Cool-season whole-plant gas exchange of exotic and native semiarid bunchgrasses

The success of invasive aridland plants may depend on their utilization of precipitation not fully exploited by native species, which could lead to seasonally altered ecosystem carbon and water fluxes. We measured volumetric soil water across 25-cm profiles (??_25cm) and springtime whole-plant water- and carbon-fluxes of the exotic Lehmann lovegrass (Eragrostis lehmanniana) and a native bunchgrass, bush muhly (Muhlenbergia porteri), following typical (55?mm in 2009) and El Ni?o-enhanced accumulations (154?mm in 2010) in a SE Arizona savanna. Across both years, ??_25cm was higher under lovegrass plots, with similar evapotranspiration (ET) between lovegrass and bush muhly plots. However, in 2010 transpiration (T) was higher in bush muhly than lovegrass, implying higher soil evaporation in lovegrass plots maintained similar ET. Net ecosystem carbon dioxide exchange (NEE) was similar between lovegrass and bush muhly plots in 2009, but was more negative in bush muhly plots following El Ni?o, indicating greater CO₂ assimilation. Ecosystem respiration (R _eco) and gross ecosystem photosynthesis (GEP) were similar between lovegrass and bush muhly plots in 2009, but were higher in bush muhly plots in 2010. As a result, lovegrass plots reduced ecosystem water-use efficiency (WUE_e?=?NEE/ET), while bush muhly WUE_e remained constant between 2009 and 2010. Concurrent whole-plant WUE (WUE_p?=?GEP/T) did not change in lovegrass plots, but increased in bush muhly plots between these years. We concluded that cool-season precipitation use is not a component of Lehmann lovegrass invasive success, but that the change in ET partitioning and attendant shifts in cool-season WUE_e may increase interannual variation in ecosystem water- and carbon-exchange dynamics in the water-limited systems it dominates.     

High Arctic Dry Heath CO2 Exchange During the Early Cold Season

Climate change may alter the terrestrial ecosystem carbon balance in the Arctic, and previous studies have emphasized the importance of cold season gas exchange when considering the annual carbon balance. Here, we examined gross ecosystem production (GEP), ecosystem respiration (R _eco) and net ecosystem exchange (NEE) during autumn at a high arctic dry open heath, over a period where air temperatures decreased from +9.8 to ?16.5°C. GEP declined by 95–100% during autumn but GEP significantly different from 0 was measured on October 8 despite sub-zero temperatures. R _eco declined by 90% and dominated NEE throughout the study as the ecosystem on all measurement days was a source of atmospheric CO₂. We estimated net September carbon losses (NEE) to be 17?g?CO₂?m^?2, emphasizing the importance of autumn in relation to annual carbon budgets. The study site has been subjected to 14 summers of water addition, and occasional pulses of nitrogen (N) addition in a fully factorial design. N addition enhanced GEP up to 17-fold during September, although there was no effect in October when GEP was very low. Summer water addition decreased autumn R _eco by up to 25%. Both N amendment and water addition decreased carbon loss, that is, increased NEE; N amendment increased NEE on all dates by 13–64% whereas water addition increased NEE by 20–54% late in September and onward, demonstrating the importance of nutrient and water availability on carbon balance in high arctic tundra, also during the autumn freeze-in.     

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.     

Nonlinear CO2 flux response to 7 years of experimentally induced permafrost thaw

Rapid Arctic warming is expected to increase global greenhouse gas concentrations as permafrost thaw exposes immense stores of frozen carbon (C) to microbial decomposition. Permafrost thaw also stimulates plant growth, which could offset C loss. Using data from 7 years of experimental Air and Soil warming in moist acidic tundra, we show that Soil warming had a much stronger effect on CO₂ flux than Air warming. Soil warming caused rapid permafrost thaw and increased ecosystem respiration (R_eco), gross primary productivity (GPP), and net summer CO₂ storage (NEE). Over 7 years R_eco, GPP, and NEE also increased in Control (i.e., ambient plots), but this change could be explained by slow thaw in Control areas. In the initial stages of thaw, R_eco, GPP, and NEE increased linearly with thaw across all treatments, despite different rates of thaw. As thaw in Soil warming continued to increase linearly, ground surface subsidence created saturated microsites and suppressed R_eco, GPP, and NEE. However R_eco and GPP remained high in areas with large Eriophorum vaginatum biomass. In general NEE increased with thaw, but was more strongly correlated with plant biomass than thaw, indicating that higher R_eco in deeply thawed areas during summer months was balanced by GPP. Summer CO₂ flux across treatments fit a single quadratic relationship that captured the functional response of CO₂ flux to thaw, water table depth, and plant biomass. These results demonstrate the importance of indirect thaw effects on CO₂ flux: plant growth and water table dynamics. Nonsummer R_eco models estimated that the area was an annual CO₂ source during all years of observation. Nonsummer CO₂ loss in warmer, more deeply thawed soils exceeded the increases in summer GPP, and thawed tundra was a net annual CO₂ source.     

