract
- In response to various stimuli, plants acquire resistance against pests and/or pathogens. Such acquired or induced resistance allows plants to rapidly adapt to their environment. Spraying the bark of mature Norway spruce (Picea abies) trees with the phytohormone methyl jasmonate (MeJA) enhances resistance to tree-killing bark beetles and their associated phytopathogenic fungi. Analysis of spruce chemical defenses and beetle colonization success suggests that MeJA treatment both directly induces immune responses and primes inducible defenses for a faster and stronger response to subsequent beetle attack. We used metabolite and transcriptome profiling to explore the mechanisms underlying MeJA-induced resistance in Norway spruce. We demonstrated that MeJA treatment caused substantial changes in the bark transcriptional response to a triggering stress (mechanical wounding). Profiling of mRNA expression showed a suite of spruce inducible defenses are primed following MeJA treatment. Although monoterpenes and diterpene resin acids increased more rapidly after wounding in MeJA-treated than control bark, expression of their biosynthesis genes did not. We suggest that priming of inducible defenses is part of a complex mixture of defense responses that underpins the increased resistance against bark beetle colonization observed in Norway spruce. This study provides the most detailed insights yet into the mechanisms underlying induced resistance in a long-lived gymnosperm.
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| 7. |
- Mageroy, Melissa H., et al.
(author)
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Priming of inducible defenses protects Norway spruce against tree‐killing bark beetles
- 2020
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In: Plant, Cell and Environment. - : John Wiley & Sons. - 0140-7791 .- 1365-3040. ; 43:2, s. 420-430
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Journal article (peer-reviewed)abstract
- Plants can form an immunological memory known as defense priming, whereby exposure to a priming stimulus enables quicker or stronger response to subsequent attack by pests and pathogens. Such priming of inducible defenses provides increased protection and reduces allocation costs of defense. Defense priming has been widely studied for short–lived model plants such as Arabidopsis, but little is known about this phenomenon in long‐lived plants like spruce. We compared the effects of pre‐treatment with sub‐lethal fungal inoculations or application of the phytohormone methyl jasmonate (MeJA) on the resistance of 48‐year‐old Norway spruce (Picea abies) trees to mass attack by a tree‐killing bark beetle beginning 35 days later. Bark beetles heavily infested and killed untreated trees, but largely avoided fungus‐inoculated trees and MeJA‐treated trees. Quantification of defensive terpenes at the time of bark beetle attack showed fungal inoculation induced 91‐fold higher terpene concentrations compared to untreated trees, while application of MeJA did not significantly increase terpenes. These results indicate that resistance in fungus‐inoculated trees is a result of direct induction of defenses while resistance in MeJA‐treated trees is due to defense priming. This work extends our knowledge of defense priming from model plants to an ecologically important tree species.
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| 8. |
- Pan, Yue, et al.
(author)
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Bark Beetle-Associated Blue-Stain Fungi Increase Antioxidant Enzyme Activities and Monoterpene Concentrations in Pinus yunnanensis
- 2018
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In: Frontiers in Plant Science. - : Frontiers Media S.A.. - 1664-462X. ; 9
-
Journal article (peer-reviewed)abstract
- Yunnan pine is the most important tree species in SW China in both economical and ecological terms, but it is often killed by pine shoot beetles (Tomicus spp.). Tomicus beetles are secondary pests in temperate regions and the aggressiveness of the beetles in SW China is considered to be due to the warm subtropical climates as well as the beetles' virulent fungal associates. Here, we assessed the virulence of three blue-stain fungi (Leptographium wushanense, L. sinense and Ophiostoma canum) associated with pine shoot beetles to Yunnan pine (Pinus yunnanensis) in SW China. Following fungal inoculation, we measured necrotic lesion lengths, antioxidant enzyme activities and monoterpene concentrations in the stem phloem of Yunnan pine. Leptographium wushanense induced twice as long lesions as L. sinense and O. canum, and all three fungi induced significantly longer lesions than sterile agar control inoculations. The activity of three tested antioxidant enzymes (peroxidase, polyphenol oxidase, and superoxide dismutase) increased after both fungal inoculation and control inoculation. However, L. wushanense and L. sinense generally caused a greater increase in enzyme activities than O. canum and the control treatment. Fungal inoculation induced stronger increases in six major monoterpenes than the control treatment, but the difference was significant only for some fungus-monoterpene combinations. Overall, our results show that L. wushanense and L. sinense elicit stronger defense responses and thus are more virulent to Yunnan pine than O. canum. The two Leptographium species may thus contribute to the aggressiveness of their beetle vectors and could damage Yunnan pine across SW China if they spread from the restricted geographical area they have been found in so far.
