Wind as a main driver of the net ecosystem carbon balance of a semiarid Mediterranean steppe in the South East of Spain

Despite the advance in our understanding of the carbon exchange between terrestrial ecosystems and the atmosphere, semiarid ecosystems have been poorly investigated and little is known about their role in the global carbon balance. We used eddy covariance measurements to determine the exchange of CO₂ between a semiarid steppe and the atmosphere over 3 years. The vegetation is a perennial grassland of Stipa tenacissima L. located in the SE of Spain. We examined diurnal, seasonal and interannual variations in the net ecosystem carbon balance (NECB) in relation to biophysical variables. Cumulative NECB was a net source of 65.7, 143.6 and 92.1 g C m^?2 yr^?1 for the 3 years studied, respectively. We separated the year into two distinctive periods: dry period and growing season. The ecosystem was a net source of CO₂ to the atmosphere, particularly during the dry period when large CO₂ positive fluxes of up to 15 μmol m^?2 s^?1 were observed in concomitance with large wind speeds. Over the growing season, the ecosystem was a slight sink or neutral with maximum rates of ?2.3 μmol m^?2 s^?1. Rainfall events caused large fluxes of CO₂ to the atmosphere and determined the length of the growing season. In this season, photosynthetic photon flux density controlled day‐time NECB just below 1000 μmol m^?2 s^?1. The analyses of the diurnal and seasonal data and preliminary geological and gas‐geochemical evaluations, including C isotopic analyses, suggest that the CO₂ released was not only biogenic but most likely included a component of geothermal origin, presumably related to deep fluids occurring in the area. These results highlight the importance of considering geological carbon sources, as well as the need to carefully interpret the results of eddy covariance partitioning techniques when applied in geologically active areas potentially affected by CO₂‐rich geofluid circulation.     

Respiration acclimation contributes to high carbon-use efficiency in a seasonally dry pine forest

Predictions of warming and drying in the Mediterranean and other regions require quantifying of such effects on ecosystem carbon dynamics and respiration. Long‐term effects can only be obtained from forests in which seasonal drought is a regular feature. We carried out measurements in a semiarid Pinus halepensis (Aleppo pine) forest of aboveground respiration rates of foliage, R_f, and stem, R_t over 3 years. Component respiration combined with ongoing biometric, net CO₂ flux [net ecosystem productivity (NEP)] and soil respiration measurements were scaled to the ecosystem level to estimate gross and net primary productivity (GPP, NPP) and carbon‐use efficiency (CUE=NPP/GPP) using 6 years data. GPP, NPP and NEP were, on average, 880, 350 and 211 g C m^?2 yr^?1, respectively. The above ground respiration made up half of total ecosystem respiration but CUE remained high at 0.4. Large seasonal variations in both R_f and R_t were not consistently correlated with seasonal temperature trends. Seasonal adjustments of respiration were observed in both the normalized rate (R₂₀) and short‐term temperature sensitivity (Q₁₀), resulting in low respiration rates during the hot, dry period. R_f in fully developed needles was highest over winter–spring, and foliage R₂₀ was correlated with photosynthesis over the year. Needle growth occurred over summer, with respiration rates in developing needles higher than the fully developed foliage at most times. R_t showed a distinct seasonal maximum in May irrespective of year, which was not correlated to the winter stem growth, but could be associated with phenological drivers such as carbohydrate re‐mobilization and cambial activity. We show that in a semiarid pine forest photosynthesis and stem growth peak in (wet) winter and leaf growth in (dry) summer, and associated adjustments of component respiration, dominated by those in R₂₀, minimize annual respiratory losses. This is likely a key for maintaining high CUE and ecosystem productivity similar to much wetter sites, and could lead to different predictions of the effect of warming and drying climate on productivity of pine forests than based on short‐term droughts.     

