Nadir Erbilgin
University of Alberta, Renewable Resources, Faculty Member
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Eriococcus spurius (Modeer) (Hemiptera: Eriociccidae) is a pest of American elm (Ulmus americana Linnaeus; Ulmaceae) trees in western North America, but no effective biological control agent has been reported. This study took place in... more
Eriococcus spurius (Modeer) (Hemiptera: Eriociccidae) is a pest of American elm (Ulmus americana Linnaeus; Ulmaceae) trees in western North America, but no effective biological control agent has been reported. This study took place in Calgary, Alberta, Canada, where urban American elms have been heavily and negatively impacted by E. spurius. Although the parasitoid Coccophagus gossypariae Gahan (Hymenoptera: Aphelinidae) was recovered from an E. spurius infested American elm in Calgary, little is known about its ecology. Prior to implementing biological control, the basic ecology of a candidate species must be understood. We documented the geographic establishment, sex ratios, and parasitism rate of C. gossyparaie on its host within the study area. Coccophagus gossypariae had established at 89% of study trees sampled and accounted for 98% of the parasitoids caught. They had a female-biased sex ratio of 13.7 females per male, and both males and females developed to maturity using the...
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ABSTRACT
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Plant pathogens can have cascading consequences on insect herbivores, though whether they alter competition among resource-sharing insect herbivores is unknown. We experimentally tested whether the infection of a plant pathogen, the... more
Plant pathogens can have cascading consequences on insect herbivores, though whether they alter competition among resource-sharing insect herbivores is unknown. We experimentally tested whether the infection of a plant pathogen, the parasitic plant dwarf mistletoe (Arceuthobium americanum), on jack pine (Pinus banksiana) altered the competitive interactions among two groups of beetles sharing the same resources: wood-boring beetles (Coleoptera: Cerambycidae) and the invasive mountain pine beetle (Dendroctonus ponderosae) (Coleoptera: Curculionidae). We were particularly interested in identifying potential mechanisms governing the direction of interactions (from competition to facilitation) between the two beetle groups. At the lowest and highest disease severity, wood-boring beetles increased their consumption rate relative to feeding levels at moderate severity. The performance (brood production and feeding) of mountain pine beetle was negatively associated with wood-boring beetle feeding and disease severity when they were reared separately. However, when both wood-boring beetles and high severity of plant pathogen infection occurred together, mountain pine beetle escaped from competition and improved its performance (increased brood production and feeding). Species-specific responses to changes in tree defense compounds and quality of resources (available phloem) were likely mechanisms driving this change of interactions between the two beetle groups. This is the first study demonstrating that a parasitic plant can be an important force in mediating competition among resource-sharing subcortical insect herbivores.
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Research Interests: Pest Management, Chemical Ecology, Biology, Pheromones, Ecology, and 15 moreIntegrated Pest Management, Great Lakes Region, Biological Sciences, Environmental Sciences, iPS, Solid Phase Microextraction, North America, Bark Beetle, PEST analysis, Predators, Population Monitoring, Pheromone, Pheromone Trap, Host Plant, and Adverse effect
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In North America, lodgepole pine is frequently subjected to attacks by various biotic disturbances that compromise its ability to defend against subsequent attacks by insect herbivores. We investigated whether infections of lodgepole pine... more
In North America, lodgepole pine is frequently subjected to attacks by various biotic disturbances that compromise its ability to defend against subsequent attacks by insect herbivores. We investigated whether infections of lodgepole pine by different pathogenic fungal species have varying effects on its defense chemistry. We selected two common pathogens: Atropellis canker and western gall rust, affecting mature lodgepole pine trees in western Canada. We also included three ophiostomoid fungi associated with the mountain pine beetle, Grosmannia clavigera, Ophiostoma montium, and Leptographium longiclavatum because symbiotic fungi are commonly used to investigate induced defenses of host trees of bark beetles. We collected phloem samples from lodgepole pine trees infected with the rust or the canker, and healthy lodgepole pine trees in the same stand. We also inoculated mature lodgepole pine trees with the three fungal symbionts and collected phloem two weeks later when the defense ...