On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm

This paper discusses the advantages and disadvantages of the different methods that separate net ecosystem exchange (NEE) into its major components, gross ecosystem carbon uptake (GEP) and ecosystem respiration (R_eco). In particular, we analyse the effect of the extrapolation of night‐time values of ecosystem respiration into the daytime; this is usually done with a temperature response function that is derived from long‐term data sets. For this analysis, we used 16 one‐year‐long data sets of carbon dioxide exchange measurements from European and US‐American eddy covariance networks. These sites span from the boreal to Mediterranean climates, and include deciduous and evergreen forest, scrubland and crop ecosystems. We show that the temperature sensitivity of R_eco, derived from long‐term (annual) data sets, does not reflect the short‐term temperature sensitivity that is effective when extrapolating from night‐ to daytime. Specifically, in summer active ecosystems the long‐term temperature sensitivity exceeds the short‐term sensitivity. Thus, in those ecosystems, the application of a long‐term temperature sensitivity to the extrapolation of respiration from night to day leads to a systematic overestimation of ecosystem respiration from half‐hourly to annual time‐scales, which can reach >25% for an annual budget and which consequently affects estimates of GEP. Conversely, in summer passive (Mediterranean) ecosystems, the long‐term temperature sensitivity is lower than the short‐term temperature sensitivity resulting in underestimation of annual sums of respiration. We introduce a new generic algorithm that derives a short‐term temperature sensitivity of R_eco from eddy covariance data that applies this to the extrapolation from night‐ to daytime, and that further performs a filling of data gaps that exploits both, the covariance between fluxes and meteorological drivers and the temporal structure of the fluxes. While this algorithm should give less biased estimates of GEP and R_eco, we discuss the remaining biases and recommend that eddy covariance measurements are still backed by ancillary flux measurements that can reduce the uncertainties inherent in the eddy covariance data.     

Seasonal and interannual variations in carbon dioxide exchange over a cropland in the North China Plain

In China, croplands account for a relatively large form of vegetation cover. Quantifying carbon dioxide exchange and understanding the environmental controls on carbon fluxes over croplands are critical in understanding regional carbon budgets and ecosystem behaviors. In this study, the net ecosystem exchange (NEE) at a winter wheat/summer maize rotation cropping site, representative of the main cropping system in the North China Plain, was continuously measured using the eddy covariance technique from 2005 to 2009. In order to interpret the abiotic factors regulating NEE, NEE was partitioned into gross primary production (GPP) and ecosystem respiration (R_eco). Daytime R_eco was extrapolated from the relationship between nighttime NEE and soil temperature under high turbulent conditions. GPP was then estimated by subtracting daytime NEE from the daytime estimates of R_eco. Results show that the seasonal patterns of the temperature responses of R_eco and light‐response parameters are closely related to the crop phenology. Daily R_eco was highly dependent on both daily GPP and air temperature. Interannual variability showed that GPP and R_eco were mainly controlled by temperature. Water availability also exerted a limit on R_eco. The annual NEE was ?585 and ?533 g C m^?2 for two seasons of 2006–2007 and 2007–2008, respectively, and the wheat field absorbed more carbon than the maize field. Thus, we concluded that this cropland was a strong carbon sink. However, when the grain harvest was taken into account, the wheat field was diminished into a weak carbon sink, whereas the maize field was converted into a weak carbon source. The observations showed that severe drought occurring during winter did not reduce wheat yield (or integrated NEE) when sufficient irrigation was carried out during spring.     

