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

Elevated 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.     

Interacting elevated CO2 and tropospheric O3 predisposes aspen (Populus tremuloides Michx.) to infection by rust (Melampsora medusae f. sp. tremuloidae)

We investigated the interaction of elevated CO₂ and/or (Ozone) O₃ on the occurrence and severity of aspen leaf rust (Melampsora medusae Thuem. f. sp. tremuloidae) on trembling aspen (Populus tremuloides Michx.). Furthermore, we examined the role of changes in leaf surface properties induced by elevated CO₂ and/or O₃ in this host–pathogen interaction. Three‐ to five‐fold increases in levels of rust infection index were found in 2 consecutive years following growing‐season‐long exposures with either O₃ alone or CO₂ + O₃ depending on aspen clone. Examination of leaf surface properties (wax appearance, wax amount, wax chemical composition, leaf surface and wettability) suggested significant effects by O₃ and CO₂ + O₃. We conclude that elevated O₃ is altering aspen leaf surfaces in such a way that it is likely predisposing the plants to increased infection by aspen leaf rust.     

Atmospheric CO2 and O3 alter competition for soil nitrogen in developing forests

Plant growth responses to rising atmospheric CO₂ and O₃ vary among genotypes and between species, which could plausibly influence the strength of competitive interactions for soil N. Ascribable to the size‐symmetric nature of belowground competition, we reasoned that differential growth responses to CO₂ and O₃ should shift as juvenile individuals mature, thereby altering competitive hierarchies and forest composition. In a 12‐year‐long forest FACE experiment, we used tracer ¹⁵N and whole‐plant N content to assess belowground competitive interactions among five Populus tremuloides genotypes, between a single P. tremuloides genotype and Betula papryrifera, as well as between the same single P. tremuloides genotype and Acer saccharum. Under elevated CO₂, the amount of soil N and ¹⁵N obtained by the P. tremuloides genotype common to each community was contingent on the nature of belowground competition. When this genotype competed with its congeners, it obtained equivalent amounts of soil N and tracer ¹⁵N under ambient and elevated CO₂; however, its acquisition of soil N under elevated CO₂ increased by a significant margin when grown in competition with B. papyrifera (+30%) and A. saccharum (+60%). In contrast, elevated O₃ had no effect on soil N and ¹⁵N acquisition by the P. tremuloides genotype common in each community, regardless of competitive interactions. Under elevated CO₂, the rank order of N acquisition among P. tremuloides genotypes shifted over time, indicating that growth responses to CO₂ change during ontogeny; this was not the case under elevated O₃. In the aspen‐birch community, the competitive advantage elevated CO₂ initially conveyed on birch diminished over time, whereas maple was a poor competitor for soil N in all regards. The extent to which elevated CO₂ and O₃ will shape the genetic structure and composition of future forests is, in part, contingent on the time‐dependent effects of belowground competition on plant growth response.     

Chemistry and decomposition of litter from Populus tremuloides Michaux grown at elevated atmospheric CO2 and varying N availability

It has been hypothesized that greater production of total nonstructural carbohydrates (TNC) in foliage grown under elevated atmospheric carbon dioxide (CO₂) will result in higher concentrations of defensive compounds in tree leaf litter, possibly leading to reduced rates of decomposition and nutrient cycling in forest ecosystems of the future. To evaluate the effects of elevated atmospheric CO₂ on litter chemistry and decomposition, we performed a 111 day laboratory incubation with leaf litter of trembling aspen (Populus tremuloides Michaux) produced at 36 Pa and 56 Pa CO₂ and two levels of soil nitrogen (N) availability. Decomposition was quantified as microbially respired CO₂ and dissolved organic carbon (DOC) in soil solution, and concentrations of nonstructural carbohydrates, N, carbon (C), and condensed tannins were monitored throughout the incubation. Growth under elevated atmospheric CO₂ did not significantly affect initial litter concentrations of TNC, N, or condensed tannins. Rates of decomposition, measured as both microbially respired CO₂ and DOC did not differ between litter produced under ambient and elevated CO₂. Total C lost from the samples was 38 mg g^?1 litter as respired CO₂ and 138 mg g^?1 litter as DOC, suggesting short‐term pulses of dissolved C in soil solution are important components of the terrestrial C cycle. We conclude that litter chemistry and decomposition in trembling aspen are minimally affected by growth under higher concentrations of CO₂.     

