Levels Of Phenolic Glycosides Research Articles

Polyploidy and growth-defense tradeoffs in natural populations of western quaking Aspen.

Polyploidy, the expression of more than two sets of chromosomes, is common in plants, and is thought to influence plant trait expression and drive plant species evolution. The degree to which polyploidy influences interactions among physiological processes such as growth and defense in natural populations through its effect on phenotypic variability is poorly understood. We link broad plant genotypic features (including polyploidy) to phenotypic expression of growth and chemical defense in natural populations of quaking aspen (Populus tremuloides) to examine patterns in resource allocation that might drive growth-defense tradeoffs. Quaking aspen are capable of rapid growth, and are also a primary food plant for a large range of herbivores, including insects and ungulates. While often diploid, aspen can exhibit polyploidy as triploid clones. We tested for the effect of genotype, cytotype (ploidy level, divided between diploids and triploids), and ramet age on relationships between growth and leaf chemistry across natural aspen clones in northern Utah. Substantial genotype variability in growth and leaf chemistry occurred across both cytotypes. Phenolic glycosides, but not condensed tannins, were negatively related to growth. Ramet age was also negatively related to growth. Phenolic glycosides were negatively related to condensed tannins, but only for the diploid clones. Triploid clones exhibited ~ 20% higher levels of phenolic glycosides than diploids. Growth in quaking aspen was likely sacrificed for the production of phenolic glycosides. Our study underscores the importance of considering polyploidy, genetic variability, and ramet age in understanding growth-defense tradeoffs in natural populations of clonal organisms, such as quaking aspen.

A metabolomic assessment of NAC154 transcription factor overexpression in field grown poplar stem wood

Several xylem-associated regulatory genes have been identified that control processes associated with wood formation in poplar. Prominent among these are the NAC domain transcription factors (NACs). Here, the putative involvement of Populus NAC154, a co-ortholog of the Arabidopsis gene SND2, was evaluated as a regulator of “secondary” biosynthetic processes in stem internode tissues by interrogating aqueous methanolic extracts from control and transgenic trees. Comprehensive untargeted metabolite profiling was accomplished with a liquid chromatography–mass spectrometry platform that utilized two different chromatographic supports (HILIC and reversed phase) and both positive and negative ionization modes. Evaluation of current and previous year tissues provided datasets for assessing the effects of NAC154 overexpression in wood maturation processes. Phenolic glycoside levels as well as those of oligolignols, sucrose and arginine were modulated with phenotypic and chemotypic traits exhibiting similar trends. Specifically, increased levels of arginine in the NAC154 overexpressing tissues supports a role for the transcription factor in senescence/dormancy-associated processes.

Root Chemistry in Populus tremuloides: Effects of Soil Nutrients, Defoliation, and Genotype

Although genetic, environmental, and G x E effects on aboveground phytochemistry have been well documented in trembling aspen (Populus tremuloides), little work has focused on the same factors affecting tissues underground. Belowground plant defenses are likely important mediators of root-feeding herbivores that can strongly influence plant fitness. We used a common garden of potted aspen trees to explore the individual and interactive effects of soil nutrient availability, foliar damage, genotype, and their interactions, on concentrations of phytochemicals in aspen roots. Our common garden experiment employed 12 aspen genotypes that were planted into either low- or high-nutrient soil environments. Half of the trees were subjected to defoliation for two successive years, while the others were protected from damage. At the end of the growing season after the second defoliation, we harvested the trees to obtain root samples for which we assessed levels of phenolic glycosides, condensed tannins, nitrogen, and starch. Phenolic glycosides were most affected by genotype, while the other root phytochemicals were most responsive to soil nutrient conditions. The effects of defoliation were observed in interaction with soil nutrient environment and/or genotype. Interestingly, the effect of defoliation on phenolic glycosides was mediated by soil nutrients, whereas the effect of defoliation on condensed tannins was observed in concert with effects of both soil nutrients and genotype. Comparison of data from this study with an earlier, related study revealed that concentrations of phenolic glycosides and condensed tannins are lower in roots than leaves, and less responsive to defoliation. That soil nutrient environment affects root phytochemical concentrations is not unexpected given the intimate association of roots and soil, but the complex interactions between soil nutrients, aboveground damage, and genotype, and their effects on root phytochemistry, are intriguing. Variation in root chemistry could have wide-reaching effects on soil microbial communities, nutrient cycling, and herbivores. Additionally, the response of phytochemicals to damage across organs can link different, spatially separated herbivores as they use different parts of the same plant resource.

