Stimulation of both photosynthesis and respiration in response to warmer and drier conditions in a boreal peatland ecosystem

Peatland ecosystems have been consistent carbon (C) sinks for millennia, but it has been predicted that exposure to warmer temperatures and drier conditions associated with climate change will shift the balance between ecosystem photosynthesis and respiration providing a positive feedback to atmospheric CO₂ concentration. Our main objective was to determine the sensitivity of ecosystem photosynthesis, respiration and net ecosystem production (NEP) measured by eddy covariance, to variation in temperature and water table depth associated with interannual shifts in weather during 2004–2009. Our study was conducted in a moderately rich treed fen, the most abundant peatland type in western Canada, in a region (northern Alberta) where peatland ecosystems are a significant landscape component. During the study, the average growing season (May–October) water depth declined approximately 38 cm, and temperature [expressed as cumulative growing degree days (GDD, March–October)] varied approximately 370 GDD. Contrary to previous predictions, both ecosystem photosynthesis and respiration showed similar increases in response to warmer and drier conditions. The ecosystem remained a strong net sink for CO₂ with an average NEP (± SD) of 189 ± 47 g C m^?2 yr^?1. The current net CO₂ uptake rates were much higher than C accumulation in peat determined from analyses of the relationship between peat age and cumulative C stock. The balance between C addition to, and total loss from, the top 0–30 cm depth (peat age range 0–70 years) of shallow peat cores averaged 43 ± 12 g C m^?2 yr^?1. The apparent long‐term average rate of net C accumulation in basal peat samples was 19–24 g C m^?2 yr^?1. The difference between current rates of net C uptake and historical rates of peat accumulation is likely a result of vegetation succession and recent increases in tree establishment and productivity.     

Nitrogen addition reduces soil respiration in a mature tropical forest in southern China

Response of soil respiration (CO₂ emission) to simulated nitrogen (N) deposition in a mature tropical forest in southern China was studied from October 2005 to September 2006. The objective was to test the hypothesis that N addition would reduce soil respiration in N saturated tropical forests. Static chamber and gas chromatography techniques were used to quantify the soil respiration, following four‐levels of N treatments (Control, no N addition; Low‐N, 5 g N m^?2 yr^?1; Medium‐N, 10 g N m^?2 yr^?1; and High‐N, 15 g N m^?2 yr^?1 experimental inputs), which had been applied for 26 months before and continued throughout the respiration measurement period. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates found in the warm and wet growing season (April–September) and the lowest rates in the dry dormant season (December–February). Soil respiration rates showed a significant positive exponential relationship with soil temperature, whereas soil moisture only affect soil respiration at dry conditions in the dormant season. Annual accumulative soil respiration was 601±30 g CO₂‐C m^?2 yr^?1 in the Controls. Annual mean soil respiration rate in the Control, Low‐N and Medium‐N treatments (69±3, 72±3 and 63±1 mg CO₂‐C m^?2 h^?1, respectively) did not differ significantly, whereas it was 14% lower in the High‐N treatment (58±3 mg CO₂‐C m^?2 h^?1) compared with the Control treatment, also the temperature sensitivity of respiration, Q₁₀ was reduced from 2.6 in the Control with 2.2 in the High‐N treatment. The decrease in soil respiration occurred in the warm and wet growing season and were correlated with a decrease in soil microbial activities and in fine root biomass in the N‐treated plots. Our results suggest that response of soil respiration to atmospheric N deposition in tropical forests is a decline, but it may vary depending on the rate of N deposition.     

