Aspen Free Air CO2 Enrichment Research Articles

Modeling forest ecosystem responses to elevated carbon dioxide and ozone using artificial neural networks

Rising atmospheric levels of carbon dioxide and ozone will impact productivity and carbon sequestration in forest ecosystems. The scale of this process and the potential economic consequences provide an incentive for the development of models to predict the types and rates of ecosystem responses and feedbacks that result from and influence of climate change. In this paper, we use phenotypic and molecular data derived from the Aspen Free Air CO2 Enrichment site (Aspen-FACE) to evaluate modeling approaches for ecosystem responses to changing conditions. At FACE, it was observed that different aspen clones exhibit clone-specific responses to elevated atmospheric levels of carbon dioxide and ozone. To identify the molecular basis for these observations, we used artificial neural networks (ANN) to examine above and below-ground community phenotype responses to elevated carbon dioxide, elevated ozone and gene expression profiles. The aspen community models generated using this approach identified specific genes and subnetworks of genes associated with variable sensitivities for aspen clones. The ANN model also predicts specific co-regulated gene clusters associated with differential sensitivity to elevated carbon dioxide and ozone in aspen species. The results suggest ANN is an effective approach to predict relevant gene expression changes resulting from environmental perturbation and provides useful information for the rational design of future biological experiments.

Elevated carbon dioxide and ozone have weak, idiosyncratic effects on herbivorous forest insect abundance, species richness, and community composition

Abstract Elevated concentrations of carbon dioxide and tropospheric ozone pose important threats to the abundance, diversity, and composition of forest arthropod communities. In turn, modification of arthropod communities may alter forest health, productivity, and ecosystem services. We studied the independent and interactive effects of elevated CO2 (eCO2) and elevated O3 (eO3) on the abundance, species richness, and community composition of herbivorous arthropods in stands of trembling aspen and paper birch at the Aspen Free Air CO2 Enrichment (FACE) site in northern Wisconsin, USA. We conducted timed, visual surveys of canopy arthropods during each of the summers of 2005, 2006, and 2007. We examined 26 983 arthropods on aspen and 8344 arthropods on birch across the fumigation treatments. Elevated CO2 and eO3 had species‐specific and temporally variable (i.e. idiosyncratic) effects on aspen and birch arthropod abundance and species richness. Weak, idiosyncratic effects of eCO2 and eO3 on herbivorous arthropod abundance and species richness did not significantly alter aspen arthropod community composition but occasionally altered birch insect community composition. Few interactive effects of CO2 and O3 were observed. Growing evidence suggests that the effects of eCO2 and eO3 on communities of insects are difficult to predict because responses are generally weak and species‐ and time‐specific. Although studies to date suggest that impacts of future atmospheres on insect community metrics are likely to be minimal, the possibility remains that effects on particularly important or susceptible species may cascade to alter trophic interactions and, ultimately, ecosystem processes.

Increased belowground carbon inputs and warming promote loss of soil organic carbon through complementary microbial responses

Current carbon cycle-climate models predict that future soil carbon storage will be determined by the balance between CO2 fertilization and warming. However, it is uncertain whether greater carbon inputs to soils with elevated CO2 will be sequestered, particularly since warming hastens soil carbon decomposition rates, and may alter the response of soils to new plant inputs. We studied the effects of elevated CO2 and warming on microbial soil carbon decomposition processes using laboratory manipulations of carbon inputs and soil temperature. We incubated soils from the Aspen Free Air CO2 Enrichment experiment, where no accumulation of soil carbon has been observed despite a decade of increased carbon inputs to soils under elevated CO2. We added isotopically-labeled sucrose to these soils in the laboratory to mimic and trace the effects of increased carbon inputs on soil organic carbon decomposition and its temperature sensitivity. Sucrose additions caused a positive priming of soil organic carbon decomposition, demonstrated by increased respiration derived from soil carbon, increased microbial abundance, and a shift in the microbial community towards faster growing microorganisms. Similar patterns were observed for elevated CO2 soils, suggesting that the priming effect was responsible for reductions in soil carbon accumulation at the site. Laboratory warming accelerated the rate of the priming effect, but the magnitude of the priming effect was not different amongst temperatures, suggesting that the priming effect was limited by substrate availability, not soil temperature. No changes in substrate use efficiency were observed with elevated CO2 or warming. The stimulatory effects of warming on the priming effect suggest that increased belowground carbon inputs from CO2 fertilization are not likely to be stored in mineral soils.

