Effects of elevated atmospheric CO2 on net ecosystem CO2 exchange of a scrub–oak ecosystem

We report the results of a 2‐year study of effects of the elevated (current ambient plus 350 μmol CO₂ mol^?1) atmospheric CO₂ concentration (C_a) on net ecosystem CO₂ exchange (NEE) of a scrub–oak ecosystem. The measurements were made in open‐top chambers (OTCs) modified to function as open gas‐exchange systems. The OTCs enclosed samples of the ecosystem (ca. 10 m² surface area) that had regenerated after a fire, 5 years before, in either current ambient or elevated C_a. Throughout the study, elevated C_a increased maximum NEE (NEE_max) and the apparent quantum yield of the NEE (φ_NEE) during the photoperiod. The magnitude of the stimulation of NEE_max, expressed per unit ground area, was seasonal, rising from 50% in the winter to 180% in the summer. The key to this stimulation was effects of elevated C_a, and their interaction with the seasonal changes in the environment, on ecosystem leaf area index, photosynthesis and respiration. The separation of these factors was difficult. When expressed per unit leaf area the stimulation of the NEE_max ranged from 7% to 60%, with the increase being dependent on increasing soil water content (W_soil). At night, the CO₂ effluxes from the ecosystem (NEE_night) were on an average 39% higher in elevated C_a. However, the increase varied between 6% and 64%, and had no clear seasonality. The partitioning of NEE_night into its belowground (R_below) and aboveground (R_above) components was carried out in the winter only. A 35% and 27% stimulation of NEE_night in December 1999 and 2000, respectively, was largely due to a 26% and 28% stimulation of R_below in the respective periods, because R_below constituted ca. 87% of NEE_night. The 37% and 42% stimulation of R_above in December 1999 and 2000, respectively, was less than the 65% and 80% stimulation of the aboveground biomass by elevated C_a at these times. An increase in the relative amount of the aboveground biomass in woody tissue, combined with a decrease in the specific rate of stem respiration of the dominant species Quercus myrtifolia in elevated C_a, was responsible for this effect. Throughout this study, elevated C_a had a greater effect on carbon uptake than on carbon loss, in terms of both the absolute flux and relative stimulation. Consequently, for this scrub–oak ecosystem carbon sequestration was greater in the elevated C_a during this 2‐year study period.     

Elevated CO2 affects photosynthetic responses in canopy pine and subcanopy deciduous trees over 10 years: a synthesis from Duke FACE

Leaf responses to elevated atmospheric CO₂ concentration (C_a) are central to models of forest CO₂ exchange with the atmosphere and constrain the magnitude of the future carbon sink. Estimating the magnitude of primary productivity enhancement of forests in elevated C_a requires an understanding of how photosynthesis is regulated by diffusional and biochemical components and up‐scaled to entire canopies. To test the sensitivity of leaf photosynthesis and stomatal conductance to elevated C_a in time and space, we compiled a comprehensive dataset measured over 10 years for a temperate pine forest of Pinus taeda, but also including deciduous species, primarily Liquidambar styraciflua. We combined over one thousand controlled‐response curves of photosynthesis as a function of environmental drivers (light, air C_a and temperature) measured at canopy heights up to 20 m over 11 years (1996–2006) to generate parameterizations for leaf‐scale models for the Duke free‐air CO₂ enrichment (FACE) experiment. The enhancement of leaf net photosynthesis (A_net) in P. taeda by elevated C_a of +200 μmol mol^?1 was 67% for current‐year needles in the upper crown in summer conditions over 10 years. Photosynthetic enhancement of P. taeda at the leaf‐scale increased by two‐fold from the driest to wettest growing seasons. Current‐year pine foliage A_net was sensitive to temporal variation, whereas previous‐year foliage A_net was less responsive and overall showed less enhancement (+30%). Photosynthetic downregulation in overwintering upper canopy pine needles was small at average leaf N (N_area), but statistically significant. In contrast, co‐dominant and subcanopy L. styraciflua trees showed A_net enhancement of 62% and no A_net–N_area adjustments. Various understory deciduous tree species showed an average A_net enhancement of 42%. Differences in photosynthetic responses between overwintering pine needles and subcanopy deciduous leaves suggest that increased C_a has the potential to enhance the mixed‐species composition of planted pine stands and, by extension, naturally regenerating pine‐dominated stands.     

