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.     

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.     

CO2 exchange and evapotranspiration across dryland ecosystems of southwestern North America

Global‐scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO₂ sink. However, such analyses are poorly constrained by measured CO₂ exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water‐limited Southwest region of North America with observed ranges in annual precipitation of 100–1000 mm, annual temperatures of 2–25°C, and records of 3–10 years (150 site‐years in total). Annual fluxes were integrated using site‐specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from ‐350 to +330 gCm^?2 across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest‐dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (R_eco) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and R_eco were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross‐site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS‐based models captured only 20–30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO₂ exchange may be up to 3–5 times larger than current estimates.     

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.     

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.     

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.     

The dependence of respiration on photosynthetic substrate supply and temperature: integrating leaf, soil and ecosystem measurements

Interactions between photosynthetic substrate supply and temperature in determining the rate of three respiration components (leaf, belowground and ecosystem respiration) were investigated within three environmentally controlled, Populus deltoides forest bays at Biosphere 2, Arizona. Over 2 months, the atmospheric CO₂ concentration and air temperature were manipulated to test the following hypotheses: (1) the responses of the three respiration components to changes in the rate of photosynthesis would differ both in speed and magnitude; (2) the temperature sensitivity of leaf and belowground respiration would increase in response to a rise in substrate availability; and, (3) at the ecosystem level, the ratio of respiration to photosynthesis would be conserved despite week‐to‐week changes in temperature. All three respiration rates responded to the CO₂ concentration‐induced changes in photosynthesis. However, the proportional change in the rate of leaf respiration was more than twice that of belowground respiration and, when photosynthesis was reduced, was also more rapid. The results suggest that aboveground respiration plays a key role in the overall response of ecosystem respiration to short‐term changes in canopy photosynthesis. The short‐term temperature sensitivity of leaf respiration, measured within a single night, was found to be affected more by developmental conditions than photosynthetic substrate availability, as the Q₁₀ was lower in leaves that developed at high CO₂, irrespective of substrate availability. However, the temperature sensitivity of belowground respiration, calculated between periods of differing air temperature, appeared to be positively correlated with photosynthetic substrate availability. At the ecosystem level, respiration and photosynthesis were positively correlated but the relationship was affected by temperature; for a given rate of daytime photosynthesis, the rate of respiration the following night was greater at 25 than 20°C. This result suggests that net ecosystem exchange did not acclimate to temperature changes lasting up to 3 weeks. Overall, the results of this study demonstrate that the three respiration terms differ in their dependence on photosynthesis and that, short‐ and medium‐term changes in temperature may affect net carbon storage in terrestrial ecosystems.     

Antecedent moisture and temperature conditions modulate the response of ecosystem respiration to elevated CO2 and warming

Terrestrial plant and soil respiration, or ecosystem respiration (R_eco), represents a major CO₂ flux in the global carbon cycle. However, there is disagreement in how R_eco will respond to future global changes, such as elevated atmosphere CO₂ and warming. To address this, we synthesized six years (2007–2012) of R_eco data from the Prairie Heating And CO₂ Enrichment (PHACE) experiment. We applied a semi‐mechanistic temperature–response model to simultaneously evaluate the response of R_eco to three treatment factors (elevated CO₂, warming, and soil water manipulation) and their interactions with antecedent soil conditions [e.g., past soil water content (SWC) and temperature (SoilT)] and aboveground factors (e.g., vapor pressure deficit, photosynthetically active radiation, vegetation greenness). The model fits the observed R_eco well (R² = 0.77). We applied the model to estimate annual (March–October) R_eco, which was stimulated under elevated CO₂ in most years, likely due to the indirect effect of elevated CO₂ on SWC. When aggregated from 2007 to 2012, total six‐year R_eco was stimulated by elevated CO₂ singly (24%) or in combination with warming (28%). Warming had little effect on annual R_eco under ambient CO₂, but stimulated it under elevated CO₂ (32% across all years) when precipitation was high (e.g., 44% in 2009, a ‘wet’ year). Treatment‐level differences in R_eco can be partly attributed to the effects of antecedent SoilT and vegetation greenness on the apparent temperature sensitivity of R_eco and to the effects of antecedent and current SWC and vegetation activity (greenness modulated by VPD) on R_eco base rates. Thus, this study indicates that the incorporation of both antecedent environmental conditions and aboveground vegetation activity are critical to predicting R_eco at multiple timescales (subdaily to annual) and under a future climate of elevated CO₂ and warming.     