基于FORCCHN模型的中国典型森林生态系统碳通量模拟

森林生态系统是陆地碳循环的重要组成部分,其固碳能力显著高于其他陆地生态系统,研究森林生态系统碳通量是认识和理解全球变化对碳循环影响的关键。碳循环模型是研究森林生态系统碳通量有效工具。以长白山温带落叶阔叶林、千烟洲亚热带常绿针叶林、鼎湖山亚热带常绿阔叶林和西双版纳热带雨林等4种中国典型森林生态系统为研究对象,利用涡度相关2003-2012年观测数据,评估FORCCHN模型对生态系统呼吸（ER）,总初级生产力（GPP）,净生态系统生产力（NEP）的模型效果。结果表明：（1） FORCCHN模型能够较好的模拟中国4种典型森林生态系统不同时间尺度的碳通量。落叶阔叶林和常绿针叶林ER和GPP的逐日变化模拟效果较好（ER的相关系数分别为0.94和0.92,GPP的相关系数分别为0.86和0.74）;（2）4种森林生态系统碳通量季节动态模拟值和观测值显著相关（P<0.01）,ER、GPP、NEP的观测值和模拟值的R²分别为0.77-0.93、0.54-0.88和0.15-0.38;模型可以很好地模拟森林生态系统不同季节碳汇（NEP>0）,碳源（NEP<0）的变化规律;（3）4种森林生态系统碳通量模拟值与观测值的年际变化有很好的吻合度,但在数值大小上存在差异,模型高估了常绿阔叶林的ER和GPP,略微低估了其他3种森林生态系统ER和GPP。     

Response of carbon fluxes to drought in a coastal plain loblolly pine forest

Full accounting of ecosystem carbon (C) pools and fluxes in coastal plain ecosystems remains less studied compared with upland systems, even though the C stocks in these systems may be up to an order of magnitude higher, making them a potentially important component in regional C cycle. Here, we report C pools and CO₂ exchange rates during three hydrologically contrasting years (i.e. 2005–2007) in a coastal plain loblolly pine plantation in North Carolina, USA. The daily temperatures were similar among the study years and to the long‐term (1971–2000) average, whereas the amount and timing of precipitation differed significantly. Precipitation was the largest in 2005 (147 mm above normal), intermediate in 2006 (48 mm below) and lowest in 2007 (486 mm below normal). The forest was a strong C sink during all years, sequestering 361 ± 67 (2005), 835 ± 55 (2006) and 724 ± 55 (2007) g C m^?2 yr^?1 according to eddy covariance measurements of net ecosystem CO₂ exchange (NEE). The interannual differences in NEE were traced to drought‐induced declines in canopy and whole tree hydraulic conductances, which declined with growing precipitation deficit and decreasing soil volumetric water content (VWC). In contrast, the interannual differences were small in gross ecosystem productivity (GEP) and ecosystem respiration (ER), both seemingly insensitive to drought. However, the drought sensitivity of GEP was masked by higher leaf area index and higher photosynthetically active radiation during the dry year. Normalizing GEP by these factors enhanced interannual differences, but there were no signs of suppressed GEP at low VWC during any given year. Although ER was very consistent across the 3 years, and not suppressed by low VWC, the total respiratory cost as a fraction of net primary production increased with annual precipitation and the contribution of heterotrophic respiration (R_h) was significantly higher during the wettest year, exceeding new litter inputs by 58%. Although the difference was smaller during the other 2 years (R_h : litterfall ratio was 1.05 in 2006 and 1.10 in 2007), the soils lost about 109 g C m^?2 yr^?1, outlining their potential vulnerability to decomposition, and pointing to potential management considerations to protect existing soil C stocks.     

