Long-term effects of elevated atmospheric CO2 on below-ground biomass and transformations to soil organic matter in grassland

We determined the effects of elevated [CO₂] on the quantity and quality of below-ground biomass and several soil organic matter pools at the conclusion of an eight-year CO₂ enrichment experiment on native tallgrass prairie. Plots in open-top chambers were exposed continuously to ambient and twice-ambient [CO₂] from early April through late October of each year. Soil was sampled to a depth of 30 cm beneath and next to the crowns of C4 grasses in these plots and in unchambered plots. Elevated [CO₂] increased the standing crops of rhizomes (87%), coarse roots (46%), and fibrous roots (40%) but had no effect on root litter (mostly fine root fragments and sloughed cortex material >500 μm). Soil C and N stocks also increased under elevated [CO₂], with accumulations in the silt/clay fraction over twice that of particulate organic matter (POM; >53 μm). The mostly root-like, light POM (density ≤1.8 Mg m^-3) appeared to turn over more rapidly, while the more amorphous and rendered heavy POM (density >1.8 Mg m^-3) accumulated under elevated [CO₂]. Overall, rhizome and root C:N ratios were not greatly affected by CO₂ enrichment. However, elevated [CO₂] increased the C:N ratios of root litter and POM in the surface 5 cm and induced a small but significant increase in the C:N ratio of the silt/clay fraction to a depth of 15 cm. Our data suggest that 8 years of CO₂ enrichment may have affected elements of the N cycle (including mineralization, immobilization, and asymbiotic fixation) but that any changes in N dynamics were insufficient to prevent significant plant growth responses. This revised version was published online in June 2006 with corrections to the Cover Date.     

Leaf and canopy responses to elevated CO2 in a pine forest under free-air CO2 enrichment

Physiological responses to elevated CO₂ at the leaf and canopy-level were studied in an intact pine (Pinus taeda) forest ecosystem exposed to elevated CO₂ using a free-air CO₂ enrichment (FACE) technique. Normalized canopy water-use of trees exposed to elevated CO₂ over an 8-day exposure period was similar to that of trees exposed to current ambient CO₂ under sunny conditions. During a portion of the exposure period when sky conditions were cloudy, CO₂-exposed trees showed minor (7%) but significant reductions in relative sap flux density compared to trees under ambient CO₂ conditions. Short-term (minutes) direct stomatal responses to elevated CO₂ were also relatively weak (5% reduction in stomatal aperture in response to high CO₂ concentrations). We observed no evidence of adjustment in stomatal conductance in foliage grown under elevated CO₂ for nearly 80 days compared to foliage grown under current ambient CO₂, so intrinsic leaf water-use efficiency at elevated CO₂ was enhanced primarily by direct responses of photosynthesis to CO₂. We did not detect statistical differences in parameters from photosynthetic responses to intercellular CO₂ (A _net-C _i curves) for Pinus taeda foliage grown under elevated CO₂ (550 mol mol^–1) for 50–80 days compared to those for foliage grown under current ambient CO₂ from similar-sized reference trees nearby. In both cases, leaf net photosynthetic rate at 550 mol mol^–1 CO₂ was enhanced by approximately 65% compared to the rate at ambient CO₂ (350 mol mol^–1). A similar level of enhancement under elevated CO₂ was observed for daily photosynthesis under field conditions on a sunny day. While enhancement of photosynthesis by elevated CO₂ during the study period appears to be primarily attributable to direct photosynthetic responses to CO₂ in the pine forest, longer-term CO₂ responses and feedbacks remain to be evaluated.     

Non-litter effects of elevated CO2 on forest floor microarthropod abundances

The arthropod assemblages of the litter and soil play significant roles in decomposition and nutrient cycling. At the FACE (Free-Air CO₂ Enrichment) site at the Duke Forest, we assessed the responses of the litter microarthropod assemblage to elevated CO₂ (200 ppm above ambient) in a loblolly pine plantation. Following the initiation of the elevated CO₂ treatment, a trend toward lower microarthropod abundance under elevated CO₂ emerged. After 18 months, the mean microarthropod abundance was 33% lower in the elevated treatment (P=0.04). The decline was evident in all microarthropod groups, but was significant only in the oribatid mites (P=0.04). Because these responses precede any changes in litter quality resulting from the CO₂ treatment, they may reflect plant-derived changes in the soil that are being conveyed into the litter layer.     

