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.     

The interacting effects of elevated atmospheric CO<Subscript>2</Subscript> concentration,drought and leaf-to-air vapour pressure deficit on ecosystem isoprene fluxes

Isoprene is the most abundant biogenic hydrocarbon released from vegetation and it plays a major role in tropospheric chemistry. Because of its link to climate change, there is interest in understanding the relationship between CO₂, water availability and isoprene emission. We explored the effect of atmospheric elevated CO₂ concentration and its interaction with vapour pressure deficit (VPD) and water stress, on gross isoprene production (GIP) and net ecosystem exchange of CO₂ (NEE) in two Populus deltoides plantations grown at ambient and elevated atmospheric CO₂ concentration in the Biosphere 2 Laboratory facility. Although GIP and NEE showed a similar response to light and temperature, their responses to CO₂ and VPD were opposite; NEE was stimulated by elevated CO₂ and depressed by high VPD, while GIP was inhibited by elevated CO₂ and stimulated by high VPD. The difference in response between isoprene production and photosynthesis was also evident during water stress. GIP was stimulated in the short term and declined only when the stress was severe, whereas NEE started to decrease from the beginning of the experiment. This contrasting response led the carbon lost as isoprene in both the ambient and the elevated CO₂ treatments to increase as water stress progressed. Our results suggest that water limitation can override the inhibitory effect of elevated CO₂ leading to increased global isoprene emissions in a climate change scenario with warmer and drier climate.     

Radon fluxes in tropical forest ecosystems of Brazilian Amazonia: night-time CO2 net ecosystem exchange derived from radon and eddy covariance methods

Radon‐222 (Rn‐222) is used as a transport tracer of forest canopy–atmosphere CO₂ exchange in an old‐growth, tropical rain forest site near km 67 of the Tapajós National Forest, Pará, Brazil. Initial results, from month‐long periods at the end of the wet season (June–July) and the end of the dry season (November–December) in 2001, demonstrate the potential of new Rn measurement instruments and methods to quantify mass transport processes between forest canopies and the atmosphere. Gas exchange rates yield mean canopy air residence times ranging from minutes during turbulent daytime hours to greater than 12 h during calm nights. Rn is an effective tracer for net ecosystem exchange of CO₂ (CO₂ NEE) during calm, night‐time hours when eddy covariance‐based NEE measurements are less certain because of low atmospheric turbulence. Rn‐derived night‐time CO₂ NEE (9.00±0.99 μmol m^?2 s^?1 in the wet season, 6.39±0.59 in the dry season) was significantly higher than raw uncorrected, eddy covariance‐derived CO₂ NEE (5.96±0.51 wet season, 5.57±0.53 dry season), but agrees with corrected eddy covariance results (8.65±1.07 wet season, 6.56±0.73 dry season) derived by filtering out lower NEE values obtained during calm periods using independent meteorological criteria. The Rn CO₂ results suggest that uncorrected eddy covariance values underestimate night‐time CO₂ loss at this site. If generalizable to other sites, these observations indicate that previous reports of strong net CO₂ uptake in Amazonian terra firme forest may be overestimated.     

Ecosystem carbon exchange in two terrestrial ecosystem mesocosms under changing atmospheric CO2 concentrations