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The environment could alter growth and resistance tradeoffs in plants by affecting the ratio of resource allocation to various competing traits. Yet, how and why functional tradeoffs change over time and space is poorly understood... more
The environment could alter growth and resistance tradeoffs in plants by affecting the ratio of resource allocation to various competing traits. Yet, how and why functional tradeoffs change over time and space is poorly understood particularly in long-lived conifer species. By establishing four common-garden test sites for five lodgepole pine populations in western Canada, combined with genomic sequencing, we revealed the decoupling pattern and genetic underpinnings of tradeoffs between height growth, drought resistance based on δ13C and dendrochronology, and metrics of pest resistance based on pest suitability ratings. Height and δ13C correlation displayed a gradient change in magnitude and/or direction along warm-to-cold test sites. All cold test sites across populations showed a positive height and δ13C relationship. However, we did not observe such a clinal correlation pattern between height or δ13C and pest suitability. Further, we found that the study populations exhibiting fu...
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While droughts, intensified by climate change, have been affecting forests worldwide, pest epidemics are a major source of uncertainty for assessing drought impacts on forest trees. Thus far, little information has documented the... more
While droughts, intensified by climate change, have been affecting forests worldwide, pest epidemics are a major source of uncertainty for assessing drought impacts on forest trees. Thus far, little information has documented the adaptability and evolvability of traits related to drought and pests simultaneously. We conducted common-garden experiments to investigate how several phenotypic traits (i.e. height growth, drought avoidance based on water-use efficiency inferred from δ 13 C and pest resistance based on defence traits) interact in five mature lodgepole pine populations established in four progeny trials in western Canada. The relevance of interpopulation variation in climate sensitivity highlighted that seed-source warm populations had greater adaptive capability than cold populations. In test sites, warming generated taller trees with higher δ 13 C and increased the evolutionary potential of height growth and δ 13 C across populations. We found, however, no pronounced grad...
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Background Genomic prediction (GP) and genome-wide association (GWA) analyses are currently being employed to accelerate breeding cycles and to identify alleles or genomic regions of complex traits in forest trees species. Here, 1490... more
Background Genomic prediction (GP) and genome-wide association (GWA) analyses are currently being employed to accelerate breeding cycles and to identify alleles or genomic regions of complex traits in forest trees species. Here, 1490 interior lodgepole pine (Pinus contorta Dougl. ex. Loud. var. latifolia Engelm) trees from four open-pollinated progeny trials were genotyped with 25,099 SNPs, and phenotyped for 15 growth, wood quality, pest resistance, drought tolerance, and defense chemical (monoterpenes) traits. The main objectives of this study were to: (1) identify genetic markers associated with these traits and determine their genetic architecture, and to compare the marker detected by single- (ST) and multiple-trait (MT) GWA models; (2) evaluate and compare the accuracy and control of bias of the genomic predictions for these traits underlying different ST and MT parametric and non-parametric GP methods. GWA, ST and MT analyses were compared using a linear transformation of gen...
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Linear regression of relationship between hyphal length and lodgepole pine killed by mountain pine beetle.
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Flight polyphenisms naturally occur as discrete or continuous traits in insects. Discrete flight polyphenisms include winged and wingless morphs, whereas continuous flight polyphenisms can take the form of short- or long-distance fliers.... more
Flight polyphenisms naturally occur as discrete or continuous traits in insects. Discrete flight polyphenisms include winged and wingless morphs, whereas continuous flight polyphenisms can take the form of short- or long-distance fliers. The mountain pine beetle (Dendroctonus ponderosae) exhibits polyphenic variation in flight distance but the consequences of this flight variation on life history strategies of beetles is unknown. This study assessed the effect of flight on two particular aspects of beetle biology: (1) an energetic trade-off between flight distance and host colonisation capacity; and (2) the relationship between flight distance and pheromone production. A 23 h flight treatment was applied to a subset of beetles using computer-linked flight mills. After flight treatment, both flown and unflown (control) beetles were given the opportunity to colonise bolts of host trees, and beetles that entered hosts were aerated to collect pheromone. A trade-off occurred between init...
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Presentation on developing and evaluating volatile organic compound (VOC) profiles for mature pine host trees and linking to beetle fitness.