Spring warming and carbon dioxide exchange over low Arctic tundra in central Canada

Tundra‐atmosphere exchanges of carbon dioxide (CO₂) and water vapour were measured near Daring Lake, Northwest Territories in the Canadian Low Arctic for 3 years, 2004–2006. The measurement period spanned late‐winter until the end of the growing period. Mean temperatures during the measurement period varied from about 2 °C less than historical average in 2004 and 2005 to 2 °C greater in 2006. Much of the added warmth in 2006 occurred at the beginning of the study, when snow melt occurred 3 weeks earlier than in the other years. Total precipitation in 2006 (163 mm) was more than double that of the driest year, 2004 (71 mm). The tundra was a net sink for CO₂ carbon in all years. Mid‐summer net ecosystem exchange of CO₂ (NEE) achieved maximum values of ?1.3 g C m^?2 day^?1 (2004) to ?1.8 g C m^?2 day^?1 (2006). Accumulated NEE values over the 109‐day period were ?32,?51 and ?61 g C m^?2 in 2004, 2005 and 2006, respectively. The larger CO₂ uptake in 2006 was attributed to the early spring coupled with warmer air and soil conditions. In 2004, CO₂ uptake was limited by the shorter growing season and mid‐summer dryness, which likely reduced ecosystem productivity. Seasonal total evapotranspiration (ET) ranged from 130 mm (2004) to 181 mm (2006) and varied in accordance with the precipitation received and with the timing of snow melt. Maximum daily ET rates ranged from 2.3 to 2.7 mm day^?1, occurring in mid July. Ecosystem water use efficiency (WUE_eco) varied slightly between years, ranging from 2.2 in the driest year to 2.5 in the year with intermediate rainfall amounts. In the wettest year, increased soil evaporation may have contributed to a lower WUE_eco (2.3). We speculate that most, if not all, of the modest growing season CO₂ sink measured at this site could be lost due to fall and winter respiration leading to the tundra being a net CO₂ source or CO₂ neutral on an annual basis. However, this hypothesis is untested as yet.     

Seasonal Net Ecosystem Carbon Exchange of a Regenerating Cutaway Bog: How Long Does it Take to Restore the C‐Sequestration Function?

We measured the net ecosystem exchange (NEE) and respiration rates and modeled the photosynthesis and respiration dynamics in a cutover bog in the Swiss Jura Mountains during one growing season at three stages of regeneration (29, 42, and 51 years after peat cutting; coded sites A, B, and C) to determine if reestablishment of Sphagnum suffices to restore the C‐sequestration function. From the younger to the older stage Sphagnum cover increased, while net primary Sphagnum production over the growing season decreased (139, 82, and, 67 g m^?2 y^?1 for A, B, and C respectively), and fen plant species were replaced by bog species. According to our NEE estimations, over the vegetation period site A was a net CO₂‐C source emitting 40 g CO₂‐C/m² while sites B and C were accumulating CO₂‐C, on average 222 and 209 g CO₂‐C/m², respectively. These differences are due to the higher respiration in site A during the summer, suggesting that early regeneration stages may be more sensitive to a warmer climate. Methane fluxes increased from site A to C in parallel with Eriophorum vaginatum cover and vascular plant leaf area. Our results show that reestablishing a Sphagnum cover is not sufficient to restore a CO₂‐sequestrating function but that after circa 50 years the ecosystem may naturally regain this function over the growing season.     

Scaling net ecosystem CO2 exchange from the community to landscape‐level at a subarctic fen

Landscape‐ and community‐level CO₂ measurements were made at a subarctic sedge fen near Churchill Manitoba during the 1997 growing season. Climatic conditions were warmer and drier than the 30‐y normal. Landscape‐scale micrometeorological measurements indicated that the wetland gained 49 g CO₂ m^?2 during the growing season. Chamber‐scale measurements from the main vegetation community types showed that small hummocks (Carex spp. sites) dominated the CO₂ exchange, yielding an effective scaling factor of 70%. Scaled parameters of two algorithms describing photosynthesis and respiration for each community type show strong similarity to those derived at the landscape level. Scaling photosynthesis, respiration, and net ecosystem CO₂ exchange from the community to landscape‐level over the season is within the maximum probable error of each methodological approach and helps substantiate the 1997 CO₂ budget. We explore the equilibrium response of net ecosystem CO₂ exchange of this fen to climatic change by examining the feedback of water table position on vegetation distribution and nitrogen availability. Based on the effective scaling factors computed for each community type, we hypothesize that a small decrease in mean water table position could nearly triple the net uptake of CO₂ at this wetland.     