Genotypic Differences and Prior Defoliation Affect Re-Growth and Phytochemistry after Coppicing in Populus tremuloides

Although considerable research has explored how tree growth and defense can be influenced by genotype, the biotic environment, and their interaction, little is known about how genotypic differences, prior defoliation, and their interactive effects persist in trees that re-grow after damage that severs their primary stem. To address these issues, we established a common garden consisting of twelve genotypes of potted aspen (Populus tremuloides) trees, and subjected half of the trees to defoliation in two successive years. At the beginning of the third year, all trees were severed at the soil surface (coppiced) and allowed to regenerate for five months. Afterwards, we counted the number of root and stump sprouts produced and measured the basal diameter (d) and height (h) of the tallest ramet in each pot. We collected leaves one and two years after the second defoliation and assessed levels of phenolic glycosides, condensed tannins, and nitrogen. In terms of re-growth, we found that the total number of sprouts produced varied by 3.6-fold among genotypes, and that prior defoliation decreased total sprout production by 24%. The size (d(2)h) of ramets, however, did not differ significantly among genotypes or defoliation classes. In terms of phytochemistry, we observed genotypic differences in concentrations of all phytochemicals assessed both one and two years after the second defoliation. Two years after defoliation, we observed effects of prior defoliation in a genotype-by-defoliation interaction for condensed tannins. Results from this study demonstrate that genotypic differences and impacts of prior defoliation persist to influence growth and defense traits in trees even after complete removal of above-ground stems, and thus likely influence productivity and plant-herbivore interactions in forests affected by natural disturbances or actively managed through coppicing.

Salicylate metabolism in Populus

Salicylate metabolism in Populus

Factors Influencing the Variability of Antioxidative Phenolic Glycosides in Salix Species

Phenolic glycosides, especially the salicylates, are important secondary metabolites in the bark of willows (Salix spp.). Because of their anti-inflammatory, analgesic, and fever-reducing properties, they are of particular interest to society. Compared to the fabrication of synthetic salicylacetylic acid, the commercial production of willow bark extracts with adequate amounts of salicylate is very difficult due to several biological and technical reasons. Therefore, one of the objectives was to identify salicylate-rich clones from three species, Salix daphnoides , Salix purpurea , and Salix pentandra , with potentially high amounts of phenolic glycosides. Three hundred different Salix clones were collected, and the chemical profiles of their bark were analyzed by HPLC. Overall, S. daphnoides clones showed the highest phenolic glycoside contents, followed by S. purpurea and S. pentandra. Second, seasonal changes of secondary compounds in willow bark were analyzed to determine the optimal harvesting time. The phenolic glycoside levels decreased over the growing season, with highest contents detected during plant dormancy. The effects of different cultivation conditions were also examined, and none of these treatments were found to have a significant effect on the phenolic glycoside content in willow bark. Biomass accumulation in the clones with grass competition was significantly lower than in the other three treatments.

Induction of Phenolic Glycosides by Quaking Aspen (Populus tremuloides) Leaves in Relation to Extrafloral Nectaries and Epidermal Leaf Mining

