Seasonal and interannual variation in carbon dioxide exchange and carbon balance in a northern temperate grassland

Net ecosystem carbon dioxide (CO₂) exchange (NEE) was measured in a northern temperate grassland near Lethbridge, Alberta, Canada for three growing seasons using the eddy covariance technique. The study objectives were to document how NEE and its major component processes—gross photosynthesis (GPP) and total ecosystem respiration (TER)—vary seasonally and interannually, and to examine how environmental and physiological factors influence the annual C budget. The greatest difference among the three study years was the amount of precipitation received. The annual precipitation for 1998 (481.7 mm) was significantly above the 1971–2000 mean (± SD, 377.9 ± 97.0 mm) for Lethbridge, whereas 1999 (341.3 mm) was close to average, and 2000 (275.5 mm) was significantly below average. The high precipitation and soil moisture in 1998 allowed a much higher GPP and an extended period of net carbon gain relative to 1999 and 2000. In 1998, the peak NEE was a gain of 5 g C m^?2 d^?1 (day 173). Peak NEE was lower and also occurred earlier in the year on days 161 (3.2 g C m^?2 d^?1) and 141 (2.4 g C m^?2 d^?1) in 1999 and 2000, respectively. Change in soil moisture was the most important ecological factor controlling C gain in this grassland ecosystem. Soil moisture content was positively correlated with leaf area index (LAI). Gross photosynthesis was strongly correlated with changes in both LAI and canopy nitrogen (N) content. Maximum GPP (A_max: value calculated from a rectangular hyperbola fitted to the relationship between GPP and incident photosynthetic photon flux density (PPFD)) was 27.5, 12.9 and 8.6 µmol m^?2 s^?1 during 1998, 1999 and 2000, respectively. The apparent quantum yield also differed among years at the time of peak photosynthetic activity, with calculated values of 0.0254, 0.018 and 0.018 during 1998, 1999 and 2000, respectively. The ecosystem accumulated a total of 111.9 g C m^?2 from the time the eddy covariance measurements were initiated in June 1998 until the end of December 2000, with most of that C gained during 1998. There was a net uptake of almost 21 g C m^?2 in 1999, whereas a net loss of 18 g C m^?2 was observed in 2000. The net uptake of C during 1999 was the combined result of slightly higher GPP (287.2 vs. 272.3 g C m^?2 year^?1) and lower TER (266.6 vs. 290.4 g C m^?2 year^?1) than occurred in 2000.     

Seasonal controls on interannual variability in carbon dioxide exchange of a near-end-of rotation Douglas-fir stand in the Pacific Northwest, 1997–2006

This study analyzes 9 years of eddy‐covariance (EC) data carried out in a Pacific Northwest Douglas‐fir (Pseudotsuga menzesii) forest (58‐year old in 2007) on the east coast of Vancouver Island, Canada, and characterizes the seasonal and interannual variability in net ecosystem productivity (NEP), gross primary productivity (GPP), and ecosystem respiration (R_e) and primary climatic controls on these fluxes. The annual values (± SD) of NEP, GPP and R_e were 357 ± 51, 2124 ± 125, and 1767 ± 146 g C m^?2 yr^?1, respectively, with ranges of 267–410, 1592–2338, and 1642–2071 g C m^?2 yr^?1, respectively. Spring to early summer (March–June) accounted for more than 80% of annual NEP while late spring to early autumn (May–August) was mainly responsible for its interannual variability (～80%). The major drivers of interannual variability in annual carbon (C) fluxes were annual and spring mean air temperatures (T_a) and water deficiency during late summer and autumn (July–October) when this Douglas‐fir forest growth was often water‐limited. Photosynthetically active radiation (Q), and the combination of Q and soil water content (θ) explained 85% and 91% of the variance of monthly GPP, respectively; and 91% and 96% of the variance of monthly R_e was explained by T_a and the combination of T_a and θ, respectively. Annual net C sequestration was high during optimally warm and normal precipitation years, but low in unusually warm or severely dry years. Excluding 1998 and 1999, the 2 years strongly affected by an El Niño/La Niña cycle, annual NEP significantly decreased with increasing annual mean T_a. Annual NEP will likely decrease whereas both annual GPP and R_e will likely increase if the future climate at the site follows a trend similar to that of the past 40 years.     