Divergent long‐term trends and interannual variation in ecosystem resource use efficiencies of a southern boreal old black spruce forest 1999–2017

Long‐term trends in ecosystem resource use efficiencies (RUEs) and their controlling factors are key pieces of information for understanding how an ecosystem responds to climate change. We used continuous eddy covariance and microclimate data over the period 1999–2017 from a 120‐year‐old black spruce stand in central Saskatchewan, Canada, to assess interannual variability, long‐term trends, and key controlling factors of gross ecosystem production (GEP) and the RUEs of carbon (CUE = net primary production [NPP]/GEP), light (LUE = GEP/absorbed photosynthetic radiation [APAR]), and water (WUE = GEP/evapotranspiration [E]). At this site, annual GEP has shown an increasing trend over the 19 years (p < 0.01), which may be attributed to rising atmospheric CO₂ concentration. Interannual variability in GEP, aside from its increasing trend, was most strongly related to spring temperatures. Associated with the significant increase in annual GEP were relatively small changes in NPP, APAR, and E, so that annual CUE showed a decreasing trend and annual LUE and WUE showed increasing trends over the 19 years. The long‐term trends in the RUEs were related to the increasing CO₂ concentration. Further analysis of detrended RUEs showed that their interannual variation was impacted most strongly by air temperature. Two‐factor linear models combining CO₂ concentration and air temperature performed well (R²~0.60) in simulating annual RUEs. LUE and WUE were positively correlated both annually and seasonally, while LUE and CUE were mostly negatively correlated. Our results showed divergent long‐term trends among CUE, LUE, and WUE and highlighted the need to account for the combined effects of climatic controls and the ‘CO₂ fertilization effect’ on long‐term variations in RUEs. Since most RUE‐based models rely primarily on one resource limitation, the observed patterns of relative change among the three RUEs may have important implications for RUE‐based modeling of C fluxes.     

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.     

Estimates of CO2 uptake and release among European forests based on eddy covariance data

The net ecosystem exchange (NEE) of forests represents the balance of gross primary productivity (GPP) and respiration (R). Methods to estimate these two components from eddy covariance flux measurements are usually based on a functional relationship between respiration and temperature that is calibrated for night‐time (respiration) fluxes and subsequently extrapolated using daytime temperature measurements. However, respiration fluxes originate from different parts of the ecosystem, each of which experiences its own course of temperature. Moreover, if the temperature–respiration function is fitted to combined data from different stages of biological development or seasons, a spurious temperature effect may be included that will lead to overestimation of the direct effect of temperature and therefore to overestimates of daytime respiration. We used the EUROFLUX eddy covariance data set for 15 European forests and pooled data per site, month and for conditions of low and sufficient soil moisture, respectively. We found that using air temperature (measured above the canopy) rather than soil temperature (measured 5 cm below the surface) yielded the most reliable and consistent exponential (Q₁₀) temperature–respiration relationship. A fundamental difference in air temperature‐based Q₁₀ values for different sites, times of year or soil moisture conditions could not be established; all were in the range 1.6–2.5. However, base respiration (R₀, i.e. respiration rate scaled to 0°C) did vary significantly among sites and over the course of the year, with increased base respiration rates during the growing season. We used the overall mean Q₁₀ of 2.0 to estimate annual GPP and R. Testing suggested that the uncertainty in total GPP and R associated with the method of separation was generally well within 15%. For the sites investigated, we found a positive relationship between GPP and R, indicating that there is a latitudinal trend in NEE because the absolute decrease in GPP towards the pole is greater than in R.     

Decrease in winter respiration explains 25% of the annual northern forest carbon sink enhancement over the last 30 years