Soil nitrogen transformations under Populus tremuloides, Betula papyrifera and Acer saccharum following 3 years exposure to elevated CO2 and O3

Increases in atmospheric CO₂ and tropospheric O₃ may affect forest N cycling by altering plant litter production and the availability of substrates for microbial metabolism. Three years following the establishment of our free‐air CO₂–O₃ enrichment experiment, plant growth has been stimulated by elevated CO₂ resulting in greater substrate input to soil; elevated O₃ has counteracted this effect. We hypothesized that rates of soil N cycling would be enhanced by greater plant productivity under elevated CO₂, and that CO₂ effects would be dampened by O₃. We found that elevated CO₂ did not alter gross N transformation rates. Elevated O₃ significantly reduced gross N mineralization and microbial biomass N, and effects were consistent among species. We also observed significant interactions between CO₂ and O₃: (i) gross N mineralization was greater under elevated CO₂ (1.0 mg N kg^?1 day^?1) than in the presence of both CO₂ and O₃ (0.5 mg N kg^?1 day^?1) and (ii) gross NH₄⁺ immobilization was also greater under elevated CO₂ (0.8 mg N kg^?1 day^?1) than under CO₂ plus O₃ (0.4 mg N kg^?1 day^?1). We used a laboratory ¹⁵N tracer method to quantify transfer of inorganic N to organic pools. Elevated CO₂ led to greater recovery of NH₄⁺‐¹⁵N in microbial biomass and corresponding lower recovery in the extractable NO₃^? pool. Elevated CO₂ resulted in a substantial increase in NO₃^?‐¹⁵N recovery in soil organic matter. We observed no O₃ main effect and no CO₂ by O₃ interaction effect on ¹⁵N recovery in any soil pool. All of the above responses were most pronounced beneath Betula papyrifera and Populus tremuloides, which have grown more rapidly than Acer saccharum. Although elevated CO₂ has increased plant productivity, the resulting increase in plant litter production has yet to overcome the influence of the pre‐existing pool of soil organic matter on soil microbial activity and rates of N cycling. Ozone reduces plant litter inputs and also appears to affect the composition of plant litter in a way that reduces microbial biomass and activity.     

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

1 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.     

Soil CO2 efflux of two silver birch clones exposed to elevated CO2 and O3 levels during three growing seasons

In the present open‐top chamber experiment, two silver birch clones (Betula pendula Roth, clone 4 and clone 80) were exposed to elevated levels of carbon dioxide (CO₂) and ozone (O₃), singly and in combination, and soil CO₂ efflux was measured 14 times during three consecutive growing seasons (1999–2001). In the beginning of the experiment, all experimental trees were 7 years old and during the experiment the trees were growing in sandy field soil and fertilized regularly. In general, elevated O₃ caused soil CO₂ efflux stimulation during most measurement days and this stimulation enhanced towards the end of the experiment. The overall soil respiration response to CO₂ was dependent on the genotype, as the soil CO₂ efflux below clone 80 trees was enhanced and below clone 4 trees was decreased under elevated CO₂ treatments. Like the O₃ impact, this clonal difference in soil respiration response to CO₂ increased as the experiment progressed. Although the O₃ impact did not differ significantly between clones, a significant time × clone × CO₂× O₃ interaction revealed that the O₃‐induced stimulation of soil respiration was counteracted by elevated CO₂ in clone 4 on most measurement days, whereas in clone 80, the effect of elevated CO₂ and O₃ in combination was almost constantly additive during the 3‐year experiment. Altogether, the root or above‐ground biomass results were only partly parallel with the observed soil CO₂ efflux responses. In conclusion, our data show that O₃ impacts may appear first in the below‐ground processes and that relatively long‐term O₃ exposure had a cumulative effect on soil CO₂ efflux. Although the soil respiration response to elevated CO₂ depended on the tree genotype as a result of which the O₃ stress response might vary considerably within a single tree species under elevated CO₂, the present experiment nonetheless indicates that O₃ stress is a significant factor affecting the carbon cycling in northern forest ecosystems.     