Wood properties of Populus and Betula in long‐term exposure to elevated CO2 and O3

We studied the interactive effects of elevated concentrations of CO2 and O3 on radial growth and wood properties of four trembling aspen (Populus tremuloides Michx.) clones and paper birch (Betula papyrifera Marsh.) saplings. The material for the study was collected from the Aspen FACE (free-air CO2 enrichment) experiment in Rhinelander (WI, USA). Trees had been exposed to four treatments [control, elevated CO2 (560 ppm), elevated O3 (1.5 times ambient) and combined CO2 + O3 ] during growing seasons 1998-2008. Most treatment responses were observed in the early phase of experiment. Our results show that the CO2- and O3-exposed aspen trees displayed a differential balance between efficiency and safety of water transport. Under elevated CO2, radial growth was enhanced and the trees had fewer but hydraulically more efficient larger diameter vessels. In contrast, elevated O3 decreased radial growth and the diameters of vessels and fibres. Clone-specific decrease in wood density and cell wall thickness was observed under elevated CO2 . In birch, the treatments had no major impacts on wood anatomy or wood density. Our study indicates that short-term impact studies conducted with young seedlings may not give a realistic view of long-term ecosystem responses.

Influence of Global Atmospheric Change on the Feeding Behavior and Growth Performance of a Mammalian Herbivore, Microtus ochrogaster

Global atmospheric change is influencing the quality of plants as a resource for herbivores. We investigated the impacts of elevated carbon dioxide (CO2) and ozone (O3) on the phytochemistry of two forbs, Solidago canadensis and Taraxacum officinale, and the subsequent feeding behavior and growth performance of weanling prairie voles (Microtus ochrogaster) feeding on those plants. Plants for the chemical analyses and feeding trials were harvested from the understory of control (ambient air), elevated CO2 (560 µl CO2 l−1), and elevated O3 (ambient × 1.5) rings at the Aspen FACE (Free Air CO2 Enrichment) site near Rhinelander, Wisconsin. We assigned individual voles to receive plants from only one FACE ring and recorded plant consumption and weanling body mass for seven days. Elevated CO2 and O3 altered the foliar chemistry of both forbs, but only female weanling voles on the O3 diet showed negative responses to these changes. Elevated CO2 increased the fiber fractions of both plant species, whereas O3 fumigation elicited strong responses among many phytochemical components, most notably increasing the carbon-to-nitrogen ratio by 40% and decreasing N by 26%. Consumption did not differ between plant species or among fumigation treatments. Male voles were unaffected by the fumigation treatments, whereas female voles grew 36% less than controls when fed O3-grown plants. These results demonstrate that global atmospheric change has the potential to affect the performance of a mammalian herbivore through changes in plant chemistry.

Can elevatedCO2and ozone shift the genetic composition of aspen (Populus tremuloides) stands?

The world's forests are currently exposed to increasing concentrations of carbon dioxide (CO2) and ozone (O3). Both pollutants can potentially exert a selective effect on plant populations. This, in turn, may lead to changes in ecosystem properties, such as carbon sequestration. Here, we report how elevated CO2 and O3 affect the genetic composition of a woody plant population via altered survival. Using data from the Aspen free-air CO2 enrichment (FACE) experiment (in which aspen clones were grown in factorial combinations of CO2 and O3), we develop a hierarchical Bayesian model of survival. We also examine how survival differences between clones could affect pollutant responses in the next generation. Our model predicts that the relative abundance of the tested clones, given equal initial abundance, would shift under either elevated CO2 or O3 as a result of changing survival rates. Survival was strongly affected by between-clone differences in growth responses. Selection could noticeably decrease O3 sensitivity in the next generation, depending on the heritability of growth responses and the distribution of seed production. The response to selection by CO2, however, is likely to be small. Our results suggest that the changing atmospheric composition could shift the genotypic composition and average pollutant responses of tree populations over moderate timescales.