Modeling to discern nitrogen fertilization impacts on carbon sequestration in a Pacific Northwest Douglas‐fir forest in the first‐postfertilization year

This study investigated how nitrogen (N) fertilization with 200 kg N ha^?1 of urea affected ecosystem carbon (C) sequestration in the first‐postfertilization year in a Pacific Northwest Douglas‐fir (Pseudotsuga menziesii) stand on the basis of multiyear eddy‐covariance (EC) and soil‐chamber measurements before and after fertilization in combination with ecosystem modeling. The approach uses a data‐model fusion technique which encompasses both model parameter optimization and data assimilation and minimizes the effects of interannual climatic perturbations and focuses on the biotic and abiotic factors controlling seasonal C fluxes using a prefertilization 9‐year‐long time series of EC data (1998–2006). A process‐based ecosystem model was optimized using the half‐hourly data measured during 1998–2005, and the optimized model was validated using measurements made in 2006 and further applied to predict C fluxes for 2007 assuming the stand was not fertilized. The N fertilization effects on C sequestration were then obtained as differences between modeled (unfertilized stand) and EC or soil‐chamber measured (fertilized stand) C component fluxes. Results indicate that annual net ecosystem productivity in the first‐post‐N fertilization year increased by～83%, from 302 ± 19 to 552 ± 36 g m^?2 yr^?1, which resulted primarily from an increase in annual gross primary productivity of～8%, from 1938 ± 22 to 2095 ± 29 g m^?2 yr^?1 concurrent with a decrease in annual ecosystem respiration (R_e) of～5.7%, from 1636 ± 17 to 1543 ± 31 g m^?2 yr^?1. Moreover, with respect to respiration, model results showed that the fertilizer‐induced reduction in R_e (～93 g m^?2 yr^?1) principally resulted from the decrease in soil respiration R_s (～62 g m^?2 yr^?1).     

Response of total night-time respiration to differences in total daily photosynthesis for leaves in a Quercus rubra L. canopy: implications for modelling canopy CO2 exchange

Measurements of photosynthesis and respiration were made on leaves in summer in a Quercus rubra L. canopy at approximately hourly intervals throughout 5 days and nights. Leaves were selected in the upper canopy in fully sunlit conditions (upper) and in the lower canopy (lower). In addition, leaves in the upper canopy were shaded (upper shaded) to decrease photosynthesis rates. The data were used to test the hypothesis that total night‐time respiration is dependent on total photosynthesis during the previous day and that the response is mediated through changes in storage in carbohydrate pools. Measurements were made on clear sunny days with similar solar irradiance and air temperature, except for the last day when temperature, especially at night, was lower than that for the previous days. Maximum rates of photosynthesis in the upper leaves (18.7 μmol m^?2 s^?1) were approximately four times higher than those in the lower leaves (4.3 μmol m^?2 s^?1) and maximum photosynthesis rates in the upper shaded leaves (8.0 μmol m^?2 s^?1) were about half those in the upper leaves. There was a strong linear relationship between total night‐time respiration and total photosynthesis during the previous day when rates of respiration were normalized to a fixed temperature of 20°C, removing the effects of temperature from this relationship. Measurements of specific leaf area, nitrogen and chlorophyll concentration and calculations of the maximum rate of carboxylation activity, V_cmax, were not significantly different between upper and upper shaded leaves 5 days after the shading treatment was started. There were small, but significant decreases in the rate of apparent maximum electron transport at saturating irradiance, J_max (P>0.05), and light use efficiency, ? (P<0.05), for upper shaded leaves compared with those for upper leaves. This suggests that the duration of shading in the experiment was sufficient to initiate changes in the electron transport, but not the carboxylation processes of photosynthesis. Support for the hypothesis was provided from analysis of soluble sugar and starch concentrations in leaves. Respiration rates in the upper shaded leaves were lower than those expected from a relationship between respiration and soluble sugar concentration for fully exposed upper and lower leaves. However, there was no similar difference in starch concentrations. This suggests that shading for the duration of several days did not affect sugar concentrations but reduced starch concentrations in leaves, leading to lower rates of respiration at night. A model was used to quantify the significance of the findings on estimated canopy CO₂ exchange for the full growing season. Introducing respiration as a function of total photosynthesis on the previous day resulted in a decrease in growing season night‐time respiration by 23% compared with the value when respiration was held constant. This highlights the need for a process‐based approach linking respiration to photosynthesis when modelling long‐term carbon exchange in forest ecosystems.     