We studied the effect of epidermal leaf mining on the leaf chemistry of quaking aspen, Populus tremuloides, during an outbreak of the aspen leaf miner, Phyllocnistis populiella, in the boreal forest of interior Alaska. Phyllocnistis populiella feeds on the epidermal cells of P. tremuloides leaves. Eleven days after the onset of leaf mining, concentrations of the phenolic glycosides tremulacin and salicortin were significantly higher in aspen leaves that had received natural levels of leaf mining than in leaves sprayed with insecticide to reduce mining damage. In a second experiment, we examined the time course of induction in more detail. The levels of foliar phenolic glycosides in naturally mined ramets increased relative to the levels in insecticide-treated ramets on the ninth day following the onset of leaf mining. Induction occurred while some leaf miner larvae were still feeding and when leaves had sustained mining over 5% of the leaf surface. Leaves with extrafloral nectaries (EFNs) had significantly higher constitutive and induced levels of phenolic glycosides than leaves lacking EFNs, but there was no difference in the ability of leaves with and without EFNs to induce phenolic glycosides in response to mining. Previous work showed that the extent of leaf mining damage was negatively related to the total foliar phenolic glycoside concentration, suggesting that phenolic glycosides deter or reduce mining damage. The results presented here demonstrate that induction of phenolic glycosides can be triggered by relatively small amounts of mining damage confined to the epidermal tissue, and that these changes in leaf chemistry occur while a subset of leaf miners are still feeding within the leaf.

Phenological and chemical differences among hybrid poplar clones (Salicaceae) varying in resistance to Cryptorhynchus lapathi (L.) (Coleoptera: Curculionidae)

Hybrid poplars with Populus maximowiczii parentage have demonstrated antixenotic resistance against adult poplar-and-willow borers, Cryptorhynchus lapathi (L.) (Coleoptera: Curculionidae), in the summer, and antibiosis against C. lapathi larvae in the spring. We examined phenological and biochemical differences among hybrid poplars with and without P. maximowiczii parentage that are resistant and susceptible, respectively, to C. lapathi, to test the hypotheses that resistance was due to physical hydraulic forces, inadequate nutrition, or abundant secondary metabolites. The hybrids with P. maximowiczii parentage flushed earlier than susceptible hybrids, but no differences were found between hybrids with respect to sap flow or bark moisture content that could explain larval mortality in the spring. Similarly, constitutive levels of nutritive compounds (sitosterol, nitrogen, linoleic and linolenic acids, and carbohydrates) were lower or not different in the most susceptible hybrid TN 302-9. Thus, all hybrids appear to be nutritionally sound. TN 302-9 contained the highest constitutive levels of condensed tannins and catechin, but overall phenolic glycoside levels (salicin and salicortin) were similar among all hybrids, with the exception of one sampling date; however, this difference would not likely explain resistance. Neither induced or constitutive polyphenol oxidase (PPO) activity nor induced levels of secondary metabolites could explain resistance. Thus, we conclude hybrids that contain P. maximowiczii parentage have a novel mechanism for resistance.

Growth–defense tradeoffs differ by gender in dioecious trembling aspen ( Populus tremuloides)

Growth–defense tradeoffs are foundational to plant defense theory, but few studies have addressed the role of plant gender in such tradeoffs. We used an experimental trial garden of mature, dioecious Populus tremuloides trees to assess gender-based differences in growth, defense, and growth–defense tradeoffs. Male trees were 9% larger than female trees in terms of diameter-at-breast height (dbh) and had levels of defense chemicals [phenolic glycosides (PGs) and condensed tannins (CTs)] similar to those of females. Although male and female aspen had roughly similar levels of growth and defense, growth–defense tradeoffs differed markedly by gender. Growth of males was not related to either PG or CT concentrations, while growth of females was positively correlated with PG concentrations and negatively correlated with CT concentrations. Higher levels of PGs, if present throughout the life of a tree, may be protective against herbivores and result in greater growth over time.