The ecosystem-level carbon uptake and respiration were measured under different CO₂ concentrations in the tropical rainforest and the coastal desert of Biosphere 2, a large enclosed facility. When the mesocosms were sealed and subjected to step-wise changes in atmospheric CO₂ between daily means of 450 and 900 μmol mol⁻¹, net ecosystem exchange (NEE) of CO₂ was derived using the diurnal changes in atmospheric CO₂ concentrations. The step-wise CO₂ treatment was effectively replicated as indicated by the high repeatability of NEE measurements under similar CO₂ concentrations over a 12-week period. In the rainforest mesocosm, daily NEE was increased significantly by the high CO₂ treatments because of much higher enhancement of canopy CO₂ assimilation relative to the increase in the nighttime ecosystem respiration under high CO₂. Furthermore, the response of daytime NEE to increasing atmospheric CO₂ in this mesocosm was not linear, with a saturation concentration of 750 μmol mol⁻¹. In the desert mesocosm, a combination of a reduction in ecosystem respiration and a small increase in canopy CO₂ assimilation in the high CO₂ treatments also enhanced daily NEE. Although soil respiration was not affected by the short-term change in atmospheric CO₂ in either mesocosm, plant dark respiration was increased significantly by the high CO₂ treatments in the rainforest mesocosm while the opposite was found in the desert mesocosm. The high CO₂ treatments increased the ecosystem light compensation points in both mesocosms. High CO₂ significantly increased ecosystem radiation use efficiency in the rainforest mesocosm, but had a much smaller effect in the desert mesocosm. The desert mesocosm showed much lower absolute response in NEE to atmospheric CO₂ than the rainforest mesocosm, probably because of the presence of C₄ plants. This study illustrates the importance of large-scale experimental research in the study of complex global change issues. Received: 30 October 1998 / Accepted: 2 December 1998     

Net ecosystem CO2 exchange in Mojave Desert shrublands during the eighth year of exposure to elevated CO2

Arid ecosystems, which occupy about 35% of the Earth's terrestrial surface area, are believed to be among the most responsive to elevated [CO₂]. Net ecosystem CO₂ exchange (NEE) was measured in the eighth year of CO₂ enrichment at the Nevada Desert Free‐Air CO₂ Enrichment (FACE) Facility between the months of December 2003–December 2004. On most dates mean daily NEE (24 h) (μmol CO₂ m^?2 s^?1) of ecosystems exposed to elevated atmospheric CO₂ were similar to those maintained at current ambient CO₂ levels. However, on sampling dates following rains, mean daily NEEs of ecosystems exposed to elevated [CO₂] averaged 23 to 56% lower than mean daily NEEs of ecosystems maintained at ambient [CO₂]. Mean daily NEE varied seasonally across both CO₂ treatments, increasing from about 0.1 μmol CO₂ m^?2 s^?1 in December to a maximum of 0.5–0.6 μmol CO₂ m^?2 s^?1 in early spring. Maximum NEE in ecosystems exposed to elevated CO₂ occurred 1 month earlier than it did in ecosystems exposed to ambient CO₂, with declines in both treatments to lowest seasonal levels by early October (0.09±0.03 μmol CO₂ m^?2 s^?1), but then increasing to near peak levels in late October (0.36±0.08 μmol CO₂ m^?2 s^?1), November (0.28±0.03 μmol CO₂ m^?2 s^?1), and December (0.54±0.06 μmol CO₂ m^?2 s^?1). Seasonal patterns of mean daily NEE primarily resulted from larger seasonal fluctuations in rates of daytime net ecosystem CO₂ uptake which were closely tied to plant community phenology and precipitation. Photosynthesis in the autotrophic crust community (lichens, mosses, and free‐living cyanobacteria) following rains were probably responsible for the high NEEs observed in January, February, and late October 2004 when vascular plant photosynthesis was low. Both CO₂ treatments were net CO₂ sinks in 2004, but exposure to elevated CO₂ reduced CO₂ sink strength by 30% (positive net ecosystem productivity=127±17 g C m^?2 yr^?1 ambient CO₂ and 90±11 g C m^?2 yr^?1 elevated CO₂, P=0.011). This level of net C uptake rivals or exceeds levels observed in some forested and grassland ecosystems. Thus, the decrease in C sequestration seen in our study under elevated CO₂– along with the extensive coverage of arid and semi‐arid ecosystems globally – points to a significant drop in global C sequestration potential in the next several decades because of responses of heretofore overlooked dryland ecosystems.     

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.     