Long-term carbon exchange in a sparse, seasonally dry tussock grassland

Rainfall and its seasonal distribution can alter carbon dioxide (CO₂) exchange and the sustainability of grassland ecosystems. Using eddy covariance, CO₂ exchange between the atmosphere and a sparse grassland was measured for 2 years at Twizel, New Zealand. The years had contrasting distributions of rain and falls (446 mm followed by 933 mm; long‐term mean=646 mm). The vegetation was sparse with total above‐ground biomass of only 1410 g m^?2. During the dry year, leaf area index peaked in spring (November) at 0.7, but it was <0.2 by early summer. The maximum daily net CO₂ uptake rate was only 1.5 g C m^?2 day^?1, and it occurred before mid‐summer in both years. On an annual basis, for the dry year, 9 g C m^?2 was lost to the atmosphere. During the wet year, 41 g C m^?2 was sequestered from the atmosphere. The net exchange rates were determined mostly by the timing and intensity of spring rainfall. The components of ecosystem respiration were measured using chambers. Combining scaled‐up measurements with the eddy CO₂ effluxes, it was estimated that 85% of ecosystem respiration emanated from the soil surface. Under well‐watered conditions, 26% of the soil surface CO₂ efflux came from soil microbial activity. Rates of soil microbial CO₂ production and net mineral‐N production were low and indicative of substrate limitation. Soil respiration declined by a factor of four as the soil water content declined from field capacity (0.21 m³ m^?3) to the driest value obtained (0.04 m³ m^?3). Rainfall after periods of drought resulted in large, but short‐lived, respiration pulses that were curvilinearly related to the increase in root‐zone water content. Coupled with the low leaf area and high root : shoot ratio, this sparse grassland had a limited capacity to sequester and store carbon. Assuming a proportionality between carbon gain and rainfall during the summer, rainfall distribution statistics suggest that the ecosystem is sustainable in the long term.     

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).     

Net ecosystem carbon dioxide exchange over grazed steppe in central Mongolia

This paper presents results of 1 year (from March 25, 2003 to March 24, 2004, 366 days) of continuous measurements of net ecosystem CO₂ exchange (NEE) above a steppe in Mongolia using the eddy covariance technique. The steppe, typical of central Mongolia, is dominated by C₃ plants adapted to the continental climate. The following two questions are addressed: (1) how do NEE and its components: gross ecosystem production (GEP) and total ecosystem respiration (R_eco) vary seasonally? (2) how do NEE, GEP, and R_eco respond to biotic and abiotic factors? The hourly minimal NEE and the hourly maximal R_eco were −3.6 and 1.2 μmol m⁻² s⁻¹, respectively (negative values denoting net carbon uptake by the canopy from the atmosphere). Peak daily sums of NEE, GEP, and R_eco were −2.3, 3.5, and 1.5 g C m⁻² day⁻¹, respectively. The annual sums of GEP, R_eco, and NEE were 179, 138, and −41 g C m⁻², respectively. The carbon removal by sheep was estimated to range between 10 and 82 g C m⁻² yr⁻¹ using four different approaches. Including these estimates in the overall carbon budget yielded net ecosystem productivity of −23 to +20 g C m⁻² yr⁻¹. Thus, within the remaining experimental uncertainty the carbon budget at this steppe site can be considered to be balanced. For the growing period (from April 23 to October 21, 2003), 26% and 53% of the variation in daily NEE and GEP, respectively, could be explained by the changes in leaf area index. Seasonality of GEP, R_eco, and NEE was closely associated with precipitation, especially in the peak growing season when GEP and R_eco were largest. Water stress was observed in late July to early August, which switched the steppe from a carbon sink to a carbon source. For the entire growing period, the light response curves of daytime NEE showed a rather low apparent quantum yield (α=−0.0047 μmol CO₂ μmol⁻¹ photons of photosynthetically active radiation). However, the α values varied with air temperature (T_a), vapor pressure deficit, and soil water content.     

