Effects of Elevated CO2 and Nitrogen Deposition on Ecosystem Carbon Fluxes on the Sanjiang Plain Wetland in Northeast China

Background

Increasing atmospheric CO₂ and nitrogen (N) deposition across the globe may affect ecosystem CO₂ exchanges and ecosystem carbon cycles. Additionally, it remains unknown how increased N deposition and N addition will alter the effects of elevated CO₂ on wetland ecosystem carbon fluxes.

Methodology/Principal Findings

Beginning in 2010, a paired, nested manipulative experimental design was used in a temperate wetland of northeastern China. The primary factor was elevated CO₂, accomplished using Open Top Chambers, and N supplied as NH₄NO₃ was the secondary factor. Gross primary productivity (GPP) was higher than ecosystem respiration (ER), leading to net carbon uptake (measured by net ecosystem CO₂ exchange, or NEE) in all four treatments over the growing season. However, their magnitude had interannual variations, which coincided with air temperature in the early growing season, with the soil temperature and with the vegetation cover. Elevated CO₂ significantly enhanced GPP and ER but overall reduced NEE because the stimulation caused by the elevated CO₂ had a greater impact on ER than on GPP. The addition of N stimulated ecosystem C fluxes in both years and ameliorated the negative impact of elevated CO₂ on NEE.

Conclusion/Significance

In this ecosystem, future elevated CO₂ may favor carbon sequestration when coupled with increasing nitrogen deposition.     

Carbon dioxide exchange over multiple temporal scales in an arid shrub ecosystem near La Paz,Baja California Sur,Mexico

Arid environments represent 30% of the global terrestrial surface, but are largely under‐represented in studies of ecosystem carbon flux. Less than 2% of all FLUXNET eddy covariance sites exist in a hot desert climate. Long‐term datasets of these regions are vital for capturing the seasonal and interannual variability that occur due to episodic precipitation events and climate change, which drive fluctuations in soil moisture and temperature patterns. The objectives of this study were to determine the meteorological variables that drive carbon flux on diel, seasonal, and annual scales and to determine how precipitation events control annual net ecosystem exchange (NEE). Patterns of NEE from 2002 to 2008 were investigated, providing a record with multiple replicates of seasons and conditions. Precipitation was extremely variable (55–339 mm) during the study period, and reduced precipitation in later years (2004–2008) appears to have resulted in annual moderate to large carbon sources (62–258 g C m^?2 yr^?1) in contrast to the previously reported sink (2002–2003). Variations in photosynthetically active radiation were found to principally drive variations in carbon uptake during the wet growing season while increased soil temperatures at a 5 cm depth stimulated carbon loss during the dry dormant season. Monthly NEE was primarily driven by soil moisture at a 5 cm depth, and years with a higher magnitude of precipitation events showed a longer growing season with annual net carbon uptake, whereas years with lower magnitude had drier soils and displayed short growing seasons with annual net carbon loss. Increased precipitation frequency was associated with increased annual NEE, which may be a function of increased microbial respiration to more small precipitation events. Annual precipitation frequency and magnitude were found to have effects on the interannual variability of NEE for up to 2 years.     

Canopy duration has little influence on annual carbon storage in the deciduous broad leaf forest

Vegetation phenology, the study of the timing and length of the terrestrial growing season and its connection to climate, is increasingly important in integrated Earth system science. Phenological variability is an excellent barometer of short‐ and long‐term climatic variability, strongly influences surface meteorology, and may influence the carbon cycle. Here, using the 1895–1993 Vegetation/Ecosystem Modelling and Analysis dataset and the Biome‐BGC terrestrial ecosystem model, we investigated the relationship between phenological metrics and annual net ecosystem exchange (NEE) of carbon. For the 1167 deciduous broad leaf forest pixels, we found that NEE was extremely weakly related to canopy duration (days from leaf appearance to complete leaf fall). Longer canopy duration, did, however, sequester more carbon if warm season precipitation was above average. Carbon uptake period (number of days with net CO₂ uptake from the atmosphere), which integrates the influence of all ecosystem states and processes, was strongly related to NEE. Results from the Harvard Forest eddy‐covariance site supported our findings. Such dramatically different results from two definitions of ‘growing season length’ highlight the potential for confusion among the many disciplines engaged in phenological research.     