Reduction of ecosystem productivity and respiration during the European summer 2003 climate anomaly: a joint flux tower, remote sensing and modelling analysis

The European CARBOEUROPE/FLUXNET monitoring sites, spatial remote sensing observations via the EOS‐MODIS sensor and ecosystem modelling provide independent and complementary views on the effect of the 2003 heatwave on the European biosphere's productivity and carbon balance. In our analysis, these data streams consistently demonstrate a strong negative anomaly of the primary productivity during the summer of 2003. FLUXNET eddy‐covariance data indicate that the drop in productivity was not primarily caused by high temperatures (‘heat stress’) but rather by limitation of water (drought stress) and that, contrary to the classical expectation about a heat wave, not only gross primary productivity but also ecosystem respiration declined by up to more than to 80 gC m⁻² month⁻¹. Anomalies of carbon and water fluxes were strongly correlated. While there are large between‐site differences in water‐use efficiency (WUE, 1–6 kg C kg⁻¹ H₂O) here defined as gross carbon uptake divided by evapotranspiration (WUE=GPP/ET), the year‐to‐year changes in WUE were small (<1 g kg⁻¹) and quite similar for most sites (i.e. WUE decreased during the year of the heatwave). Remote sensing data from MODIS and AVHRR both indicate a strong negative anomaly of the fraction of absorbed photosynthetically active radiation in summer 2003, at more than five standard deviations of the previous years. The spatial differentiation of this anomaly follows climatic and land‐use patterns: Largest anomalies occur in the centre of the meteorological anomaly (central Western Europe) and in areas dominated by crops or grassland. A preliminary model intercomparison along a gradient from data‐oriented models to process‐oriented models indicates that all approaches are similarly describing the spatial pattern of ecosystem sensitivity to the climatic 2003 event with major exceptions in the Alps and parts of Eastern Europe, but differed with respect to their interannual variability.     

Warmer and drier conditions stimulate respiration more than photosynthesis in a boreal peatland ecosystem: Analysis of automatic chambers and eddy covariance measurements

Continuous half‐hourly net CO₂ exchange measurements were made using nine automatic chambers in a treed fen in northern Alberta, Canada from June–October in 2005 and from May–October in 2006. The 2006 growing season was warmer and drier than in 2005. The average chamber respiration rates normalized to 10 °C were much higher in 2006 than in 2005, while calculations of the temperature sensitivity (Q₁₀) values were similar in the two years. Daytime average respiration values were lower than the corresponding, temperature‐corrected respiration rates calculated from night‐time chamber measurements. From June to September, the season‐integrated estimates of chamber photosynthesis and respiration were 384 and 590 g C m^?2, respectively in 2006, an increase of 100 and 203 g C m^?2 over the corresponding values in 2005. The season‐integrated photosynthesis and respiration rates obtained using the eddy covariance technique, which included trees and a tall shrub not present in the chambers, were 720 and 513 g C m^?2, respectively, in 2006, an increase of 50 and 125 g C m^?2 over the corresponding values in 2005. While both photosynthesis and respiration rates were higher in the warmer and drier conditions of 2006, the increase in respiration was more than twice the increase in photosynthesis.     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

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.     

Diurnal, seasonal and annual variation in the net ecosystem CO2 exchange of a desert shrub community (Sarcocaulescent) in Baja California, Mexico

Estimates of net ecosystem exchange (NEE) of CO₂ have been measured on a variety of ecosystems world wide including grasslands, savannahs, boreal, pine, deciduous, Mediterranean and tropical rain forests as well as arctic tundra. While there have been numerous comparisons between net primary productivity of arid and semiarid grasslands and shrublands, notably lacking are estimates of NEE with a few exceptions. The objective of this study was to characterize the seasonal and annual carbon flux of a desert shrub ecosystem using the eddy covariance technique to determine the sensitivity of the system to the timing and varying amounts of precipitation. Measurements began in July of 2001, a year with 339 mm of rainfall, considerably above the long‐term average of 174 mm and preceded by 2 years of below average rainfall (50–62 mm). Over the 2 complete years of measurements, precipitation was 147 and 197 mm in 2002 and 2003, respectively. In all years, the majority of the precipitation fell between August and September. The site was a sink of ?39 g C m^?2 yr^?1 in 2002 with a relatively strong uptake in the early part of the year and reduced uptake after the suboptimal rainfall in September. This contrasts with 2003 when the ecosystem took up ?52 g C m^?2 yr^?1 concentrated in the fall after significant rain in August and September. Likely, extremely low rainfall years would result in a carbon loss while a strengthening of the typical winter secondary peak in precipitation (notably absent in the 2 years of measurements) may extend uptake into the spring resulting in more carbon accumulation. The system appears to be buffered against variations in annual rainfall attributed to water storage in the stems and roots.     