Modelling carbon balances of coastal arctic tundra under changing climate

Rising air temperatures are believed to be hastening heterotrophic respiration (R_h) in arctic tundra ecosystems, which could lead to substantial losses of soil carbon (C). In order to improve confidence in predicting the likelihood of such loss, the comprehensive ecosystem model ecosys was first tested with carbon dioxide (CO₂) fluxes measured over a tundra soil in a growth chamber under various temperatures and soil‐water contents (θ). The model was then tested with CO₂ and energy fluxes measured over a coastal arctic tundra near Barrow, Alaska, under a range of weather conditions during 1998–1999. A rise in growth chamber temperature from 7 to 15 °C caused large, but commensurate, rises in respiration and CO₂ fixation, and so no significant effect on net CO₂ exchange was modelled or measured. An increase in growth chamber θ from field capacity to saturation caused substantial reductions in respiration but not in CO₂ fixation, and so an increase in net CO₂ exchange was modelled and measured. Long daylengths over the coastal tundra at Barrow caused an almost continuous C sink to be modelled and measured during most of July (2–4 g C m^?2 d^?1), but shortening daylengths and declining air temperatures caused a C source to be modelled and measured by early September (～1 g C m^?2 d^?1). At an annual time scale, the coastal tundra was modelled to be a small C sink (4 g C m^?2 y^?1) during 1998 when average air temperatures were 4 °C above normal, and a larger C sink (16 g C m^?2 y^?1) during 1999 when air temperatures were close to long‐term normals. During 100 years under rising atmospheric CO₂ concentration (C_a), air temperature and precipitation driven by the IS92a emissions scenario, modelled R_h rose commensurately with net primary productivity (NPP) under both current and elevated rates of atmospheric nitrogen (N) deposition, so that changes in soil C remained small. However, methane (CH₄) emissions were predicted to rise substantially in coastal tundra with IS92a‐driven climate change (from ～20 to ～40 g C m^?2 y^?1), causing a substantial increase in the emission of CO₂ equivalents. If the rate of temperature increase hypothesized in the IS92a emissions scenario had been raised by 50%, substantial losses of soil C (～1 kg C m^?2) would have been modelled after 100 years, including additional emissions of CH₄.     

Effects of elevated CO2 and temperature on photosynthesis and leaf traits of an understory dwarf bamboo in subalpine forest zone,China

The dwarf bamboo (Fargesia rufa Yi), growing understory in subalpine dark coniferous forest, is one of the main foods for giant panda, and it influences the regeneration of subalpine coniferous forests in southwestern China. To investigate the effects of elevated CO₂, temperature and their combination, the dwarf bamboo plantlets were exposed to two CO₂ regimes (ambient and double ambient CO₂ concentration) and two temperatures (ambient and +2.2°C) in growth chambers. Gas exchange, leaf traits and carbohydrates concentration were measured after the 150‐day experiment. Elevated CO₂ significantly increased the net photosynthetic rate (A_net), intrinsic water‐use efficiency (WUE_i) and carbon isotope composition (δ¹³C) and decreased stomatal conductance (g_s) and total chlorophyll concentration based on mass (Chl_m) and area (Chl_a). On the other hand, elevated CO₂ decreased specific leaf area (SLA), which was increased by elevated temperature. Elevated CO₂ also increased foliar carbon concentration based on mass (C_m) and area (C_a), nitrogen concentration based on area (N_a), carbohydrates concentration (i.e. sucrose, sugar, starch and non‐structural carbohydrates) and the slope of the A_net–N_a relationship. However, elevated temperature decreased C_m, C_a and N_a. The combination of elevated CO₂ and temperature hardly affected SLA, C_m, C_a, N_m, N_a, Chl_m and Chl_a. Variables A_net and N_a had positive linear relationships in all treatments. Our results showed that photosynthetic acclimation did not occur in dwarf bamboo at elevated CO₂ and it could adjust physiology and morphology to enable the capture of more light, to increase WUE and improve nutritional conditions.     