On the relationship between water table depth and water vapor and carbon dioxide fluxes in a minerotrophic fen

The focus of this study is the relationship between water table depth (WTD) and water vapor [evapotranspiration (ET)] and carbon dioxide [CO₂; net ecosystem exchange (NEE)] fluxes in a fen in western Canada. We analyzed hydrological and eddy covariance measurements from four snow‐free periods (2003–2006) with contrasting meteorological conditions to establish the link between daily WTD and ET and gross ecosystem CO₂ exchange (GEE) and ecosystem respiration (R_eco; NEE=R_eco?GEE), respectively: 2003 was warm and dry, 2004 was cool and wet, and 2005 and 2006 were both wet. In 2003, the water table (WT) was below the ground surface. In 2004, the WT rose above the ground surface, and in 2005 and 2006, the WT stayed well above the ground surface. There were no significant differences in total ET (～316 mm period^?1), but total NEE was significantly different (2003: 8 g C m^?2 period^?1; 2004: ?139 g C m^?2 period^?1; 2005: ?163 g C m^?2 period^?1; 2006: ?195 g C m^?2 period^?1), mostly due to differences in total GEE (2003: 327 g C m^?2 period^?1; 2004: 513 g C m^?2 period^?1; 2005: 411 g C m^?2 period^?1; 2006: 556 g C m^?2 period^?1). Variation in ET is mostly explained by radiation (67%), and the contribution of WTD is only minor (33%). WTD controls the compensating contributions of different land surface components, resulting in similar total ET regardless of the hydrological conditions. WTD and temperature each contribute about half to the explained variation in GEE up to a threshold ponding depth, below which temperature alone is the key explanatory variable. WTD is only of minor importance for the variation in R_eco, which is mainly controlled by temperature. Our study implies that future peatland modeling efforts explicitly consider topographic and hydrogeological influences on WTD.     

Grassland to shrubland state transitions enhance carbon sequestration in the northern Chihuahuan Desert

The replacement of native C₄‐dominated grassland by C₃‐dominated shrubland is considered an ecological state transition where different ecological communities can exist under similar environmental conditions. These state transitions are occurring globally, and may be exacerbated by climate change. One consequence of the global increase in woody vegetation may be enhanced ecosystem carbon sequestration, although the responses of arid and semiarid ecosystems may be highly variable. During a drier than average period from 2007 to 2011 in the northern Chihuahuan Desert, we found established shrubland to sequester 49 g C m^?2 yr^?1 on average, while nearby native C₄ grassland was a net source of 31 g C m^?2 yr^?1 over this same period. Differences in C exchange between these ecosystems were pronounced – grassland had similar productivity compared to shrubland but experienced higher C efflux via ecosystem respiration, while shrubland was a consistent C sink because of a longer growing season and lower ecosystem respiration. At daily timescales, rates of carbon exchange were more sensitive to soil moisture variation in grassland than shrubland, such that grassland had a net uptake of C when wet but lost C when dry. Thus, even under unfavorable, drier than average climate conditions, the state transition from grassland to shrubland resulted in a substantial increase in terrestrial C sequestration. These results illustrate the inherent tradeoffs in quantifying ecosystem services that result from ecological state transitions, such as shrub encroachment. In this case, the deleterious changes to ecosystem services often linked to grassland to shrubland state transitions may at least be partially offset by increased ecosystem carbon sequestration.     