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How carbohydrate reserves in conifers respond to drought and bark beetle attacks are poorly understood. We investigated changes in carbohydrate reserves and carbon‐dependent diterpene defences in ponderosa pine trees that were... more
How carbohydrate reserves in conifers respond to drought and bark beetle attacks are poorly understood. We investigated changes in carbohydrate reserves and carbon‐dependent diterpene defences in ponderosa pine trees that were experimentally subjected to two levels of drought stress (via root trenching) and two types of biotic challenge treatments (pheromone‐induced bark beetle attacks or inoculations with crushed beetles that include beetle‐associated fungi) for two consecutive years. Our results showed that trenching did not influence carbohydrates, whereas both biotic challenges reduced amounts of starch and sugars of trees. However, only the combined trenched‐bark beetle attacked trees depleted carbohydrates and died during the first year of attacks. While live trees contained higher carbohydrates than dying trees, amounts of constitutive and induced diterpenes produced did not vary between live and beetle‐attacked dying trees, respectively. Based on these results we propose tha...
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Phytopathogenic ophiostomatoid fungi are common associates of bark beetles and contribute to beetle-associated mortality of trees. Mountain pine beetle outbreaks in Canada are facilitating novel associations between its vectored fungi... more
Phytopathogenic ophiostomatoid fungi are common associates of bark beetles and contribute to beetle-associated mortality of trees. Mountain pine beetle outbreaks in Canada are facilitating novel associations between its vectored fungi (Grosmannia clavigera, Leptographium longiclavatum and Ophiostoma montium) and jack pine. How the induced defense-related metabolite responses of jack and lodgepole pines vary in response to the fungi is unknown. Understanding this variation is important to clarifying pine susceptibility to and the physiological impacts of infection. We used a comparative metabolite profiling approach to investigate the defense-related signaling, carbon utilization/mobilization, and synthesis responses of both pines to the fungi. Both pine species largely exhibited similar metabolite responses to the fungi. The magnitude of pine metabolite responses positively reflected pathogen virulence. Our findings indicate that pines can recognize and metabolomically respond to no...
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Studies with conifer-infesting bark beetles commonly use bolts cut from trees to evaluate the effects of host tree quality on various aspects of insect biology. Yet, whether host quality changes between live trees and bolts cut from these... more
Studies with conifer-infesting bark beetles commonly use bolts cut from trees to evaluate the effects of host tree quality on various aspects of insect biology. Yet, whether host quality changes between live trees and bolts cut from these trees has not been assessed. Particularly, changes in concentrations of defense chemicals (such as monoterpenes) and nutrients (such as nitrogen and carbon) have not been compared between live trees and their cut bolts. To determine whether monoterpene and nutrient concentrations differ after cutting, jack pine (Pinus banksiana Lamb.) trees in Lac La Biche (Alberta) were selected and sampled for phloem tissue. Then, these trees were harvested into two bolts per tree and stored at 4 °C for 3 and 6 months. Phloem was sampled from both live trees and bolts 3 and 6 months after storage. We found that major monoterpenes of jack pine were higher in phloem from bolts than from live trees. Storage time did not affect the results. Furthermore, some nutrient...
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Warming climate is allowing tree-killing bark beetles to expand their ranges and access naïve and semi-naïve conifers. Conifers respond to attack using complex mixtures of chemical defenses that can impede beetle success, but beetles... more
Warming climate is allowing tree-killing bark beetles to expand their ranges and access naïve and semi-naïve conifers. Conifers respond to attack using complex mixtures of chemical defenses that can impede beetle success, but beetles exploit some compounds for host location and communication. Outcomes of changing relationships will depend on concentrations and compositions of multiple host compounds, which are largely unknown. We analyzed constitutive and induced chemistries of Dendroctonus ponderosae's primary historical host, Pinus contorta, and P. albicaulis, a high-elevation species whose encounters with this beetle are transitioning from intermittent to continuous. We quantified multiple classes of terpenes, phenolics, carbohydrates, and minerals. Pinus contorta had higher constitutive allocation to, and generally stronger inducibility of, compounds that resist these beetle-fungal complexes. Pinus albicaulis contained higher proportions of specific monoterpenes that enhance...