Impacts of Hurricane Frances on Florida scrub-oak ecosystem processes: defoliation, net CO2 exchange and interactions with elevated CO2

Hurricane disturbances have profound impacts on ecosystem structure and function, yet their effects on ecosystem CO₂ exchange have not been reported. In September 2004, our research site on a fire‐regenerated scrub‐oak ecosystem in central Florida was struck by Hurricane Frances with sustained winds of 113 km h⁻¹ and wind gusts as high as 152 km h⁻¹. We quantified the hurricane damage on this ecosystem resulting from defoliation: we measured net ecosystem CO₂ exchange, the damage and recovery of leaf area, and determined whether growth in elevated carbon dioxide concentration in the atmosphere (C_a) altered this disturbance. The hurricane decreased leaf area index (LAI) by 21%, which was equal to 60% of seasonal variation in canopy growth during the previous 3 years, but stem damage was negligible. The reduction in LAI led to a 22% decline in gross primary production (GPP) and a 25% decline in ecosystem respiration (R_e). The compensatory declines in GPP and R_e resulted in no significant change in net ecosystem production (NEP). Refoliation began within a month after the hurricane, although this period was out of phase with the regular foliation period, and recovered 20% of the defoliation loss within 2.5 months. Full recovery of LAI, ecosystem CO₂ assimilation, and ecosystem respiration did not occur until the next growing season. Plants exposed to elevated C_a did not sustain greater damage, nor did they recover faster than plants grown under ambient C_a. Thus, our results indicate that hurricanes capable of causing significant defoliation with negligible damage to stems have negligible effects on NEP under current or future CO₂‐enriched environment.     

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.     

Modelling assimilation and intercellular CO2 from measured conductance: a synthesis of approaches

A spectrum of models that estimate assimilation rate A from intercellular carbon dioxide concentration (C_i) and measured stomatal conductance to CO₂ (g_c) were investigated using leaf‐level gas exchange measurements. The gas exchange measurements were performed in a uniform loblolly pine stand (Pinus taeda L.) using the Free Air CO₂ Enrichment (FACE) facility under ambient and elevated atmospheric CO₂ for 3 years. These measurements were also used to test a newly proposed framework that combines basic properties of the A–C_i curve with a Fickian diffusion transport model to predict the relationship between C_i/C_a and g_c, where C_a is atmospheric carbon dioxide concentration. The widely used Ball–Berry model and five other models as well as the biochemical model proposed by Farquhar et al. (1980) were also reformulated to express variations in C_i/C_a as a function of their corresponding driving mechanisms. To assess the predictive capabilities of these approaches, their respective parameters were estimated from independent measurements of long‐term stable carbon isotope determinations (δ¹³C), meteorological variables, and ensemble A–C_i curves. All eight approaches reproduced the measured A reasonably well, in an ensemble sense, from measured water vapour conductance and modeled C_i/C_a. However, the scatter in the instantaneous A estimates was sufficiently large for both ambient and elevated C_a to suggest that other transient processes were not explicitly resolved by all eight parameterizations. An important finding from our analysis is that added physiological complexity in modeling C_i/C_a (when g_c is known) need not always translate to increased accuracy in predicting A. Finally, the broader utility of these approaches to estimate assimilation and net ecosystem exchange is discussed in relation to elevated atmospheric CO₂.     

Exposure to an enriched CO2 atmosphere alters carbon assimilation and allocation in a pine forest ecosystem

We linked a leaf-level CO₂ assimilation model with a model that accounts for light attenuation in the canopy and measurements of sap-flux-based canopy conductance into a new canopy conductance-constrained carbon assimilation (4C-A) model. We estimated canopy CO₂ uptake (A_nC) at the Duke Forest free-air CO₂ enrichment (FACE) study. Rates of A_nC estimated from the 4C-A model agreed well with leaf gas exchange measurements (A_net) in both CO₂ treatments. Under ambient conditions, monthly sums of net CO₂ uptake by the canopy (A_nC) were 13% higher than estimates based on eddy-covariance and chamber measurements. Annual estimates of A_nC were only 3% higher than carbon (C) accumulations and losses estimated from ground-based measurements for the entire stand. The C budget for the Pinus taeda component was well constrained (within 1% of ground-based measurements). Although the closure of the C budget for the broadleaf species was poorer (within 20%), these species are a minor component of the forest. Under elevated CO₂, the C used annually for growth, turnover, and respiration balanced only 80% of the A_nC. Of the extra 700 g C m⁻² a⁻¹ (1999 and 2000 average), 86% is attributable to surface soil CO₂ efflux. This suggests that the production and turnover of fine roots was underestimated or that mycorrhizae and rhizodeposition became an increasingly important component of the C balance. Under elevated CO₂, net ecosystem production increased by 272 g C m⁻² a⁻¹: 44% greater than under ambient CO₂. The majority (87%) of this C was sequestered in a moderately long-term C pool in wood, with the remainder in the forest floor–soil subsystem.     

