System-level adjustments to elevated CO2 in model spruce ecosystems

Atmospheric carbon dioxide enrichment and increasing nitrogen deposition are often predicted to increase forest productivity based on currently available data for isolated forest tree seedlings or their leaves. However, it is highly uncertain whether such seedling responses will scale to the stand level. Therefore, we studied the effects of increasing CO₂ (280, 420 and 560 μL L^-1) and increasing rates of wet N deposition (0, 30 and 90 kg ha^-1 y^-1) on whole stands of 4-year-old spruce trees (Picea abies). One tree from each of six clones, together with two herbaceous understory species, were established in each of nine 0.7 m² model ecosystems in nutrient poor forest soil and grown in a simulated montane climate for two years. Shoot level light-saturated net photosynthesis measured at growth CO₂ concentrations increased with increasing CO₂, as well as with increasing N deposition. However, predawn shoot respiration was unaffected by treatments. When measured at a common CO₂ concentration of 420 μL L^-1 37% down-regulation of photosynthesis was observed in plants grown at 560 μL CO₂ L^-1. Length growth of shoots and stem diameter were not affected by CO₂ or N deposition. Bud burst was delayed, leaf area index (LAI) was lower, needle litter fall increased and soil CO₂ efflux increased with increasing CO₂. N deposition had no effect on these traits. At the ecosystem level the rate of net CO₂ exchange was not significantly different between CO₂ and N treatments. Most of the responses to CO₂ studied here were nonlinear with the most significant differences between 280 and 420 μL CO₂ L^-1 and relatively small changes between 420 and 560 μL CO₂ L^-1. Our results suggest that the lack of above-ground growth responses to elevated CO₂ is due to the combined effects of physiological down-regulation of photosynthesis at the leaf level, allometric adjustment at the canopy level (reduced LAI), and increasing strength of below-ground carbon sinks. The non-linearity of treatment effects further suggests that major responses of coniferous forests to atmospheric CO₂ enrichment might already be under way and that future responses may be comparatively smaller.     

Variations of ecosystem gas exchange in the rain forest mesocosm at Biosphere 2 in response to elevated CO2

The effects of elevated CO₂ on tropical ecosystems were studied in the artificial rain forest mesocosm at Biosphere 2, a large-scale and ecologically diverse experimental facility located in Oracle, Arizona. The ecosystem responses were assessed by comparing the whole-system net gas exchange (NEE) upon changing CO₂ levels from 900 to 450 ppmV. The day-NEE was significantly higher in the elevated CO₂ treatment. In both experiments, the NEE rates were similar to values observed in natural analogue systems. Variations in night-NEE, reflecting both soil CO₂ efflux and plants respiration, covaried with temperature but showed no clear correlation with atmospheric CO₂ levels. After correcting for changes in CO₂ efflux we show that the rain forest net photosynthesis increased in response to increasing atmospheric CO₂. The photosynthetic enhancement was expressed in higher quantum yields, maximum assimilation rates and radiation use efficiency. The results suggest that photosynthesis in large tropical trees is CO₂ sensitive, at least following short exposures of days to weeks. Taken at face value, the data suggest that as a result of anthropogenic emissions of CO₂, tropical rain forests may shift out of steady state, and become a carbon sink at least for short periods. However, a better understanding of the unique conditions and phenomena in Biosphere 2 is necessary before these results are broadly useful.     