A simple method for testing leaf responses of tall tropical forest trees to elevated CO2

The effects of atmospheric CO₂ enrichment on mature trees in their natural environment are largely unknown. Here we present a new, and inexpensive technique which can be used in situ to address some key physiological questions related to the CO₂ problem. Small, light-weight cups mounted on the lower side of rigid leaves at the top of tall tropical forest trees were supplied with CO₂-enriched air derived from a low-technology air mixing device utilizing forest floor CO₂ evolution. We present the scientific rationale for such field experiments, technical details, an assessment of potential cup artifacts and first results illustrating effects of elevated CO₂ on stomata and carbohydrate accumulation in the canopies of mature trees.     

Effects of elevated CO2 and nitrogen on growth ofPoa pratensis (L.)

The growth responses of a grass,Poa pratensis, to elevated CO₂ and nitrogen were investigated. Light-saturated photosynthetic rate per unit leaf area increased with exposure to elevated CO₂, while dry weight did not respond to increased CO₂. Patterns of biomass allocation within plants, including leaf area, leaf area ratio, specific leaf area, and root to shoot ratios, were not altered by elevated CO₂, but changed considerably with N treatment Shoot and whole-plant tissue N concentrations were significantly diluted by elevated CO₂ (Tukey test, P < 0.05). Total N content did not differ significantly among CO₂ treatments. The absence of a concomitant increase in N uptake under elevated CO₂ may have caused a dilution in plant tissue [N], probably negating the positive effects of increased photosynthesis on biomass accumulation.     

Elevated CO<Subscript>2</Subscript> reduces sap flux in mature deciduous forest trees

We enriched in CO₂ the canopy of 14 broad-leaved trees in a species-rich, ca. 30-m-tall forest in NW Switzerland to test whether elevated CO₂ reduces water use in mature forest trees. Measurements of sap flux density (J_S) were made prior to CO₂ enrichment (summer 2000) and throughout the first whole growing season of CO₂ exposure (2001) using the constant heat-flow technique. The short-term responses of sap flux to brief (1.5–3 h) interruptions of CO₂ enrichment were also examined. There were no significant a priori differences in morphological and physiological traits between trees which were later exposed to elevated CO₂ (n=14) and trees later used as controls (n=19). Over the entire growing season, CO₂ enrichment resulted in an average 10.7% reduction in mean daily J_S across all species compared to control trees. Responses were most pronounced in Carpinus, Acer, Prunus and Tilia, smaller in Quercus and close to zero in Fagus trees. The J_S of treated trees significantly increased by 7% upon transient exposure to ambient CO₂ concentrations at noon. Hence, responses of the different species were, in the short term, similar in magnitude to those observed over the whole season (though opposite because of the reversed treatment). The reductions in mean J_S of CO₂-enriched trees were high (22%) under conditions of low evaporative demand (vapour pressure deficit, VPD <5 hPa) and small (2%) when mean daily VPD was greater than 10 hPa. During a relatively dry period, the effect of elevated CO₂ on J_S even appeared to be reversed. These results suggest that daily water savings by CO₂-enriched trees may have accumulated to a significantly improved water status by the time when control trees were short of soil moisture. Our data indicate that the magnitude of CO₂ effects on stand transpiration will depend on rainfall regimes and the relative abundance of the different species, being more pronounced under humid conditions and in stands dominated by species such as Carpinus and negligible in mono-specific Fagus forests.     