Decomposition of Betula papyrifera leaf litter under the independent and interactive effects of elevated CO2 and O3

Litter decay dynamics of paper birch (Betula papyrifera) were assessed at the Aspen free‐air CO₂ enrichment (FACE) facility in northern Wisconsin, USA. Leaf litter was decomposed for 12 months under factorial combinations of 360 vs. 560 μL CO₂ L^?1, crossed with 36 vs. 55 nL O₃ L^?1. To differentiate between substrate quality and environment effects, litterbags were placed in their Native Plots of origin or transplanted into the other treatments. CO₂ enrichment, regardless of O₃ concentration, produced poorer quality litter (high C/N, lignin/N and condensed tannins) than did ambient CO₂ (low C/N, lignin/N and condensed tannins). Substrate quality differences were reflected in the mass loss rates (k‐values), which were high for litter generated under ambient CO₂ (0.887 year^?1) and low for litter generated under elevated CO₂ (0.674 year^?1). The rate‐retarding effects of CO₂ enrichment were neither alleviated nor exacerbated by O₃ exposure. Decay rates varied, however, depending on whether litter was placed back into its plot of origin or transplanted to Common Gardens. The results of this study are species specific, but they have important implications for understanding the processes regulating storage of fixed C and the release of CO₂ from northern forest ecosystems.     

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.     

Below-ground responses of silver birch trees exposed to elevated CO2 and O3 levels during three growing seasons

Field‐growing silver birch (Betula pendula Roth) clones (clone 4 and 80) were exposed to elevated CO₂ and O₃ in open‐top chambers for three consecutive growing seasons (1999–2001). At the beginning of the OTC experiment, all trees were 7 years old. We studied the single and interaction effects of CO₂ and O₃ on silver birch below‐ground carbon pools (i.e. effects on fine roots and mycorrhizas, soil microbial communities and sporocarp production) and also assessed whether there are any clonal differences in these below‐ground CO₂ and O₃ responses. The total mycorrhizal infection level of both clones was stimulated by elevated CO₂ alone and elevated O₃ alone, but not when elevated CO₂ was used in fumigation in combination with elevated O₃. In both clones, elevated CO₂ affected negatively light brown/orange mycorrhizas, while its effect on other mycorrhizal morphotypes was negligible. Elevated O₃, instead, clearly decreased the proportions of black and liver‐brown mycorrhizas and increased that of light brown/orange mycorrhizas. Elevated O₃ had a tendency to decrease standing fine root mass and sporocarp production as well, both of these O₃ effects mainly manifesting in clone 4 trees. CO₂ and O₃ treatment effects on soil microbial community composition (PLFA, 2‐ and 3‐OH‐FA profiles) were negligible, but quantitative PLFA data showed that in 2001 the PLFA fungi : bacteria‐ratio of clone 80 trees was marginally increased because of elevated CO₂ treatments. This study shows that O₃ effects were most clearly visible at the mycorrhizal root level and that some clonal differences in CO₂ and O₃ responses were observable in the below‐ground carbon pools. In conclusion, the present data suggests that CO₂ effects were minor, whereas increasing tropospheric O₃ levels can be an important stress factor in northern birch forests, as they might alter mycorrhizal morphotype assemblages, mycorrhizal infection rates and sporocarp production.     