Carbon dioxide exchange above a Mediterranean C3/C4 grassland during two climatologically contrasting years

Eddy‐covariance measurements of net ecosystem carbon exchange (NEE) were carried out above a grazed Mediterranean C3/C4 grassland in southern Portugal, during two hydrological years, 2004–2005 and 2005–2006, of contrasting rainfall. Here, we examine the seasonal and interannual variation in NEE and its major components, gross primary production (GPP) and ecosystem respiration (R_eco), in terms of the relevant biophysical controls. The first hydrological year was dry, with total precipitation 45% below the long‐term mean (669 mm) and the second was normal, with total precipitation only 12% above the long‐term mean. The drought conditions during the winter and early spring of the dry year limited grass production and the leaf area index (LAI) was very low. Hence, during the peak of the growth period, the maximum daily rate of NEE and the light‐use and water‐use efficiencies were approximately half of those observed in the normal year. In the summer of 2006, the warm‐season C4 grass, Cynodon dactylon L., exerted an evident positive effect on NEE by converting the ecosystem into a carbon sink after strong rain events and extending the carbon sequestration for several days, after the end of senescence of the C3 grasses. On an annual basis, the GPP and NEE were 524 and 49 g C m^?2, respectively, for the dry year, and 1261 and ?190 g C m^?2 for the normal year. Therefore, the grassland was a moderate net source of carbon to the atmosphere, in the dry year, and a considerable net carbon sink, in the normal year. In these 2 years of experiment the total amount of precipitation was the main factor determining the interannual variation in NEE. In terms of relevant controls, GPP and NEE were strongly related to incident photosynthetic photon flux density on short‐term time scales. Changes in LAI explained 84% and 77% of the variation found in GPP and NEE, respectively. Variations in R_eco were mainly controlled by canopy photosynthesis. After each grazing event, the reduction in LAI affected negatively the NEE.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Effects of simulated drought on the carbon balance of Everglades short‐hydroperiod marsh

Hydrology drives the carbon balance of wetlands by controlling the uptake and release of CO₂ and CH₄. Longer dry periods in between heavier precipitation events predicted for the Everglades region, may alter the stability of large carbon pools in this wetland's ecosystems. To determine the effects of drought on CO₂ fluxes and CH₄ emissions, we simulated changes in hydroperiod with three scenarios that differed in the onset rate of drought (gradual, intermediate, and rapid transition into drought) on 18 freshwater wetland monoliths collected from an Everglades short‐hydroperiod marsh. Simulated drought, regardless of the onset rate, resulted in higher net CO₂ losses net ecosystem exchange (NEE) over the 22‐week manipulation. Drought caused extensive vegetation dieback, increased ecosystem respiration (R_eco), and reduced carbon uptake gross ecosystem exchange (GEE). Photosynthetic potential measured by reflective indices (photochemical reflectance index, water index, normalized phaeophytinization index, and the normalized difference vegetation index) indicated that water stress limited GEE and inhibited R_eco. As a result of drought‐induced dieback, NEE did not offset methane production during periods of inundation. The average ratio of net CH₄ to NEE over the study period was 0.06, surpassing the 100‐year greenhouse warming compensation point for CH₄ (0.04). Drought‐induced diebacks of sawgrass (C₃) led to the establishment of the invasive species torpedograss (C₄) when water was resupplied. These changes in the structure and function indicate that freshwater marsh ecosystems can become a net source of CO₂ and CH₄ to the atmosphere, even following an extended drought. Future changes in precipitation patterns and drought occurrence/duration can change the carbon storage capacity of freshwater marshes from sinks to sources of carbon to the atmosphere. Therefore, climate change will impact the carbon storage capacity of freshwater marshes by influencing water availability and the potential for positive feedbacks on radiative forcing.     

Productivity of forests in the Eurosiberian boreal region and their potential to act as a carbon sink –- a synthesis