Rain events decrease boreal peatland net CO2 uptake through reduced light availability

Boreal peatlands store large amounts of carbon, reflecting their important role in the global carbon cycle. The short‐term exchange and the long‐term storage of atmospheric carbon dioxide (CO₂) in these ecosystems are closely associated with the permanently wet surface conditions and are susceptible to drought. Especially, the single most important peat forming plant genus, Sphagnum, depends heavily on surface wetness for its primary production. Changes in rainfall patterns are expected to affect surface wetness, but how this transient rewetting affects net ecosystem exchange of CO₂ (NEE) remains unknown. This study explores how the timing and characteristics of rain events during photosynthetic active periods, that is daytime, affect peatland NEE and whether rain event associated changes in environmental conditions modify this response (e.g. water table, radiation, vapour pressure deficit, temperature). We analysed an 11‐year time series of half‐hourly eddy covariance and meteorological measurements from Degerö Stormyr, a boreal peatland in northern Sweden. Our results show that daytime rain events systematically decreased the sink strength of peatlands for atmospheric CO₂. The decrease was best explained by rain associated reduction in light, rather than by rain characteristics or drought length. An average daytime growing season rain event reduced net ecosystem CO₂ uptake by 0.23–0.54 gC m^?2. On an annual basis, this reduction of net CO₂ uptake corresponds to 24% of the annual net CO₂ uptake (NEE) of the study site, equivalent to a 4.4% reduction of gross primary production (GPP) during the growing season. We conclude that reduced light availability associated with rain events is more important in explaining the NEE response to rain events than rain characteristics and changes in water availability. This suggests that peatland CO₂ uptake is highly sensitive to changes in cloud cover formation and to altered rainfall regimes, a process hitherto largely ignored.     

Partitioning net ecosystem carbon exchange with isotopic fluxes of CO2

Because biological and physical processes alter the stable isotopic composition of atmospheric CO₂, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of ¹³CO₂ above terrestrial ecosystems can potentially be used to separate net ecosystem CO₂ exchange (NEE) into its component fluxes, net photosynthetic assimilation (F_A) and ecosystem respiration (F_R). In this paper theory is developed to partition measured NEE into F_A and F_R, using measurements of fluxes of CO₂ and ¹³CO₂, and isotopic composition of respired CO₂ and forest air. The theory is then applied to fluxes measured (or estimated, for ¹³CO₂) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in ¹³CO₂ fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in F_A and F_R were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night‐time NEE and temperature (a standard technique which provides alternate estimates of F_R and F_A). The light response curve for photosynthesis (F_A vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation‐weighted discrimination against ¹³CO₂ for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole‐canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of ¹³CO₂ flux are needed to assess the validity of the assumptions used and provide defensible isotope‐based estimates of the component fluxes of net ecosystem exchange.     

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%.     

Elevated CO2 and temperature impacts on different components of soil CO2 efflux in Douglas-fir terracosms

Although numerous studies indicate that increasing atmospheric CO₂ or temperature stimulate soil CO₂ efflux, few data are available on the responses of three major components of soil respiration [i.e. rhizosphere respiration (root and root exudates), litter decomposition, and oxidation of soil organic matter] to different CO₂ and temperature conditions. In this study, we applied a dual stable isotope approach to investigate the impact of elevated CO₂ and elevated temperature on these components of soil CO₂ efflux in Douglas-fir terracosms. We measured both soil CO₂ efflux rates and the ¹³C and ¹⁸O isotopic compositions of soil CO₂ efflux in 12 sun-lit and environmentally controlled terracosms with 4-year-old Douglas fir seedlings and reconstructed forest soils under two CO₂ concentrations (ambient and 200 ppmv above ambient) and two air temperature regimes (ambient and 4 °C above ambient). The stable isotope data were used to estimate the relative contributions of different components to the overall soil CO₂ efflux. In most cases, litter decomposition was the dominant component of soil CO₂ efflux in this system, followed by rhizosphere respiration and soil organic matter oxidation. Both elevated atmospheric CO₂ concentration and elevated temperature stimulated rhizosphere respiration and litter decomposition. The oxidation of soil organic matter was stimulated only by increasing temperature. Release of newly fixed carbon as root respiration was the most responsive to elevated CO₂, while soil organic matter decomposition was most responsive to increasing temperature. Although some assumptions associated with this new method need to be further validated, application of this dual-isotope approach can provide new insights into the responses of soil carbon dynamics in forest ecosystems to future climate changes.     