Decomposition of soybean grown under elevated concentrations of CO2 and O3

A critical global climate change issue is how increasing concentrations of atmospheric CO₂ and ground‐level O₃ will affect agricultural productivity. This includes effects on decomposition of residues left in the field and availability of mineral nutrients to subsequent crops. To address questions about decomposition processes, a 2‐year experiment was conducted to determine the chemistry and decomposition rate of aboveground residues of soybean (Glycine max (L.) Merr.) grown under reciprocal combinations of low and high concentrations of CO₂ and O₃ in open‐top field chambers. The CO₂ treatments were ambient (370 μmol mol^?1) and elevated (714 μmol mol^?1) levels (daytime 12 h averages). Ozone treatments were charcoal‐filtered air (21 nmol mol^?1) and nonfiltered air plus 1.5 times ambient O₃ (74 nmol mol^?1) 12 h day^?1. Elevated CO₂ increased aboveground postharvest residue production by 28–56% while elevated O₃ suppressed it by 15–46%. In combination, inhibitory effects of added O₃ on biomass production were largely negated by elevated CO₂. Plant residue chemistry was generally unaffected by elevated CO₂, except for an increase in leaf residue lignin concentration. Leaf residues from the elevated O₃ treatments had lower concentrations of nonstructural carbohydrates, but higher N, fiber, and lignin levels. Chemical composition of petiole, stem, and pod husk residues was only marginally affected by the elevated gas treatments. Treatment effects on plant biomass production, however, influenced the content of chemical constituents on an areal basis. Elevated CO₂ increased the mass per square meter of nonstructural carbohydrates, phenolics, N, cellulose, and lignin by 24–46%. Elevated O₃ decreased the mass per square meter of these constituents by 30–48%, while elevated CO₂ largely ameliorated the added O₃ effect. Carbon mineralization rates of component residues from the elevated gas treatments were not significantly different from the control. However, N immobilization increased in soils containing petiole and stem residues from the elevated CO₂, O₃, and combined gas treatments. Mass loss of decomposing leaf residue from the added O₃ and combined gas treatments was 48% less than the control treatment after 20 weeks, while differences in decomposition of petiole, stem, and husk residues among treatments were minor. Decreased decomposition of leaf residues was correlated with lower starch and higher lignin levels. However, leaf residues only comprised about 20% of the total residue biomass assayed so treatment effects on mass loss of total aboveground residues were relatively small. The primary influence of elevated atmospheric CO₂ and O₃ concentrations on decomposition processes is apt to arise from effects on residue mass input, which is increased by elevated CO₂ and suppressed by O₃.     

Interannual climatic variation mediates elevated CO2 and O3 effects on forest growth

We analyzed growth data from model aspen (Populus tremuloides Michx.) forest ecosystems grown in elevated atmospheric carbon dioxide ([CO₂]; 518 μL L^?1) and ozone concentrations ([O₃]; 1.5 × background of 30–40 nL L^?1 during daylight hours) for 7 years using free‐air CO₂ enrichment technology to determine how interannual variability in present‐day climate might affect growth responses to either gas. We also tested whether growth effects of those gasses were sustained over time. Elevated [CO₂] increased tree heights, diameters, and main stem volumes by 11%, 16%, and 20%, respectively, whereas elevated ozone [O₃] decreased them by 11%, 8%, and 29%, respectively. Responses similar to these were found for stand volume and basal area. There were no growth responses to the combination of elevated [CO₂+O₃]. The elevated [CO₂] growth stimulation was found to be decreasing, but relative growth rates varied considerably from year to year. Neither the variation in annual relative growth rates nor the apparent decline in CO₂ growth response could be explained in terms of nitrogen or water limitations. Instead, growth responses to elevated [CO₂] and [O₃] interacted strongly with present‐day interannual variability in climatic conditions. The amount of photosynthetically active radiation and temperature during specific times of the year coinciding with growth phenology explained 20–63% of the annual variation in growth response to elevated [CO₂] and [O₃]. Years with higher photosynthetic photon flux (PPF) during the month of July resulted in more positive growth responses to elevated [CO₂] and more negative growth responses to elevated [O₃]. Mean daily temperatures during the month of October affected growth in a similar fashion the following year. These results indicate that a several‐year trend of increasingly cloudy summers and cool autumns were responsible for the decrease in CO₂ growth response.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Elevated CO2 influences the responses of two birch species to soil moisture: implications for forest community structure