Topographic and climatic controls on soil respiration in six temperate mixed-hardwood forest slopes, Korea

To better understand the effects of local topography and climate on soil respiration, we conducted field measurements and soil incubation experiments to investigate various factors influencing spatial and temporal variations in soil respiration for six mixed‐hardwood forest slopes in the midst of the Korean Peninsula. Soil respiration and soil water content (SWC) were significantly greater (P=0.09 and 0.003, respectively) on north‐facing slopes compared to south‐facing slopes, while soil temperature was not significantly different between slopes (P>0.5). At all sites, soil temperature was the primary factor driving temporal variations in soil respiration (r²=0.84–0.96) followed by SWC, which accounted for 30% of soil respiration spatial and temporal variability. Results from both field measurements and incubation experiments indicate that variations in soil respiration due to aspect can be explained by a convex‐shaped function relating SWC to normalized soil respiration rates. Annual soil respiration estimates (1070–1246 g C m^?2 yr^?1) were not closely related to mean annual air temperatures among sites from different climate regimes. When soils from each site were incubated at similar temperatures in a laboratory, respiration rates for mineral soils from wetter and cooler sites were significantly higher than those for the drier and warmer sites (n=4, P<0.01). Our results indicate that the application of standard temperature‐based Q₁₀ models to estimate soil respiration rates for larger geographic areas covering different aspects or climatic regimes are not adequate unless other factors, such as SWC and total soil nitrogen, are considered in addition to soil temperature.     

Partitioning of ecosystem respiration of CO2 released during land‐use transition from temperate agricultural grassland to Miscanthus × giganteus

Conversion of large areas of agricultural grassland is inevitable if European and UK domestic production of biomass is to play a significant role in meeting demand. Understanding the impact of these land‐use changes on soil carbon cycling and stocks depends on accurate predictions from well‐parameterized models. Key considerations are cultivation disturbance and the effect of autotrophic root input stimulation on soil carbon decomposition under novel biomass crops. This study presents partitioned parameters from the conversion of semi‐improved grassland to Miscanthus bioenergy production and compares the contribution of autotrophic and heterotrophic respiration to overall ecosystem respiration of CO₂ in the first and second years of establishment. Repeated measures of respiration from within and without root exclusion collars were used to produce time‐series model integrations separating live root inputs from decomposition of grass residues ploughed in with cultivation of the new crop. These parameters were then compared to total ecosystem respiration derived from eddy covariance sensors. Average soil surface respiration was 13.4% higher in the second growing season, increasing from 2.9 to 3.29 g CO₂‐C m^?2 day^?1. Total ecosystem respiration followed a similar trend, increasing from 4.07 to 5.4 g CO₂‐C m^?2 day^?1. Heterotrophic respiration from the root exclusion collars was 32.2% lower in the second growing season at 1.20 g CO₂‐C m^?2 day^?1 compared to the previous year at 1.77 g CO₂‐C m^?2 day^?1. Of the total respiration flux over the two‐year time period, aboveground autotrophic respiration plus litter decomposition contributed 38.46% to total ecosystem respiration while belowground autotrophic respiration and stimulation by live root inputs contributed 46.44% to soil surface respiration. This figure is notably higher than mean figures for nonforest soils derived from the literature and demonstrates the importance of crop‐specific parameterization of respiration models.     

Differential responses of production and respiration to temperature and moisture drive the carbon balance across a climatic gradient in New Mexico

Southwestern North America faces an imminent transition to a warmer, more arid climate, and it is critical to understand how these changes will affect the carbon balance of southwest ecosystems. In order to test our hypothesis that differential responses of production and respiration to temperature and moisture shape the carbon balance across a range of spatio‐temporal scales, we quantified net ecosystem exchange (NEE) of CO₂ and carbon storage across the New Mexico Elevational Gradient, which consists of six eddy‐covariance sites representing biomes ranging from desert to subalpine conifer forest. Within sites, hotter and drier conditions were associated with an increasing advantage of respiration relative to production such that daily carbon uptake peaked at intermediate temperatures – with carbon release often occurring on the hottest days – and increased with soil moisture. Across sites, biotic adaptations modified but did not override the dominant effects of climate. Carbon uptake increased with decreasing temperature and increasing precipitation across the elevational gradient; NEE ranged from a source of ～30 g C m^?2 yr^?1 in the desert grassland to a sink of ～350 g C m^?2 yr^?1 in the subalpine conifer forest. Total aboveground carbon storage increased dramatically with elevation, ranging from 186 g C m^?2 in the desert grassland to 26 600 g C m^?2 in the subalpine conifer forest. These results make sense in the context of global patterns in NEE and biomass storage, and support that increasing temperature and decreasing moisture shift the carbon balance of ecosystems in favor of respiration, such that the potential for ecosystems to sequester and store carbon is reduced under hot and/or dry conditions. This implies that projected climate change will trigger a substantial net release of carbon in these New Mexico ecosystems (～3 Gt CO₂ statewide by the end of the century), thereby acting as a positive feedback to climate change.     