Temperate forest responses to carbon dioxide, temperature and nitrogen: a model analysis

The ITE Edinburgh Forest Model, which describes diurnal and seasonal changes in the pools and fluxes of C, N and water in a fully coupled forest–soil system, was parametrized to simulate a managed conifer plantation in upland Britain. The model was used to examine (i) the transient effects on forest growth of an IS92a scenario of increasing [CO₂] and temperature over two future rotations, and (ii) the equilibrium (sustainable) effects of all combinations of increases in [CO₂] from 350 to 550 and 750 μmol mol^?1, mean annual temperature from 7.5 to 8.5 and 9.5°C and annual inputs of 20 or 40 kg N ha^?1. Changes in underlying processes represented in the model were then used to explain the responses. Eight conclusions were supported by the model for this forest type and climate.

• 1 Increasing temperatures above 3°C alone may cause forest decline owing to water stress.
• 2 Elevated [CO₂] can protect trees from water stress that they may otherwise suffer in response to increased temperature.
• 3 In N-limiting conditions, elevated [CO₂] can increase allocation to roots with little increase in leaf area, whereas in N-rich conditions elevated [CO₂] may not increase allocation to roots and generally increases leaf area.
• 4 Elevated [CO₂] can decrease water use by forests in N-limited conditions and increase water use in N-rich conditions.
• 5 Elevated [CO₂] can increase forest productivity even in N-limiting conditions owing to increased N acquisition and use efficiency.
• 6 Rising temperatures (along with rising [CO₂]) may increase or decrease forest productivity depending on the supply of N and changes in water stress.
• 7 Gaseous losses of N from the soil can increase or decrease in response to elevated [CO₂] and temperature.
• 8 Projected increases in [CO₂] and temperature (IS92a) are likely to increase net ecosystem productivity and hence C sequestration in temperate forests.

     

Modeled interactive effects of precipitation, temperature, and [CO2] on ecosystem carbon and water dynamics in different climatic zones

Interactive effects of multiple global change factors on ecosystem processes are complex. It is relatively expensive to explore those interactions in manipulative experiments. We conducted a modeling analysis to identify potentially important interactions and to stimulate hypothesis formulation for experimental research. Four models were used to quantify interactive effects of climate warming (T), altered precipitation amounts [doubled (DP) and halved (HP)] and seasonality (SP, moving precipitation in July and August to January and February to create summer drought), and elevated [CO₂] (C) on net primary production (NPP), heterotrophic respiration (R_h), net ecosystem production (NEP), transpiration, and runoff. We examined those responses in seven ecosystems, including forests, grasslands, and heathlands in different climate zones. The modeling analysis showed that none of the three‐way interactions among T, C, and altered precipitation was substantial for either carbon or water processes, nor consistent among the seven ecosystems. However, two‐way interactive effects on NPP, R_h, and NEP were generally positive (i.e. amplification of one factor's effect by the other factor) between T and C or between T and DP. A negative interaction (i.e. depression of one factor's effect by the other factor) occurred for simulated NPP between T and HP. The interactive effects on runoff were positive between T and HP. Four pairs of two‐way interactive effects on plant transpiration were positive and two pairs negative. In addition, wet sites generally had smaller relative changes in NPP, R_h, runoff, and transpiration but larger absolute changes in NEP than dry sites in response to the treatments. The modeling results suggest new hypotheses to be tested in multifactor global change experiments. Likewise, more experimental evidence is needed for the further improvement of ecosystem models in order to adequately simulate complex interactive processes.     