Based on review and original data, this synthesis investigates carbon pools and fluxes of Siberian and European forests (600 and 300 million ha, respectively). We examine the productivity of ecosystems, expressed as positive rate when the amount of carbon in the ecosystem increases, while (following micrometeorological convention) downward fluxes from the atmosphere to the vegetation (NEE = Net Ecosystem Exchange) are expressed as negative numbers. Productivity parameters are Net Primary Productivity (NPP=whole plant growth), Net Ecosystem Productivity (NEP = CO₂ assimilation minus ecosystem respiration), and Net Biome Productivity (NBP = NEP minus carbon losses through disturbances bypassing respiration, e.g. by fire and logging). Based on chronosequence studies and national forestry statistics we estimate a low average NPP for boreal forests in Siberia: 123 gC m^–2 y^–1. This contrasts with a similar calculation for Europe which suggests a much higher average NPP of 460 gC m^–2 y^–1 for the forests there. Despite a smaller area, European forests have a higher total NPP than Siberia (1.2–1.6 vs. 0.6–0.9 × 10¹⁵ gC region^–1 y^–1). This arises as a consequence of differences in growing season length, climate and nutrition. For a chronosequence of Pinus sylvestris stands studied in central Siberia during summer, NEE was most negative in a 67-y old stand regenerating after fire (– 192 mmol m^–2 d^–1) which is close to NEE in a cultivated forest of Germany (– 210 mmol m^–2 d^–1). Considerable net ecosystem CO₂-uptake was also measured in Siberia in 200- and 215-y old stands (NEE:174 and – 63 mmol m^–2 d^–1) while NEP of 7- and 13-y old logging areas were close to the ecosystem compensation point. Two Siberian bogs and a bog in European Russia were also significant carbon sinks (– 102 to – 104 mmol m^–2 d^–1). Integrated over a growing season (June to September) we measured a total growing season NEE of – 14 mol m^–2 summer^–1 (– 168 gC m^–2 summer^–1) in a 200-y Siberian pine stand and – 5 mol m^–2 summer^–1 (– 60 gC m^–2 summer^–1) in Siberian and European Russian bogs. By contrast, over the same period, a spruce forest in European Russia was a carbon source to the atmosphere of (NEE: + 7 mol m^–2 summer^–1 = + 84 gC m^–2 summer^–1). Two years after a windthrow in European Russia, with all trees being uplifted and few successional species, lost 16 mol C m^–2 to the atmosphere over a 3-month in summer, compared to the cumulative NEE over a growing season in a German forest of – 15.5 mol m^–2 summer^–1 (– 186 gC m^–2 summer^–1; European flux network annual averaged – 205 gC m^–2 y^–1). Differences in CO₂-exchange rates coincided with differences in the Bowen ratio, with logging areas partitioning most incoming radiation into sensible heat whereas bogs partitioned most into evaporation (latent heat). Effects of these different surface energy exchanges on local climate (convective storms and fires) and comparisons with the Canadian BOREAS experiment are discussed. Following a classification of disturbances and their effects on ecosystem carbon balances, fire and logging are discussed as the main processes causing carbon losses that bypass heterotrophic respiration in Siberia. Following two approaches, NBP was estimated to be only about 13–16 mmol m^–2 y^–1 for Siberia. It may reach 67 mmol m^–2 y^–1 in North America, and about 140–400 mmol m^–2 y^–1 in Scandinavia. We conclude that fire speeds up the carbon cycle, but that it results also in long-term carbon sequestration by charcoal formation. For at least 14 years after logging, regrowth forests remain net sources of CO₂ to the atmosphere. This has important implications regarding the effects of Siberian forest management on atmospheric concentrations. For many years after logging has taken place, regrowth forests remain weaker sinks for atmospheric CO₂ than are nearby old-growth forests.     

Consequences of wildfire on ecosystem CO2 and water vapour fluxes in the Great Basin