Cultivation of a perennial grass for bioenergy on a boreal organic soil – carbon sink or source?

The area under the cultivation of perennial bioenergy crops on organic soils in the northern countries is fast increasing. To understand the impact of reed canary grass (RCG, Phalaris arundinaceae L.) cultivation on the carbon dioxide (CO₂) balance of an organic soil, net ecosystem CO₂ exchange (NEE) was measured for four years in a RCG cultivated cutover peatland in eastern Finland using the eddy covariance technique. There were striking differences among the years in the annual precipitation. The annual precipitation was higher during 2004 and 2007 and lower during 2005 and 2006 than the 1971–2000 regional mean. During wet growing seasons, moderate temperatures, high surface soil moisture and low evaporative demand favoured high CO₂ uptake. During dry seasons, owing to soil moisture and atmospheric stress, photosynthetic activity was severely restricted. The CO₂ uptake [gross primary productivity (GPP)] was positively correlated with soil moisture, air temperature and inversely with vapour pressure deficit. Total ecosystem respiration (TER) increased with increasing soil temperature but decreased with increasing soil moisture. The relative responses of GPP and TER to moisture stress were different. While changes in TER for a given change in soil moisture were moderate, variations in GPP were drastic. Also, the seasonal variations in TER were not as conspicuous as those in GPP implying that GPP is the primary regulator of the interannual variability in NEE in this ecosystem. The ecosystem accumulated a total of 398 g C m^?2 from the beginning of 2004 until the end of 2007. It retained some carbon during a wet year such as 2004 even after accounting for the loss of carbon in the form of harvested biomass. Based on this CO₂ balance analysis, RCG cultivation is found to be a promising after‐use option on an organic soil.     

Interannual variability in net CO2 exchange of a native tallgrass prairie

Year‐round eddy covariance flux measurements were made in a native tallgrass prairie in north‐central Oklahoma, USA during 1997–2000 to quantify carbon exchange and its interannual variability. This prairie is dominated by warm season C₄ grasses. The soil is a relatively shallow silty clay loam underlined with a heavy clay layer and a limestone bedrock. During the study period, the prairie was burned in the spring of each year, and was not grazed. In 1997 there was adequate soil moisture through the growing season, but 1998 had two extended periods of substantially low soil moisture (with concurrent high air temperatures and vapor pressure deficits), one early and one later in the growing season. There was also moisture stress in 1999, but it was less severe and occurred later in the season. The annual net ecosystem CO₂ exchange, NEE (before including carbon loss during the burn) was 274, 46 and 124 g C m ^{? 2} yr ^{? 1} in 1997, 1998, and 1999, respectively (flux toward the surface is positive), and the associated variation seemed to mirror the severity of moisture stress. We also examined integrated values of NEE during different periods (e.g. day/night; growing season/senescence). Annually integrated carbon dioxide uptake during the daytime showed the greatest variability from year to year, and was primarily linked to the severity of moisture stress. Carbon loss during nighttime was a significant part of the annual daytime NEE, and was fairly stable from year to year. When carbon loss during the burn (estimated from pre‐ and post‐burn biomass samples) was incorporated in the annual NEE, the prairie was found to be approximately carbon neutral (i.e. net carbon uptake/release was near zero) in years with no moisture stress (1997) or with some stress late in the season (1999). During a year with severe moisture stress early in the season (1998), the prairie was a net source of carbon. It appears that moisture stress (severity as well as timing of occurrence) was a dominating factor regulating the annual carbon exchange of the prairie.     

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.     