Carbon balance in tussock tundra under ambient and elevated atmospheric CO2

Summary Whole ecosystem CO₂ flux under ambient (340 l/l) and elevated (680 l/l) CO₂ was measured in situ in Eriophorum tussock tundra on the North Slope of Alaska. Elevated CO₂ resulted in greater carbon acquisition than control treatments and there was a net loss of CO₂ under ambient conditions at this upland tundra site. These measurements indicate a current loss of carbon from upland tundra, possibly the result of recent climatic changes. Elevated CO₂ for the duration of one growing season appeared to delay the onset of dormancy and resulted in approximately 10 additional days of positive ecosystem flux. Homeostatic adjustment of ecosystem CO₂ flux (sum of species' response) was apparent by the third week of exposure to elevated CO₂. Ecosystem dark respiration rates were not significantly higher at elevated CO₂ levels. Rapid homeostatic adjustment to elevated CO₂ may limit carbon uptake in upland tundra. Abiotic factors were evaluated as predictors of ecosystem CO₂ flux. For chambers exposed to ambient and elevated CO₂ levels for the duration of the growing season, seasonality (Julian day) was the best predictor of ecosystem CO₂ flux at both ambient and elevated CO₂ levels. Light (PAR), soil temperature, and air temperature were also predictive of seasonal ecosystem flux, but only at elevated CO₂ levels. At any combination of physical conditions, flux of the elevated CO₂ treatment was greater than that at ambient. In short-term manipulations of CO₂, tundra exposed to elevated CO₂ had threefold greater carbon gain, and had one half the ecosystem level, light compensation point when compared to ambient CO₂ treatments. Elevated CO₂-acclimated tundra had twofold greater carbon gain compared to ambient treatments, but there was no difference in ecosystem level, light compensation point between elevated and ambient CO₂ treatments. The predicted future increases in cloudiness could substantially decrease the effect of elevated atmospheric CO₂ on net ecosystem carbon budget. These analyses suggest little if any long-term stimulation of ecosystem carbon acquisition by increases in atmospheric CO₂.     

Compensatory responses of CO2 exchange and biomass allocation and their effects on the relative growth rate of ponderosa pine in different CO2 and temperature regimes

Increases in the concentration of atmospheric carbon dioxide may have a fertilizing effect on plant growth by increasing photosynthetic rates and therefore may offset potential growth decreases caused by the stress associated with higher temperatures and lower precipitation. However, plant growth is determined both by rates of net photosynthesis and by proportional allocation of fixed carbon to autotrophic tissue and heterotrophic tissue. Although CO₂ fertilization may enhance growth by increasing leaf-level assimilation rates, reallocation of biomass from leaves to stems and roots in response to higher concentrations of CO₂ and higher temperatures may reduce whole-plant assimilation and offset photosynthetic gains. We measured growth parameters, photosynthesis, respiration, and biomass allocation of Pinus ponderosa seedlings grown for 2 months in 2×2 factorial treatments of 350 or 650 bar CO₂ and 10/25° C or 15/30° C night/day temperatures. After 1 month in treatment conditions, total seedling biomass was higher in elevated CO₂, and temperature significantly enhanced the positive CO₂ effect. However, after 2 months the effect of CO₂ on total biomass decreased and relative growth rates did not differ among CO₂ and temperature treatments over the 2-month growth period even though photosynthetic rates increased 7% in high CO₂ treatments and decreased 10% in high temperature treatments. Additionally, CO₂ enhancement decreased root respiration and high temperatures increased shoot respiration. Based on CO₂ exchange rates, CO₂ fertilization should have increased relative growth rates (RGR) and high temperatures should have decreased RGR. Higher photosynthetic rates caused by CO₂ fertilization appear to have been mitigated during the second month of exposure to treatment conditions by a 3% decrease in allocation of biomass to leaves and a 9% increase in root:shoot ratio. It was not clear why diminished photosynthetic rates and increased respiration rates at high temperatures did not result in lower RGR. Significant diametrical and potentially compensatory responses of CO₂ exchange and biomass allocation and the lack of differences in RGR of ponderosa pine after 2 months of exposure of high CO₂ indicate that the effects of CO₂ fertilization and temperature on whole-plant growth are determined by complex shifts in biomass allocation and gas exchange that may, for some species, maintain constant growth rates as climate and atmospheric CO₂ concentrations change. These complex responses must be considered together to predict plant growth reactions to global atmospheric change, and the potential of forest ecosystems to sequester larger amounts of carbon in the future.     