Scaling up evolutionary responses to elevated CO2: lessons from Arabidopsis

Results from norm of reaction studies and selection experiments indicate that elevated CO₂ will act as a selective agent on natural plant populations, especially for C₃ species that are most sensitive to changes in atmospheric CO₂ concentration. Evolutionary responses to CO₂ may alter plant physiology, development rate, growth, and reproduction in ways that cannot be predicted from single generation studies. Moreover, ecological and evolutionary changes in plant communities will have a range of consequences at higher spatial scales and may cause substantial deviations from ecosystem level predictions based on short‐term responses to elevated CO₂. Therefore, steps need to be taken to identify the plant traits that are most likely to evolve at elevated CO₂, and to understand how these changes may affect net primary productivity within ecosystems. These processes may range in scale from molecular and physiological changes that occur among genotypes at the individual and population levels, to changes in community‐ and ecosystem‐level productivity that result from the integrative effects of different plant species evolving simultaneously. In this review, we (1) synthesize recent studies investigating the role of atmospheric CO₂ as a selective agent on plants, (2) discuss possible control points during plant development that may change in response to selection at elevated CO₂ with an emphasis at the primary molecular level, and (3) provide a quantitative framework for scaling the evolutionary effects of CO₂ on plants in order to determine changes in community and ecosystem productivity. Furthermore, this review points out that studies integrating the effects of plant evolution in response to elevated CO₂ are lacking, and therefore more attention needs be devoted to this issue among the global change research community.     

Carbon balance of young birch trees grown in ambient and elevated atmospheric CO2 concentrations

We constructed a carbon budget for young birch trees grown in ambient and elevated CO₂ concentrations over their fourth year of growth. The annual total of net leaf photosynthesis was 110% more in elevated CO₂ than in ambient CO₂. However, the trees in elevated CO₂ grew only 59% more biomass than the trees in ambient CO₂ over the year. Modelling studies showed that larger loss of carbon from fine-root production and growth of the root-associated mycorrhiza by the trees in elevated CO₂ probably accounted for all the remaining difference in net photosynthesis between the two treatments. Our modelling also showed that the fraction of net photosynthate consumed by respiration of nonleaf tissue was similar in the two CO₂ treatments, and was 26% and 24% for trees in ambient and elevated CO₂, respectively. Trees in elevated CO₂ had 43% more leaves, and produced 110% more net photosynthate than trees in ambient CO₂, even though the maximum rate of carboxylation per unit leaf nitrogen decreased by 21%. Sensitivity studies showed that down-regulation reduced the annual net photosynthetic production of the trees in elevated CO₂ by only 6%. Direct effects of higher CO₂ on photosynthesis and greater leaf area of the trees in elevated CO₂ increased the net photosynthesis of the trees by 68% and 60%, respectively; and together accounted for most of the difference in net photosynthesis between the two treatments.     

Atmospheric CO2 enrichment increases growth and photosynthesis of Pinus taeda: a 4 year experiment in the field

Forest trees are major components of the terrestrial biome and their response to rising atmospheric CO₂ plays a prominent role in the global carbon cycle. In this study, loblolly pine seedlings were planted in the field in recently disturbed soil of high fertility, and CO₂ partial pressures were maintained at ambient CO₂ (Amb) and elevated CO₂ (Amb + 30 Pa) for 4 years. The objective of the study was to measure seasonal and long-term responses in growth and photosynthesis of loblolly pine exposed to elevated CO₂ under ambient field conditions of precipitation, light, temperature and nutrient availability. Loblolly pine trees grown in elevated CO₂ produced 90% more biomass after four growing seasons than did trees grown in ambient CO₂. This large increase in final biomass was primarily due to a 217% increase in leaf area in the first growing season which resulted in much higher relative growth rates for trees grown in elevated CO₂. Although there was not a sustained effect of elevated CO₂ on relative growth rate after the first growing season, absolute production of biomass continued to increase each year in trees grown in elevated CO₂ as a consequence of the compound interest effect of increased leaf area on the production of more new leaf area and more biomass. Allometric analyses of biomass allocation patterns demonstrated size-dependent shifts in allocation, but no direct effects of elevated CO₂ on partitioning of biomass. Leaf photosynthetic rates were always higher in trees grown in elevated CO₂, but these differences were greater in the summer (60–130% increase) than in the winter (14–44% increase), reflecting strong seasonal effects of temperature on photosynthesis. Our results suggest that seasonal variation in the relative photosynthetic response to elevated CO₂ will occur in natural ecosystems, but total non-structural carbohydrate (TNC) levels in leaves indicate that this variation may not always be related to sink activity. Despite indications of canopy-level adjustments in carbon assimilation, enhanced levels of leaf photosynthesis coupled with increased total leaf area indicate that net carbon assimilation for the whole tree was greater for trees grown under elevated CO₂ compared with ambient CO₂. If the large growth enhancement observed in loblolly pine were maintained after canopy closure, then these trees could be a large sink for fossil carbon emitted to the atmosphere and produce a negative feedback on atmospheric CO₂.     