Net ecosystem exchange of CO2 with rapidly changing high Arctic landscapes

High Arctic landscapes are expansive and changing rapidly. However, our understanding of their functional responses and potential to mitigate or enhance anthropogenic climate change is limited by few measurements. We collected eddy covariance measurements to quantify the net ecosystem exchange (NEE) of CO₂ with polar semidesert and meadow wetland landscapes at the highest latitude location measured to date (82°N). We coupled these rare data with ground and satellite vegetation production measurements (Normalized Difference Vegetation Index; NDVI) to evaluate the effectiveness of upscaling local to regional NEE. During the growing season, the dry polar semidesert landscape was a near‐zero sink of atmospheric CO₂ (NEE: ?0.3 ± 13.5 g C m^?2). A nearby meadow wetland accumulated over 300 times more carbon (NEE: ?79.3 ± 20.0 g C m^?2) than the polar semidesert landscape, and was similar to meadow wetland NEE at much more southerly latitudes. Polar semidesert NEE was most influenced by moisture, with wetter surface soils resulting in greater soil respiration and CO₂ emissions. At the meadow wetland, soil heating enhanced plant growth, which in turn increased CO₂ uptake. Our upscaling assessment found that polar semidesert NDVI measured on‐site was low (mean: 0.120–0.157) and similar to satellite measurements (mean: 0.155–0.163). However, weak plant growth resulted in poor satellite NDVI–NEE relationships and created challenges for remotely detecting changes in the cycling of carbon on the polar semidesert landscape. The meadow wetland appeared more suitable to assess plant production and NEE via remote sensing; however, high Arctic wetland extent is constrained by topography to small areas that may be difficult to resolve with large satellite pixels. We predict that until summer precipitation and humidity increases enough to offset poor soil moisture retention, climate‐related changes to productivity on polar semideserts may be restricted.     

Scaling-up knowledge of growing-season net ecosystem exchange for long-term assessment of North Dakota grasslands under the Conservation Reserve Program

Scaling‐up knowledge of land‐atmosphere net ecosystem exchange (NEE) from a single experimental site to numerous perennial grass fields in the Northern Great Plains (NGP) requires appropriate scaling protocols. We addressed this problem using synoptic data available from the Landsat sensor for 10 growing seasons (April 15 to September 30) over a North Dakota field‐site, where we continuously measured CO₂ exchange using a Bowen Ratio Energy Balance (BREB) system. NEE observed during the growing season at our field‐site from 1997 to 2006 vacillated with drought and deluge, with net carbon (C) losses to the atmosphere in 2006. We used stepwise linear regression with 10 years of Landsat and NEE data to construct and validate a model for estimating grassland growing‐season NEE from field to landscape scales. Eighty‐nine percent of the variability in NEE was explained by year, live biomass, carbon : nitrogen ratio, day of image acquisition, and annual precipitation. We then applied this model on 20 620 ha of North Dakota perennial grass fields enrolled in the Conservation Reserve Program (CRP), including 1272 fields east of the Missouri River and 165 fields west‐river. Growing‐season NEE for individual CRP fields was highly variable from 1997 to 2006, ranging from ?366 to 692 g C m^?2 growing season^?1. Mean annual growing‐season fluxes over 10 years for CRP fields located east‐river and west‐river were 317 g C m^?2 growing season^?1 and 239 g C m^?2 growing season^?1, respectively. Average cumulative growing‐season NEE modeled for fields east‐ and west‐river diverged from one another in 2002–2006, when west‐river fields received < 70% of the long‐term annual average precipitation during these years. Results indicate assessment of conservation practices on grassland CO₂ exchange during the growing season can be remotely estimated at field and landscape scales under variable environmental conditions and should be followed up with similar, spatially explicit investigations of NEE during the dormant season.     

Variability in exchange of CO2 across 12 northern peatland and tundra sites

Many wetland ecosystems such as peatlands and wet tundra hold large amounts of organic carbon (C) in their soils, and are thus important in the terrestrial C cycle. We have synthesized data on the carbon dioxide (CO₂) exchange obtained from eddy covariance measurements from 12 wetland sites, covering 1–7 years at each site, across Europe and North America, ranging from ombrotrophic and minerotrophic peatlands to wet tundra ecosystems, spanning temperate to arctic climate zones. The average summertime net ecosystem exchange of CO₂ (NEE) was highly variable between sites. However, all sites with complete annual datasets, seven in total, acted as annual net sinks for atmospheric CO₂. To evaluate the influence of gross primary production (GPP) and ecosystem respiration (R_eco) on NEE, we first removed the artificial correlation emanating from the method of partitioning NEE into GPP and R_eco. After this correction neither R_eco (P= 0.162) nor GPP (P= 0.110) correlated significantly with NEE on an annual basis. Spatial variation in annual and summertime R_eco was associated with growing season period, air temperature, growing degree days, normalized difference vegetation index and vapour pressure deficit. GPP showed weaker correlations with environmental variables as compared with R_eco, the exception being leaf area index (LAI), which correlated with both GPP and NEE, but not with R_eco. Length of growing season period was found to be the most important variable describing the spatial variation in summertime GPP and R_eco; global warming will thus cause these components to increase. Annual GPP and NEE correlated significantly with LAI and pH, thus, in order to predict wetland C exchange, differences in ecosystem structure such as leaf area and biomass as well as nutritional status must be taken into account.     