Aim Winter snow has been suggested to regulate terrestrial carbon (C) cycling by modifying microclimate, but the impacts of change in snow cover on the annual C budget at a large scale are poorly understood. Our aim is to quantify the C balance under changing snow depth. Location Non‐permafrost region of the northern forest area. Methods Here, we used site‐based eddy covariance flux data to investigate the relationship between depth of snow cover and ecosystem respiration (R_eco) during winter. We then used the Biome‐BGC model to estimate the effect of reductions in winter snow cover on the C balance of northern forests in the non‐permafrost region. Results According to site observations, winter net ecosystem C exchange (NEE) ranged from 0.028 to 1.53 gC·m^?2·day^?1, accounting for 44 ± 123% of the annual C budget. Model simulation showed that over the past 30 years, snow‐driven change in winter C fluxes reduced non‐growing season CO₂ emissions, enhancing the annual C sink of northern forests. Over the entire study area, simulated winter R_eco significantly decreased by 0.33 gC·m^?2·day^?1·year^?1 in response to decreasing depth of snow cover, which accounts for approximately 25% of the simulated annual C sink trend from 1982 to 2009. Main conclusion Soil temperature is primarily controlled by snow cover rather than by air temperature as snow serves as an insulator to prevent chilling impacts. A shallow snow cover has less insulation potential, causing colder soil temperatures and potentially lower respiration rates. Both eddy covariance analysis and model‐simulated results show that both R_eco and NEE are significantly and positively correlated with variation in soil temperature controlled by variation in snow depth. Overall, our results highlight that a decrease in winter snow cover restrains global warming as less C is emitted to the atmosphere.     

The growth respiration component in eddy CO2 flux from a Quercus ilex mediterranean forest

Ecosystem respiration, arising from soil decomposition as well as from plant maintenance and growth, has been shown to be the most important component of carbon exchange in most terrestrial ecosystems. The goal of this study was to estimate the growth component of whole‐ecosystem respiration in a Mediterranean evergreen oak (Quercus ilex) forest over the course of 3 years. Ecosystem respiration (R_eco) was determined from night‐time carbon dioxide flux (F_c) using eddy correlation when friction velocity (u_*) was greater than 0.35 m s^?1 We postulated that growth respiration could be evaluated as a residual after removing modeled base R_eco from whole‐ecosystem R_eco during periods when growth was most likely occurring. We observed that the model deviated from the night‐time F_c‐based R_eco during the period from early February to early July with the largest discrepancies occurring at the end of May, coinciding with budburst when active aboveground growth and radial growth increment are greatest. The highest growth respiration rates were observed in 2001 with daily fluxes reaching up to 4 g C m^?2. The cumulative growth respiration for the entire growth period gave total carbon losses of 170, 208, and 142 g C m^?2 for 1999, 2001, and 2002, respectively. Biochemical analysis of soluble carbohydrates, starch, cellulose, hemicellulose, proteins, lignin, and lipids for leaves and stems allowed calculation of the total construction costs of the different growth components, which yielded values of 154, 200, and 150 g C for 3 years, respectively, corresponding well to estimated growth respiration. Estimates of both leaf and stem growth showed very large interannual variation, although average growth respiration coefficients and average yield of growth processes were fairly constant over the 3 years and close to literature values. The time course of the growth respiration may be explained by the growth pattern of leaves and stems and by cambial activity. This approach has potential applications for interpreting the effects of climate variation, disturbances, and management practices on growth and ecosystem respiration.     

When vegetation change alters ecosystem water availability

The combined effects of vegetation and climate change on biosphere–atmosphere water vapor (H₂O) and carbon dioxide (CO₂) exchanges are expected to vary depending, in part, on how biotic activity is controlled by and alters water availability. This is particularly important when a change in ecosystem composition alters the fractional covers of bare soil, grass, and woody plants so as to influence the accessibility of shallower vs. deeper soil water pools. To study this, we compared 5 years of eddy covariance measurements of H₂O and CO₂ fluxes over a riparian grassland, shrubland, and woodland. In comparison with the surrounding upland region, groundwater access at the riparian sites increased net carbon uptake (NEP) and evapotranspiration (ET), which were sustained over more of the year. Among the sites, the grassland used less of the stable groundwater resource, and increasing woody plant density decoupled NEP and ET from incident precipitation (P), resulting in greater exchange rates that were less variable year to year. Despite similar gross patterns, how groundwater accessibility affected NEP was more complex than ET. The grassland had higher respiration (R_eco) costs. Thus, while it had similar ET and gross carbon uptake (GEP) to the shrubland, grassland NEP was substantially less. Also, grassland carbon fluxes were more variable due to occasional flooding at the site, which both stimulated and inhibited NEP depending upon phenology. Woodland NEP was large, but surprisingly similar to the less mature, sparse shrubland, even while having much greater GEP. Woodland R_eco was greater than the shrubland and responded strongly and positively to P, which resulted in a surprising negative NEP response to P. This is likely due to the large accumulation of carbon aboveground and in the surface soil. These long‐term observations support the strong role that water accessibility can play when determining the consequences of ecosystem vegetation change.     

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.     