Performance of the invasive weevil Polydrusus sericeus is influenced by atmospheric CO2 and host species

Natural forest systems constitute a major portion of the world's land area, and are subject to the potentially negative effects of both global climate change and invasion by exotic insects. A suite of invasive weevils has become established in the northern hardwood forests of North America. How these insects will respond to increasing CO2 or O3 is unknown. The present study examined the effects of elevated atmospheric CO2 and O3 on the invasive weevil Polydrusus sericeus Schaller at the Aspen Free Air CO2 Enrichment (FACE) site near Rhinelander, Wisconsin. A performance assay was conducted in the laboratory during the summer of 2007 using mated pairs of P. sericeus fed a combination of aspen, birch and maple foliage. We recorded leaf area consumption, oviposition and adult longevity. We also conducted visual abundance surveys in the field from 2004 to 2007 on aspen and birch at Aspen FACE. Elevated CO2, but not O3, significantly affected P. sericeus performance. Female, but not male, longevity was reduced under elevated CO2. Polydrusus sericeus also produced fewer eggs under elevated CO2 conditions compared with ambient conditions. Adult P. sericeus strongly preferred birch over both aspen and maple, regardless of fumigation treatment. The effects of elevated CO2 on P. sericeus populations at Aspen FACE were minimal, and varied among years and host tree species. Polydrusus sericeus abundance was significantly greater on birch than aspen. Over the long term, elevated CO2 may reduce adult female longevity and fecundity of P. sericeus. Further studies are needed to evaluate how this information may scale to ecosystem impacts.

Soil carbon and nitrogen mineralization following deposition of insect frass and greenfall from forests under elevated CO2 and O3

Elevated CO2 and O3 alter tree quality and the quality of herbivore inputs, such as frass, to forest soil. Altered quality or quantity of herbivore inputs to the forest floor can have large impacts on below- ground processes. We collected green leaves and frass from whitemarked tussock moth caterpillars from aspen-birch stands at the Aspen Free Air CO2 Enrichment (FACE) site near Rhinelander, WI, USA. Small or large quantities of frass, greenfall, or a 1:1 ratio of frass and greenfall were added to microcosms for each FACE treatment (control, +CO2 ,+ O 3, +CO2+O3). We measured initial frass and greenfall quality, and recorded microbial respiration, and nitrate leaching over 40 days. Elevated carbon dioxide (eCO2) and tropospheric ozone (eO3) significantly altered the carbon, nitrogen, and condensed tannin content of insect frass and green leaves. Although FACE treatments affected input quality, they had minimal effect on microbial respiration and no effect on nitrogen leaching. In contrast, input quantity substantially influenced microbial respiration and nitrate leaching. Respiratory carbon loss and nitrate immobilization were nearly double in microcosms receiving large amounts of herbivore inputs than those receiving no herbivore inputs. Small amounts of herbivore inputs, however, did not significantly alter microbial respiration or immobilization, suggest- ing that effects of herbivore inputs on soil processes will be detected only at moderate to high herbivory/ input levels. These results suggest that subtle changes in frass and greenfall quality may not affect soil nutrient cycling. In contrast, environmental change induced increases in insect population size or frass and greenfall inputs to the soil may substantially impact nutrient cycling.

Productivity and community structure of ectomycorrhizal fungal sporocarps under increased atmospheric CO2 and O3

Sporocarp production is essential for ectomycorrhizal fungal recombination and dispersal, which influences fungal community dynamics. Increasing atmospheric carbon dioxide (CO2) and ozone (O3) affect host plant carbon gain and allocation, which may in turn influence ectomycorrhizal sporocarp production if the carbon available to the ectomycorrhizal fungus is dependant upon the quantity of carbon assimilated by the host. We measured sporocarp production of ectomycorrhizal fungi over 4 years at the Aspen FACE (free air CO2 enrichment) site, which corresponded to stand ages seven to 10 years. Total mean sporocarp biomass was greatest under elevated CO2, regardless of O3 concentration, while it was generally lowest under elevated O3 with ambient CO2. Community composition differed significantly among the treatments, with less difference in the final year of the study. Whether this convergence was due to succession or environmental factors is uncertain. CO2 and O3 affect ectomycorrhizal sporocarp productivity and community composition, with likely effects on dispersal, colonization and sporocarp-dependent food webs.