On the difference in the net ecosystem exchange of CO2 between deciduous and evergreen forests in the southeastern United States

The southeastern United States is experiencing a rapid regional increase in the ratio of pine to deciduous forest ecosystems at the same time it is experiencing changes in climate. This study is focused on exploring how these shifts will affect the carbon sink capacity of southeastern US forests, which we show here are among the strongest carbon sinks in the continental United States. Using eight‐year‐long eddy covariance records collected above a hardwood deciduous forest (HW) and a pine plantation (PP) co‐located in North Carolina, USA, we show that the net ecosystem exchange of CO₂ (NEE) was more variable in PP, contributing to variability in the difference in NEE between the two sites (ΔNEE) at a range of timescales, including the interannual timescale. Because the variability in evapotranspiration (ET) was nearly identical across the two sites over a range of timescales, the factors that determined the variability in ΔNEE were dominated by those that tend to decouple NEE from ET. One such factor was water use efficiency, which changed dramatically in response to drought and also tended to increase monotonically in nondrought years (P < 0.001 in PP). Factors that vary over seasonal timescales were strong determinants of the NEE in the HW site; however, seasonality was less important in the PP site, where significant amounts of carbon were assimilated outside of the active season, representing an important advantage of evergreen trees in warm, temperate climates. Additional variability in the fluxes at long‐time scales may be attributable to slowly evolving factors, including canopy structure and increases in dormant season air temperature. Taken together, study results suggest that the carbon sink in the southeastern United States may become more variable in the future, owing to a predicted increase in drought frequency and an increase in the fractional cover of southern pines.     

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.     

Spatial and Temporal Variability in Growing-Season Net Ecosystem Carbon Dioxide Exchange at a Large Peatland in Ontario,Canada

We measured net ecosystem exchange of carbon dioxide (CO2) (NEE) during wet and dry summers (2000 and 2001) across a range of plant communities at Mer Bleue, a large peatland near Ottawa, southern Ontario, Canada. Wetland types included ombrotrophic bog hummocks and hollows, mineral-poor fen, and beaver pond margins. NEE was significantly different among the sites in both years, but rates of gross photosynthesis did not vary spatially even though species composition at the sites was variable. Soil respiration rates were very different across sites and dominated interannual variability in summer NEE within sites. During the dry summer of 2001, net CO2 uptake was significantly smaller, and most locations switched from a net sink to a source of CO2 under a range of levels of photosynthetically active radiation (PAR). The wetter areas--poor fen and beaver pond margin--had the largest rates of CO2 uptake and smallest rates of respiratory loss during the dry summer. Communities dominated by ericaceous shrubs (bog sites) maintained similar rates of gross photosynthesis between years; by contrast, the sedge-dominated areas (fen sites) showed signs of early senescence under drought conditions. Water table position was the strongest control on respiration in the drier summer, whereas surface peat temperature explained most of the variability in the wetter summer. Q 10 temperature-respiration quotients averaged 1.6 to 2.2. The ratio between maximum photosynthesis and respiration ranged from 3.7:1 in the poor fen to 1.2:1 at some bog sites; it declined at all sites in the drier summer owing to greater respiration rates relative to photosynthesis in evergreen shrub sites and a change in both processes in sedge sites. Our ability to predict ecosystem responses to changing climate depends on a more complete understanding of the factors that control NEE across a range of peatland plant communities.     