Increased fire frequency in the Great Basin of North America's intermountain West has led to large‐scale conversion of native sagebrush (Artemisia tridentata Nutt.) communities to postfire successional communities dominated by native and non‐native annual species during the last century. The consequences of this conversion for basic ecosystem functions, however, are poorly understood. We measured net ecosystem CO₂ exchange (NEE) and evapotranspiration (ET) during the first two dry years after wildfire using a 4‐m diameter (16.4 m³) translucent static chamber (dome), and found that both NEE and ET were higher in a postfire successional ecosystem (?0.9–2.6 µ mol CO₂ m^?2 s^?1 and 0.0–1.0 mmol H₂O m^?2 s^?2, respectively) than in an adjacent intact sagebrush ecosystem (?1.2–2.3 µ mol CO₂ m^?2 s^?1 and ?0.1–0.8 mmol H₂O m^?2 s^?2, respectively) during relatively moist periods. Higher NEE in the postfire ecosystem appears to be due to lower rates of above‐ground plant respiration while higher ET appears to be caused by higher surface soil temperatures and increased soil water recharge after rains. These patterns disappeared or were reversed, however, when the conditions were drier. Daily net ecosystem productivity (NEP; g C m^?2 d^?1), derived from multiple linear regressions of measured fluxes with continuously measured climate variables, was very small (close to zero) throughout most of the year. The wintertime was an exception in the intact sagebrush ecosystem with C losses exceeding C gains leading to negative NEP while C balance of the postfire ecosystem remained near zero. Taken together, our results indicate that wildfire‐induced conversion of native sagebrush steppe to ecosystems dominated by herbaceous annual species may have little effect on C balance during relatively dry years (except in winter months) but may stimulate water loss immediately following fires.     

Productivity overshadows temperature in determining soil and ecosystem respiration across European forests

This paper presents CO₂ flux data from 18 forest ecosystems, studied in the European Union funded EUROFLUX project. Overall, mean annual gross primary productivity (GPP, the total amount of carbon (C) fixed during photosynthesis) of these forests was 1380 ± 330 gC m^?2 y^?1 (mean ±SD). On average, 80% of GPP was respired by autotrophs and heterotrophs and released back into the atmosphere (total ecosystem respiration, TER = 1100 ± 260 gC m^?2 y^?1). Mean annual soil respiration (SR) was 760 ± 340 gC m^?2 y^?1 (55% of GPP and 69% of TER). Among the investigated forests, large differences were observed in annual SR and TER that were not correlated with mean annual temperature. However, a significant correlation was observed between annual SR and TER and GPP among the relatively undisturbed forests. On the assumption that (i) root respiration is constrained by the allocation of photosynthates to the roots, which is coupled to productivity, and that (ii) the largest fraction of heterotrophic soil respiration originates from decomposition of young organic matter (leaves, fine roots), whose availability also depends on primary productivity, it is hypothesized that differences in SR among forests are likely to depend more on productivity than on temperature. At sites where soil disturbance has occurred (e.g. ploughing, drainage), soil espiration was a larger component of the ecosystem C budget and deviated from the relationship between annual SR (and TER) and GPP observed among the less‐disturbed forests. At one particular forest, carbon losses from the soil were so large, that in some years the site became a net source of carbon to the atmosphere. Excluding the disturbed sites from the present analysis reduced mean SR to 660 ± 290 gC m^?2 y^?1, representing 49% of GPP and 63% of TER in the relatively undisturbed forest ecosystems.     

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.     

Changes in carbon storage and fluxes in a chronosequence of ponderosa pine

Forest development following stand‐replacing disturbance influences a variety of ecosystem processes including carbon exchange with the atmosphere. On a series of ponderosa pine (Pinius ponderosa var. Laws.) stands ranging from 9 to> 300 years in central Oregon, USA, we used biological measurements to estimate carbon storage in vegetation and soil pools, net primary productivity (NPP) and net ecosystem productivity (NEP) to examine variation with stand age. Measurements were made on plots representing four age classes with three replications: initiation (I, 9–23 years), young (Y, 56–89 years), mature (M, 95–106 years), and old (O, 190–316 years) stands typical of the forest type in the region. Net ecosystem productivity was lowest in the I stands (?124 g C m^?2 yr^?1), moderate in Y stands (118 g C m^?2 yr^?1), highest in M stands (170 g C m^?2 yr^?1), and low in the O stands (35 g C m^?2 yr^?1). Net primary productivity followed similar trends, but did not decline as much in the O stands. The ratio of fine root to foliage carbon was highest in the I stands, which is likely necessary for establishment in the semiarid environment, where forests are subject to drought during the growing season (300–800 mm precipitation per year). Carbon storage in live mass was the highest in the O stands (mean 17.6 kg C m^?2). Total ecosystem carbon storage and the fraction of ecosystem carbon in aboveground wood mass increased rapidly until 150–200 years, and did not decline in older stands. Forest inventory data on 950 ponderosa pine plots in Oregon show that the greatest proportion of plots exist in stands ～ 100 years old, indicating that a majority of stands are approaching maximum carbon storage and net carbon uptake. Our data suggests that NEP averages ～ 70 g C m^?2 year^?1 for ponderosa pine forests in Oregon. About 85% of the total carbon storage in biomass on the survey plots exists in stands greater than 100 years, which has implications for managing forests for carbon sequestration. To investigate variation in carbon storage and fluxes with disturbance, simulation with process models requires a dynamic parameterization for biomass allocation that depends on stand age, and should include a representation of competition between multiple plant functional types for space, water, and nutrients.     