Browse Quality in Quaking Aspen (Populus tremuloides): Effects of Genotype, Nutrients, Defoliation, and Coppicing

The consequences of interactions among genetic, ontogenetic, and environmental factors for the quality of winter-dormant tissues as food for browsing herbivores is poorly understood. We conducted two sequential common garden studies to assess the impacts of intraspecific genetic variation, nutrient availability, prior defoliation, and ontogenetic stage on the chemical quality of winter-dormant tissue in quaking aspen (Populus tremuloides Michx.). In the first study, saplings of 12 aspen genotypes were grown under low and high soil nutrient conditions, with or without two successive seasons of defoliation. Quantity and quality of current year's twig growth were assessed. Twig production varied among genotypes and declined under low nutrient availability, but showed little response to prior defoliation. Chemical quality of sapling twigs varied substantially among genotypes, and in response to nutrient availability and prior defoliation. Overall, browse quality improved (nitrogen levels increased while phenolic glycoside and condensed tannin levels decreased) after defoliation. Growth and chemical variables exhibited low to moderate clonal repeatability (broad sense heritability) values. Our second study employed the same 12 genotypes, grown under high-nutrient conditions and with or without two seasons of defoliation. The trees were coppiced to produce root sprouts, which were chemically assessed 1 yr later. Rejuvenation via coppicing led to increased levels of nitrogen, phenolic glycosides (salicortin), and tannins in root sprouts, and the magnitude of change varied among aspen genotypes. Signatures of defoliation nearly 2 yr earlier persisted in terms of elevated levels of phenolic glycosides in root sprouts of previously defoliated trees. Aspen forests likely present browsing herbivores with chemically heterogeneous environments because of the interactions of genetic, ontogenetic, and environmental factors that vary over space and time.

Tri‐trophic effects of plant defenses: chickadees consume caterpillars based on host leaf chemistry

Few studies have addressed how plant chemical defenses that directly affect herbivores in turn affect consumption patterns of vertebrates at higher trophic levels. We studied how variable foliar chemistry of trembling aspen (Populus tremuloides Michx.) affects the diet preferences of an avian insectivore feeding on an introduced herbivore, the gypsy moth (Lymantria dispar L.).Black‐capped chickadees (Poecile atricapilla) were offered paired choices of gypsy moth caterpillars feeding on one of three genotypes of aspen that differed in chemical composition. Chickadees chose to eat caterpillars fed aspen foliage with low levels of both condensed tannins and phenolic glycosides, or caterpillars fed foliage with high levels of tannins and low levels of phenolic glycosides, over caterpillars fed foliage with low levels of condensed tannins and high levels of phenolic glycosides. In addition, diet choices of the birds were affected by their previous experience. These findings are consistent with the “extended phenotype” concept, in that genetically‐based chemical traits in an ecologically dominant plant influence the feeding behavior of third trophic level organisms, whose efficacy as regulators of herbivore populations may in turn be modified.

Genotype and environment determine allocation to and costs of resistance in quaking aspen

Although genetic variability and resource availability both influence plant chemical composition, little is known about how these factors interact to modulate costs of resistance, expressed as negative correlations between growth and defense. We evaluated genotype x environment effects on foliar chemistry and growth of quaking aspen (Populus tremuloides) by growing multiple aspen genotypes under variable conditions of light and soil nutrient availability in a common garden. Foliage was analyzed for levels of nitrogen, phenolic glycosides and condensed tannins. Bioassays of leaf quality were conducted with fourth-stadium gypsy moth (Lymantria dispar) larvae. Results revealed strong effects of plant genotype, light availability and nutrient availability; the importance of each factor depended upon compound type. For example, tannin concentrations differed little among genotypes and across nutrient regimes under low light conditions, but markedly so under high light conditions. Phenolic glycoside concentrations, in contrast, were largely determined by genotype. Variation in phenolic glycoside concentrations among genotypes was the most important factor affecting gypsy moth performance. Gypsy moth biomass and development time were negatively and positively correlated, respectively, with phenolic glycoside levels. Allocation to phenolic glycosides appeared to be costly in terms of growth, but only under resource-limiting conditions. Context-dependent trade-offs help to explain why costs of allocation to resistance are often difficult to demonstrate.