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Research Interests: Chemistry, Forestry, Forest Ecology And Management, Biology, Forest Ecology, and 15 moreBiological Sciences, Concentration, Environmental Sciences, Factorial Design, Efficiency, Fertilization, Soil management, Allocation, Stand Development, Indexation, Mountain Pine Beetle, Carbon Sink, Fertilisation, Pinus Contorta, and Basal Area
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Waring and Pitman suggested that high growth efficiency (GE) (i.e. basal area increment/sapwood area - considered a measure of tree vigor) is related to the ability of lodgepole pine to defend against mountain pine beetles (Dendroctonus... more
Waring and Pitman suggested that high growth efficiency (GE) (i.e. basal area increment/sapwood area - considered a measure of tree vigor) is related to the ability of lodgepole pine to defend against mountain pine beetles (Dendroctonus ponderosae Hopkins) (MPB). In this study we explore the relationship between GE, and other measures of vigor such as tree size, live crown ratio, growth increment, stem, foliage nutrients and root carbohydrate reserves. We then compare these measures of vigor with characteristics of tree defense such as density of resin ducts, monoterpenes and size of phloem lesions after inoculation with Grossmania clavigera (a blue stain fungus associated with MPB). In a field experiment we tested the suggested relationship between vigor and defense by thinning and fertilization in ten pure lodgepole pine stands in central Alberta. In the third summer after the fertilization and thinning treatment we inoculated trees with the blue stain fungus. The combination of fertilization and thinning increased GE and resin duct density but negatively impacted starch reserves. Lesion size increased with tree size in control plots, however, this relationship was lost when trees were fertilized and thinned. This suggests that treatments to increase growth rate also increased constitutive defenses in larger trees
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Operational forest monitoring often requires fine-detail information in the form of an orthomosaic, created by stitching overlapping nadir images captured by aerial platforms such as drones. RGB drone sensors are commonly used for... more
Operational forest monitoring often requires fine-detail information in the form of an orthomosaic, created by stitching overlapping nadir images captured by aerial platforms such as drones. RGB drone sensors are commonly used for low-cost, high-resolution imaging that is conducive to effective orthomosaicking, but only capture visible light. Thermal sensors, on the other hand, capture long-wave infrared radiation, which is useful for early pest detection among other applications. However, these lower-resolution images suffer from reduced contrast and lack of descriptive features for successful orthomosaicking, leading to gaps or swirling artifacts in the orthomosaic. To tackle this, we propose a thermal orthomosaicking workflow that leverages simultaneously acquired RGB images. The latter are used for producing a surface mesh via structure from motion, while thermal images are only used to texture this mesh and yield a thermal orthomosaic. Prior to texturing, RGB-thermal image pair...
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Interactions between mutualistic bark beetles and ophiostomatoid fungi have received considerable attention in recent years. Studies have shown how volatile organic compounds emitted from mutualist fungi affect the behaviors of several... more
Interactions between mutualistic bark beetles and ophiostomatoid fungi have received considerable attention in recent years. Studies have shown how volatile organic compounds emitted from mutualist fungi affect the behaviors of several bark beetle species. However, we currently lack sufficient knowledge regarding whether bark beetle pheromones can influence mutualist fungi. Here, we measured growth and biomass of two mutualistic fungi of the mountain pine beetle in response to headspace of a beetle pheromone ( trans -verbenol), a blend of host tree volatiles, the combination of both, or control (no volatile source) in vitro experiments consisting of a nitrogen-based medium. The surface area and ergosterol content of the mycelia were used as surrogates for fungal growth and biomass respectively. We found that both growth and biomass of Grosmannia clavigera and Ophiostoma montium were greater in medium exposed to any type of volatile sources than the control. While growth and ergosterol content of G. clavigera were highest in the combination treatment, there were no differences in growth or biomass among the types of volatiles introduced for O. montium . These results suggest that both mutualistic fungi can utilize both bark beetle pheromone and host tree volatiles as nutrient sources. Overall, these results support the on-going studies on the role of volatile organic compounds mediating mutualistic bark beetle-fungi interactions.
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Seedling performance data Project: Inoculation of seedlings alters fungal communities but does not improve their performance in beetle-killed sites Created by: Josh Wasyliw Date: 2020-11-20 ID_site Site identifier; each site contains 2... more
Seedling performance data Project: Inoculation of seedlings alters fungal communities but does not improve their performance in beetle-killed sites Created by: Josh Wasyliw Date: 2020-11-20 ID_site Site identifier; each site contains 2 subplots; each subplot contains 4 quads Quad 1-4; three of quads were planted with seedlings receiving an inoculum treatment, and there was an extra quad that was seeded but no seedlings collected Subplot 2 subplots, A and B, each split into four quads Direction position of the quad Treatment One of three: Control, soil inoculum from non-attacked or attacked stands Soil_type From Evan Fellrath's survey, coded as silty or sandy Seedlings_alive_2018 Number of surviving seedlings in 2018 Seedlings_alive_2019 Number of surviving seedlings in 2019 Seedlings_planted Number of seedlings planted in 2017 Survival Seedlings alive 2019/ Seedlings planted Fresh_mass g, total biomass of pooled seedlings per quad Dry_mass g, total biomass of pooled seedlings pe...