The effect of charcoal production on carbon cycling in African biomes

Using biomass for charcoal production in sub-Saharan Africa (SSA) may change carbon stock dynamics and lead to irreversible changes in the carbon balance, yet we have little understanding of whether these dynamics vary by biome in this region. Currently, charcoal production contributes up to 7% of yearly deforestation in tropical regions, with carbon emissions corresponding to 71.2 million tonnes of CO₂ and 1.3 million tonnes of CH₄. With a projected increased demand for charcoal in the coming decades, even low harvest rates may throw the carbon budget off-balance due to legacy effects. Here, we parameterized the dynamic global vegetation model LPJ-GUESS for six SSA biomes and examined the effect of charcoal production on net ecosystem exchange (NEE), carbon stock sizes and recovery time for tropical rain forest, montane forest, moist savanna, dry savanna, temperate grassland and semi-desert. Under historical charcoal regimes, tropical rain forests and montane forests transitioned from net carbon sinks to net sources, that is, mean cumulative NEE from −3.56 ± 2.59 kg C/m² to 2.46 ± 3.43 kg C/m² and −2.73 ± 2.80 kg C/m² to 1.87 ± 4.94 kg C/m² respectively. Varying charcoal production intensities resulted in tropical rain forests showing at least two times higher carbon losses than the other biomes. Biome recovery time varied by carbon stock, with tropical and montane forests taking about 10 times longer than the fast recovery observed for semi-desert and temperate grasslands. Our findings show that high biomass biomes are disproportionately affected by biomass harvesting for charcoal, and even low harvesting rates strongly affect vegetation and litter carbon and their contribution to the carbon budget. Therefore, the prolonged biome recoveries imply that current charcoal production practices in SSA are not sustainable, especially in tropical rain forests and montane forests, where we observe longer recovery for vegetation and litter carbon stocks.     

The annual cycles of CO2 and H2O exchange over a northern mixed forest as observed from a very tall tower

We present the annual patterns of net ecosystem‐atmosphere exchange (NEE) of CO₂ and H2O observed from a 447 m tall tower sited within a mixed forest in northern Wisconsin, USA. The methodology for determining NEE from eddy‐covariance flux measurements at 30, 122 and 396 m above the ground, and from CO₂ mixing ratio measurements at 11, 30, 76, 122, 244 and 396 m is described. The annual cycle of CO₂ mixing ratio in the atmospheric boundary layer (ABL) is also discussed, and the influences of local NEE and large‐scale advection are estimated. During 1997 gross ecosystem productivity (947?18 g C m^?2 yr^?1), approximately balanced total ecosystem respiration (963±19 g C m^?2 yr^?1), and NEE of CO₂ was close to zero (16±19 g C m^?2 yr^?1 emitted into the atmosphere). The error bars represent the standard error of the cumulative daily NEE values. Systematic errors are also assessed. The identified systematic uncertainties in NEE of CO₂ are less than 60 g C m^?2 yr^?1. The seasonal pattern of NEE of CO₂ was highly correlated with leaf‐out and leaf‐fall, and soil thaw and freeze, and was similar to purely deciduous forest sites. The mean daily NEE of CO₂ during the growing season (June through August) was ?1.3 g C m^?2 day^?1, smaller than has been reported for other deciduous forest sites. NEE of water vapor largely followed the seasonal pattern of NEE of CO₂, with a lag in the spring when water vapor fluxes increased before CO₂ uptake. In general, the Bowen ratios were high during the dormant seasons and low during the growing season. Evapotranspiration normalized by potential evapotranspiration showed the opposite pattern. The seasonal course of the CO₂ mixing ratio in the ABL at the tower led the seasonal pattern of NEE of CO₂ in time: in spring, CO₂ mixing ratios began to decrease prior to the onset of daily net uptake of CO₂ by the forest, and in fall mixing ratios began to increase before the forest became a net source for CO₂ to the atmosphere. Transport as well as local NEE of CO₂ are shown to be important components of the ABL CO₂ budget at all times of the year.     