Combined effects of elevated CO2 and air temperature on carbon assimilation of Pinus taeda trees

Branches of 22-year-old loblolly pine (Pinus taeda, L.) trees growing in a plantation were exposed to ambient CO₂, ambient + 165 μmol mol^?1 CO₂ or ambient + 330 μmol mol^?1 CO₂ concentrations in combination with ambient or ambient + 2°C air temperatures for 3 years. Field measurements in the third year indicated that net carbon assimilation was enhanced in the elevated CO₂ treatments in all seasons. On the basis of A/C_i, curves, there was no indication of photosynthetic down-regulation. Branch growth and leaf area also increased significantly in the elevated CO₂ treatments. The imposed 2°C increase in air temperature only had slight effects on net assimilation and growth. Compared with the ambient CO₂ treatment, rates of net assimilation were ～1·6 times greater in the ambient + 165 μmol mol^?1 CO₂ treatment and 2·2 times greater in the ambient + 330 μmol mol^?1 CO₂ treatment. These ratios did not change appreciably in measurements made in all four seasons even though mean ambient air temperatures during the measurement periods ranged from 12·6 to 28·2°C. This indicated that the effect of elevated CO₂ concentrations on net assimilation under field conditions was primarily additive. The results also indicated that the effect of elevated CO₂ (+ 165 or + 330 μmol mol^?1) was much greater than the effect of a 2°C increase in air temperature on net assimilation and growth in this species.     

Invasive insects impact forest carbon dynamics

Invasive insects can impact ecosystem functioning by altering carbon, nutrient, and hydrologic cycles. In this study, we used eddy covariance to measure net CO₂ exchange with the atmosphere (NEE), and biometric measurements to characterize net ecosystem productivity (NEP) in oak‐ and pine‐dominated forests that were defoliated by Gypsy moth (Lymantria dispar L.) in the New Jersey Pine Barrens. Three years of data were used to compare C dynamics; 2005 with minimal defoliation, 2006 with partial defoliation of the canopy and understory in a mixed stand, and 2007 with complete defoliation of an oak‐dominated stand, and partial defoliation of the mixed and pine‐dominated stands. Previous to defoliation in 2005, annual net CO₂ exchange (NEE_yr) was estimated at ?187, ?137 and ?204 g C m^?2 yr^?1 at the oak‐, mixed‐, and pine‐dominated stands, respectively. Annual NEP estimated from biometric measurements was 108%, 100%, and 98% of NEE_yr in 2005 for the oak‐, mixed‐, and pine‐dominated stands, respectively. Gypsy moth defoliation strongly reduced fluxes in 2006 and 2007 compared with 2005; NEE_yr was ?122, +103, and ?161 g C m^?2 yr^?1 in 2006, and +293, +129, and ?17 g C m^?2 yr^?1 in 2007 at the oak‐, mixed‐, and pine‐dominated stands, respectively. At the landscape scale, Gypsy moths defoliated 20.2% of upland forests in 2007. We calculated that defoliation in these upland forests reduced NEE_yr by 41%, with a 55% reduction in the heavily impacted oak‐dominated stands. ‘Transient’ disturbances such as insect defoliation, nonstand replacing wildfires, and prescribed burns are major factors controlling NEE across this landscape, and when integrated over time, may explain much of the patterning of aboveground biomass and forest floor mass in these upland forests.     

Direct and indirect climate change effects on carbon dioxide fluxes in a thawing boreal forest–wetland landscape

In the sporadic permafrost zone of northwestern Canada, boreal forest carbon dioxide (CO₂) fluxes will be altered directly by climate change through changing meteorological forcing and indirectly through changes in landscape functioning associated with thaw‐induced collapse‐scar bog (‘wetland’) expansion. However, their combined effect on landscape‐scale net ecosystem CO₂ exchange (NEE_LAND), resulting from changing gross primary productivity (GPP) and ecosystem respiration (ER), remains unknown. Here, we quantify indirect land cover change impacts on NEE_LAND and direct climate change impacts on modeled temperature‐ and light‐limited NEE_LAND of a boreal forest–wetland landscape. Using nested eddy covariance flux towers, we find both GPP and ER to be larger at the landscape compared to the wetland level. However, annual NEE_LAND (?20 g C m^?2) and wetland NEE (?24 g C m^?2) were similar, suggesting negligible wetland expansion effects on NEE_LAND. In contrast, we find non‐negligible direct climate change impacts when modeling NEE_LAND using projected air temperature and incoming shortwave radiation. At the end of the 21st century, modeled GPP mainly increases in spring and fall due to reduced temperature limitation, but becomes more frequently light‐limited in fall. In a warmer climate, ER increases year‐round in the absence of moisture stress resulting in net CO₂ uptake increases in the shoulder seasons and decreases during the summer. Annually, landscape net CO₂ uptake is projected to decline by 25 ± 14 g C m^?2 for a moderate and 103 ± 38 g C m^?2 for a high warming scenario, potentially reversing recently observed positive net CO₂ uptake trends across the boreal biome. Thus, even without moisture stress, net CO₂ uptake of boreal forest–wetland landscapes may decline, and ultimately, these landscapes may turn into net CO₂ sources under continued anthropogenic CO₂ emissions. We conclude that NEE_LAND changes are more likely to be driven by direct climate change rather than by indirect land cover change impacts.     