Urea additions and defoliation affect plant responses to elevated CO2 in a C3 grass from Yellowstone National Park

A common grass from Yellowstone National Park, Stipa occidentalis, was grown in a factorial experiment to determine if its response to the direct effects of elevated CO₂ would be affected by defoliation, and urea additions simulating the N in a urine hit. Plants were grown in tall pots (to mimic rooting depth in the field) in growth chambers under elevated (700 ppm) and ambient (370 ppm) CO₂, were defoliated or left undefoliated, and given N-supply rates based on field mineralization rates (untreated) or with an additional 40 g N/m². Growth increases in response to elevated CO₂ were largest when plants remained unclipped and received urea additions, and were found primarily in crowns and roots (storage organs). Aboveground biomass, which is the part of the plant consumed by grazing mammals, was not affected by elevated CO₂. The elevated CO₂ treatment caused a reduction in leaf percent N. However, there was a significant interaction between the CO₂ and urea treatments, resulting in a larger difference in leaf percent N between urea-treated and control plants under elevated than under ambient CO₂. Hence, elevations in atmospheric CO₂ may cause an increase in the amount of urine-hit-induced spatial variability in temperate grasslands. Since food quantity remained largely unchanged in response to elevated CO₂, and forage N content went down, grazing mammals may be negatively affected by increases in atmospheric CO₂.     

Studying the effect of elevated CO2 in the open in a naturally enriched environment in Central Italy

A gas vents area was recently localized in Central Italy. The gas emitted from the vents is composed by 92% of carbon dioxide and this produces an anomaly in the composition of the atmosphere over an area of about 2 ha. Atmospheric carbon dioxide concentration was measured by means of an infrared gas analyzer and diffusion tubes in several points and for some days within the area. Measurements revealed that the site can be at least divided into three sub-areas having increasing CO₂ concentration in the air. A preliminary analysis of natural vegetation in the area was conducted by counting stomatal and epidermal cells number and measuring guard cell size on leaves of several oak trees growing both near and far away from the vents. This analysis suggested that elevated CO₂ may have reduced the size of guard cells leaving stomatal density and stomatal index unaltered.     

Time-dependent responses of soil CO2 efflux components to elevated atmospheric [CO2] and temperature in experimental forest mesocosms

We previously used dual stable isotope techniques to partition soil CO₂ efflux into three source components (rhizosphere respiration, litter decomposition, and soil organic matter (SOM) oxidation) using experimental chambers planted with Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] seedlings. The components responded differently to elevated CO₂ (ambient + 200 mol mol^–1) and elevated temperature (ambient + 4 °C) treatments during the first year. Rhizosphere respiration increased most under elevated CO₂, and SOM oxidation increased most under elevated temperature. However, many studies show that plants and soil processes can respond to altered climates in a transient way. Herein, we extend our analysis to 2 years to evaluate the stability of the responses of the source components. Total soil CO₂ efflux increased significantly under elevated CO₂ and elevated temperature in both years (1994 and 1995), but the enhancement was much less in 1995. Rhizosphere respiration increased less under elevated temperature in 1995 compared with 1994. Litter decomposition also tended to increase comparatively less in 1995 under elevated CO₂, but was unresponsive to elevated temperature between years. In contrast, SOM oxidation was similar under elevated CO₂ in the 2 years. Less SOM oxidation occurred under elevated temperature in 1995 compared with 1994. Our results indicate that temporal variations can occur in CO₂ production by the sources. The variations likely involve responses to antecedent physical disruption of the soil and physiological processes.     

Effects of elevated CO2 and nitrogen on nutrient uptake in ponderosa pine seedlings

This paper reports on the results of a controlled-environment study on the effects of CO₂ (370, 525, and 700 mol mol^-1) and N [0, 200, and 400 g N g soil^-1 as (NH₄)SO₄] on ponderosa pine (Pinus ponderosa) seedlings. Based upon a review of the literature, we hypothesized that N limitations would not prevent a growth response to elevated CO₂. The hypothesis was not supported under conditions of extreme N deficiency (no fertilizer added to a very poor soil), but was supported when N limitations were less severe but still suboptimal (lower rate of fertilization). The growth increases in N-fertilized seedlings occurred mainly between 36 and 58 weeks without any additional N uptake. Thus, it appeared that elevated CO₂ allowed more efficient use of internal N reserves in the previously-fertilized seedlings, whereas internal N reserves in the unfertilized seedlings were insufficient to allow this response. Uptake rates of other nutrients were generally proportional to growth. Nitrogen treatment caused reductions in soil exchangeable K⁺, Ca²⁺, and Mg²⁺ (presumably because of nitrification and NO₃ ^- leaching) but increases in extractable P (presumably due to stimulation of phosphatase activity).The results of this and other seedling studies show that elevated CO₂ causes a reduction in tissue N concentration, even under N-rich conditions. The unique response of N is consistent with the hypothesis that the efficiency of Rubisco increases with elevated CO₂. These results collectively have significant implications for the response of mature, N-deficient forests to evevated CO₂.     