Induced resistance in the indeterminate growth of aspen (Populus tremuloides)

Studies of induction in trees have examined rapid induced resistance (RIR) or delayed induced resistance (DIR), but have not examined induction that occurs in leaves produced by indeterminately growing trees subsequent to, but in the same season as, damage. We refer to induction that occurs during this time period as intermediate-delayed induced resistance (IDIR). We assessed the influences of genetic and environmental factors, and their interactions, on temporal and spatial variation in induction and on tradeoffs between induced and constitutive levels of resistance in indeterminately growing saplings of aspen (Populus tremuloides). We utilized a common garden of 12 aspen genotypes experiencing two levels of defoliation and two levels of soil nutrients. We assessed concentrations of phenolic glycosides and condensed tannins in damaged leaf remnants collected 1 week after defoliation to examine rapid and local induction, and in undamaged leaves produced 8 weeks after defoliation to assess intermediate-delayed and systemic induction. In general, tannins showed RIR, while phenolic glycosides expressed IDIR. For both classes of allelochemicals, we found high estimates of broad-sense heritability and genetic variation in both induced and constitutive levels. Genetic variation may be maintained by both direct costs of allelochemicals and by costs of inducibility (phenotypic plasticity). Such costs may drive the tradeoff exhibited between induced and constitutive levels of phenolic glycosides. IDIR may be important in reducing total-season tissue loss by providing augmented resistance against late summer herbivores in trees that have experienced damage earlier in the season. Herbivore-resistant compensatory growth is especially beneficial to young trees growing in competitive environments.

Cottonwood Leaf Beetle (Coleoptera: Chrysomelidae) Performance in Relation to Variable Phytochemistry in Juvenile Aspen (Populus tremuloidesMichx.)

Abstract Larval performance of the cottonwood leaf beetle, Chrysomela scripta F., was evaluated in relation to genetic variation in phytochemical characteristics among first year micropropagated ramets of five aspen clones (Populus tremuloides Michx.). Foliage from the juvenile ramets used in this experiment exhibited moderate variation in nitrogen, phenolic glycosides, and condensed tannin concentrations among clones, and overall, had very high levels of phenolic glycosides (15–22% dry weight) and low levels of condensed tannins (4–6% dry weight). Results from performance assays indicate that genetic differences among aspen clones resulted in only marginal differences in larval performance of this specialist leaf beetle. Although tannin levels were quite low in the juvenile trees, larval growth rate was reduced by 30% with increasing condensed tannin concentrations (R2 = 0.209). Recent evidence suggests that aspen undergoes ontogenetic shifts in foliar concentrations of secondary metabolites resulting in...

Foliar quality influences tree-herbivore-parasitoid interactions: effects of elevated CO2, O3, and plant genotype.

This study examined the effects of carbon dioxide (CO2)-, ozone (O3)-, and genotype-mediated changes in quaking aspen (Populus tremuloides) chemistry on performance of the forest tent caterpillar (Malacosoma disstria) and its dipteran parasitoid (Compsilura concinnata) at the Aspen Free-Air CO2 Enrichment (FACE) site. Parasitized and non-parasitized forest tent caterpillars were reared on two aspen genotypes under elevated levels of CO2 and O3, alone and in combination. Foliage was collected for determination of the chemical composition of leaves fed upon by forest tent caterpillars during the period of endoparasitoid larval development. Elevated CO2 decreased nitrogen levels but had no effect on concentrations of carbon-based compounds. In contrast, elevated O3 decreased nitrogen and phenolic glycoside levels, but increased concentrations of starch and condensed tannins. Foliar chemistry also differed between aspen genotypes. CO2, O3, genotype, and their interactions altered forest tent caterpillar performance, and differentially so between sexes. In general, enriched CO2 had little effect on forest tent caterpillar performance under ambient O3, but reduced performance (for insects on one aspen genotype) under elevated O3. Conversely, elevated O3 improved forest tent caterpillar performance under ambient, but not elevated, CO2. Parasitoid larval survivorship decreased under elevated O3, depending upon levels of CO2 and aspen genotype. Additionally, larval performance and masses of mature female parasitoids differed between aspen genotypes. These results suggest that host-parasitoid interactions in forest systems may be altered by atmospheric conditions anticipated for the future, and that the degree of change may be influenced by plant genotype.