高浓度CO2对树木生理生态的影响研究进展

大气CO2浓度升高已成为世界范围内的重要环境问题。CO2浓度升高势必会对植物的生理生态变化产生重要影响。综述了国内外有关高浓度CO2对树木生理生态影响研究的最新进展,具体包括高浓度CO2对树木生长发育、光合和呼吸作用、抗氧化系统、树木代谢物质、挥发性有机化合物以及树木凋落物等方面的影响。高浓度CO2一般会促进树木地上植株的生长和发育,但也因树种差异而有所不同。最新研究表明,高浓度CO2促进了树木细根周转,树木根系生长在大气CO2浓度升高条件下表现为促进作用,这种作用加快了全球森林生态系统的C循环。高浓度CO2虽然在一定程度上促进树木光合速率的增加,但长期熏蒸也往往会发生光合驯化,这种现象产生的生理学机制目前仍无定论。高浓度CO2对树木呼吸作用尤其是根系呼吸的影响将是未来研究的重点和难点。高浓度CO2一般会提高树木抗氧化酶活性与抗氧化剂含量,但不同树种响应高浓度CO2的过程和机理也有所差异。研究表明,高浓度CO2一般对树木凋落物的分解产生不利影响,但也因树种而异。需要强调的是,目前关于树木地下部分、树木对高浓度CO2的适应机理和重要过程(碳氮水耦合及基因调控等)以及多个树种包括不同类型树种及不同品种之间比较研究较少;关于某一重要生理生态机制(如根系生理代谢)尤其是多个生态因子复合条件下缺乏长期深入的研究。在此基础上给出了大气CO2浓度升高下树木生理生态学研究的未来发展方向,包括高CO2浓度条件下树木根系生理代谢及机制、树木碳氮水耦合的生理过程及机制、不同生态因子复合作用对树木生理影响机制以及树木分子作用机理等方面的研究。这些研究不仅将丰富森林树木应对未来气候变化的有关科学理论,也为全球气候变化背景下实现森林树种生态功能的优化选择及森林生态系统的可持续发展与经营提供重要的生理生态学理论依据和参考。     

Increased leaf reflectance in tropical trees under elevated CO2

Globally increasing atmospheric CO₂ concentrations are known to affect many aspects of plant physiology and development; however, little attention has been given to leaf and canopy optical properties. Three tropical trees in the Leguminosae, an important canopy tree family in many tropical forests, responded similarly to an experimental doubling of CO₂ partial pressure with a 9–23% increase in spectral leaf reflectance to light in the visible (400–700 nm) waveband. Decreased leaf chlorophyll content under elevated CO₂ may explain part of the observed increase in reflectance. However, analyses that statistically corrected for chlorophyll content effects on reflectance still indicated a significant CO₂ effect. This results, in conjunction with the spectral pattern of the response, suggests that the primary mechanism is increased optical masking of chlorophyll under elevated CO₂. The magnitude of the increase in leaf reflectance is sufficient to suggest that increased canopy reflectance of tropical forests (and possibly other terrestrial ecosystems) may be an important negative feedback in the response of global net radiative climate forcing to increasing atmospheric CO₂.     

Thirty years of in situ tree growth under elevated CO2: a model for future forest responses?