Carbon exchange of a mature,naturally regenerated pine forest in north Florida

We used eddy covariance and biomass measurements to quantify the carbon (C) dynamics of a naturally regenerated longleaf pine/slash pine flatwoods ecosystem in north Florida for 4 years, July 2000 to June 2002 and 2004 to 2005, to quantify how forest type, silvicultural intensity and environment influence stand‐level C balance. Precipitation over the study periods ranged from extreme drought (July 2000–June 2002) to above‐average precipitation (2004 and 2005). After photosynthetic photon flux density (PPFD), vapor pressure deficit (VPD) >1.5 kPa and air temperature <10 °C were important constraints on daytime half‐hourly net CO₂ exchange (NEE_day) and reduced the magnitude of midday CO₂ exchange by >5 μmol CO₂ m^?2 s^?1. Analysis of water use efficiency indicated that stomatal closure at VPD>1.5 kPa moderated transpiration similarly in both drought and wet years. Night‐time exchange (NEE_night) was an exponential function of air temperature, with rates further modulated by soil moisture. Estimated annual net ecosystem production (NEP) was remarkably consistent among the four measurement years (range: 158–192 g C m^?2 yr^?1). In comparison, annual ecosystem C assimilation estimates from biomass measurements between 2000 and 2002 ranged from 77 to 136 g C m^?2 yr^?1. Understory fluxes accounted for approximately 25–35% of above‐canopy NEE over 24‐h periods, and 85% and 27% of whole‐ecosystem fluxes during night and midday (11:00–15:00 hours) periods, respectively. Concurrent measurements of a nearby intensively managed slash pine plantation showed that annual NEP was three to four times greater than that of the Austin Cary Memorial Forest, highlighting the importance of silviculture and management in regulating stand‐level C budgets.     

Effects of elevated atmospheric CO2 on fine root production and activity in an intact temperate forest ecosystem

We investigated the effects of elevated atmospheric CO₂ concentrations (ambient + 200 ppm) on fine root production and soil carbon dynamics in a loblolly pine (Pinus taeda) forest subject to free‐air CO₂ enrichment (FACE) near Durham, NC (USA). Live fine root mass (LFR) showed less seasonal variation than dead fine root mass (DFR), which was correlated with seasonal changes in soil moisture and soil temperature. LFR mass increased significantly (by 86%) in the elevated CO₂ treatment, with an increment of 37 g(dry weight) m^?2 above the control plots after two years of CO₂ fumigation. There was no long‐term increment in DFR associated with elevated CO₂, but significant seasonal accumulations of DFR mass occurred during the summer of the second year of fumigation. Overall, root net primary production (RNPP) was not significantly different, but annual carbon inputs were 21.7 gC m^?2 y^?1 (68%) higher in the elevated CO₂ treatment compared to controls. Specific root respiration was not altered by the CO₂ treatment during most of the year; however, it was significantly higher by 21% and 13% in September 1997 and May 1998, respectively, in elevated CO₂. We did not find statistically significant differences in the C/N ratio of the root tissue, root decomposition or phosphatase activity in soil and roots associated with the treatment. Our data show that the early response of a loblolly pine forest ecosystem subject to CO₂ enrichment is an increase in its fine root population and a trend towards higher total RNPP after two years of CO₂ fumigation.     