The dependence of respiration on photosynthetic substrate supply and temperature: integrating leaf, soil and ecosystem measurements

Interactions between photosynthetic substrate supply and temperature in determining the rate of three respiration components (leaf, belowground and ecosystem respiration) were investigated within three environmentally controlled, Populus deltoides forest bays at Biosphere 2, Arizona. Over 2 months, the atmospheric CO₂ concentration and air temperature were manipulated to test the following hypotheses: (1) the responses of the three respiration components to changes in the rate of photosynthesis would differ both in speed and magnitude; (2) the temperature sensitivity of leaf and belowground respiration would increase in response to a rise in substrate availability; and, (3) at the ecosystem level, the ratio of respiration to photosynthesis would be conserved despite week‐to‐week changes in temperature. All three respiration rates responded to the CO₂ concentration‐induced changes in photosynthesis. However, the proportional change in the rate of leaf respiration was more than twice that of belowground respiration and, when photosynthesis was reduced, was also more rapid. The results suggest that aboveground respiration plays a key role in the overall response of ecosystem respiration to short‐term changes in canopy photosynthesis. The short‐term temperature sensitivity of leaf respiration, measured within a single night, was found to be affected more by developmental conditions than photosynthetic substrate availability, as the Q₁₀ was lower in leaves that developed at high CO₂, irrespective of substrate availability. However, the temperature sensitivity of belowground respiration, calculated between periods of differing air temperature, appeared to be positively correlated with photosynthetic substrate availability. At the ecosystem level, respiration and photosynthesis were positively correlated but the relationship was affected by temperature; for a given rate of daytime photosynthesis, the rate of respiration the following night was greater at 25 than 20°C. This result suggests that net ecosystem exchange did not acclimate to temperature changes lasting up to 3 weeks. Overall, the results of this study demonstrate that the three respiration terms differ in their dependence on photosynthesis and that, short‐ and medium‐term changes in temperature may affect net carbon storage in terrestrial ecosystems.     

Wood CO2 efflux in a primary tropical rain forest

The balance between photosynthesis and plant respiration in tropical forests may substantially affect the global carbon cycle. Woody tissue CO₂ efflux is a major component of total plant respiration, but estimates of ecosystem‐scale rates are uncertain because of poor sampling in the upper canopy and across landscapes. To overcome these problems, we used a portable scaffolding tower to measure woody tissue CO₂ efflux from ground level to the canopy top across a range of sites of varying slope and soil phosphorus content in a primary tropical rain forest in Costa Rica. The objectives of this study were to: (1) determine whether to use surface area, volume, or biomass for modeling and extrapolating wood CO₂ efflux, (2) determine if wood CO₂ efflux varied seasonally, (3) identify if wood CO₂ efflux varied by functional group, height in canopy, soil fertility, or slope, and (4) extrapolate wood CO₂ efflux to the forest. CO₂ efflux from small diameter woody tissue (<10 cm) was related to surface area, while CO₂ efflux from stems >10 cm was related to both surface area and volume. Wood CO₂ efflux showed no evidence of seasonality over 2 years. CO₂ efflux per unit wood surface area at 25° (F_A) was highest for the N‐fixing dominant tree species Pentaclethra macroloba, followed by other tree species, lianas, then palms. Small diameter F_A increased steeply with increasing height, and large diameter F_A increased with diameter. Soil phosphorus and slope had slight, but complex effects on F_A. Wood CO₂ efflux per unit ground area was 1.34±0.36 μmol m^?2 s^?1, or 508±135 g C m^?2 yr^?1. Small diameter wood, only 15% of total woody biomass, accounted for 70% of total woody tissue CO₂ efflux from the forest; while lianas, only 3% of total woody biomass, contributed one‐fourth of the total wood CO₂ efflux.     