Rising concentrations of atmospheric carbon dioxide have been predicted to stimulate the growth of forest trees. However, long-term effects on trees growing to maturity and to canopy closure while exposed to elevated CO₂ have never been examined. We compared tree ring chronologies of Mediterranean Quercus ilex which have been continuously exposed to elevated CO₂ (around 650 μmol mol^–1) since they were seedlings, near two separate natural CO₂ springs with those from trees at nearby ambient-CO₂‘control’ sites. Trees grown under high CO₂ for 30 years (1964–93) showed a 12% greater final radial stem width than those growing at the ambient-CO₂ control sites. However, this stimulation was largely due to responses when trees were young. By the time trees were 25–30 y old the annual difference in tree ring width between low and high CO₂ grown trees had disappeared. At any given tree age, elevated CO₂ had a relatively greater positive effect on tree ring width in years with a dry spring compared to years with more rainfall between April and May. This indicates a beneficial effect of elevated CO₂ on tree water relations under drought stress. Our data suggest that the early regeneration phase of forest stands can be accelerated in CO₂-enriched atmospheres and that maximum biomass per land area may be reached sooner than under lower CO₂ concentrations. In our study, high CO₂ grown Q. ilex trees reached the same stem basal area at the age of 26 y as control trees at 29 y, i.e. three years earlier (faster turnover of carbon?). Reliable predictions of the future development of forests need to account for the variable responses of trees over their entire lifetime. Such responses to elevated CO₂ can presently only be assessed at such unique field sites.     

Tree responses to rising CO2 in field experiments: implications for the future forest

The need to assess the role of forests in the global cycling of carbon and how that role will change as the atmospheric concentration of CO₂ increases has spawned many experiments over a range of scales. Experiments using open-top chambers have been established at many sites to test whether the short-term responses of tree seedlings described in controlled environments would be sustained over several growing seasons under field conditions. Here we review the results of those experiments, using the framework of the interacting cycles of carbon, water and nutrients, because that is the framework of the ecosystem models that are being used to address the decades-long response of forests. Our analysis suggests that most of what was learned in seedling studies was qualitatively correct. The evidence from field-grown trees suggests a continued and consistent stimulation of photosynthesis of about 60% for a 300 p.p.m. increase in [CO₂], and there is little evidence of the long-term loss of sensitivity to CO₂ that was suggested by earlier experiments with tree seedlings in pots. Despite the importance of respiration to a tree's carbon budget, no strong scientific consensus has yet emerged concerning the potential direct or acclimation response of woody plant respiration to CO₂ enrichment. The relative effect of CO₂ on above-ground dry mass was highly variable and greater than that indicated by most syntheses of seedling studies. Effects of CO₂ concentration on static measures of response are confounded with the acceleration of ontogeny observed in elevated CO₂. The trees in these open-top chamber experiments were in an exponential growth phase, and the large growth responses to elevated CO₂ resulted from the compound interest associated with an increasing leaf area. This effect cannot be expected to persist in a closed-canopy forest where growth potential is constrained by a steady-state leaf area index. A more robust and informative measure of tree growth in these experiments is the annual increment in wood mass per unit leaf area, which increased 27% in elevated CO₂. There is no support for the conclusion from many studies of seedlings that root-to-shoot ratio is increased by elevated CO₂; the production of fine roots may be enhanced, but it is not clear that this response would persist in a forest. Foliar nitrogen concentrations were lower in CO₂-enriched trees, but to a lesser extent than was indicated in seedling studies and only when expressed on a leaf mass basis. The prediction that leaf litter C/N ratio would increase was not supported in field experiments. Also contrasting with seedling studies, there is little evidence from the field studies that stomatal conductance is consistently affected by CO₂; however, this is a topic that demands more study. Experiments with trees in open-top chambers under field conditions have provided data on longer-term, larger-scale responses of trees to elevated CO₂ under field conditions, confirmed some of the conclusions from previous seedling studies, and challenged other conclusions. There remain important obstacles to using these experimental results to predict forest responses to rising CO₂, but the studies are valuable nonetheless for guiding ecosystem model development and revealing the critical questions that must be addressed in new, larger-scale CO₂ experiments.     