Importance of changing CO2, temperature, precipitation, and ozone on carbon and water cycles of an upland-oak forest: incorporating experimental results into model simulations

Observed responses of upland‐oak vegetation of the eastern deciduous hardwood forest to changing CO₂, temperature, precipitation and tropospheric ozone (O₃) were derived from field studies and interpreted with a stand‐level model for an 11‐year range of environmental variation upon which scenarios of future environmental change were imposed. Scenarios for the year 2100 included elevated [CO₂] and [O₃] (+385 ppm and +20 ppb, respectively), warming (+4°C), and increased winter precipitation (+20% November–March). Simulations were run with and without adjustments for experimentally observed physiological and biomass adjustments. Initial simplistic model runs for single‐factor changes in CO₂ and temperature predicted substantial increases (+191% or 508 g C m^?2 yr^?1) or decreases (?206% or ?549 g C m^?2 yr^?1), respectively, in mean annual net ecosystem carbon exchange (NEE_a≈266±23 g C m^?2 yr^?1 from 1993 to 2003). Conversely, single‐factor changes in precipitation or O₃ had comparatively small effects on NEE_a (0% and ?35%, respectively). The combined influence of all four environmental changes yielded a 29% reduction in mean annual NEE_a. These results suggested that future CO₂‐induced enhancements of gross photosynthesis would be largely offset by temperature‐induced increases in respiration, exacerbation of water deficits, and O₃‐induced reductions in photosynthesis. However, when experimentally observed physiological adjustments were included in the simulations (e.g. acclimation of leaf respiration to warming), the combined influence of the year 2100 scenario resulted in a 20% increase in NEE_a not a decrease. Consistent with the annual model's predictions, simulations with a forest succession model run for gradually changing conditions from 2000 to 2100 indicated an 11% increase in stand wood biomass in the future compared with current conditions. These model‐based analyses identify critical areas of uncertainty for multivariate predictions of future ecosystem response, and underscore the importance of long term field experiments for the evaluation of acclimation and growth under complex environmental scenarios.     

Seasonal soil‐surface carbon fluxes from the root systems of young Pinus radiata trees growing at ambient and elevated CO2 concentration

Elevated atmospheric CO₂ concentration may result in increased below‐ground carbon allocation by trees, thereby altering soil carbon cycling. Seasonal estimates of soil surface carbon flux were made to determine whether carbon losses from Pinus radiata trees growing at elevated CO₂ concentration were higher than those at ambient CO₂ concentration, and whether this was related to increased fine root growth. Monthly soil surface carbon flux density (f) measurements were made on plots with trees growing at ambient (350) and elevated (650 μmol mol^?1) CO₂ concentration in large open‐top chambers. Prior to planting the soil carbon concentration (0.1%) and f (0.28 μmol m^?2 s^?1 at 15 °C) were low. A function describing the radial pattern of f with distance from tree stems was used to estimate the annual carbon flux from tree plots. Seasonal estimates of fine root production were made from minirhizotrons and the radial distribution of roots compared with radial measurements of f. A one‐dimensional gas diffusion model was used to estimate f from soil CO₂ concentrations at four depths. For the second year of growth, the annual carbon flux from the plots was 1671 g y^?1 and 1895 g y^?1 at ambient and elevated CO₂ concentrations, respectively, although this was not a significant difference. Higher f at elevated CO₂ concentration was largely explained by increased fine root biomass. Fine root biomass and stem production were both positively related to f. Both root length density and f declined exponentially with distance from the stem, and had similar length scales. Diurnal changes in f were largely explained by changes in soil temperature at a depth of 0.05 m. Ignoring the change of f with increasing distance from tree stems when scaling to a unit ground area basis from measurements with individual trees could result in under‐ or overestimates of soil‐surface carbon fluxes, especially in young stands when fine roots are unevenly distributed.     

Salinity and sea level mediate elevated CO2 effects on C3–C4 plant interactions and tissue nitrogen in a Chesapeake Bay tidal wetland

Elevated atmospheric carbon dioxide concentrations ([CO₂]) generally increase plant photosynthesis in C₃ species, but not in C₄ species, and reduce stomatal conductance in both C₃ and C₄ plants. In addition, tissue nitrogen concentration ([N]) often fails to keep pace with enhanced carbon gain under elevated CO₂, particularly in C₃ species. While these responses are well documented in many species, implications for plant growth and nutrient cycling in native ecosystems are not clear. Here we present data on 18 years of measurement of above and belowground biomass, tissue [N] and total standing crop of N for a Scirpus olneyi‐dominated (C₃ sedge) community, a Spartina patens‐dominated (C₄ grass) community and a C₃–C₄‐mixed species community exposed to ambient and elevated (ambient +340 ppm) atmospheric [CO₂] in natural salinity and sea level conditions of a Chesapeake Bay wetland. Increased biomass production (shoots plus roots) under elevated [CO₂] in the S. olneyi‐dominated community was sustained throughout the study, averaging approximately 35%, while no significant effect of elevated [CO₂] was found for total biomass in the C₄‐dominated community. We found a significant decline in C₄ biomass (correlated with rising sea level) and a concomitant increase in C₃ biomass in the mixed community. This shift from C₄ to C₃ was accelerated by the elevated [CO₂] treatment. The elevated [CO₂] stimulation of total biomass accumulation was greatest during rainy, low salinity years: the average increase above the ambient treatment during the three wettest years (1994, 1996, 2003) was 2.9 t ha⁻¹ but in the three driest years (1995, 1999, 2002), it was 1.2 t ha⁻¹. Elevated [CO₂] depressed tissue [N] in both species, but especially in the S. olneyi where the relative depression was positively correlated with salinity and negatively related with the relative enhancement of total biomass production. Thus, the greatest amount of carbon was added to the S. olneyi‐dominated community during years when shoot [N] was reduced the most, suggesting that the availability of N was not the most or even the main limitation to elevated [CO₂] stimulation of carbon accumulation in this ecosystem.     