Dependence of carbon sequestration on the differential responses of ecosystem photosynthesis and respiration to rain pulses in a semiarid steppe

Precipitation pulses play an important role in regulating ecosystem carbon exchange and balance of semiarid steppe ecosystems. It has been predicted that the frequency of extreme rain events will increase in the future, especially in the arid and semiarid regions. We hypothesize that large rain pulses favor carbon sequestration, while small ones cause more carbon release in the semiarid steppes. To understand the potential response in carbon sequestration capacity of semiarid steppes to the changes in rain pulse size, we conducted a manipulative experiment with five simulated rain pulse sizes (0, 5, 10, 25, and 75 mm) in Inner Mongolia steppe. Our results showed that both gross ecosystem productivity (GEP) and ecosystem respiration (R_e) responded rapidly (within 24 h) to rain pulses and the initial response time was independent of pulse size. However, the time of peak GEP was 1–3 days later than that of R_e, which depended on pulse size. Larger pulses caused greater magnitude and longer duration of variations in GEP and R_e. Differences in the response time of microbes and plants to wetting events constrained the response pattern of heterotrophic (R_h) and autotrophic (R_a) components of R_e following a rain event. R_h contributed more to the increase of R_e in the early stage of rain pulse response, while R_a played an more important role later, and determined the duration of pulse response, especially for large rain events of >10 mm. The distinct responses of ecosystem photosynthesis and respiration to increasing pulse sizes led to a threshold in rain pulse size between 10 and 25 mm, above which post wetting responses favored carbon sequestration. The disproportionate increase of the primary productivity of higher plants, compared with those in the activities of microbial decomposers to larger pulse events suggests that the carbon sequestration capacity of Inner Mongolia steppes will be sensitive to changes in precipitation size distribution rather than just precipitation amount.     

Impact of past and present land‐management on the C‐balance of a grassland in the Swiss Alps

Grasslands cover about 40% of the ice‐free global terrestrial surface, but their quantitative importance in global carbon exchange with the atmosphere is still highly uncertain, and thus their potential for carbon sequestration remains speculative. Here, we report on CO₂ exchange of an extensively used mountain hay meadow and pasture in the Swiss pre‐Alps on high‐organic soils (7–45% C by mass) over a 3‐year period (18 May 2002–20 September 2005), including the European summer 2003 heat‐wave period. During all 3 years, the ecosystem was a net source of CO₂ (116–256 g C m^?2 yr^?1). Harvests and grazing cows (mostly via C export in milk) further increased these C losses, which were estimated at 355 g C m^?2 yr^?1 during 2003 (95% confidence interval 257–454 g C m^?2 yr^?1). Although annual carbon losses varied considerably among years, the CO₂ budget during summer 2003 was not very different from the other two summers. However, and much more importantly, the winter that followed the warm summer of 2003 observed a significantly higher carbon loss when there was snow (133±6 g C m^?2) than under comparable conditions during the other two winters (73±5 and 70±4 g C m^?2, respectively). The continued annual C losses can most likely be attributed to the long‐term effects of drainage and peat exploitation that began 119 years ago, with the last significant drainage activities during the Second World War around 1940. The most realistic estimate based on depth profiles of ash content after combustion suggests that there is an 500–910 g C m^?2 yr^?1 loss associated with the decomposition of organic matter. Our results clearly suggest that putting efforts into preserving still existing carbon stocks may be more successful than attempts to increase sequestration rates in such high‐organic mountain grassland soils.     

Ecosystem responses to water and nitrogen amendment in a California grassland

The world's ecosystems are experiencing simultaneous changes in the supply of multiple limiting resources. Two of these, water and nitrogen (N) can strongly limit grassland production and can affect community composition and biogeochemical cycles in different ways. Grassland ecosystems in California may be particularly vulnerable to current and predicted changes in precipitation and N deposition, and ecosystem responses to potential interactive effects of water and N are not well understood. Here, we show strong colimitation of plant production resulting from factorial addition of water and N. In addition, water and N addition in combination led to increased dominance of the two most abundant grass species, while N addition regardless of water availability led to decreased species diversity. Late season carbon (C) flux response to water addition depended on N. Only plots that received additional water, but not N, still showed net ecosystem C uptake at the end of the experiment. Our results suggest that grassland ecosystem response to N deposition will be strongly dependent on future precipitation patterns.     