Simulated annual carbon fluxes of grassland ecosystems in extremely arid conditions

In order to understand how changes in climate and land cover affect carbon cycles and structure and function of regional grassland ecosystems, we developed a grassland landscape productivity model, proposed an approach that combined both process-based modeling and spatial analysis with field measurements, and provided an example of semiarid region in Inner Mongolia, China, in extremely arid conditions. The modeled monthly mean and total net primary productivity, and monthly and annual mean respiration rates for Leymus chinensis steppes during the growing seasons in 2002 were mostly within the normal varying ranges of measured values, or similar to the field measurements, conducted in the similarly arid conditions. And the modeled total net ecosystem productivity (NEP) for L. chinensis steppes and Stipa grandis steppes were both between the lower and the higher measurements or within modeled multi-annual data by the other model. The modeled total NEP was 1.91 g C/m²/year over the entire study region. It indicated that if human disturbances were not considered, carbon budget over the entire study region during the growing seasons was nearly in balance or weak carbon sink even under extremely arid conditions. However, the modeled NEP spatially greatly varied not only over the entire study region (−48.28–52.09 g C/m²/year), but also among different land cover types. The modeled results also showed that there were obvious seasonal variations in carbon fluxes, mainly caused by leaf area index; and annual precipitation was the major limiting factor for the obvious spatial patterns of carbon fluxes from the east to the west. The modeled results also revealed the influence of extreme drought on carbon fluxes. The study provides an effective approach to derive useful information about carbon fluxes for different land cover types, and thus can instruct regional land-use planning and resource management based on carbon storage conditions.     

CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Carbon dioxide and water vapor exchange in a warm temperate grassland

Grasslands cover about 40% of the ice-free global terrestrial surface, but their contribution to local and regional water and carbon fluxes and sensitivity to climatic perturbations such as drought remains uncertain. Here, we assess the direction and magnitude of net ecosystem carbon exchange (NEE) and its components, ecosystem carbon assimilation (A _c) and ecosystem respiration (R _E), in a southeastern United States grassland ecosystem subject to periodic drought and harvest using a combination of eddy-covariance measurements and model calculations. We modeled A _c and evapotranspiration (ET) using a big-leaf canopy scheme in conjunction with ecophysiological and radiative transfer principles, and applied the model to assess the sensitivity of NEE and ET to soil moisture dynamics and rapid excursions in leaf area index (LAI) following grass harvesting. Model results closely match eddy-covariance flux estimations on daily, and longer, time steps. Both model calculations and eddy-covariance estimates suggest that the grassland became a net source of carbon to the atmosphere immediately following the harvest, but a rapid recovery in LAI maintained a marginal carbon sink during summer. However, when integrated over the year, this grassland ecosystem was a net C source (97 g C m^–2 a^–1) due to a minor imbalance between large A _c (–1,202 g C m^–2 a^–1) and R _E (1,299 g C m^–2 a^–1) fluxes. Mild drought conditions during the measurement period resulted in many instances of low soil moisture (<0.2 m³m^–3), which influenced A _c and thereby NEE by decreasing stomatal conductance. For this experiment, low had minor impact on R _E. Thus, stomatal limitations to A _c were the primary reason that this grassland was a net C source. In the absence of soil moisture limitations, model calculations suggest a net C sink of –65 g C m^–2 a^–1 assuming the LAI dynamics and physiological properties are unaltered. These results, and the results of other studies, suggest that perturbations to the hydrologic cycle are key determinants of C cycling in grassland ecosystems.     