Elevated CO2 and tree fecundity: the role of tree size, interannual variability, and population heterogeneity

Long‐term population effects of changes in atmospheric CO₂ will be largely determined by reproductive effort. Our research objectives were to quantify variability in seed production and rate of maturation among individual Pinus taeda L. (Pinaceae) trees growing in elevated CO₂ (ambient plus 200 μL L^?1) since 1996. Estimating tree fecundity in nature is frustrated by the difficulty of counting seeds from individual trees and the need for long‐term data. We have used a hierarchical Bayes approach to model individual tree fecundity, accounting for the complexity of experimentation in a natural setting over multiple years. The study presented here demonstrates large variability in natural fecundity rates and contributes to our understanding of how both interannual variation and population heterogeneity influence elevated CO₂ effects. We found that trees growing under elevated CO₂ matured earlier and produced more seeds and cones per unit basal area than ambient grown trees. By 2004, trees grown in high CO₂ had produced an average 300 more seeds per tree than ambient grown trees. Although there was a trend toward decreasing mean CO₂ effect (difference in fecundity between elevated and ambient treatments) over time, the hierarchical analysis indicates that this decrease comes from the emergence of a few highly fecund ambient grown trees by 2002, rather than acclimation or downregulation among the fumigated trees. The most important effect of increased CO₂ in forest ecosystems may be the increase in fecundity reported here. Although biomass responses can sometimes be large, the increase in fecundity can have long‐term impacts on forest dynamics that transcend the current generation.     

Responses of soil biota to elevated atmospheric carbon dioxide

Increasing concentrations of atmospheric CO₂ could have dramatic effects upon terrestrial ecosystems including changes in ecosystem structure, nutrient cycling rates, net primary production, C source-sink relationships and successional patterns. All of these potential changes will be constrained to some degree by below ground processes and mediated by responses of soil biota to indirect effects of CO₂ enrichment. A review of our current state of knowledge regarding responses of soil biota is presented, covering responses of mycorrhizae, N-fixing bacteria and actinomycetes, soil microbiota, plant pathogens, and soil fauna. Emphasis will be placed on consequences to biota of increasing C input through the rhizosphere and resulting feedbacks to above ground systems. Rising CO₂ may also result in altered nutrient concentrations of plant litter, potentially changing decomposition rates through indirect effects upon decomposer communities. Thus, this review will also cover current information on decomposition of litter produced at elevated CO₂. Summary Predictably, the responses of soil biota to CO₂ enrichment and the degree of experimental emphasis on them increase with proximity to, and intimacy with, roots. Symbiotic associations are all stimulated to some degree. Total plant mycorrhization increases with elevated CO₂. VAM fungi increase proportionately with fine root length/mass increase. ECM fungi, however, exhibit greater colonization per unit root length/mass at elevated CO₂ than at current atmospheric levels. Total N-fixation per plant increases in all species examined, although the mechanisms of increase, as well as the eventual benefit to the host relative to N uptake may vary. Microbial responses are unclear. The assumption that changes in root exudation will drive increased mineralization and facilitate nutrient uptake should be examined experimentally, in light of recent models. Microbial results to date suggest that metabolic activity (measured as changes in process rates) is stimulated by root C input, rather than population size (measured by cell or colony counts). Insufficient evidence exists to predict responses of either soil-borne plant pathogens or soil fauna (i.e., food web responses). These are areas requiring attention, the first for its potential to limit ecosystem production through disease and the second because of its importance to nutrient cycling processes. Preliminary data on foliar litter decomposition suggests that neither nutrient ratios nor decomposition rates will be affected by rising CO₂. This is another important area that may be better understood as the number of longer term studies with more realistic CO₂ exposures increase. Evidence continues to mount that C fixation increases with CO₂ enrichment and that the bulk of this C enters the belowground component of ecosystems. The global fate and effects of this additional C may affect all hierarchical levels, from organisms to ecosystems, and will be largely determined by responses of soil biota.     

A branch bag technique for simultaneous CO2 enrichment and assimilation measurements on beech (Fagus sylvatica L.)

A cheap CO₂ enrichment system was designed to perform continuous gas exchange measurements of branches of mature European beech trees (Fagus sylvatica L.). Branches were grown at ambient (350 cm³ m^-3) and elevated CO₂ (700cm³ m^-3) during the whole 1992 leafy period. Leaks resulting from airtightness defaults in the system appeared to be low enough to measure accurately net CO₂ assimilation and transpiration rates during the day. However, the CO₂ exchange rates during the night (respiration) were too low to allow accurate measurements. Elevated CO₂ had a great effect on the net assimilation rate of branches via its influence on both the C₃ photosynthetic pathway and the shade-tolerance of beech trees (85% increase). The A/Ca curves showed no acclimation effect to high CO₂, both control and enriched branches increasing their net assimilation in the same way. The decrease of net assimilation rates in mature leaves was similar for both control and enriched branches. The pattern of daily transpiration rates remained the same for both control and enriched branches, hence we can assume that there was no visible CO₂ effect on stomata.     