Effects of nitrogen supply on the acclimation of photosynthesis to elevated CO2

A common observation in plants grown in elevated CO₂ concentration is that the rate of photosynthesis is lower than expected from the dependence of photosynthesis upon CO₂ concentration in single leaves of plants grown at present CO₂ concentration. Furthermore, it has been suggested that this apparent down regulation of photosynthesis may be larger in leaves of plants at low nitrogen supply than at higher nitrogen supply. However, the available data are rather limited and contradictory. In this paper, particular attention is drawn to the way in which whole plant growth response to N supply constitutes a variable sink strength for carbohydrate usage and how this may affect photosynthesis. The need for further studies of the acclimation of photosynthesis at elevated CO₂ in leaves of plants whose N supply has resulted in well-defined growth rate and sink activity is emphasised, and brief consideration is made of how this might be achieved.Abbreviations A rate of CO₂ assimilation - C_i internal CO₂ concentration - PCR photosynthetic carbon reduction - Rubisco Ribulose 1,5-bisphosphate carboxylase/oxygenase - RuBP ribulose 1,5-bisphosphate     

二氧化碳储存通量对森林生态系统碳收支的影响

涡度相关系统观测高度以下的CO₂储存通量对准确评价森林生态系统与大气间净CO₂交换量（NEE）有着重要的影响.本研究以长白山阔叶红松林为研究对象,利用2003年的涡度相关观测数据以及CO₂浓度廓线数据,分析了CO₂储存通量的变化规律及其对碳收支过程的影响.结果表明：涡度相关观测高度以下的CO₂储存通量具有典型的日变化特征,其最大变化量出现在大气稳定与不稳定层结转换期.利用涡度相关系统观测的单点CO₂浓度变化方法与利用CO₂浓度廓线方法计算的CO₂储存通量差异不显著.忽略CO₂储存通量,在半小时尺度上会造成对夜间和白天的NEE分别低估25%和19%,在日和年尺度上,会对NEE低估10%和25%;忽略CO₂储存通量,会低估Michaelis-Menten光响应方程及Lloyd-Taylor呼吸方程的参数,并且对表观初始量子效率α和参考呼吸R_ref的低估最大;忽略CO₂储存通量,在半小时、日及年尺度上,均会对总光合作用（GPP）和生态系统呼吸（R_e）低估约20%.     

Effects of CO2 and nutrient enrichment on tissue quality of two California annuals

The effects of CO₂ enrichment and soil nutrient status on tissue quality were investigated and related to the potential effect on growth and decomposition. Two California annuals, Avena fatua and Plantago erecta, were grown at ambient and ambient plus 35 Pa atmospheric CO₂ in nutrient unamended and amended serpentine soil. Elevated CO₂ led to significantly increased Avena shoot nitrogen concentrations in the nutrient amended treatment. It also led to decreased lignin concentrations in Avena roots in both nutrient treatments, and in Plantago shoots and roots with nutrient addition. Concentrations of total nonstructural carbohydrate (TNC) and carbon did not change with elevated CO₂ in either species. As a consequence of increased biomass accumulation, increased CO₂ led to larger total pools of TNC, lignin, total carbon, and total nitrogen in Avena with nutrient additions. Doubling CO₂ had no significant effect on Plantago. Given the limited changes in the compounds related to decomposibility and plant growth, effects of increased atmospheric CO₂ mediated through tissue composition on Avena and Plantago are likely to be minor and depend on site fertility. This study suggests that other factors such as litter moisture, whether or not litter is on the ground, and biomass allocation among roots and shoots, are likely to be more important in this California grassland ecosystem. CO₂ could influence those directly as well as indirectly.     