Modelling night‐time ecosystem respiration by a constrained source optimization method

One of the main challenges to quantifying ecosystem carbon budgets is properly quantifying the magnitude of night‐time ecosystem respiration. Inverse Lagrangian dispersion analysis provides a promising approach to addressing such a problem when measured mean CO₂ concentration profiles and nocturnal velocity statistics are available. An inverse method, termed ‘Constrained Source Optimization’ or CSO, which couples a localized near‐field theory (LNF) of turbulent dispersion to respiratory sources, is developed to estimate seasonal and annual components of ecosystem respiration. A key advantage to the proposed method is that the effects of variable leaf area density on flow statistics are explicitly resolved via higher‐order closure principles. In CSO, the source distribution was computed after optimizing key physiological parameters to recover the measured mean concentration profile in a least‐square fashion. The proposed method was field‐tested using 1 year of 30‐min mean CO₂ concentration and CO₂ flux measurements collected within a 17‐year‐old (in 1999) even‐aged loblolly pine (Pinus taeda L.) stand in central North Carolina. Eddy‐covariance flux measurements conditioned on large friction velocity, leaf‐level porometry and forest‐floor respiration chamber measurements were used to assess the performance of the CSO model. The CSO approach produced reasonable estimates of ecosystem respiration, which permits estimation of ecosystem gross primary production when combined with daytime net ecosystem exchange (NEE) measurements. We employed the CSO approach in modelling annual respiration of above‐ground plant components (c. 214 g C m^?2 year^?1) and forest floor (c. 989 g C m^?2 year^?1) for estimating gross primary production (c. 1800 g C m^?2 year^?1) with a NEE of c. 605 g C m^?2 year^?1 for this pine forest ecosystem. We conclude that the CSO approach can utilise routine CO₂ concentration profile measurements to corroborate forest carbon balance estimates from eddy‐covariance NEE and chamber‐based component flux measurements.     

Severe drought effects on ecosystem CO2 and H2O fluxes at three Mediterranean evergreen sites: revision of current hypotheses?

Eddy covariance and sapflow data from three Mediterranean ecosystems were analysed via top‐down approaches in conjunction with a mechanistic ecosystem gas‐exchange model to test current assumptions about drought effects on ecosystem respiration and canopy CO₂/H₂O exchange. The three sites include two nearly monospecific Quercus ilex L. forests – one on karstic limestone (Puéchabon), the other on fluvial sand with access to ground water (Castelporziano) – and a typical mixed macchia on limestone (Arca di Noè). Estimates of ecosystem respiration were derived from light response curves of net ecosystem CO₂ exchange. Subsequently, values of ecosystem gross carbon uptake were computed from eddy covariance CO₂ fluxes and estimates of ecosystem respiration as a function of soil temperature and moisture. Bulk canopy conductance was calculated by inversion of the Penman‐Monteith equation. In a top‐down analysis, it was shown that all three sites exhibit similar behaviour in terms of their overall response to drought. In contrast to common assumptions, at all sites ecosystem respiration revealed a decreasing temperature sensitivity ( Q ₁₀) in response to drought. Soil temperature and soil water content explained 70–80% of the seasonal variability of ecosystem respiration. During the drought, light‐saturated ecosystem gross carbon uptake and day‐time averaged canopy conductance declined by up to 90%. These changes were closely related to soil water content. Ecosystem water‐use efficiency of gross carbon uptake decreased during the drought, regardless whether evapotranspiration from eddy covariance or transpiration from sapflow had been used for the calculation. We evidence that this clearly contrasts current models of canopy function which predict increasing ecosystem water‐use efficiency (WUE) during the drought. Four potential explanations to those results were identified (patchy stomatal closure, changes in physiological capacities of photosynthesis, decreases in mesophyll conductance for CO₂, and photoinhibition), which will be tested in a forthcoming paper. It is suggested to incorporate the new findings into current biogeochemical models after further testing as this will improve estimates of climate change effects on (semi)arid ecosystems' carbon balances.     

Carbon sequestration in boreal jack pine stands following harvesting

A large area of boreal jack pine (Pinus banksiana Lamb.) forest in Canada is recovering from clear‐cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO₂ exchange was measured using the eddy‐covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was ?137±11, 19±16, 73±28, and 22±30 g C m^?2 yr^?1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m^?2 yr^?1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration (R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2‐year‐old stand. R was most sensitive to soil temperature for the 90‐year‐old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year‐round flux measurements from 2000 to 2005, the 10‐year stand had larger interannual variability in GEP and R than the 90‐year stand. Interannual variability in NEP was driven primarily by early‐growing‐season temperature and growing‐season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration.     