Increased levels of atmospheric CO₂ may alter the structure and composition of plant communities by affecting how species respond to their physical and biological environment. We investigated how elevated CO₂ influenced the response of paper birch ( Betula papyrifera Marsh.) and yellow birch (Betula alleghaniensis Britt.) seedlings to variation in soil moisture. Seedlings were grown for four months on a soil moisture gradient, individually and in mixed species stands, in controlled environment facilities at ambient (375 μL L^–1) and elevated (700 μL L^–1) atmospheric CO₂. For both individually and competitively grown paper birch seedlings, there was a greater CO₂ growth enhancement for seedlings watered less frequently than for well-watered seedlings. This differential change in CO₂ responsiveness across the moisture gradient reduced the difference in seedling growth between high and low water levels and effectively broadened the regeneration niche of paper birch. In contrast, for yellow birch seedlings, elevated CO₂ only produced a significant growth enhancement at the wet end of the soil moisture gradient, and increased the size difference between seedlings at the two ends of the gradient. Gas exchange measurements showed that paper birch seedlings were more sensitive than yellow birch seedlings to declines in soil moisture, and that elevated CO₂ reduced this sensitivity. Additionally, elevated CO₂ improved survival of yellow birch seedlings growing in competition with paper birch in dry stands. Thus, elevated CO₂ may influence regeneration patterns of paper birch and yellow birch on sites of differing soil moisture. In the future, as atmospheric CO₂ levels rise, growth of paper birch seedlings and survival of yellow birch seedlings may be enhanced on xeric sites, while yellow birch may show improved growth on mesic sites.     

Concentrations of CO2 and O2 in floodwater and in internodal lacunae of floating rice growing at 1–2 metre water depths

Abstract. Environment and plant measurements were made to determine what factors may limit growth of deepwater and floating rice plants during partial or complete submergence. Field surveys included measurements of temperature, pH, light, O₂ and CO₂ in floodwater in Thailand. In addition, measurements were made of O₂ and CO₂ concentrations inside internodal lacunae of deepwater and floating rice growing at 0.5–2.0 m water depths. The bulk of measurements were taken during periods when the changes in water level were less than 50 mm d^?1. In the 0–0.02 m surface layer of floodwater at any location there were large changes in oxygen concentrations over diurnal cycles: there were decreases during the night down to 0.02–0.18 mol m^?3 O₂ at 0600 h and increases during the day to 0.13–0.28 mol m^?3 O₂ at 1500 h (0.28 mol m^?3 being 120% of the O₂ concentration of air saturated water at 30°C). During the day oxygen concentrations decreased with increasing water depth; concentrations just above the soil surface were occasionally zero. Most of this gradient disappeared during the night, and at dawn the 0.6 m surface layer of water had uniform low O₂ concentrations. O₂ concentrations were also measured during flash floods in Thailand. In contrast to the conditions with only small increases in water level, the O₂ concentrations in the water during flash floods were more uniform with depth and changed little over a diurnal cycle, the O₂ ranging between 0.14–0.19 mol m^?3. In most locations floodwater contained 0.2–1.9 mol m^?3 CO₂ and 0.7–1.6 mol m^?3 bicarbonate; however, in a location with acid sulphate soil CO₂ was only 0.05–0.2 mol m^?3, and bicarbonate concentrations were several fold lower. Concentrations of CO₂ in floodwater increased with increasing water depth. O₂ and CO₂ concentrations inside internodal lacunae of rice were determined in the field when water depth were 1–2 m. Concentrations of O₂ in internodes at the water surface were 16–20%, and decreased to 10% and 5% at 0.8 and 1.8 m water depth respectively. There was no diurnal cycle in O₂ concentrations inside internodes. In contrast, CO₂ concentrations in the lacunae increased with water depth and ranged from 1–3% in internodes at the water surface to 5–10% in internodes at 1.8 m water depth. There was evidence for a diurnal cycle in CO₂ concentrations in the basal internode near the soil surface, CO₂ increased during the day and decreased during the night. The above data are used to show that there is little or no relationship between gas concentrations in floodwater and internodal lacunae of rice plants. Results are discussed in relation to O₂ supply to submerged portions of rice and metabolism of these tissues at low O₂ concentrations.     