PRODUCTIVITY OF THE FILAMENTOUS ALGA PITHOPHORA OEDOGONIA (CHLOROPHYTA) IN SURREY LAKE,INDIANA1

Biomass, akinete numbers, net photosynthesis, and respiration of Pithophora oedogonia were monitored over two growing seasons in shallow Surrey Lake, Indiana. Low rates of photosynthesis occurred from late fall to early spring and increased to maximum levels in late spring to summer (29–39 mgO₂·g^?1 dry wt·h^?1). Areal biomass increased following the rise in photosynthesis and peaked in autumn (163–206g dry wt·m^?2). Photosynthetic rates were directly correlated with temperature, nitrogen, and phosphorus over the entire annual cycle and during the growing season. Differences in photosynthetic activity and biomass between the two growing seasons (1980 and 1981) were apparently related to higher, early spring temperatures and higher levels of NO₃-N and PO₄-P in 1981. Laboratory investigations of temperature and light effects on Pithophora photosynthesis and respiration indicated that these processes were severely inhibited below 15°C. The highest P_max value occurred at 35°C (0.602 μmol O₂·mg^?1 chl a·min^?1). Rates of dark respiration did not increase above 25°C thus contributing to a favorable balance of photosynthetic production to respiratory utilization at high temperatures. Light was most efficiently utilized at 15°C as indicated by minimum values of I_k(47 μE·m^?2·s^?1) and I_c (6 μE·m^?2·s^?1). Comparison of P. oedogonia and Cladophora glomerata indicated that the former was more tolerant of temperatures above 30°C. Pithophora's tolerance of high temperature and efficient use of low light intensity appear to be adaptive to conditions found within the dense, floating algal mats and the shallow littoral areas inhabited by this filamentous alga.     

Partitioning of river metabolism identifies phytoplankton as a major contributor in the regulated Murray River (Australia)

1. River metabolism was measured over an annual cycle at three sites distributed along a 1000 km length of the lowland Murray River, Australia. 2. Whole system metabolism was measured using water column changes in dissolved oxygen concentrations while planktonic and benthic metabolism were partitioned using light‐dark bottles and benthic chambers. 3. Annual gross primary production (GPP) ranged from 775 to 1126 g O₂ m^?2 year^?1 which in comparison with rivers of similar physical characteristics is moderately productive. 4. Community respiration (CR) ranged from 872 to 1284 g O₂ m^?2 year^?1 so that annual net ecosystem production (NEP) was near zero, suggesting photosynthesis and respiration were balanced and that allochthonous organic carbon played a minor role in fuelling metabolism. 5. Planktonic rates of gross photosynthesis and respiration were similar to those of the total channel, indicating that plankton were responsible for much of the observed metabolism. 6. Respiration rates correlated with phytoplankton standing crop (estimated as the sum of GPP plus the chlorophyll concentration in carbon units), yielding a specific respiration rate of ?1.1 g O₂ g C^?1 day^?1. The respiration rate was equivalent to 19% of the maximum rate of phytoplankton photosynthesis, which is typical of diatoms. 7. The daily GPP per unit phytoplankton biomass correlated with the mean irradiance of the water column giving a constant carbon specific photon fixation rate of 0.35 gO₂ g Chl a^?1 day^?1 per μmole photons m^?2 s^?1 (ca. 0.08 per mole photons m^?2 on a carbon basis) indicating that light availability determined daily primary production. 8. Annual phytoplankton net production (NP) estimates at two sites indicated 25 and 36 g C m^?2 year^?1 were available to support riverine food webs, equivalent to 6% and 11% of annual GPP. 9. Metabolised organic carbon was predominantly derived from phytoplankton and was fully utilised, suggesting that food‐web production was restricted by the energy supply.     