Antecedent moisture and seasonal precipitation influence the response of canopy-scale carbon and water exchange to rainfall pulses in a semi-arid grassland

The influences of prior monsoon-season drought (PMSD) and the seasonal timing of episodic rainfall ('pulses') on carbon and water exchange in water-limited ecosystems are poorly quantified. *In the present study, we estimated net ecosystem exchange of CO(2) (NEE) and evapotranspiration (ET) before, and for 15 d following, experimental irrigation in a semi-arid grassland during June and August 2003. Rainout shelters near Tucson, Arizona, USA, were positioned on contrasting soils (clay and sand) and planted with native (Heteropogon contortus) or non-native invasive (Eragrostis lehmanniana) C4 bunchgrasses. Plots received increased ('wet') or decreased ('dry') monsoon-season (July-September) rainfall during 2002 and 2003. Following a June 2003 39-mm pulse, species treatments had similar NEE and ET dynamics including 15-d integrated NEE (NEE(pulse)). Contrary to predictions, PMSD increased net C uptake during June in plots of both species. Greater flux rates after an August 2003 39-mm pulse reflected biotic activity associated with the North American Monsoon. Furthermore, August NEE(pulse) and ecosystem pulse-use efficiency (PUE(e) = NEE(pulse)/ET(pulse)) was greatest in Heteropogon plots. PMSD and rainfall seasonal timing may interact with bunchgrass invasions to alter NEE and ET dynamics with consequences for PUE(e) in water-limited ecosystems.     

A long-term record of carbon exchange in a boreal black spruce forest: means, responses to interannual variability, and decadal trends

We present a decadal (1994–2004) record of carbon dioxide flux in a 160‐year‐old black spruce forest/veneer bog complex in central Manitoba, Canada. The ecosystem shifted from a source (+41 g C m⁻², 1995) to a sink (−21 g C m⁻², 2004) of CO₂ over the decade, with an average net carbon balance near zero. Annual mean temperatures increased 1–2° during the period, consistent with the decadal trend across the North American boreal biome. We found that ecosystem carbon exchange responded strongly to air temperature, moisture status, potential evapotranspiration, and summertime solar radiation. The seasonal cycle of ecosystem respiration significantly lagged that of photosynthesis, limited by the rate of soil thaw and the slow drainage of the soil column. Factors acting over long time scales, especially water table depth, strongly influenced the carbon budget on annual time scales. Net uptake was enhanced and respiration inhibited by multiple years of rainfall in excess of evaporative demand. Contrary to expectations, we observed no correlation between longer growing seasons and net uptake, possibly because of offsetting increases in ecosystem respiration. The results indicate that the interactions between soil thaw and water table depth provide critical controls on carbon exchange in boreal forests underlain by peat, on seasonal to decadal time scales, and these factors must be simulated in terrestrial biosphere models to predict response of these regions to future climate.     

Response of plant species richness and primary productivity in shrublands along a north–south gradient in Europe to seven years of experimental warming and drought: reductions in primary productivity in the heat and drought year of 2003

We used a nonintrusive field experiment carried out at six sites – Wales (UK), Denmark (DK), the Netherlands (NL), Hungary (HU), Sardinia (Italy – IT), and Catalonia (Spain – SP) – along a climatic and latitudinal gradient to examine the response of plant species richness and primary productivity to warming and drought in shrubland ecosystems. The warming treatment raised the plot daily temperature by ca. 1 °C, while the drought treatment led to a reduction in soil moisture at the peak of the growing season that ranged from 26% at the SP site to 82% in the NL site. During the 7 years the experiment lasted (1999–2005), we used the pin‐point method to measure the species composition of plant communities and plant biomass, litterfall, and shoot growth of the dominant plant species at each site. A significantly lower increase in the number of species pin‐pointed per transect was found in the drought plots at the SP site, where the plant community was still in a process of recovering from a forest fire in 1994. No changes in species richness were found at the other sites, which were at a more mature and stable state of succession and, thus less liable to recruitment of new species. The relationship between annual biomass accumulation and temperature of the growing season was positive at the coldest site and negative at the warmest site. The warming treatment tended to increase the aboveground net primary productivity (ANPP) at the northern sites. The relationship between annual biomass accumulation and soil moisture during the growing season was not significant at the wettest sites, but was positive at the driest sites. The drought treatment tended to reduce the ANPP in the NL, HU, IT, and SP sites. The responses to warming were very strongly related to the Gaussen aridity index (stronger responses the lower the aridity), whereas the responses to drought were not. Changes in the annual aboveground biomass accumulation, litterfall, and, thus, the ANPP, mirrored the interannual variation in climate conditions: the most outstanding change was a decrease in biomass accumulation and an increase in litterfall at most sites during the abnormally hot year of 2003. Species richness also tended to decrease in 2003 at all sites except the cold and wet UK site. Species‐specific responses to warming were found in shoot growth: at the SP site, Globularia alypum was not affected, while the other dominant species, Erica multiflora, grew 30% more; at the UK site, Calluna vulgaris tended to grow more in the warming plots, while Empetrum nigrum tended to grow less. Drought treatment decreased plant growth in several studied species, although there were some species such as Pinus halepensis at the SP site or C. vulgaris at the UK site that were not affected. The magnitude of responses to warming and drought thus depended greatly on the differences between sites, years, and species and these multiple plant responses may be expected to have consequences at ecosystem and community level. Decreases in biodiversity and the increase in E. multiflora growth at the SP site as a response to warming challenge the assumption that sensitivity to warming may be less well developed at more southerly latitudes; likewise, the fact that one of the studied shrublands presented negative ANPP as a response to the 2003 heat wave also challenges the hypothesis that future climate warming will lead to an enhancement of plant growth and carbon sequestration in temperate ecosystems. Extreme events may thus change the general trend of increased productivity in response to warming in the colder sites.     