<b>Abstract</b><br/>The current unprecedented outbreak of mountain pine beetle (Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests of western Canada has resulted in a landscape consisting of a mosaic of... more
<b>Abstract</b><br/>The current unprecedented outbreak of mountain pine beetle (Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests of western Canada has resulted in a landscape consisting of a mosaic of forest stands at different stages of mortality. Within forest stands, understory communities are the reservoir of the majority of plant species diversity and influence the composition of future forests in response to disturbance. Although changes to stand composition following beetle outbreaks are well documented, information on immediate responses of forest understory plant communities is limited. The objective of this study was to examine the effects of D. ponderosae-induced tree mortality on initial changes in diversity and productivity of understory plant communities. We established a total of 110 1-m2 plots across eleven mature lodgepole pine forests to measure changes in understory diversity and productivity as a function of tree mortality and below ground resource availability across multiple years. Overall, understory community diversity and productivity increased across the gradient of increased tree mortality. Richness of herbaceous perennials increased with tree mortality as well as soil moisture and nutrient levels. In contrast, the diversity of woody perennials did not change across the gradient of tree mortality. Understory vegetation, namely herbaceous perennials, showed an immediate response to improved growing conditions caused by increases in tree mortality. How this increased pulse in understory richness and productivity affects future forest trajectories in a novel system is unknown.
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Intra-specific variation in conifers has been extensively studied with respect to defense against herbivores and pathogens. While studies have shown the ability of individual or specific mixtures of compounds to influence insects and... more
Intra-specific variation in conifers has been extensively studied with respect to defense against herbivores and pathogens. While studies have shown the ability of individual or specific mixtures of compounds to influence insects and microbes, research testing biologically relevant mixtures of defense compounds reflecting intra-specific variation amongst tree populations to enemy complexes is needed. We characterized the variations in lodgepole pine monoterpenes from a progeny trial in western Canada and grouped trees in four clusters using their monoterpene profiles. We then selected 11 representative families across four clusters and amended their entire monoterpene profiles (with the exception of β-phellandrene) in media to determine how representative families affect the performance of the mountain pine beetle or its fungal symbiont. We placed adult beetles or inoculated fungus on the amended media and measured beetle performance and fungal growth as a proxy to host suitability. We found that different clusters or families differentially influenced beetle or fungal responses. However, monoterpene profiles of trees suitable to the beetle or the fungus were dissimilar. These outcomes reflect a co-evolutionary arms-race between the host and the bark beetle-fungus complex, which has resulted in the production of complementary defense metabolites among different pine populations to enhance tree survival. This article is protected by copyright. All rights reserved.
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The mountain pine beetle, Dendroctonus ponderosae, has infested over ~16 Mha of pine forests in British Columbia killing >50% of mature lodgepole pine, Pinus contorta, trees in affected stands. At present, it is functionally an... more
The mountain pine beetle, Dendroctonus ponderosae, has infested over ~16 Mha of pine forests in British Columbia killing >50% of mature lodgepole pine, Pinus contorta, trees in affected stands. At present, it is functionally an invasive species in Alberta, killing and reproducing in evolutionarily naïve populations of lodgepole pine (P. contorta), novel jack pine (P. banksiana), and their hybrids. The entomopathogenic fungus Beauveria bassiana has shown some potential as a biocontrol agent of several bark beetle species. In this study, nine isolates of B. bassiana were examined for insect virulence characteristics, including conidiation rate, pigmentation, and infection rate in laboratory-reared D. ponderosae, to assess for their potential as biocontrol agents. The strains were categorized into three phenotypic groups based on pigmentation, conidial density, and myceliation rate. Virulence screening utilizing insect-based agar medium (D. ponderosae and European honeybee Apis mellifera carcasses) revealed no difference in selection of fungal growth. However, infection studies on D. ponderosae and A. mellifera showed contrasting results. In vivo A. mellifera infection model revealed ~5% mortality, representing the natural death rate of the hive population, whereas laboratory-reared D. ponderosae showed 100% mortality and mycosis. The LT50 (median lethal time 50) ranges from 2 to 5 ± 0.33 days, and LT100 ranges from 4 to 6 ± 0.5 days. We discuss the selective advantages of the three phenotypic groups in terms of virulence, pigmentation, conidial abundance, and tolerance to abiotic factors like UV and host tree monoterpenes. These results can further provide insights into the development of several phenotypically diverse B. bassiana strains in controlling the spread of the invasive D. ponderosae in Western Canada. KEY POINTS: • Three B. bassiana morphotype groups have been demonstrated to kill D. ponderosae. • A range of effective lethal times (LT50 and LT100) was established against D. ponderosae. • Variable tolerance to UV light and pine monoterpenes were observed in B. bassiana.