Forest and agricultural land-use-dependent CO2 exchange in Thuringia, Germany

Eddy covariance was used to measure the net CO₂ exchange (NEE) over ecosystems differing in land use (forest and agriculture) in Thuringia, Germany. Measurements were carried out at a managed, even‐aged European beech stand (Fagus sylvatica, 70–150 years old), an unmanaged, uneven‐aged mixed beech stand in a late stage of development (F. sylvatica, Fraxinus excelsior, Acer pseudoplantanus, and other hardwood trees, 0–250 years old), a managed young Norway spruce stand (Picea abies, 50 years old), and an agricultural field growing winter wheat in 2001, and potato in 2002. Large contrasts were found in NEE rates between the land uses of the ecosystems. The managed and unmanaged beech sites had very similar net CO₂ uptake rates (～?480 to ?500 g C m^?2 yr^?1). Main differences in seasonal NEE patterns between the beech sites were because of a later leaf emergence and higher maximum leaf area index at the unmanaged beech site, probably as a result of the species mix at the site. In contrast, the spruce stand had a higher CO₂ uptake in spring but substantially lower net CO₂ uptake in summer than the beech stands. This resulted in a near neutral annual NEE (?4 g C m^?2 yr^?1), mainly attributable to an ecosystem respiration rate almost twice as high as that of the beech stands, despite slightly lower temperatures, because of the higher elevation. Crops in the agricultural field had high CO₂ uptake rates, but growing season length was short compared with the forest ecosystems. Therefore, the agricultural land had low‐to‐moderate annual net CO₂ uptake (?34 to ?193 g C m^?2), but with annual harvest taken into account it will be a source of CO₂ (+97 to +386 g C m^?2). The annually changing patchwork of crops will have strong consequences on the regions' seasonal and annual carbon exchange. Thus, not only land use, but also land‐use history and site‐specific management decisions affect the large‐scale carbon balance.     

Elevated CO<Subscript>2</Subscript> mitigates the adverse effects of drought on daytime net ecosystem CO<Subscript>2</Subscript> exchange and photosynthesis in a Florida scrub-oak ecosystem

Drought is a normal, recurrent feature of climate. In order to understand the potential effect of increasing atmospheric CO₂ concentration (C _a) on ecosystems, it is essential to determine the combined effects of drought and elevated C _a (EC) under field conditions. A severe drought occurred in Central Florida in 1998 when precipitation was 88 % less than the average between 1984 and 2002. We determined daytime net ecosystem CO₂ exchange (NEE) before, during, and after the drought in the Florida scrub-oak ecosystem exposed to doubled C _a in open-top chamber since May 1996. We measured diurnal leaf net photosynthetic rate (P _N) of Quercus myrtifolia Willd, the dominant species, during and after the drought. Drought caused a midday depression in NEE and P _N at ambient CO₂ concentration (AC) and EC. EC mitigated the midday depression in NEE by about 60 % compared to AC and the effect of EC on leaf P _N was similar to its effect on NEE. Growth in EC lowered the sensitivity of NEE to air vapor pressure deficit under drought. Thus EC would help the scrub-oak ecosystem to survive the consequences of the effects of rising atmospheric CO₂ on climate change, including increased frequency of drought, while simultaneously sequestering more anthropogenic carbon.     

Simulated mechanisms of soil N feedback on the forest CO2 response

An improved understanding of the response of forest ecosystems to elevated levels of CO₂ in the atmosphere is crucial because atmospheric CO₂ concentration continues to increase at an accelerating rate and forests are an important sink in the global carbon cycle. Several CO₂‐enrichment experiments have now been running for more than 10 years, with highly variable short‐term results after the first decade. Responses to rising [CO₂] over the next few decades will depend on several plant and ecosystem feedbacks that are inadequately understood. In this study, we conduct a sensitivity analysis, within the context of the simulated CO₂ response, using a new version of the G'DAY ecosystem model, with an improved decomposition submodel, applied to a nitrogen‐limited Norway spruce forest site in the north of Sweden. The new decomposition model incorporates important modifications to soil processes, including some that constitute negative feedbacks on an ecosystem's growth response to elevated [CO₂]. The sensitivity analysis reveals key parameters and processes that are important for the simulated CO₂ response on the short term and others that are more important on the long term. A process that has a strong impact on the short‐term response is a change in decomposer composition, potentially in response to altered litter quality. Parameters that become increasingly important in the long term are carbon allocation to root exudates that are directly or indirectly associated with atmospheric N₂ fixation, and the rate of humification of soil organic matter. We identify factors intrinsic to species and site (microbes and resources) and ecosystem nutrient supply that determine the duration of the enhanced simulated growth response to elevated [CO₂].     