CO2 enrichment in a maturing pine forest: are CO2 exchange and water status in the canopy affected?

A _net, leaf net CO₂ assimilation
c_a, CO₂ concentration of air surrounding a leaf
c_i, leaf intercellular CO₂ concentration
Δ, ¹³C isotope discrimination
δ¹³C, relative stable carbon isotope content
?, ratio of A_net at c_a = 560μmol mol^–1 to A_net at c_a = 360 μmol mol^–1
FACE, free-air CO₂ enrichment
g_w, stomatal conductance to water vapour
Π_i, initial leaf osmotic potential
R_t, relative water content at incipient turgor loss
Ψ_l, xylem water potential of leaves
Ψ_m, soil matric potential

Elevated CO₂ is expected to reduce forest water use as a result of CO₂-induced stomatal closure, which has implications for ecosystem-scale phenomena controlled by water availability. Leaf-level CO₂ and H₂O exchange responses and plant and soil water relations were examined in a maturing loblolly pine (Pinus taeda L.) stand in a free-air CO₂ enrichment (FACE) experiment in North Carolina, USA to test if these parameters were affected by elevated CO₂. Current-year foliage in the canopy was continuously exposed to elevated CO₂ (ambient CO₂+200μmol mol^–1) in free-air during needle growth and development for up to 400 d. Photosynthesis in upper canopy foliage was stimulated by 50–60% by elevated CO₂ compared with ambient controls. This enhancement was similar in current-year, ambient-grown foliage temporarily measured at elevated CO₂ compared with long-term elevated CO₂ grown foliage. Significant photosynthetic enhancement by CO₂ was maintained over a range of conditions except during peak drought. There was no evidence of water savings in elevated CO₂ plots in FACE compared to ambient plots under drought and non-drought conditions. This was supported by evidence from three independent measures. First, stomatal conductance was not significantly different in elevated CO₂ versus ambient trees of P. taeda. Calculations of time-integrated c_i/c_a ratios from analysis of foliar δ¹³C showed that these ratios were maintained in foliage under elevated CO₂. Second, soil moisture was not significantly different between ambient and elevated CO₂ plots during drought. Third, pre-dawn and mid-day leaf water potentials were also unaffected by the seasonal CO₂ exposure, as were tissue osmotic potentials and turgor loss points. Together the results strongly support the hypothesis that maturing P. taeda trees have low stomatal responsiveness to elevated CO₂. Elevated CO₂ effects on water relations in loblolly pine-dominated forest ecosystems may be absent or small apart from those mediated by leaf area. Large photosynthetic enhancements in the upper canopy of P. taeda by elevated CO₂ indicate that this maturing forest may have a large carbon sequestration capacity with limiting water supply.     

Integration of CO2 flux and remotely-sensed data for primary production and ecosystem respiration analyses in the Northern Great Plains: potential for quantitative spatial extrapolation