Growing season net ecosystem CO2 exchange of two desert ecosystems with alkaline soils in Kazakhstan

Central Asia is covered by vast desert ecosystems, and the majority of these ecosystems have alkaline soils. Their contribution to global net ecosystem CO₂ exchange (NEE) is of significance simply because of their immense spatial extent. Some of the latest research reported considerable abiotic CO₂ absorption by alkaline soil, but the rate of CO₂ absorption has been questioned by peer communities. To investigate the issue of carbon cycle in Central Asian desert ecosystems with alkaline soils, we have measured the NEE using eddy covariance (EC) method at two alkaline sites during growing season in Kazakhstan. The diurnal course of mean monthly NEE followed a clear sinusoidal pattern during growing season at both sites. Both sites showed significant net carbon uptake during daytime on sunny days with high photosynthetically active radiation (PAR) but net carbon loss at nighttime and on cloudy and rainy days. NEE has strong dependency on PAR and the response of NEE to precipitation resulted in an initial and significant carbon release to the atmosphere, similar to other ecosystems. These findings indicate that biotic processes dominated the carbon processes, and the contribution of abiotic carbon process to net ecosystem CO₂ exchange may be trivial in alkaline soil desert ecosystems over Central Asia.     

Sensitivity of Temperate Desert Steppe Carbon Exchange to Seasonal Droughts and Precipitation Variations in Inner Mongolia,China

Arid grassland ecosystems have significant interannual variation in carbon exchange; however, it is unclear how environmental factors influence carbon exchange in different hydrological years. In this study, the eddy covariance technique was used to investigate the seasonal and interannual variability of CO₂ flux over a temperate desert steppe in Inner Mongolia, China from 2008 to 2010. The amounts and times of precipitation varied significantly throughout the study period. The precipitation in 2009 (186.4 mm) was close to the long-term average (183.9±47.6 mm), while the precipitation in 2008 (136.3 mm) and 2010 (141.3 mm) was approximately a quarter below the long-term average. The temperate desert steppe showed carbon neutrality for atmospheric CO₂ throughout the study period, with a net ecosystem carbon dioxide exchange (NEE) of −7.2, −22.9, and 26.0 g C m⁻² yr⁻¹ in 2008, 2009, and 2010, not significantly different from zero. The ecosystem gained more carbon in 2009 compared to other two relatively dry years, while there was significant difference in carbon uptake between 2008 and 2010, although both years recorded similar annual precipitation. The results suggest that summer precipitation is a key factor determining annual NEE. The apparent quantum yield and saturation value of NEE (NEE_sat) and the temperature sensitivity coefficient of ecosystem respiration (R_eco) exhibited significant variations. The values of NEE_sat were −2.6, −2.9, and −1.4 µmol CO₂ m⁻² s⁻¹ in 2008, 2009, and 2010, respectively. Drought suppressed both the gross primary production (GPP) and R_eco, and the drought sensitivity of GPP was greater than that of R_eco. The soil water content sensitivity of GPP was high during the dry year of 2008 with limited soil moisture availability. Our results suggest the carbon balance of this temperate desert steppe was not only sensitive to total annual precipitation, but also to its seasonal distribution.     

Long‐term impacts of agricultural practices and climatic variability on carbon storage in a permanent pasture

Intra‐ and interannual variability of precipitation can lead to major modifications of grassland production and carbon storage capacity. Greater understanding of how climatic variability affects net CO₂ exchange [i.e. net ecosystem exchange (NEE)] of grazed grasslands is important to adapt grassland management and reduce risks of carbon losses. Since 2002, we continuously measured NEE (i.e. eddy covariance technique) on an upland grassland site (7 ha), divided in two paddocks grazed by heifers (intensive: 1 LSU ha^?1 yr^?1, 213 kg N ha^?1 yr^?1 and extensive: 0.5 LSU ha^?1 yr^?1, no fertilization). For years with dry and warm growing seasons (i.e. 2003, 2005 and 2008), absolute annual NEE was higher in the intensive paddock compared with the extensive paddock. The opposite was observed during years of ample seasonal rainfall and soil moisture (i.e. 2004, 2006 and 2007). Contrasted management led to two distinct plant communities being different in leaf area index (LAI), soil bulk density and soil water holding capacity. Differences in annual NEEs could thus be assigned to interactions between in carbon and water fluxes during dry and wet growth periods. Dry growth periods led to a reduction in weekly gross primary productivity (GPP) in the extensively managed paddock, whereas the GPP was maintained in the intensive paddock. In turn, during wet growth periods, GPP was similar in both paddocks, whereas N amendment and frequent defoliation significantly increased ecosystem respiration in the intensive paddock, presumably through a higher heterotrophic respiration following on a better C substrate quality and availability (rhizodeposition and senescent fine roots). In the extensive paddock, where plant cover was denser (reducing soil temperature) and less decomposable, C losses through heterotrophic respiration were comparatively smaller under wet conditions. Our results demonstrate that grassland subjected to a moderately intensive management could be more resilient in terms of carbon storage during drought and heat waves, presumably because of a trade‐off between heterotrophic and autotrophic respiration.     