Responses of CAM species to increasing atmospheric CO2 concentrations

Crassulacean acid metabolism (CAM) species show an average increase in biomass productivity of 35% in response to a doubled atmospheric CO₂ concentration. Daily net CO₂ uptake is similarly enhanced, reflecting in part an increase in chlorenchyma thickness and accompanied by an even greater increase in water‐use efficiency. The responses of net CO₂ uptake in CAM species to increasing atmospheric CO₂ concentrations are similar to those for C₃ species and much greater than those for C₄ species. Increases in net daily CO₂ uptake by CAM plants under elevated atmospheric CO₂ concentrations reflect increases in both Rubisco‐mediated daytime CO₂ uptake and phosphoenolpyruvate carboxylase (PEPCase)‐mediated night‐time CO₂ uptake, the latter resulting in increased nocturnal malate accumulation. Chlorophyll contents and the activities of Rubisco and PEPCase decrease under elevated atmospheric CO₂, but the activated percentage for Rubisco increases and the K_M(HCO₃ ^? ) for PEPCase decreases, resulting in more efficient photosynthesis. Increases in root:shoot ratios and the formation of additional photosynthetic organs, together with increases in sucrose‐Pi synthase and starch synthase activity in these organs under elevated atmospheric CO₂ concentrations, decrease the potential feedback inhibition of photosynthesis. Longer‐term studies for several CAM species show no downward acclimatization of photosynthesis in response to elevated atmospheric CO₂ concentrations. With increasing temperature and drought duration, the percentage enhancement of daily net CO₂ uptake caused by elevated atmospheric CO₂ concentrations increases. Thus net CO₂ uptake, productivity, and the potential area for cultivation of CAM species will be enhanced by the increasing atmospheric CO₂ concentrations and the increasing temperatures associated with global climate change.     

Seventeen years of elevated CO2 exposure in a Chesapeake Bay Wetland: sustained but contrasting responses of plant growth and CO2 uptake

Increased atmospheric CO₂ concentration (Ca) produces a short‐term stimulation of photosynthesis and plant growth across terrestrial ecosystems. However, the long‐term response remains uncertain and is thought to depend on environmental constraints. In the longest experiment on natural ecosystem response to elevated Ca, we measured the shoot‐density, biomass and net CO₂ exchange (NEE) responses to elevated Ca from 1987 to 2003 in a Scirpus olneyi wetland sedge community of the Chesapeake Bay, MD, USA. Measurements were conducted in five replicated open‐top chambers per CO₂ treatment (ambient and elevated). In addition, unchambered control plots were monitored for shoot density. Responses of daytime NEE, Scirpus plant biomass and shoot density to elevated Ca were positive for any single year of the 17‐year period of study. Daytime NEE stimulation by elevated Ca rapidly dropped from 80% at the onset of the experiment to a long‐term stimulation average of about 35%. Shoot‐density stimulation by elevated Ca increased linearly with duration of exposure (r²=0.89), exceeding 120% after 17 years. Although of lesser magnitude, the shoot biomass response to elevated Ca was similar to that of the shoot density. Daytime NEE response to elevated Ca was not explained by the duration of exposure, but negatively correlated with salinity of the marsh, indicating that this elevated‐Ca response was decreased by water‐related stress. By contrast, circumstantial evidence suggested that salinity stress increased the stimulation of shoot density by elevated Ca, which highlights the complexity of the interaction between water‐related stresses and plant community responses to elevated Ca. Notwithstanding the effects of salinity stress, we believe that the most important finding of the present research is that a species response to elevated Ca can continually increase when this species is under stress and declining in its natural environment. This is particularly important because climate changes associated with elevated Ca are likely to increase environmental stresses on numerous species and modify their present distribution. Our results point to an increased resilience to change under elevated Ca when plants are exposed to adverse environmental conditions.