Transcriptomic comparison in the leaves of two aspen genotypes having similar carbon assimilation rates but different partitioning patterns under elevated [CO2

This study compared the leaf transcription profiles, physiological characteristics and primary metabolites of two Populus tremuloides genotypes (clones 216 and 271) known to differ in their responses to long-term elevated [CO2] (e[CO2]) at the Aspen free-air CO2 enrichment site near Rhinelander, WI, USA. The physiological responses of these clones were similar in terms of photosynthesis, stomatal conductance and leaf area index under e[CO2], yet very different in terms of growth enhancement (0-10% in clone 216; 40-50% in clone 271). Although few genes responded to long-term exposure to e[CO2], the transcriptional activity of leaf e[CO2]-responsive genes was distinctly different between the clones, differentially impacting multiple pathways during both early and late growing seasons. An analysis of transcript abundance and carbon/nitrogen biochemistry suggested that the CO2-responsive clone (271) partitions carbon into pathways associated with active defense/response to stress, carbohydrate/starch biosynthesis and subsequent growth. The CO2-unresponsive clone (216) partitions carbon into pathways associated with passive defense (e.g. lignin, phenylpropanoid) and cell wall thickening. This study indicates that there is significant variation in expression patterns between different tree genotypes in response to long-term exposure to e[CO2]. Consequently, future efforts to improve productivity or other advantageous traits for carbon sequestration should include an examination of genetic variability in CO2 responsiveness.

The Influence of Chemistry, Production and Community Composition on Leaf Litter Decomposition Under Elevated Atmospheric CO2 and Tropospheric O3 in a Northern Hardwood Ecosystem

We examined the effects of elevated CO2 and O3 and their interaction on leaf litter chemistry and decomposition in pure stands of aspen (Populus tremuloides) and mixed stands of birch (Betula papyrifera) and aspen at the Aspen Free Air CO2 Enrichment (FACE) experiment. A 935-day in situ incubation study was performed using litterbags filled with naturally senesced leaf litter. We found that elevated CO2 had no overall effects on litter decomposition rates, whereas elevated O3 reduced litter mass loss (−13%) in the first year. The effect of O3 on mass loss disappeared in the second year. For aspen litter but not mixed birch-aspen litter, decomposition rates were negatively correlated with initial concentrations of condensed tannins and phenolics. Most soluble components (94% of soluble sugars, 99% of condensed tannins, and 91% of soluble phenolics) and any treatment effects on their initial concentrations disappeared rapidly. However, the mean residence time (MRT) of birch-aspen litter (3.1 years) was significantly lower than that of aspen litter (4.8 years). Further, because of variation in total litterfall, total litter mass, C, lignin and N remaining in the ecosystem was highest under elevated CO2 and lowest under elevated O3 during the incubation period. Our results indicate that elevated CO2 and O3 can alter short-term litter decomposition dynamics, but longer-term effects will depend more on indirect effects mediated through changes in forest community composition. Treatment effects on soluble components are likely to influence cyclical microbial processes and carbon pulses in the ecosystem only when coupled with increased (CO2) or decreased (O3) litter inputs.

Elevated atmospheric carbon dioxide and ozone alter forest insect abundance and community composition

Abstract. 1 Human-induced climate changes threaten the health of forest ecosystems. In particular, carbon dioxide (CO2) and tropospheric ozone (O3) will likely have significant but opposing impacts on forests and their associated insect communities. Compared with other animal groups, insect communities are expected to be especially sensitive to changes in global climate. 2 This study examined the effects of elevated CO2 and O3 (eCO2 and eO2) individually and in combination on the abundance, diversity and composition of forest insect communities. Insects were sampled using yellow pan traps in an aggrading aspen-birch forest at the Aspen Free Air CO2 Enrichment (FACE) site in northern Wisconsin, USA. We trapped for 24 h every 10–15 days throughout the summers (June to September) of 2000–2003. 3 We examined 47 415 insects from 4 orders and 83 families. Elevated CO2 reduced abundance of phloem-feeding herbivores and increased abundance of chewing herbivores, although results were not statistically significant. Enriched CO2 increased numbers of some parasitoids. The effects of eO3 on insect abundance were generally opposite those of eCO2. No significant differences in arthropod family richness were found among treatments. However, eCO2, eO3, or both significantly affected insect community composition in all years. 4 Carbon dioxide and tropospheric ozone have the potential to alter significantly forest insect communities. Feeding guild may strongly influence insect response to environmental change and may provide the best opportunity to generalise for conservation efforts. Because insect communities influence forest health and ecosystem services, continued research on their response to global change is critically important to forest management and conservation.