Effect of CO2 enrichment and nitrogen availability on resource acquisition and resource allocation in a grass,Bromus mollis

Summary The effects of CO₂ enrichment on the growth, biomass partitioning, photosynthetic rates, and leaf nitrogen concentration of a grass, Bromus mollis (C₃), were investigated at a favorable and a low level of nitrogen availability. Despite increases in root: shoot ratios, leaf nitrogen concentrations were decreased under CO₂ enrichment at both nitrogen levels. For the low-nitrogen treatment, this resulted in lower photosynthetic rates measured at 650 l/l for the CO₂-enriched plants, compared to photosynthetic rates measured at 350 l/l for the non-enriched plants. At higher nitrogen availability, photosynthetic rates of plants grown and measured at 650 l/l were greater than photosynthetic rates of the non-enriched plants measured at 350 l/l. Water use efficiency, however, was increased in enriched plants at both nitrogen levels. CO₂ enrichment stimulated vegetative growth at both high and low nitrogen during most of the vegetative growth phase but, at the end of the experiment, total biomass of the high and low CO₂ treatments did not differ for plants grown at low nitrogen availability. While not statistically significant, CO₂ tended to stimulate seed production at high nitrogen and to decrease it at low nitrogen.     

Effects of low and elevated CO2 on C3 and C4 annuals

Abutilon theophrasti (C₃) and Amaranthus retroflexus (C₄), were grown from seed at four partial pressures of CO₂: 15 Pa (below Pleistocene minimum), 27 Pa (pre-industrial), 35 Pa (current), and 70 Pa (future) in the Duke Phytotron under high light, high nutrient, and wellwatered conditions to evaluate their photosynthetic response to historic and future levels of CO₂. Net photosynthesis at growth CO₂ partial pressures increased with increasing CO₂ for C₃ plants, but not C₄ plants. Net photosynthesis of Abutilon at 15 Pa CO₂ was 70% less than that of plants grown at 35 Pa CO₂, due to greater stomatal and biochemical limitations at 15 Pa CO₂. Relative stomatal limitation (RSL) of Abutilon at 15 Pa CO₂ was nearly 3 times greater than at 35 Pa CO₂. A photosynthesis model was used to estimate ribulose-1,5-bisphosphate carboxylase (rubisco) activity (Vc_max), electron transport mediated RuBP regeneration capacity (J _max), and phosphate regeneration capacity (PiRC) in Abutilon from net photosynthesis versus intercellular CO₂ (A–C _i) curves. All three component processes decreased by approximately 25% in Abutilon grown at 15 Pa compared with 35 Pa CO₂. Abutilon grown at 15 Pa CO₂ had significant reductions in total rubisco activity (25%), rubisco content (30%), activation state (29%), chlorophyll content (39%), N content (32%), and starch content (68%) compared with plants grown at 35 Pa CO₂. Greater allocation to rubisco relative to light reaction components and concomitant decreases in J _max and PiRC suggest co-regulation of biochemical processes occurred in Abutilon grown at 15 Pa CO₂. There were no significant differences in photosynthesis or leaf properties in Abutilon grown at 27 Pa CO₂ compared with 35 Pa CO₂, suggesting that the rise in CO₂ since the beginning of the industrial age has had little effect on the photosynthetic performance of Abutilon. For Amaranthus, limitations of photosynthesis were balanced between stomatal and biochemical factors such that net photosynthesis was similar in all CO₂ treatments. Differences in photosynthetic response to growth over a wide range of CO₂ partial pressures suggest changes in the relative performance of C₃ and C₄ annuals as atmospheric CO₂ has fluctuated over geologic time.     