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

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

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

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

A decade of boreal rich fen greenhouse gas fluxes in response to natural and experimental water table variability

Rich fens are common boreal ecosystems with distinct hydrology, biogeochemistry and ecology that influence their carbon (C) balance. We present growing season soil chamber methane emission (F_CH4), ecosystem respiration (ER), net ecosystem exchange (NEE) and gross primary production (GPP) fluxes from a 9‐years water table manipulation experiment in an Alaskan rich fen. The study included major flood and drought years, where wetting and drying treatments further modified the severity of droughts. Results support previous findings from peatlands that drought causes reduced magnitude of growing season F_CH4, GPP and NEE, thus reducing or reversing their C sink function. Experimentally exacerbated droughts further reduced the capacity for the fen to act as a C sink by causing shifts in vegetation and thus reducing magnitude of maximum growing season GPP in subsequent flood years by ~15% compared to control plots. Conversely, water table position had only a weak influence on ER, but dominant contribution to ER switched from autotrophic respiration in wet years to heterotrophic in dry years. Droughts did not cause inter‐annual lag effects on ER in this rich fen, as has been observed in several nutrient‐poor peatlands. While ER was dependent on soil temperatures at 2 cm depth, F_CH4 was linked to soil temperatures at 25 cm. Inter‐annual variability of deep soil temperatures was in turn dependent on wetness rather than air temperature, and higher F_CH4 in flooded years was thus equally due to increased methane production at depth and decreased methane oxidation near the surface. Short‐term fluctuations in wetness caused significant lag effects on F_CH4, but droughts caused no inter‐annual lag effects on F_CH4. Our results show that frequency and severity of droughts and floods can have characteristic effects on the exchange of greenhouse gases, and emphasize the need to project future hydrological regimes in rich fens.     

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.     

Environmental variation is directly responsible for short- but not long-term variation in forest-atmosphere carbon exchange

Tower‐based eddy covariance measurements of forest‐atmosphere carbon dioxide (CO₂) exchange from many sites around the world indicate that there is considerable year‐to‐year variation in net ecosystem exchange (NEE). Here, we use a statistical modeling approach to partition the interannual variability in NEE (and its component fluxes, ecosystem respiration, R_eco, and gross photosynthesis, P_gross) into two main effects: variation in environmental drivers (air and soil temperature, solar radiation, vapor pressure deficit, and soil water content) and variation in the biotic response to this environmental forcing (as characterized by the model parameters). The model is applied to a 9‐year data set from the Howland AmeriFlux site, a spruce‐dominated forest in Maine, USA. Gap‐filled flux measurements at this site indicate that the forest has been sequestering, on average, 190 g C m⁻² yr⁻¹, with a range from 130 to 270 g C m⁻² yr⁻¹. Our fitted model predicts somewhat more uptake (mean 270 g C m⁻² yr⁻¹), but interannual variation is similar, and wavelet variance analyses indicate good agreement between tower measurements and model predictions across a wide range of timescales (hours to years). Associated with the interannual variation in NEE are clear differences among years in model parameters for both R_eco and P_gross. Analysis of model predictions suggests that, at the annual time step, about 40% of the variance in modeled NEE can be attributed to variation in environmental drivers, and 55% to variation in the biotic response to this forcing. As model predictions are aggregated at longer timescales (from individual days to months to calendar year), variation in environmental drivers becomes progressively less important, and variation in the biotic response becomes progressively more important, in determining the modeled flux. There is a strong negative correlation between modeled annual P_gross and R_eco (r=−0.93, P≤0.001); two possible explanations for this correlation are discussed. The correlation promotes homeostasis of NEE: the interannual variation in modeled NEE is substantially less than that for either P_gross or R_eco