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Global change has altered nitrogen availability in boreal forest soils. As ectomycorrhizal fungi play critical ecological functions, shifts in their abundance and community composition must be considered in the response of forests to... more
Global change has altered nitrogen availability in boreal forest soils. As ectomycorrhizal fungi play critical ecological functions, shifts in their abundance and community composition must be considered in the response of forests to changes in nitrogen availability. Furthermore, ectomycorrhizas are symbiotic, so the response of ectomycorrhizal fungi to nitrogen cannot be understood in isolation of their plant partners. Most previous studies, however, neglect to measure the response of host trees to nitrogen addition simultaneously with that of fungal communities. In addition to being one‐sided, most of these studies have also been conducted in coniferous forests. Deciduous and “dual‐mycorrhizal” tree species, namely those that form ecto‐ and arbuscular mycorrhizas, have received little attention despite being widespread in the boreal forest. We applied nitrogen (30 kg ha−1 year−1) for 13 years to stands dominated by aspen (Populus tremuloides Michx.) and hypothesized that tree stem radial growth would increase, ectomycorrhizal fungal biomass would decrease, ectomycorrhizal fungal community composition would shift, and the abundance of arbuscular mycorrhizal (AM) fungi would increase. Nitrogen addition initially increased stem radial growth of aspen, but it was not sustained at the time we characterized their mycorrhizas. After 13 years, the abundance of fungi possessing extramatrical hyphae, or “high‐biomass” ectomycorrhizas, doubled. No changes occurred in ectomycorrhizal and AM fungal community composition, or in ecto‐ and AM abundance measured as root colonization. This dual‐mycorrhizal tree species did not shift away from ectomycorrhizal fungal dominance with long‐term nitrogen input. The unexpected increase in high‐biomass ectomycorrhizal fungi with nitrogen addition may be due to increased carbon allocation to their fungal partners by growth‐limited trees. Given the focus on conifers in past studies, reconciling results of plant–mycorrhizal fungal relationships in stands of deciduous trees may demand a broader view on the impacts of nitrogen addition on the structure and function of boreal forests.
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Climate change has a large influence on plant functional and phenotypic traits including plant primary and secondary metabolites. One well-established approach to investigating the variation in plant metabolites involves studying plant... more
Climate change has a large influence on plant functional and phenotypic traits including plant primary and secondary metabolites. One well-established approach to investigating the variation in plant metabolites involves studying plant populations along elevation and latitude gradients. We considered how two space-for-time climate change gradients (elevation and latitude) influence carbohydrate reserves (soluble sugars, starches) and secondary metabolites (monoterpenes, diterpene resin acids) of lodgepole pine trees in western Canada. We were particularly interested in the relationship of terpenes and carbohydrates with a wide range of tree, site, and climatic factors. We found that only elevation had a strong influence on the expression of both terpenes and carbohydrates of trees. Specifically, as elevation increased, concentrations of monoterpenes and diterpenes generally increased and soluble sugars (glucose, sucrose, total sugars) decreased. In contrast, latitude had no impact on either of terpenes or carbohydrates. Furthermore, we found a positive relationship between concentrations of starch and total terpenes and diterpenes in the elevation study; whereas neither starches nor sugars were correlated to terpenes in the latitude study. Similarly, both terpenes and carbohydrates had a much greater number of significant correlations to site characteristics such as slope, basal area index, and sand basal area, in the elevational than in the latitude study. Overall, these results support the conclusion that both biotic and abiotic factors likely drive the patterns of primary and secondary metabolite profiles of lodgepole pine along geographical gradients. Also, presence of a positive relationship between terpenes and starches suggests an interaction between primary ad secondary metabolites of lodgepole pine trees.