Subalpine grassland carbon dioxide fluxes indicate substantial carbon losses under increased nitrogen deposition,but not at elevated ozone concentration

Ozone (O₃) and nitrogen (N) deposition affect plant carbon (C) dynamics and may change ecosystem C‐sink/‐source properties. We studied effects of increased background [O₃] (up to [ambient] × 2) and increased N deposition (up to +50 kg ha^?1 a^?1) on mature, subalpine grassland during the third treatment year. During 10 days and 13 nights, distributed evenly over the growth period of 2006, we measured ecosystem‐level CO₂ exchange using a static cuvette. Light dependency of gross primary production (GPP) and temperature dependency of ecosystem respiration rates (R_eco) were established. Soil temperature, soil water content, and solar radiation were monitored. Using R_eco and GPP values, we calculated seasonal net ecosystem production (NEP), based on hourly averages of global radiation and soil temperature. Differences in NEP were compared with differences in soil organic C after 5 years of treatment. The high [O₃] had no effect on aboveground dry matter productivity (DM), but seasonal mean rates of both R_eco and GPP decreased ca. 8%. NEP indicated an unaltered growing season CO₂–C balance. High N treatment, with a +31% increase in DM, mean R_eco increased ca. 3%, but GPP decreased ca. 4%. Consequently, seasonal NEP yielded a 53.9 g C m^?2 (±22.05) C loss compared with control. Independent of treatment, we observed a negative NEP of 146.4 g C m^?2 (±15.3). Carbon loss was likely due to a transient management effect, equivalent to a shift from pasture to hay meadow and a drought effect, specific to the 2006 summer climate. We argue that this resulted from strongly intensified soil microbial respiration, following mitigation of nutrient limitation. There was no interaction between O₃ and N treatments. Thus, during the 2006 growing season, the subalpine grassland lost >2% of total topsoil organic C as respired CO₂, with increased N deposition responsible for one‐third of that loss.     

Seasonal CO2 exchange patterns of developing peach (Prunus persica) fruits in response to temperature, light and CO2 concentration

CO₂ exchange rates per unit dry weight, measured in the field on attached fruits of the late-maturing Cal Red peach cultivar, at 1200 μmol photons m^?2S^?1 and in dark, and photosynthetic rates, calculated by the difference between the rates of CO₂ evolution in light and dark, declined over the growing season. Calculated photosynthetic rates per fruit increased over the season with increasing fruit dry matter, but declined in maturing fruits apparently coinciding with the loss of chlorophyll. Slight net fruit photosynthetic rates ranging from 0. 087 ± 0. 06 to 0. 003 ± 0. 05 nmol CO₂ (g dry weight)^?1 S^?1 were measured in midseason under optimal temperature (15 and 20°C) and light (1200 μmol photons m^?2 S^?1) conditions. Calculated fruit photosynthetic rates per unit dry weight increased with increasing temperatures and photon flux densities during fruit development. Dark respiration rates per unit dry weight doubled within a temperature interval of 10°C; the mean seasonal O₁₀ value was 2. 03 between 20 and 30°C. The highest photosynthetic rates were measured at 35°C throughout the growing season. Since dark respiration rates increased at high temperatures to a greater extent than CO₂ exchange rates in light, fruit photosynthesis was apparently stimulated by high internal CO₂ concentrations via CO₂ refixation. At 15°C, fruit photosynthetic rates tended to be saturated at about 600 μmol photons m^?2 S^?1. Young peach fruits responded to increasing ambient CO₂ concentrations with decreasing net CO₂ exchange rates in light, but more mature fruits did not respond to increases in ambient CO₂. Fruit CO₂ exchange rates in the dark remained fairly constant, apparently uninfluenced by ambient CO₂ concentrations during the entire growing season. Calculated fruit photosynthetic rates clearly revealed the difference in CO₂ response of young and mature peach fruits. Photosynthetic rates of younger peach fruits apparently approached saturation at 370 μl CO₂1^?2. In CO₂ free air, fruit photosynthesis was dependent on CO₂ refixation since CO₂ uptake by the fruits from the external atmosphere was not possible. The difference in photosynthetic rates between fruits in CO₂-free air and 370 μl CO₂ 1^?1 indicated that young peach fruits were apparently able to take up CO₂ from the external atmosphere. CO₂ uptake by peach fruits contributed between 28 and 16% to the fruit photosynthetic rate early in the season, whereas photosynthesis in maturing fruits was supplied entirely by CO₂ refixation.     