Diurnal, seasonal and annual variation in net ecosystem CO2 exchange of an alpine shrubland on Qinghai-Tibetan plateau

Thus far, grassland ecosystem research has mainly been focused on low‐lying grassland areas, whereas research on high‐altitude grassland areas, especially on the carbon budget of remote areas like the Qinghai‐Tibetan plateau is insufficient. To address this issue, flux of CO₂ were measured over an alpine shrubland ecosystem (37°36′N, 101°18′E; 325 above sea level [a. s. l.]) on the Qinghai‐Tibetan Plateau, China, for 2 years (2003 and 2004) with the eddy covariance method. The vegetation is dominated by formation Potentilla fruticosa L. The soil is Mol–Cryic Cambisols. To interpret the biotic and abiotic factors that modulate CO₂ flux over the course of a year we decomposed net ecosystem CO₂ exchange (NEE) into its constituent components, and ecosystem respiration (R_eco). Results showed that seasonal trends of annual total biomass and NEE followed closely the change in leaf area index. Integrated NEE were ?58.5 and ?75.5 g C m^?2, respectively, for the 2003 and 2004 years. Carbon uptake was mainly attributed from June, July, August, and September of the growing season. In July, NEE reached seasonal peaks of similar magnitude (4–5 g C m^?2 day^?1) each of the 2 years. Also, the integrated night‐time NEE reached comparable peak values (1.5–2 g C m^?2 day^?1) in the 2 years of study. Despite the large difference in time between carbon uptake and release (carbon uptake time < release time), the alpine shrubland was carbon sink. This is probably because the ecosystem respiration at our site was confined significantly by low temperature and small biomass and large day/night temperature difference and usually soil moisture was not limiting factor for carbon uptake. In general, R_eco was an exponential function of soil temperature, but with season‐dependent values of Q₁₀. The temperature‐dependent respiration model failed immediately after rain events, when large pulses of R_eco were observed. Thus, for this alpine shrubland in Qinghai‐Tibetan plateau, the timing of rain events had more impact than the total amount of precipitation on ecosystem R_eco and NEE.     

干旱对陆地生态系统生产力的影响

该文综述了干旱对陆地生态系统生产力的影响,分析了其影响机制,并总结了植被对干旱的响应与适应及其机理机制。干旱通过抑制光合作用来降低陆地生态系统总初级生产力,干旱还可以降低生态系统的自养呼吸和异养呼吸。同时干旱还可以通过影响其它干扰形式来间接影响陆地生态系统生产力,如增加火干扰的发生频率和强度,增加植物的死亡率,增加病虫害的发生等。在生态系统水平上干旱可以降低碳固定,减弱碳汇功能,甚至把生态系统从碳汇改变成碳源。目前生态系统水平上的干旱影响研究主要通过两种方法实现,一种是模型模拟,另一种就是大型模拟实验。作为陆地生态系统生产力的实现者,在干旱胁迫条件下,植物也会采取积极的适应策略以减弱干旱对生态系统生产力的影响,其适应策略主要分以下3种:在一些周期性发生干旱的地区,植物会调整生长期以避开干旱或通过休眠来减弱干旱所造成的伤害;还有一些植物会通过调节体内的代谢过程,改变一些生理特性来抵御干旱;而长期生活在干旱条件下的植物则通过进化来改变了自身的生理生化代谢过程,形成耐旱机制。目前,植物对干旱响应的分子学机制,以及生态系统水平上对干旱的响应和适应仍然是薄弱的领域,也必然成为未来研究的重点。