Environmental and Vegetation Drivers of Seasonal CO2 Fluxes in a Sub-arctic Forest–Mire Ecotone

Unravelling the role of structural and environmental drivers of gross primary productivity (GPP) and ecosystem respiration (R _eco) in highly heterogeneous tundra is a major challenge for the upscaling of chamber-based CO₂ fluxes in Arctic landscapes. In a mountain birch woodland-mire ecotone, we investigated the role of LAI (and NDVI), environmental factors (microclimate, soil moisture), and microsite type across tundra shrub plots (wet hummocks, dry hummocks, dry hollows) and lichen hummocks, in controlling net ecosystem CO₂ exchange (NEE). During a growing season, we measured NEE fluxes continuously, with closed dynamic chambers, and performed multiple fits (one for each 3-day period) of a simple light and temperature response model to hourly NEE data. Tundra shrub plots were largely CO₂ sinks, as opposed to lichen plots, although fluxes were highly variable within microsite type. For tundra shrub plots, microsite type did not influence photosynthetic parameters but it affected basal (that is, temperature-normalized) ecosystem respiration (R ₀). PAR-normalized photosynthesis (P ₆₀₀) increased with air temperature and declined with increasing vapor pressure deficit. R ₀ declined with soil moisture and showed an apparent increase with temperature, which may underlie a tight link between GPP and R _eco. NDVI was a good proxy for LAI, maximum P ₆₀₀ and maximum R ₀ of shrub plots. Cumulative CO₂ fluxes were strongly correlated with LAI (NDVI) but we observed a comparatively low GPP/LAI in dry hummocks. Our results broadly agree with the reported functional convergence across tundra vegetation, but here we show that the role of decreased productivity in transition zones and the influence of temperature and water balance on seasonal CO₂ fluxes in sub-Arctic forest–mire ecotones cannot be overlooked.     

A distinct seasonal pattern of the ratio of soil respiration to total ecosystem respiration in a spruce-dominated forest

Annual budgets and fitted temperature response curves for soil respiration and ecosystem respiration provide useful information for partitioning annual carbon budgets of ecosystems, but they may not adequately reveal seasonal variation in the ratios of these two fluxes. Soil respiration (R_s) typically contributes 30–80% of annual total ecosystem respiration (R_eco) in forests, but the temporal variation of these ratios across seasons has not been investigated. The objective of this study was to investigate seasonal variation in the R_s/R_eco ratio in a mature forest dominated by conifers at Howland, ME, USA. We used chamber measurements of R_s and tower‐based eddy covariance measurements of R_eco. The R_s/R_eco ratio reached a minimum of about 0.45 in the early spring, gradually increased through the late spring and early summer, leveled off at about 0.65 for the summer, and then increased again to about 0.8 in the autumn. A spring pulse of aboveground respiration presumably causes the springtime minimum in this ratio. Soil respiration ‘catches up’ as the soils warm and as root growth presumably accelerates in the late spring, causing the R_s/R_eco ratios to increase. The summertime plateau of R_s/R_eco ratios is consistent with summer drought suppressing R_s that would otherwise be increasing, based on increasing soil temperature alone, thus causing the R_s/R_eco ratios to not increase as soils continue to warm. Declining air temperatures and litter fall apparently contribute to increased R_s/R_eco ratios in the autumn. Differences in phenology of growth of aboveground and belowground plant tissues, mobilization and use of stored substrates within woody plants, seasonal variation in photosynthate and litter substrates, and lags between temperature changes of air and soil contribute to a distinct seasonal pattern of R_s/R_eco ratios.     

Rapid Recovery of Gross Production and Respiration in a Mesic Mountain Big Sagebrush Ecosystem Following Prescribed Fire

The impact of land management actions such as prescribed fire remains a key uncertainty in understanding the spatiotemporal patterns of carbon cycling in the Western USA. We therefore quantified carbon exchange and aboveground carbon stocks following a prescribed fire in a mountain big sagebrush ecosystem located in the northern Great Basin, USA. Specifically, we examined the changes in plant functional type, leaf area index, standing aboveground carbon stocks, net ecosystem production (NEP), gross ecosystem production (GEP), and ecosystem-level respiration (R_eco) for 2 years before and 7 of 9 years after a prescribed fire. Post-burn GEP and R_eco exceeded pre-burn GEP and R_eco within 2 years and remained elevated. The variation in GEP and R_eco provided no evidence of a large and prolonged net efflux of carbon in the 9 years after the fire. Rather, NEP indicated the site was a sink before and after the fire, with little change in sink strength associated with the burn. Re-sprouting and recruitment of grasses and forbs drove the post-burn increase in GEP. Woody shrub growth was the dominant control on aboveground biomass accumulation after fire, with shrub aboveground biomass reaching ~ 11% of pre-burn biomass after 5 years. The rapid recovery of GEP and the growth of mid-successional shrubs suggest ecosystem-level carbon fluxes and stocks can recover rapidly after fire in mesic mountain big sagebrush ecosystems.     