Effects of genotype, nutrient availability, and defoliation on aspen phytochemistry and insect performance.

Genetic and environmental variability, and their interactions, influence phytochemical composition and, in turn, herbivore performance. We evaluated the independent and interactive effects of plant genotype, nutrient availability, and defoliation on the foliar chemistry of quaking aspen (Populus tremuloides) and consequences for performance of gypsy moths (Lymantria dispar). Saplings of four genotypes were grown under two conditions of nutrient availability and subjected to three levels of artificial defoliation. Concentrations of all secondary and primary metabolites evaluated responded to at least one or more of the experimental treatments. Of the secondary metabolites, phenolic glycosides were affected strongly by genotype, less so by nutrient availability, and not induced by defoliation. Condensed tannins were strongly dependent upon genotype, soil nutrient availability, and their interaction, and, in contrast to phenolic glycosides, were induced by artificial defoliation. Of the primary metabolites, foliar nitrogen was affected by genotype and soil nutrient availability. Starch concentrations were affected by genotype, nutrient availability, defoliation and interactions among these factors. Foliar water content responded to genotype, nutrient availability, and defoliation, and the effect of nutrient availability depended on genotype. Herbivore performance on these plants was strongly influenced by plant genotype and soil nutrient availability, but much less so by defoliation. Although several of the compound types (condensed tannins, starch, and water) responded to defoliation, quantitative variation in these compounds did not contribute to substantive changes in herbivore performance. Rather, the primary source of variation in insect performance was due to plant genotype (phenolic glycoside levels), while nutrient availability (foliar nitrogen levels) was of secondary importance. These results suggest that genetic variation in aspen plays a major role in determining patterns of insect performance, whereas environmental variation, such as was tested, here is of negligible importance.

Effects of Phenolic Glycosides and Protein on Gypsy Moth (Lepidoptera: Lymantriidae) and Forest Tent Caterpillar (Lepidoptera: Lasiocampidae) Performance and Detoxication Activities

Levels of phenolic glycosides and protein influence the quality of aspen leaves to herbivorous insects, and vary in relation to genetic and environmental factors. This research was conducted to assess the independent and interactive effects of phenolic glycosides and protein on the performance and detoxication enzyme activities of gypsy moths, Lymantria dispar (L.), and forest tent caterpillars, Malacosoma disstria Hubner. We fed fourth-stadium larvae aspen leaves supplemented with 0, 2, or 4% (wet weight) phenolic glycosides and 0 or 5% (wet weight) casein. We measured stadium duration, growth and consumption rates, and food conversion efficiencies. In addition, we measured the activities of three midgut enzymes likely involved in the metabolism of phenolic glycosides: b-glucosidase, esterase, and glutathione transferase. Phenolic glycosides re- duced performance of both insect species in terms of increased developmental time and decreased growth rates. Casein supplementation increased growth rates of gypsy moth larvae but slightly reduced growth rates of forest tent caterpillars. Phenolic glycosides and protein did not interactively influence stadium duration or growth rates. b-glucosidase activities declined for both insect species when reared on diets with phenolic glycosides. Esterase activities were induced by phenolic glycosides only in gypsy moths, whereas glutathione transferase activities were induced by phenolic glycosides in both species. Casein supplementation had little influence on enzyme activities, and phenolic glycosides and protein interactively affected only forest tent caterpillar esterase activity.