Fungal community composition and function after long‐term exposure of northern forests to elevated atmospheric CO2 and tropospheric O3

The long‐term effects of rising atmospheric carbon dioxide (CO₂) and tropospheric O₃ concentrations on fungal communities in soil are not well understood. Here, we examine fungal community composition and the activities of cellobiohydrolase and N‐acetylglucosaminidase (NAG) after 10 years of exposure to 1.5 times ambient levels of CO₂ and O₃ in aspen and aspen–birch forest ecosystems, and compare these results to earlier studies in the same long‐term experiment. The forest floor community was dominated by saprotrophic fungi, and differed slightly between plant community types, as did NAG activity. Elevated CO₂ and O₃ had small but significant effects on the distribution of fungal genotypes in this horizon, and elevated CO₂ also lead to an increase in the proportion of Sistotrema spp. within the community. Yet, although cellobiohydrolase activity was lower in the forest floor under elevated O₃, it was not affected by elevated CO₂. NAG was also unaffected. The soil community was dominated by ectomycorrhizal species. Both CO₂ and O₃ had a minor effect on the distribution of genotypes; however, phylogenetic analysis indicated that under elevated O₃Cortinarius and Inocybe spp. increased in abundance and Laccaria and Tomentella spp. declined. Although cellobiohydrolase activity in soil was unaffected by either CO₂ or O₃, NAG was higher (～29%) under CO₂ in aspen–birch, but lower (～18%) under aspen. Time series analysis indicated that CO₂ increased cellulolytic enzyme activity during the first 5 years of the experiment, but that the magnitude of this effect diminished over time. NAG activity also showed strong early stimulation by elevated CO₂, but after 10 years this effect is no longer evident. Elevated O₃ appears to have variable stimulatory and repressive effects depending on the soil horizon and time point examined.     

Immobilization, stabilization and remobilization of nitrogen in forest soils at elevated CO2: a 15N and 13C tracer study

The fate of immobilized N in soils is one of the great uncertainties in predicting C sequestration at increased CO₂ and N deposition. In a dual isotope tracer experiment (¹³C, ¹⁵N) within a 4‐year CO₂ enrichment (+200 ppm_v) study with forest model ecosystems, we (i) quantified the effects of elevated CO₂ on the partitioning of N; (ii) traced immobilized N into physically separated pools of soil organic matter (SOM) with turnover rates known from their ¹³C signals; and (iii) estimated the remobilization and thus, the bio‐availability of newly sequestered C and N. (1) CO₂ enrichment significantly decreased NO₃^? concentrations in soil waters and export from 1.5 m deep lysimeters by 30–80%. Consequently, elevated CO₂ increased the overall retention of N in the model ecosystems. (2) About 60–80% of added ¹⁵NH₄¹⁵NO₃ were retained in soils. The clay fraction was the greatest sink for the immobilized ¹⁵N sequestering 50–60% of the total new soil N. SOM associated with clay contained only 25% of the total new soil C pool and had small C/N ratios (<13), indicating that it consists of humified organic matter with a relatively slow turn over rate. This implies that added ¹⁵N was mainly immobilized in stable mineral‐bound SOM pools. (3) Incubation of soils for 1 year showed that the remobilization of newly sequestered N was three to nine times smaller than that of newly sequestered C. Thus, inorganic inputs of N were stabilized more effectively in soils than C. Significantly less newly sequestered N was remobilized from soils previously exposed to elevated CO₂. In summary, our results show firstly that a large fraction of inorganic N inputs becomes effectively immobilized in relative stable SOM pools and secondly that elevated CO₂ can increase N retention in soils and hence it may tighten N cycling and diminish the risk of nitrate leaching to groundwater.     