Ecological functioning in grass–shrub Mediterranean ecosystems measured by eddy covariance

Climate change may alter ecosystem functioning, as assessed via the net carbon (C) exchange (NEE) with the atmosphere, composed of the biological processes photosynthesis (GPP) and respiration (R _eco). In addition, in semi-arid Mediterranean ecosystems, a significant fraction of respired CO₂ is stored in the vadose zone and emitted afterwards by subsoil ventilation (VE), contributing also to NEE. Such conditions complicate the prediction of NEE for future change scenarios. To evaluate the possible effects of climate change on annual NEE and its underlying processes (GPP, R _eco and VE) we present, over a climate/altitude range, the annual and interannual variability of NEE, GPP, R _eco and VE in three Mediterranean sites. We found that annual NEE varied from a net source of around 130 gC m^?2 in hot and arid lowlands to a net sink of similar magnitude for alpine meadows (above 2,000 m a.s.l) that are less water stressed. Annual net C fixation increased because of increased GPP during intermittent and several growth periods occurring even during winter, as well as due to decreased VE. In terms of interannual variability, the studied subalpine site behaved as a neutral C sink (from emission of 49 to fixation of 30 gC m^?2 year^?1), with precipitation as the main factor controlling annual GPP and R _eco. Finally, the importance of VE as 0–23 % of annual NEE is highlighted, indicating that this process could shift some Mediterranean ecosystems from annual C sinks to sources.     

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.     

Separation of net ecosystem exchange into assimilation and respiration using a light response curve approach: critical issues and global evaluation

The measured net ecosystem exchange (NEE) of CO₂ between the ecosystem and the atmosphere reflects the balance between gross CO₂ assimilation [gross primary production (GPP)] and ecosystem respiration (R_eco). For understanding the mechanistic responses of ecosystem processes to environmental change it is important to separate these two flux components. Two approaches are conventionally used: (1) respiration measurements made at night are extrapolated to the daytime or (2) light–response curves are fit to daytime NEE measurements and respiration is estimated from the intercept of the ordinate, which avoids the use of potentially problematic nighttime data. We demonstrate that this approach is subject to biases if the effect of vapor pressure deficit (VPD) modifying the light response is not included. We introduce an algorithm for NEE partitioning that uses a hyperbolic light response curve fit to daytime NEE, modified to account for the temperature sensitivity of respiration and the VPD limitation of photosynthesis. Including the VPD dependency strongly improved the model's ability to reproduce the asymmetric diurnal cycle during periods with high VPD, and enhances the reliability of R_eco estimates given that the reduction of GPP by VPD may be otherwise incorrectly attributed to higher R_eco. Results from this improved algorithm are compared against estimates based on the conventional nighttime approach. The comparison demonstrates that the uncertainty arising from systematic errors dominates the overall uncertainty of annual sums (median absolute deviation of GPP: 47 g C m^?2 yr^?1), while errors arising from the random error (median absolute deviation: ～2 g C m^?2 yr^?1) are negligible. Despite site‐specific differences between the methods, overall patterns remain robust, adding confidence to statistical studies based on the FLUXNET database. In particular, we show that the strong correlation between GPP and R_eco is not spurious but holds true when quasi‐independent, i.e. daytime and nighttime based estimates are compared.     