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| 9. |
- Pan, Yue, et al.
(author)
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Ophiostomatales (Ascomycota) associated with Tomicus species in southwestern China with an emphasis on Ophiostoma canum
- 2020
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In: Journal of Forestry Research. - : Springer. - 1007-662X .- 1993-0607. ; 31:6, s. 2549-2562
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Journal article (peer-reviewed)abstract
- Ophiostomatalean fungi may facilitate bark beetle colonization and reproduction. In the present study of the fungal community associated with bark beetle spe-cies belonging to Tomicus in Yunnan, China, six ophios-tomatalean fungi (Ophiostoma canum, O. ips, O. tingens, Leptographium yunnanense, Leptographium sp. 1 and Lep-tographium sp. 2) were isolated from the beetles or their galleries; O. canum was the most common fungal species. The distribution of O. canum was associated with stands heavily damaged by Tomicus species and a higher percentage of valid galleries of Tomicus yunnanensis and T. minor in Yunnan pine (Pinus yunnanensis). After inoculation of Yun-nan pine with the fungus, a phloem reaction zone formed and monoterpenes accumulated in the phloem. These results suggested that O. canum was pathogenic to Yunnan pine and that the wide distribution of the fungus might be beneficial to reproduction of pine shoot beetles in Yunnan pine. How-ever, because the reaction zone and monoterpene accumu-lation were mild, fungal damage of Yunnan pine might be limited. A more integrated study considering all the fungal species should be done to better understand the interactions among bark beetles, blue-stain fungi, and the tree hosts in the region.
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| 10. |
- Puentes, Adriana, et al.
(author)
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Variation in Methyl Jasmonate-Induced Defense Among Norway Spruce Clones and Trade-Offs in Resistance Against a Fungal and an Insect Pest
- 2021
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In: Frontiers in Plant Science. - : Frontiers Media SA. - 1664-462X. ; 12
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Journal article (peer-reviewed)abstract
- An essential component of plant defense is the change that occurs from a constitutive to an induced state following damage or infection. Exogenous application of the plant hormone methyl jasmonate (MeJA) has shown great potential to be used as a defense inducer prior to pest exposure, and could be used as a plant protection measure. Here, we examined (1) the importance of MeJA-mediated induction for Norway spruce (Picea abies) resistance against damage by the pine weevil Hylobius abietis, which poses a threat to seedling survival, and infection by the spruce bark beetle-associated blue-stain fungus Endoconidiophora polonica, (2) genotypic variation in MeJA-induced defense (terpene chemistry), and (3) correlations among resistance to each pest. In a semi-field experiment, we exposed rooted-cuttings from nine different Norway spruce clones to insect damage and fungal infection separately. Plants were treated with 0, 25, or 50 mM MeJA, and planted in blocks where only pine weevils were released, or in a separate block in which plants were fungus-inoculated or not (control group). As measures of resistance, stem area debarked and fungal lesion lengths were assessed, and as a measure of defensive capacity, terpene chemistry was examined. We found that MeJA treatment increased resistance to H. abietis and E. polonica, but effects varied with clone. Norway spruce clones that exhibited high constitutive resistance did not show large changes in area debarked or lesion length when MeJA-treated, and vice versa. Moreover, insect damage negatively correlated with fungal infection. Clones receiving little pine weevil damage experienced larger lesion lengths, and vice versa, both in the constitutive and induced states. Changes in absolute terpene concentrations occurred with MeJA treatment (but not on proportional terpene concentrations), however, variation in chemistry was mostly explained by differences between clones. We conclude that MeJA can enhance protection against H. abietis and E. polonica, but the extent of protection will depend on the importance of constitutive and induced resistance for the Norway spruce clone in question. Trade-offs among resistances do not necessarily hinder the use of MeJA, as clones that are constitutively more resistant to either pest, should show greater MeJA-induced resistance against the other.
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| 11. |
- Zhao, Tao, 1969-, et al.