Wood properties of two silver birch clones exposed to elevated CO2 and O3

Effects of elevated carbon dioxide (CO₂) and ozone (O₃) on wood properties of two initially 7‐year‐old silver birch (Betula pendula Roth) clones were studied after a fumigation during three growing seasons. Forty trees, representing two fast‐growing clones (4 and 80), were exposed in open‐top chambers to the following treatments: outside control, chamber control, 2 × ambient [CO₂], 2 × ambient [O₃] and 2 × ambient [CO₂]+2 × ambient [O₃]. After the 3‐year exposure, the trees were felled and wood properties were analyzed. The treatments affected both stem wood structure and chemistry. Elevated [CO₂] increased annual ring width, and concentrations of extractives and starch, and decreased concentrations of cellulose and gravimetric lignin. Elevated O₃ decreased vessel percentage and increased cell wall percentage in clone 80. In vessel percentage, elevated CO₂ ameliorated the O₃‐induced decrease. In clone 4, elevated O₃ decreased nitrogen concentration of wood. The two clones had different wood properties. In clone 4, the concentrations of extractives, starch, soluble sugars and nitrogen were greater than in clone 80, while in clone 80 the concentrations of cellulose and acid‐soluble lignin were higher. Clone 4 also had slightly longer fibres, greater vessel lumen diameter and vessel percentage than clone 80, while in clone 80 cell wall percentage was greater. Our results show that wood properties of young silver birch trees were altered under elevated CO₂ in both clones, whereas the effects of O₃ depended on clone.     

Nitrogen cycling in microcosms of yellow birch exposed to elevated CO2: simultaneous positive and negative below-ground feedbacks

This study investigated simultaneous plant and soil feedbacks on growth enhancement with elevated [CO₂] within microcosms of yellow birch (Betula alleghaniensis Britt.) in the second year of growth. Understanding the integrated responses of model ecosystems may provide key insight into the potential net nutrient feedbacks on [CO₂] growth enhancements in temperate forests. We measured the net biomass production, C:N ratios, root architecture, and mycorrhizal responses of yellow birch, in situ rates gross nitrogen mineralization and the partitioning of available NH₄⁺ between yellow birch and soil microbes. Elevated atmospheric [CO₂] resulted in significant alterations in the cycling of N within the microcosms. Plant C/N ratios were significantly increased, gross mineralization and NH₄⁺ consumption rates were decreased, and relative microbial uptake of NH₄⁺ was increased, representing a suite of N cycling negative feedbacks on N availability. However, increased C/N ratios may also be a mechanism which allows plants to maintain higher growth with a constant or reduced N supply. Total plant N content was increased with elevated [CO₂], suggesting that yellow birch had successfully increased their ability to acquire nutrients during the first year of growth. However, plant uptake rates of NH₄⁺ had decreased in the second year. This discrepancy implies that, in this study, nitrogen uptake showed a trend through ontogeny of decreasing enhancement under elevated [CO₂]. The reduced N mineralization and relatively increased N immobilization are a potential feedback which may drive this ontogenetic trend. This study has demonstrated the importance of using an integrated approach to exploring potential nutrient-cycling feedbacks in elevated [CO₂].     

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

Abstract

• 1 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.).
• 2 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.
• 3 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.
• 4 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.
• 5 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.