Arctic browning: Impacts of extreme climatic events on heathland ecosystem CO2 fluxes

Extreme climatic events are among the drivers of recent declines in plant biomass and productivity observed across Arctic ecosystems, known as “Arctic browning.” These events can cause landscape‐scale vegetation damage and so are likely to have major impacts on ecosystem CO₂ balance. However, there is little understanding of the impacts on CO₂ fluxes, especially across the growing season. Furthermore, while widespread shoot mortality is commonly observed with browning events, recent observations show that shoot stress responses are also common, and manifest as high levels of persistent anthocyanin pigmentation. Whether or how this response impacts ecosystem CO₂ fluxes is not known. To address these research needs, a growing season assessment of browning impacts following frost drought and extreme winter warming (both extreme climatic events) on the key ecosystem CO₂ fluxes Net Ecosystem Exchange (NEE), Gross Primary Productivity (GPP), ecosystem respiration (R_eco) and soil respiration (R_soil) was carried out in widespread sub‐Arctic dwarf shrub heathland, incorporating both mortality and stress responses. Browning (mortality and stress responses combined) caused considerable site‐level reductions in GPP and NEE (of up to 44%), with greatest impacts occurring at early and late season. Furthermore, impacts on CO₂ fluxes associated with stress often equalled or exceeded those resulting from vegetation mortality. This demonstrates that extreme events can have major impacts on ecosystem CO₂ balance, considerably reducing the carbon sink capacity of the ecosystem, even where vegetation is not killed. Structural Equation Modelling and additional measurements, including decomposition rates and leaf respiration, provided further insight into mechanisms underlying impacts of mortality and stress on CO₂ fluxes. The scale of reductions in ecosystem CO₂ uptake highlights the need for a process‐based understanding of Arctic browning in order to predict how vegetation and CO₂ balance will respond to continuing climate change.     

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.     

The fate of assimilated carbon during drought: impacts on respiration in Amazon rainforests

Interannual variations in CO₂ exchange across Amazonia, as deduced from atmospheric inversions, correlate with El Niño occurrence. They are thought to result from changes in net ecosystem exchange and fire incidence that are both related to drought intensity. Alterations to net ecosystem production (NEP) are caused by changes in gross primary production (GPP) and ecosystem respiration (R_eco). Here, we analyse observations of the components of R_eco (leaves, live and dead woody tissue, and soil) to provide first estimates of changes in R_eco during short-term (seasonal to interannual) moisture limitation. Although photosynthesis declines if moisture availability is limiting, leaf dark respiration is generally maintained, potentially acclimating upwards in the longer term. If leaf area is lost, then short-term canopy-scale respiratory effluxes from wood and leaves are likely to decline. Using a moderate short-term drying scenario where soil moisture limitation leads to a loss of 0.5 m² m⁻² yr⁻¹ in leaf area index, we estimate a reduction in respiratory CO₂ efflux from leaves and live woody tissue of 1.0 (±0.4) t C ha⁻¹ yr⁻¹. Necromass decomposition declines during drought, but mortality increases; the median mortality increase following a strong El Niño is 1.1% (n=46 tropical rainforest plots) and yields an estimated net short-term increase in necromass CO₂ efflux of 0.13–0.18 t C ha⁻¹ yr⁻¹. Soil respiration is strongly sensitive to moisture limitation over the short term, but not to associated temperature increases. This effect is underestimated in many models but can lead to estimated reductions in CO₂ efflux of 2.0 (±0.5) t C ha⁻¹ yr⁻¹. Thus, the majority of short-term respiratory responses to drought point to a decline in R_eco, an outcome that contradicts recent regional-scale modelling of NEP. NEP varies with both GPP and R_eco but robust moisture response functions are clearly needed to improve quantification of the role of R_eco in influencing regional-scale CO₂ emissions from Amazonia.     

Small-scale hydrological variation determines landscape CO2 fluxes in the high Arctic

We explored the influence of small-scale spatial variation in soil moisture on CO₂ fluxes in the high Arctic. Of five sites forming a hydrological gradient, CO₂ was emitted from the three driest sites and only the wettest site was a net sink of CO₂. Soil moisture was a good predictor of net ecosystem exchange (NEE). Higher gross ecosystem photosynthesis (GEP) was linked to higher bryophyte biomass and activity in response to the moisture conditions. Ecosystem respiration (R _e) rates increased with soil moisture until the soil became anaerobic and then R _e decreased. At well-drained sites R _e was driven by GEP, suggesting substrate and moisture limitation of soil respiration. We propose that spatial variability in soil moisture is a primary driver of NEE.     