Seasonal and annual respiration of a ponderosa pine ecosystem

The net ecosystem exchange of CO₂ between forests and the atmosphere, measured by eddy covariance, is the small difference between two large fluxes of photosynthesis and respiration. Chamber measurements of soil surface CO₂ efflux (F_s), wood respiration (F_w) and foliage respiration (F_f) help identify the contributions of these individual components to net ecosystem exchange. Models developed from the chamber data also provide independent estimates of respiration costs. We measured CO₂ efflux with chambers periodically in 1996–97 in a ponderosa pine forest in Oregon, scaled these measurements to the ecosystem, and computed annual totals for respiration by component. We also compared estimated half-hourly ecosystem respiration at night (F_nc) with eddy covariance measurements. Mean foliage respiration normalized to 10 °C was 0.20 μmol m^–2 (hemi-leaf surface area) s^–1, and reached a maximum of 0.24 μmol m^–2 HSA s^–1 between days 162 and 208. Mean wood respiration normalized to 10 °C was 5.9 μmol m^–3 sapwood s^–1, with slightly higher rates in mid-summer, when growth occurs. There was no significant difference (P > 0.10) between wood respiration of young (45 years) and old trees (250 years). Soil surface respiration normalized to 10 °C ranged from 0.7 to 3.0 μmol m^–2 (ground) s^–1 from days 23 to 329, with the lowest rates in winter and highest rates in late spring. Annual CO₂ flux from soil surface, foliage and wood was 683, 157, and 54 g C m^–2 y^–1, with soil fluxes responsible for 76% of ecosystem respiration. The ratio of net primary production to gross primary production was 0.45, consistent with values for conifer sites in Oregon and Australia, but higher than values reported for boreal coniferous forests. Below-ground carbon allocation (root turnover and respiration, estimated as F_s– litterfall carbon) consumed 61% of GPP; high ratios such as this are typical of sites with more water and nutrient constraints. The chamber estimates were moderately correlated with change in CO₂ storage in the canopy (F_stor) on calm nights (friction velocity u* < 0.25 m s^–1; R² = 0.60); F_stor was not significantly different from summed chamber estimates. On windy nights (u* > 0.25 m s^–1), the sum of turbulent flux measured above the canopy by eddy covariance and F_stor was only weakly correlated with summed chamber estimates (R² = 0.14); the eddy covariance estimates were lower than chamber estimates by 50%.     

Measurements of gross and net ecosystem productivity and water vapour exchange of a Pinus ponderosa ecosystem,and an evaluation of two generalized models

Net ecosystem productivity (NEP), net primary productivity (NPP), and water vapour exchange of a mature Pinus ponderosa forest (44°30′ N, 121°37′ W) growing in a region subject to summer drought were investigated along with canopy assimilation and respiratory fluxes. This paper describes seasonal and annual variation in these factors, and the evaluation of two generalized models of carbon and water balance (PnET‐II and 3‐PG) with a combination of traditional measurements of NPP, respiration and water stress, and eddy covariance measurements of above‐and below‐canopy CO₂ and water vapour exchange. The objective was to evaluate the models using two years of traditional and eddy covariance measurements, and to use the models to help interpret the relative importance of processes controlling carbon and water vapour exchange in a water‐limited pine ecosystem throughout the year. PnET‐II is a monthly time‐step model that is driven by nitrogen availability through foliar N concentration, and 3‐PG is a monthly time‐step quantum‐efficiency model constrained by extreme temperatures, drought, and vapour pressure deficits. Both models require few parameters and have the potential to be applied at the watershed to regional scale. There was 2/3 less rainfall in 1997 than in 1996, providing a challenge to modelling the water balance, and consequently the carbon balance, when driving the models with the two years of climate data, sequentially. Soil fertility was not a key factor in modelling processes at this site because other environmental factors limited photosynthesis and restricted projected leaf area index to ～1.6. Seasonally, GEP and LE were overestimated in early summer and underestimated through the rest of the year. The model predictions of annual GEP, NEP and water vapour exchange were within 1–39% of flux measurements, with greater disparity in 1997 because soil water never fully recharged. The results suggest that generalized models can provide insights to constraints on productivity on an annual basis, using a minimum of site data.