Independent, Interactive, and Species-Specific Responses of Leaf Litter Decomposition to Elevated CO2 and O3 in a Northern Hardwood Forest

The future capacity of forest ecosystems to sequester atmospheric carbon is likely to be influenced by CO2-mediated shifts in nutrient cycling through changes in litter chemistry, and by interactions with pollutants like O3. We evaluated the independent and interactive effects of elevated CO2 (560 l ll )1 ) and O3 (55 nl l l )1 )o n leaf litter decomposition in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) at the Aspen free air CO2 enrichment (FACE) site (Wisconsin, USA). Fumigation treatments consisted of replicated ambient, +CO2, +O3, and +CO2 +O 3 FACE rings. We followed mass loss and litter chemistry over 23 months, using reciprocally transplanted litterbags to separate substrate quality from environment effects. Aspen decayed more slowly than birch across all treatment conditions, and changes in decomposition dynamics of both species were driven by shifts in substrate quality rather than by fumigation environment. Aspen litter produced under elevated CO2 decayed more slowly than litter produced under ambient CO2, and this effect was exacerbated by elevated O3. Similarly, birch litter produced under elevated CO2 also decayed more slowly than litter produced under ambient CO2. In contrast to results for aspen, however, elevated O3 accelerated birch decay under ambient CO2, but decelerated decay under enriched CO2. Changes in decomposition rates (k-values) were due to CO2- and O3-mediated shifts in litter quality, particularly levels of carbohydrates, nitrogen, and tannins. These results suggest that in early-successional forests of the future, elevated concentrations of CO2 will likely reduce leaf litter decomposition, although the magnitude of effect will vary among species and in response to interactions with tropospheric O3.

Effects of elevated concentrations of atmospheric CO2 and tropospheric O3 on leaf litter production and chemistry in trembling aspen and paper birch communities

Human activities are increasing the concentrations of atmospheric carbon dioxide ([CO2]) and tropospheric ozone ([O3]), potentially leading to changes in the quantity and chemical quality of leaf litter inputs to forest soils. Because the quality and quantity of labile and recalcitrant carbon (C) compounds influence forest productivity through changes in soil organic matter content, characterizing changes in leaf litter in response to environmental change is critical to understanding the effects of global change on forests. We assessed the independent and combined effects of elevated [CO2] and elevated [O3] on foliar litter production and chemistry in aspen (Populus tremuloides Michx.) and birch-(Betula papyrifera Marsh.) aspen communities at the Aspen free-air CO2 enrichment (FACE) experiment in Rhinelander, WI. Litter was analyzed for concentrations of C, nitrogen (N), soluble sugars, lipids, lignin, cellulose, hemicellulose and C-based defensive compounds (soluble phenolics and condensed tannins). Concentrations of these chemical compounds in naturally senesced litter were similar in aspen and birch-aspen communities among treatments, except for N, the C:N ratio and lipids. Elevated [CO2] significantly increased C:N (+8.7%), lowered mean litter N concentration (-10.7%) but had no effect on the concentrations of soluble sugars, soluble phenolics and condensed tannins. Elevated [CO2] significantly increased litter biomass production (+33.3%), resulting in significant increases in fluxes of N, soluble sugars, soluble phenolics and condensed tannins to the soil. Elevated [O3] significantly increased litter concentrations of soluble sugars (+78.1%), soluble phenolics (+53.1%) and condensed tannins (+77.2%). There were no significant effects of elevated [CO2] or elevated [O3] on the concentrations of individual C structural carbohydrates (cellulose, hemicellulose and lignin). Elevated [CO2] significantly increased cellulose (+37.4%) input to soil, whereas elevated [O3] significantly reduced hemicellulose and lignin inputs to soil (-22.3 and -31.5%, respectively). The small changes in litter chemistry in response to elevated [CO2] and tropospheric [O3] that we observed, combined with changes in litter biomass production, could significantly alter the inputs of N, soluble sugars, condensed tannins, soluble phenolics, cellulose and lignin to forest soils in the future.