Changes in net photosynthesis and growth of Pinus eldarica seedlings in response to atmospheric CO2 enrichment

Pinus eldarica L. trees, rooted in the natural soil of an agricultural field at Phoenix, Arizona, were grown from the seedling stage in clear-plastic-wall open-top enclosures maintained at four different atmospheric CO₂ concentrations for 15 months. Light response functions were determined for one tree from each treatment by means of whole-tree net CO₂ exchange measurements at the end of this period, after which rates of carbon assimilation of an ambient-treatment tree were measured across a range of atmospheric CO₂ concentrations. The first of these data sets incorporates the consequences of both the CO₂-induced enhancement of net photosynthesis per unit needle area and the CO₂-induced enhancement of needle area itself (due primarily to the production of more needles), whereas the second data set reflects only the first of these effects. Hence the division of the normalized results of the first data set by the normalized results of the second set yields a representation of the increase in whole-tree net photosynthesis due to enhanced needle production caused by atmospheric CO₂ enrichment. In the solitary trees we studied, the relative contribution of this effect increased rapidly with the CO₂ concentration of the air to increase whole-tree net photosynthesis by nearly 50% at a CO₂ concentration approximately 300 μmol mol⁻¹ above ambient.     

Does the growth response of woody plants to elevated CO2 increase with temperature? A model‐oriented meta‐analysis

The temperature dependence of the reaction kinetics of the Rubisco enzyme implies that, at the level of a chloroplast, the response of photosynthesis to rising atmospheric CO₂ concentration (C_a) will increase with increasing air temperature. Vegetation models incorporating this interaction predict that the response of net primary productivity (NPP) to elevated CO₂ (eC_a) will increase with rising temperature and will be substantially larger in warm tropical forests than in cold boreal forests. We tested these model predictions against evidence from eC_a experiments by carrying out two meta‐analyses. Firstly, we tested for an interaction effect on growth responses in factorial eC_a × temperature experiments. This analysis showed a positive, but nonsignificant interaction effect (95% CI for above‐ground biomass response = ?0.8, 18.0%) between eC_a and temperature. Secondly, we tested field‐based eC_a experiments on woody plants across the globe for a relationship between the eC_a effect on plant biomass and mean annual temperature (MAT). This second analysis showed a positive but nonsignificant correlation between the eC_a response and MAT. The magnitude of the interactions between CO₂ and temperature found in both meta‐analyses were consistent with model predictions, even though both analyses gave nonsignificant results. Thus, we conclude that it is not possible to distinguish between the competing hypotheses of no interaction vs. an interaction based on Rubisco kinetics from the available experimental database. Experiments in a wider range of temperature zones are required. Until such experimental data are available, model predictions should aim to incorporate uncertainty about this interaction.     

Ocean acidification and rising temperatures may increase biofilm primary productivity but decrease grazer consumption

Climate change may cause ecosystems to become trophically restructured as a result of primary producers and consumers responding differently to increasing CO₂ and temperature. This study used an integrative approach using a controlled microcosm experiment to investigate the combined effects of CO₂ and temperature on key components of the intertidal system in the UK, biofilms and their consumers (Littorina littorea). In addition, to identify whether pre-exposure to experimental conditions can alter experimental outcomes we explicitly tested for differential effects on L. littorea pre-exposed to experimental conditions for two weeks and five months. In contrast to predictions based on metabolic theory, the combination of elevated temperature and CO₂ over a five-week period caused a decrease in the amount of primary productivity consumed by grazers, while the abundance of biofilms increased. However, long-term pre-exposure to experimental conditions (five months) altered this effect, with grazing rates in these animals being greater than in animals exposed only for two weeks. We suggest that the structure of future ecosystems may not be predictable using short-term laboratory experiments alone owing to potentially confounding effects of exposure time and effects of being held in an artificial environment over prolonged time periods. A combination of laboratory (physiology responses) and large, long-term experiments (ecosystem responses) may therefore be necessary to adequately predict the complex and interactive effects of climate change as organisms may acclimate to conditions over the longer term.     