Aim Extrapolation of tower CO₂ fluxes will be greatly facilitated if robust relationships between flux components and remotely sensed factors are established. Long‐term measurements at five Northern Great Plains locations were used to obtain relationships between CO₂ fluxes and photosynthetically active radiation (Q), other on‐site factors, and Normalized Difference Vegetation Index (NDVI) from the SPOT VEGETATION data set. Location CO₂ flux data from the following stations and years were analysed: Lethbridge, Alberta 1998–2001; Fort Peck, MT 2000, 2002; Miles City, MT 2000–01; Mandan, ND 1999–2001; and Cheyenne, WY 1997–98. Results Analyses based on light‐response functions allowed partitioning net CO₂ flux (F) into gross primary productivity (P_g) and ecosystem respiration (R_e). Weekly averages of daytime respiration, γ_day, estimated from light responses were closely correlated with weekly averages of measured night‐time respiration, γ_night (R² 0.64 to 0.95). Daytime respiration tended to be higher than night‐time respiration, and regressions of γ_day on γ_night for all sites were different from 1 : 1 relationships. Over 13 site‐years, gross primary production varied from 459 to 2491 g CO₂ m^?2 year^?1, ecosystem respiration from 996 to 1881 g CO₂ m^?2 year^?1, and net ecosystem exchange from ?537 (source) to +610 g CO₂ m^?2 year^?1 (sink). Maximum daily ecological light‐use efficiencies, ?_d,max = P_g/Q, were in the range 0.014 to 0.032 mol CO₂ (mol incident quanta)^?1. Main conclusions Ten‐day average P_g was significantly more highly correlated with NDVI than 10‐day average daytime flux, P_d (R² = 0.46 to 0.77 for P_g‐NDVI and 0.05 to 0.58 for P_d‐NDVI relationships). Ten‐day average R_e was also positively correlated with NDVI, with R² values from 0.57 to 0.77. Patterns of the relationships of P_g and R_e with NDVI and other factors indicate possibilities for establishing multivariate functions allowing scaling‐up local fluxes to larger areas using GIS data, temporal NDVI, and other factors.     

Elevated atmospheric CO2 stimulates aboveground biomass in a fire-regenerated scrub-oak ecosystem

The effect of elevated atmospheric CO₂ concentration (C_a) on the aboveground biomass of three oak species, Quercus myrtifolia, Q. geminata, and Q. chapmanii, was estimated nondestructively using allometric relationships between stem diameter and aboveground biomass after four years of experimental treatment in a naturally fire‐regenerated scrub‐oak ecosystem. After burning a stand of scrub‐oak vegetation, re‐growing plants were exposed to either current ambient (379 µL L^?1 CO₂) or elevated (704 µL L^?1 CO₂) C_a in 16 open‐top chambers over a four‐year period, and measurements of stem diameter were carried out annually on all oak shoots within each chamber. Elevated C_a significantly increased aboveground biomass, expressed either per unit ground area or per shoot; elevated C_a had no effect on shoot density. The relative effect of elevated C_a on aboveground biomass increased each year of the study from 44% (May 96–Jan 97), to 55% (Jan 97–Jan 98), 66% (Jan 98–Jan 99), and 75% (Jan 99–Jan 00). The effect of elevated C_a was species specific: elevated C_a significantly increased aboveground biomass of the dominant species, Q. myrtifolia, and tended to increase aboveground biomass of Q. chapmanii, but had no effect on aboveground biomass of the subdominant, Q. geminata. These results show that rising atmospheric CO₂ has the potential to stimulate aboveground biomass production in ecosystems dominated by woody species, and that species‐specific growth responses could, in the long term, alter the composition of the scrub‐oak community.     

Dynamic changes of canopy-scale mesophyll conductance to CO₂ diffusion of sunflower as affected by CO₂ concentration and abscisic acid

Leaf‐level measurements have shown that mesophyll conductance (g_m) can vary rapidly in response to CO₂ and other environmental factors, but similar studies at the canopy‐scale are missing. Here, we report the effect of short‐term variation of CO₂ concentration on canopy‐scale g_m and other CO₂ exchange parameters of sunflower (Helianthus annuus L.) stands in the presence and absence of abscisic acid (ABA) in their nutrient solution. g_m was estimated from gas exchange and on‐line carbon isotope discrimination (Δ_obs) in a ¹³CO₂/¹²CO₂ gas exchange mesocosm. The isotopic contribution of (photo)respiration to stand‐scale Δ_obs was determined with the experimental approach of Tcherkez et al. Without ABA, short‐term exposures to different CO₂ concentrations (C_a 100 to 900 µmol mol^?1) had little effect on canopy‐scale g_m. But, addition of ABA strongly altered the CO₂‐response: g_m was high (approx. 0.5 mol CO₂ m^?2 s^?1) at C_a < 200 µmol mol^?1 and decreased to <0.1 mol CO₂ m^?2 s^?1 at C_a >400 µmol mol^?1. In the absence of ABA, the contribution of (photo)respiration to stand‐scale Δ_obs was high at low C_a (7.2‰) and decreased to <2‰ at C_a > 400 µmol mol^?1. Treatment with ABA halved this effect at all C_a.