Effects of CO2 and light on tree phytochemistry and insect performance

Direct and interactive effects of CO2 and light on tree phytochemistry and insect fitness parameters were examined through experimental manipulations of plant growth conditions and performance of insect bioassays. Three species of deciduous trees (quaking aspen, Populus tremuloides; paper birch, Betula papyrifera; sugar maple, Acer saccharum) were grown under ambient (387±8 μL/L) and elevated (696±2 μL/L) levels of atmospheric CO2, with low and high light availability (375 and 855 μmol×m−2×s−1 at solar noon). Effects on the population and individual performance of a generalist phytophagous insect, the white‐marked tussock moth (Orgyia leucostigma) were evaluated. Caterpillars were reared on experimental trees for the duration of the larval stage, and complementary short‐term (fourth instar) feeding trials were conducted with insects fed detached leaves. Phytochemical analyses demonstrated strong effects of both CO2 and light on all foliar nutritional variables (water, starch and nitrogen). For all species, enriched CO2 decreased water content and increased starch content, especially under high light conditions. High CO2 availability reduced levels of foliar nitrogen, but effects were species specific and most pronounced for high light aspen and birch. Analyses of secondary plant compounds revealed that levels of phenolic glycosides (salicortin and tremulacin) in aspen and condensed tannins in birch and maple were positively influenced by levels of both CO2 and light. In contrast, levels of condensed tannins in aspen were primarily affected by light, whereas levels of ellagitannins and gallotannins in maple responded to light and CO2, respectively. The long‐term bioassays showed strong treatment effects on survival, development time, and pupal mass. In general, CO2 effects were pronounced in high light and decreased along the gradient aspen birch maple. For larvae reared on high light aspen, enriched CO2 resulted in 62% fewer survivors, with increased development time, and reduced pupal mass. For maple‐fed insects, elevated CO2 levels had negative effects on survival and pupal mass in low light. For birch, the only negative CO2 effects were observed in high light, where female larvae showed prolonged development. Fourth instar feeding trials demonstrated that low food conversion efficiency reduced insect performance. Elevated levels of CO2 significantly reduced total consumption, especially by insects on high light aspen and low light maple. This research demonstrates that effects of CO2 on phytochemistry and insect performance can be strongly light‐dependent, and that plant responses to these two environmental variables differ among species. Overall, increased CO2 availability appeared to increase the defensive capacity of early‐successional species primarily under high light conditions, and of late‐successional species under low light conditions. Due to the interactive effects of tree species, light, CO2, and herbivory, community composition of forests may change in the future.

Phytochemical Variation in Quaking Aspen: Effects on Gypsy Moth Susceptibility to Nuclear Polyhedrosis Virus

The performance of gypsy moths (Lymantria dispar) feeding on quaking aspen (Populus tremuloides) is strongly influenced by host foliar chemistry and susceptibility to a nuclear polyhedrosis virus (LdNPV), but the relationship of susceptibility to chemistry is poorly understood. We investigated the effects of genetic and resource-mediated variation in phytochemistry on viral pathogenicity. Trees were grown in pots in a common garden. Disks were punched from aspen leaves, inoculated with LdNPV and fed to third instars. Additional leaves were analyzed for levels of nitrogen, starch, phenolic glycosides, and condensed tannins. Despite marked variation among trees in levels of phenolic glycosides and tannins, we observed minimal variation in larval susceptibility to LdNPV. Viral pathogenicity was only weakly (inversely) correlated with tannin concentrations in one of two experiments. These results suggest that differential defoliation of aspen by gypsy moths in the field is due to the direct effects of host chemistry on larval performance rather than to the indirect effects of host chemistry on efficacy of this natural enemy.

Light environment alters response to ozone stress in seedlings of Acer saccharum Marsh, and hybrid Populus L.: III. Consequences for performance of gypsy moth.

Both light conditions and ozone fumigation alter the chemical composition of tree foliage and are thus likely to influence tree-insect interactions. We investigated the direct and interactive effects of light environment and ozone exposure on the performance of gypsy moth (Lymantria dispar L.) larvae reared on hybrid poplar (Populus tristis Fisch. ×P. balsamifera L. cv. Tristis) and sugar maple (Acer saccharum Marsh). We used a split-plot experimental design (light nested within ozone) and fourth-instar bioassays to calculate standard indices of insect growth and feeding performance. For insects fed poplar, consumption, growth and processing efficiencies were affected more by light environment than by ozone. Larvae ate and grew less on high-light foliage, responses attributable to higher levels of phenolic glycosides in those leaves. For insects fed maple, no significant effects of light, ozone, or light x ozone were observed. These results demonstrate that light environment and ozone pollution can alter the dynamics of interactions between trees and associated insects and that responses are species-specific.