(author)
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Convergent evolution of semiochemicals across Kingdoms : bark beetles and their fungal symbionts
- 2019
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In: The ISME Journal. - : Nature Publishing Group. - 1751-7362 .- 1751-7370. ; 13:6, s. 1535-1545
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Journal article (peer-reviewed)abstract
- Convergent evolution of semiochemical use in organisms from different Kingdoms is a rarely described phenomenon. Tree-killing bark beetles vector numerous symbiotic blue-stain fungi that help the beetles colonize healthy trees. Here we show for the first time that some of these fungi are able to biosynthesize bicyclic ketals that are pheromones and other semiochemicals of bark beetles. Volatile emissions of five common bark beetle symbionts were investigated by gas chromatography-mass spectrometry. When grown on fresh Norway spruce bark the fungi emitted three well-known bark beetle aggregation pheromones and semiochemicals (exo-brevicomin, endo-brevicomin and trans-conophthorin) and two structurally related semiochemical candidates (exo-1,3-dimethyl-2,9-dioxabicyclo[3.3.1]nonane and endo-1,3-dimethyl-2,9-dioxabicyclo[3.3.1]nonane) that elicited electroantennogram responses in the spruce bark beetle Ips typographus. When grown on malt agar with C-13 D-Glucose, the fungus Grosmannia europhioides incorporated C-13 into exo-brevicomin and trans-conophthorin. The enantiomeric compositions of the fungus-produced ketals closely matched those previously reported from bark beetles. The production of structurally complex bark beetle pheromones by symbiotic fungi indicates cross-kingdom convergent evolution of signal use in this system. This signaling is susceptible to disruption, providing potential new targets for pest control in conifer forests and plantations.
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| 12. |
- Zhao, Tao, 1969-, et al.
(author)
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Fungal associates of the tree-killing bark beetle, Ips typographus, vary in virulence, ability to degrade conifer phenolics and influence bark beetle tunning behavior
- 2019
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In: Fungal ecology. - Amsteradam : Elsevier. - 1754-5048 .- 1878-0083. ; 38, s. 71-79
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Journal article (peer-reviewed)abstract
- The bark beetle Ips typographus carries numerous fungi that could be assisting the beetle in colonizing live Norway spruce (Picea abies) trees. Phenolic defenses in spruce phloem are degraded by the beetle's major tree-killing fungus Endoconidiophora polonica, but it is unknown if other beetle associates can also catabolize these compounds. We compared the ability of five fungi commonly associated with I. typographus to degrade phenolic compounds in Norway spruce phloem. Grosmannia penicillata and Grosmannia europhioides were able to degrade stilbenes and flavonoids faster than E. polonica and grow on minimal growth medium with spruce bark constituents as the only nutrients. Furthermore, beetles avoided medium amended with phenolics but marginally preferred medium colonized by fungi. Taken together our results show that different bark beetle-associated fungi have complementary roles in degrading host metabolites and thus might improve this insect's persistence in well defended host tissues.
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| 13. |
- Zhao, Tao, 1969-, et al.
(author)
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Fungal Symbionts of the Spruce Bark Beetle Synthesize the Beetle Aggregation Pheromone 2-Methyl-3-buten-2-ol
- 2015
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In: Journal of Chemical Ecology. - : Springer Science+Business Media B.V.. - 0098-0331 .- 1573-1561. ; 41, s. 848-852
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Journal article (peer-reviewed)abstract
- Tree-killing bark beetles depend on aggregation pheromones to mass-attack their host trees and overwhelm their resistance. The beetles are always associated with phytopathogenic ophiostomatoid fungi that probably assist in breaking down tree resistance, but little is known about if or how much these fungal symbionts contribute to the beetles’ aggregation behavior. In this study, we determined the ability of four major fungal symbionts of the spruce bark beetle Ips typographus to produce beetle aggregation pheromones. The fungi were incubated on Norway spruce Picea abies bark, malt agar, or malt agar amended with 0.5 % 13C glucose. Volatiles present in the headspace of each fungus were analyzed for 7 days after incubation using a SPME autosampler coupled to a GC/MS. Two Grosmannia species (G. penicillata and G. europhioides) produced large amounts of 2-methyl-3-buten-2-ol (MB), the major component in the beetles’ aggregation pheromone blend, when growing on spruce bark or malt agar. Grosmannia europhioides also incorporated 13C glucose into MB, demonstrating that the fungi can synthesize MB de novo using glucose as a carbon source. This is the first clear evidence that fungal symbionts of bark beetles can produce components in the aggregation pheromone blend of their beetle vectors. This provides new insight into the possible ecological roles of fungal symbionts in bark beetle systems and may deepen our understanding of species interactions and coevolution in these important biological systems.
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