Leaf gas exchange and nitrogen dynamics of N2-fixing, field-grown Alnus glutinosa under elevated atmospheric CO2

Few studies have investigated the effects of elevated CO₂ on the physiology of symbiotic N₂-fixing trees. Tree species grown in low N soils at elevated CO₂ generally show a decline in photosynthetic capacity over time relative to ambient CO₂ controls. This negative adjustment may be due to a reallocation of leaf N away from the photosynthetic apparatus, allowing for more efficient use of limiting N. We investigated the effect of twice ambient CO₂ on net CO₂ assimilation (A), photosynthetic capacity, leaf dark respiration, and leaf N content of N2-fixing Alnus glutinosa (black alder) grown in field open top chambers in a low N soil for 160 d. At growth CO₂, A was always greater in elevated compared to ambient CO₂ plants. Late season A vs. internal leaf p(CO₂) response curves indicated no negative adjustment of photosynthesis in elevated CO₂ plants. Rather, elevated CO₂ plants had 16% greater maximum rate of CO₂ fixation by Rubisco. Leaf dark respiration was greater at elevated CO₂ on an area basis, but unaffected by CO₂ on a mass or N basis. In elevated CO₂ plants, leaf N content (μg N cm^?2) increased 50% between Julian Date 208 and 264. Leaf N content showed little seasonal change in ambient CO₂ plants. A single point acetylene reduction assay of detached, nodulated root segments indicated a 46% increase in specific nitrogenase activity in elevated compared to ambient CO₂ plants. Our results suggest that N₂-fixing trees will be able to maintain high A with minimal negative adjustment of photosynthetic capacity following prolonged exposure to elevated CO₂ on N-poor soils.     

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.     

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.     

Photosynthetic capacity of loblolly pine (Pinus taeda L.) trees during the first year of carbon dioxide enrichment in a forest ecosystem

Our objective was to assess the photosynthetic responses of loblolly pine trees (Pinus taeda L.) during the first full growth season (1997) at the Brookhaven National Lab/Duke University Free Air CO₂ Enrichment (FACE) experiment. Gas exchange, fluorescence characteristics, and leaf biochemistry of ambient CO₂ (control) needles and ambient + 20 Pa CO₂ (elevated) needles were examined five times during the year. The enhancement of photosynthesis by elevated CO₂ in mature loblolly pine trees varied across the season and was influenced by abiotic and biotic factors. Photosynthetic enhancement by elevated CO₂ was strongly correlated with leaf temperature. The magnitude of photosynthetic enhancement was zero in March but was as great as 52% later in the season. In March, reduced sink demand and lower temperatures resulted in lower net photosynthesis, lower carboxylation rates and higher excess energy dissipation from the elevated CO₂ needles than from control needles. The greatest photosynthetic enhancement by CO₂ enrichment was observed in July during a period of high temperature and low precipitation, and in September during recovery from this period of low precipitation. In July, loblolly pine trees in the control rings exhibited lower net photosynthetic rates, lower maximum rates of photosynthesis at saturating CO₂ and light, lower values of carboxylation and electron transport rates (modelled from A–C_i curves), lower total Rubisco activity, and lower photochemical quenching of fluorescence in comparison to other measurement periods. During this period of low precipitation trees in the elevated CO₂ rings exhibited reduced net photosynthesis and photochemical quenching of fluorescence, but there was little effect on light- and CO₂-saturated rates of photosynthesis, modelled rates of carboxylation or electron transport, or Rubisco activity. These first-year data will be used to compare with similar measurements from subsequent years of the FACE experiment in order to determine whether photosynthetic acclimation to CO₂ occurs in these canopy loblolly pine trees growing in a forest ecosystem.