Effects of elevated concentrations of atmospheric CO2 and tropospheric O3 on decomposition of fine roots

Rising atmospheric carbon dioxide (CO2) concentration ([CO2]) could alter terrestrial carbon (C) cycling by affecting plant growth, litter chemistry and decomposition. How the concurrent increase in tropospheric ozone (O3) concentration ([O3]) will interact with rising atmospheric [CO2] to affect C cycling is unknown. A major component of carbon cycling in forests is fine root production, mortality and decomposition. To better understand the effects of elevated [CO2] and [O3] on the dynamics of fine root C, we conducted a combined field and laboratory incubation experiment to monitor decomposition dynamics and changes in fine root litter chemistry. Free-air CO2 enrichment (FACE) technology at the FACTS-II Aspen FACE project in Rhinelander, Wisconsin, elevated [CO2] (535 microl 1-1) and [O3] (53 nl 1-1) in intact stands of pure trembling aspen (Populus tremuloides Michx.) and in mixed stands of trembling aspen plus paper birch (Betula papyrifera Marsh.) and trembling aspen plus sugar maple (Acer saccharum Marsh.). We hypothesized that the trees would react to increased C availability (elevated [CO2]) by increasing allocation to C-based secondary compounds (CBSCs), thereby decreasing rates of decomposition. Because of its lower growth potential, we reasoned this effect would be greatest in the aspen-maple community relative to the aspen and aspen-birch communities. As a result of decreased C availability, we expected elevated [O3] to counteract shifts in C allocation induced by elevated [CO2]. Concentrations of CBSCs were rarely significantly affected by the CO2 and O3 treatments in decomposing fine roots. Rates of microbial respiration and mass loss from fine roots were unaffected by the treatments, although the production of dissolved organic C differed among communities. We conclude that elevated [CO2] and [O3] induce only small changes in fine root chemistry that are insufficient to significantly influence fine root decomposition. If changes in soil C cycling occur in the future, they will most likely be brought about by changes in litter production.

CO2 and O3 effects on host plant preferences of the forest tent caterpillar (Malacosoma disstria)

AbstractElevated levels of CO2 and O3 affect plant growth and phytochemistry, which in turn can alter physiological performance of associated herbivores. Little is known, however, about how generalist insect herbivores respond behaviorally to CO2‐ and O3‐mediated changes in their host plants. This research examined the effects of elevated CO2 and O3 levels on host plant preferences and consumption of forest tent caterpillar (FTC, Malacosoma disstria Hbn.) larvae. Dual choice feeding assays were performed with foliage from birch (Betula papyrifera Marsh.) and aspen (Populus tremuloides Michx., genotypes 216 and 259). Trees were grown at the Aspen Free Air CO2 Enrichment (FACE) facility near Rhinelander, WI, USA, and had been exposed to ambient or elevated concentrations of CO2 and/or O3. Levels of nutritional and secondary compounds were quantified through phytochemical analyses. The results showed that elevated O3 levels increased FTC larval preferences for birch compared with aspen, whereas elevated CO2 levels had the opposite effect. In assays with the two aspen genotypes, addition of both CO2 and O3 caused a shift in feeding preferences from genotype 259 to genotype 216. Consumption was unaffected by experimental treatments in assays comparing aspen and birch, but were increased for larvae given high O3 foliage in the aspen genotype assays. Elevated levels of CO2 and O3 altered tree phytochemistry, but did not explain shifts in feeding preferences. The results demonstrate that increased levels of CO2 and O3 can alter insect host plant preferences both between and within tree species. Also, consequences of altered host quality (e.g., compensatory consumption) may be buffered by partial host shifts in situations when alternative plant species are available. Environmentally induced changes in host plant preferences may have the potential to alter the distribution of herbivory across plant genotypes and species, as well as competitive interactions among them.