Spatial heterogeneity in ecosystem structure and productivity in a moist Kenyan savanna

Topographic variations and disturbances are key sources of spatial heterogeneity in the ecosystem and may influence its functioning, productivity, and carbon (C) storage. In water controlled ecosystems, structural and functional heterogeneity become distinct during drought when the ecosystem processes are operating at their limits. We examined spatial heterogeneity arising from grazing, abandoned cropland, presence of Acacia trees, and termite mounds (termitaria). Soil water content (SWC) was significantly (P < 0.05) higher in termitaria and fenced (un-grazed) plots. Higher soil nitrogen (N) content occurred in the Acacia, termitaria, and fenced plots while total biomass was highest in the fenced plot. The termitaria plots showed the highest net ecosystem CO₂ exchange (NEE), ecosystem respiration (R _eco), and gross primary production (GPP) and were the only plots that were net CO₂ sinks. Except in fenced plots, maximum GPP was positively correlated with SWC and green biomass in all the other plots. Green biomass and R _eco were positively correlated with SWC. Shifting cultivation (abandoned farmlands) negatively affected soil quality, ecosystem CO₂ assimilation, and productivity. Removal of grazing (cattle) from the ecosystem negatively influenced GPP, while the presence of termitaria and Acacia trees facilitated soil water and N availability and ecosystem productivity. We concluded that soil water availability was responsible for most of the localized differences in the savanna and has a strong influence on ecosystem C capture and storage. We recommend that future studies on savanna productivity and ecosystem CO₂ fluxes should consider heterogeneity in the ecosystem in order to avoid bias and increase the accuracy of any estimates made.     

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.     

Summer rain pulses may stimulate a CO2 release rather than absorption in desert halophyte communities

Background and aims

The response of plants and soil to rain pulses determines seasonal variations in the exchange of materials and energy at the ecosystem scale in arid and semi-arid regions. We assessed how the ecosystem carbon exchange (NEE) of desert halophyte communities of different plant functional-types responds to summer precipitation pulses in Tamarix and Haloxylon communities.

Methods

Plant water status, photosynthetic gas exchange, soil respiration and net ecosystem carbon exchange were measured to test the hypothesis that high physiological sensitivity may induce a greater changes in NEE resulting from the summer precipitation pulses in Haloxylon community.

Results

Plant water status and photosynthetic assimilation did not differ before and after summer precipitation pulses in either community. In contrast, soil respiration and NEE responded strongly to summer precipitation events in both communities. At the ecosystem level, precipitation pulses induced a pulse of CO₂ release, rather than absorption. The NEE response to summer precipitation was less in the deep-rooted Tamarix community, compared to the shallow-rooted Haloxylon community, which was even converted into a carbon source after summer precipitation inputs. As a result, the effect of summer precipitation inputs on soil respiration was more important than the plant (carbon assimilation) response in determining the ecosystem response to episodic precipitation pulses.     

Ecosystem respiration and its controlling factors in a coniferous and broad-leaved mixed forest in Dinghushan, China

Accurate estimation of ecosystem respiration (R_eco) in forest ecosysteM_s is critical for validating terrestrial carbon models. Continuous eddy covariance measuremenT_s of R_eco were conducted in a coniferous and broad-leaved mixed forest located in Dinghushan Nature Reserve of southern China. R_eco was estimated and the controlling environmental factors were analyzed based on two years' data from 2003 to 2004. Major resulT_s included that: (1) R_eco was affected by soil temperature, soil moisture, canopy air temperature and humidity, where soil temperature at 5 cm depth was the dominant factor. (2) The exponential equation, Van't Hoff equation, Arrhenius equation and Lyold-Talor equation can be used to describe the relationship between R_eco and temperature factors with similar statistical significance, while Lyold-Talor equation was the most sensitive to the temperature index (Q₁₀). (3) The multiplicative model driven by soil temperature (T_s) and soil moisture (M_s) was more corresponsive to R_eco, which explained that there were more R_eco variations than Lyold-Talor equation, both for higher and lower M_s. However, there was no statistical difference between the two models. (4) Annually accumulated R_eco of the mixed forest in 2003 was estimated as 1100–1135.6 gC m^?2 a^?1 by using daytime data, which was 12%–25% higher than R_eco (921–975 gC m^?2 a^?1) estimated by using nighttime data. The resulT_s suggested that using daytime data to estimate R_eco can avoid the common underestimation problem caused by using eddy covariance methods. The study provides a basic method for further study on accurate estimation of net ecosystem CO₂ exchange (NEE) in the coniferous and broad-leaved mixed forest in southern China.