Carbon sequestration in a high-elevation, subalpine forest

We studied net ecosystem CO₂ exchange (NEE) dynamics in a high‐elevation, subalpine forest in Colorado, USA, over a two‐year period. Annual carbon sequestration for the forest was 6.71 mol C m^?2 (80.5 g C m^?2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m^?2 (57.6 g C m^?2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO₂ uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing‐season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid‐summer moisture was adequate (as in the first year of the study) or lower mid‐day photosynthesis rates when mid‐summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO₂ uptake during both the first 30 days of the growing season and the mid‐summer months. The reduction in CO₂ uptake during the first 30 days of the second year was due to an earlier‐than‐normal spring warm‐up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO₂ uptake during the mid‐summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day‐to‐day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather. The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier‐than‐normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm‐up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm‐up. Furthermore, the strong influence of mid‐summer precipitation on CO₂ uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.     

LIGHT AND TEMPERATURE AS FACTORS REGULATING SEASONAL GROWTH AND DISTRIBUTION OF ULOTHRIX ZONATA (ULVOPHYCEAE)1

Ulothrix zonata (Weber and Mohr) Kütz. is an unbranched filamentous green alga found in rocky littoral areas of many northern lakes. Field observations of its seasonal and spatial distribution indicated that it should have a low temperature and a high irradiance optimum for net photosynthesis, and at temperatures above 10°C it should show an increasingly unfavorable energy balance. Measurements of net photosynthesis and respiration were made at 56 combinations of light and temperature. Optimum conditions were 5°C and 1100 μE·m^?2·s^?1 at which net photosynthesis was 16.8 mg O₂·g^?1·h^?1. As temperature increased above 5° C optimum irradiance decreased to 125 μE·m^?2·s^?1 at 30°C. Respiration rates increased with both temperature and prior irradiance. Light-enhanced respiration rates were significantly greater than dark respiration rates following irradiance exposures of 125 μE·m^?2·s^?1 or greater. Polynomials were fitted to the data to generate response surfaces. Polynomial equations represent statistical models which can accurately predict photosynthesis and respiration for inclusion in ecosystem models.     

Variation in the carbon and oxygen isotope composition of plant biomass and its relationship to water‐use efficiency at the leaf‐ and ecosystem‐scales in a northern Great Plains grassland

Measurements of the carbon (δ¹³C_m) and oxygen (δ¹⁸O_m) isotope composition of C₃ plant tissue provide important insights into controls on water‐use efficiency. We investigated the causes of seasonal and inter‐annual variability in water‐use efficiency in a grassland near Lethbridge, Canada using stable isotope (leaf‐scale) and eddy covariance measurements (ecosystem‐scale). The positive relationship between δ¹³C_m and δ¹⁸O_m values for samples collected during 1998–2001 indicated that variation in stomatal conductance and water stress‐induced changes in the degree of stomatal limitation of net photosynthesis were the major controls on variation in δ¹³C_m and biomass production during this time. By comparison, the lack of a significant relationship between δ¹³C_m and δ¹⁸O_m values during 2002, 2003 and 2006 demonstrated that water stress was not a significant limitation on photosynthesis and biomass production in these years. Water‐use efficiency was higher in 2000 than 1999, consistent with expectations because of greater stomatal limitation of photosynthesis and lower leaf c_i/ca during the drier conditions of 2000. Calculated values of leaf‐scale water‐use efficiency were 2–3 times higher than ecosystem‐scale water‐use efficiency, a difference that was likely due to carbon lost in root respiration and water lost during soil evaporation that was not accounted for by the stable isotope measurements.     

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.