Aphid individual performance may not predict population responses to elevated CO2 or O3

AbstractChanges in atmospheric composition affect plant quality and herbivore performance. We used the Aspen Free Air CO2 Enrichment (FACE) facility to investigate the impacts of elevated carbon dioxide (CO2) and ozone (O3) on the performance of the aphid Cepegillettea betulaefoliae Granovsky feeding on paper birch (Betula papyrifera Marsh.). In Year 1, we simultaneously measured individual performance and population growth rates, and in Year 2 we surveyed natural aphid, predator and parasitoid populations throughout the growing season. Aphid growth and development (relative growth rate (RGR), development time, adult weight, embryo number and the birth weight of newborn nymphs) were unaffected by CO2 and O3. Aphid fecundity decreased on trees grown at elevated CO2, O3 and CO2+O3. Neither nymphal performance nor adult size were reliable indicators of future fecundity at elevated CO2 and/or O3. Aphid populations protected from natural enemies were unaffected by elevated CO2, but increased significantly at elevated O3. Individual fecundity in elevated CO2 and O3 atmospheres did not predict population growth rates, probably because of changes in the strength of intraspecific competition or the ability of the aphids to induce nutrient sinks. Natural aphid, predator and parasitoids populations (Year 2) showed few significant responses to CO2 and O3, although CO2 and O3 did affect the timing of aphid and natural enemy peak abundance. Elevated CO2 and O3 affected aphid and natural enemy populations independently: no CO2× O3 interactions were observed. We conclude that: (1) aphid individual performance did not predict population responses to CO2 and O3 and (2) elevated CO2 and O3 atmospheres are unlikely to affect C. betulaefoliae populations in the presence of natural enemy communities.

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.

Responses of trembling aspen (Populus tremuloides) phytochemistry and aspen blotch leafminer (Phyllonorycter tremuloidiella) performance to elevated levels of atmospheric CO2 and O3

Abstract1 This research was conducted at the Aspen FACE (Free Air CO2 Enrichment) site located in northern Wisconsin, U.S.A. where trembling aspen (Populus tremuloides Michaux) trees were exposed to one of four atmospheric treatments: elevated carbon dioxide (CO2; 560 µL/L), elevated ozone (O3; ambient × 1.5), elevated CO2 and O3, or ambient air. We evaluated the effects of these fumigants on aspen foliar quality and the performance of aspen blotch leafminer (Phyllonorycter tremuloidiella Braun).2 CO2 and O3 each affected foliar quality, with the major changes consisting of an 11% reduction in nitrogen under elevated CO2 and a 20% reduction in tremulacin under elevated O3. In the CO2 + O3 treatment, nitrogen levels were reduced by 15% and CO2 ameliorated the O3‐mediated reduction in tremulacin levels.3 Phyllonorycter tremuloidiella were allowed to colonize trees naturally. Elevated CO2 and O3 reduced colonization rates by 42 and 49% relative to ambient CO2 and O3, respectively. The only effect of fumigation treatments on larval performance occurred under elevated O3, where male development time and larval consumption increased by 8 and 28%, respectively, over insects reared under ambient O3.4 These data demonstrate that the individual and combined effects of CO2 and O3 can alter aspen foliar chemistry and that these alterations in foliar chemistry produce little to no change in larval performance. However, both CO2 and O3 greatly reduced oviposition. In order to ascertain the full effects of CO2 and O3 on insect performance, future studies should address both population‐ and individual‐level characteristics.

Response of quaking aspen genotypes to enriched CO2: foliar chemistry and tussock moth performance

Abstract Genetic variation in the phytochemical responses of plants to CO2 enrichment is likely to alter trophic dynamics, and to shift intraspecific selection pressures on plant populations. We evaluated the independent and interactive effects of atmospheric CO2 and quaking aspen (Populus tremuloides Michx.) genotype on chemical composition of foliage and performance of the whitemarked tussock moth (Orgyia leucostigma J. E. Sm.). This research was conducted at the Aspen FACE (Free Air CO2 Enrichment) site in northern Wisconsin, U.S.A. Leaf samples were collected periodically from each of three genetically variable aspen genotypes growing under ambient and elevated CO2, and analysed for levels of primary and secondary metabolites. Tussock moth larvae were reared in situ on experimental trees, and development times and pupal masses were recorded. Foliar chemical composition varied among aspen genotypes and in response to CO2 enrichment. However, chemical responses of trees to elevated CO2 were generally consistent across genotypes. Larval development times varied among host genotypes and increased slightly for insects on high‐CO2 plants. Enriched CO2 tended to reduce insect pupal masses, particularly for females on one of the three aspen genotypes. CO2 × genotype interactions observed for plant chemistry and insect performance in this study with a small number of genotypes are probably too few, and too weak, to shift selection pressures in aspen populations. These results differ, however, from earlier work in which more substantial CO2 × genotype interactions were observed for plant chemistry.