Canopy N and P dynamics of a southeastern US pine forest under elevated CO2

Forest production is strongly nutrient limited throughout the southeastern US. If nutrient limitations constrain plant acquisition of essential resources under elevated CO₂, reductions in the mass or nutrient content of forest canopies could constrain C assimilation from the atmosphere. We tested this idea by quantifying canopy biomass, foliar concentrations of N and P, and the total quantity of N and P in a loblolly pine (Pinus taeda) canopy subject to 4 years of free-air CO₂ enrichment. We also used N:P ratios to detect N versus P limitation to primary production under elevated CO₂. Canopy biomass was significantly higher under elevated CO₂ during the first 4 years of this experiment. Elevated CO₂ significantly reduced the concentration of N in loblolly pine foliage (5% relative to ambient CO₂) but not P. Despite the slight reduction foliage N concentrations, there were significant increases in canopy N and P contents under elevated CO₂. Foliar N:P ratios were not altered by elevated CO₂ and were within a range suggesting forest production is N limited not P limited. Despite the clear limitation of NPP by N under ambient and elevated CO₂ at this site, there is no evidence that the mass of N or P in the canopy is declining through the first 4 years of CO₂ fumigation. As a consequence, whole-canopy C assimilation is strongly stimulated by elevated CO₂ making this forest a larger net C sink under elevated CO₂ than under ambient CO₂. We discuss the potential for future decreases in canopy nutrient content as a result of limited changes in the size of the plant-available pools of N under elevated CO₂.     

Responses of temporal distribution of gastropods to individual and combined effects of elevated CO2 and N deposition in annual grassland

Terrestrial gastropods have been shown to exert major impacts on the plant structure and species composition of temperate grasslands and other terrestrial plant communities. In order to develop predictions of plant community responses to changing environments, it is critical to understand how factors structuring plant communities will be influenced by global changes. Nevertheless, little is known about the potential for the size and abundance of gastropods to be altered by the individual and combined effects of elevated atmospheric CO₂ and increased deposition of N. Previous work suggests that responses of herbivore abundance to these global changes seem likely to depend, in part, upon the changes in the quantity of plant biomass available for consumption, and changes in the nutritional quality of this plant material for gastropod survival, development, and reproduction. Therefore, in this study we investigated the effect of elevated CO₂ and increased N deposition on gastropod size and abundance, as well as the effects on plant community production and N status. Elevated CO₂ depressed the size of gastropods early in the growing season by 29% to 42%, increased mid-season gastropod abundance by 38% to 43%, and depressed abundance late in the growing season by 21% to 29%. These changes in gastropod size and abundance were due largely to the modification of plant tissue quality and quantity. Increased N deposition, in contrast, influenced neither gastropod abundance nor per capita biomass during any part of the growing season. Neither elevated CO₂ nor increased N deposition disrupted the temporal synchrony between plant production and slug abundance.     

Daily and seasonal CO2 exchange in Scots pine grown under elevated O3 and CO2: experiment and simulation

Starting in early spring of 1994, naturally regenerated, 30-year-old Scots pine (Pinus sylvestris L.) trees were grown in open-top chambers and exposed in situ to doubled ambient O₃,doubled ambient CO₂ and a combination of O₃ and CO₂ from 15 April to 15 September. To investigate daily and seasonal responses of CO₂ exchange to elevated O₃ and CO₂, the CO₂ exchange of shoots was measured continuously by an automatic system for measuring gas exchange during the course of one year (from 1 Januray to 31 December 1996). A process-based model of shoot photosynthesis was constructed to quantify modifications in the intrinsic capacity of photosynthesis and stomatal conductance by simulating the daily CO₂ exchange data from the field. Results showed that on most days of the year the model simulated well the daily course of shoot photosynthesis. Elevated O₃ significantly decreased photosynthetic capacity and stomatal conductance during the whole photosynthetic period. Elevated O₃ also led to a delay in onset of photosynthetic recovery in early spring and an increase in the sensitivity of photosynthesis to environmental stress conditions. The combination of elevated O₃ and CO₂ had an effect on photosynthesis and stomatal conductance similar to that of elevated O₃ alone, but significantly reduced the O₃-induced depression of photosynthesis. Elevated CO₂ significantly increased the photosynthetic capacity of Scots pine during the main growing season but slightly decreased it in early spring and late autumn. The model calculation showed that, compared to the control treatment, elevated O₃ alone and the combination of elevated O₃ and CO₂ decreased the annual total of net photosynthesis per unit leaf area by 55% and 38%, respectively. Elevated CO₂ increased the annual total of net photosynthesis by 13%.