Physiological and environmental regulation of interannual variability in CO2 exchange on rangelands in the western United States

For most ecosystems, net ecosystem exchange of CO₂ (NEE) varies within and among years in response to environmental change. We analyzed measurements of CO₂ exchange from eight native rangeland ecosystems in the western United States (58 site‐years of data) in order to determine the contributions of photosynthetic and respiratory (physiological) components of CO₂ exchange to environmentally caused variation in NEE. Rangelands included Great Plains grasslands, desert shrubland, desert grasslands, and sagebrush steppe. We predicted that (1) week‐to‐week change in NEE and among‐year variation in the response of NEE to temperature, net radiation, and other environmental drivers would be better explained by change in maximum rates of ecosystem photosynthesis (A_max) than by change in apparent light‐use efficiency (α) or ecosystem respiration at 10 °C (R₁₀) and (2) among‐year variation in the responses of NEE, A_max, and α to environmental drivers would be explained by changes in leaf area index (LAI). As predicted, NEE was better correlated with A_max than α or R₁₀ for six of the eight rangelands. Week‐to‐week variation in NEE and physiological parameters correlated mainly with time‐lagged indices of precipitation and water‐related environmental variables, like potential evapotranspiration, for desert sites and with net radiation and temperature for Great Plains grasslands. For most rangelands, the response of NEE to a given change in temperature, net radiation, or evaporative demand differed among years because the response of photosynthetic parameters (A_max, α) to environmental drivers differed among years. Differences in photosynthetic responses were not explained by variation in LAI alone. A better understanding of controls on canopy photosynthesis will be required to predict variation in NEE of rangeland ecosystems.     

Functional diversity of photosynthesis during drought in a model tropical rainforest – the contributions of leaf area, photosynthetic electron transport and stomatal conductance to reduction in net ecosystem carbon exchange

The tropical rainforest mesocosm within the Biosphere 2 Laboratory, a model system of some 110 species developed over 12 years under controlled environmental conditions, has been subjected to a series of comparable drought experiments during 2000–2002. In each study, the mesocosm was subjected to a 4–6 week drought, with well‐defined rainfall events before and after the treatment. Ecosystem CO₂ uptake rate (A_eco) declined 32% in response to the drought, with changes occurring within days and being reversible within weeks, even though the deeper soil layers did not become significantly drier and leaf‐level water status of most large trees was not greatly affected. The reduced A_eco during the drought reflected both morphological and physiological responses. It is estimated that the drought‐induced 32% reduction of A_eco has three principal components: (1) leaf fall increased two‐fold whereas leaf expansion growth of some canopy dominants declined to 60%, leading to a 10% decrease in foliage coverage of the canopy. This might be the main reason for the persistent reduction of A_eco after rewatering. (2) The maximum photosynthetic electron transport rate at high light intensities in remaining leaves was reduced to 71% for three of the four species measured, even though no chronic photo‐inhibition occurred. (3) Stomata closed, leading to a reduced ecosystem water conductance to water vapour (33% of pre‐drought values), which not only reduced ecosystem carbon uptake rate, but may also have implications for water and energy budgets of tropical ecosystems. Additionally, individual rainforest trees responded differently, expressing different levels of stress and stress avoiding mechanisms. This functional diversity renders the individual response heterogeneous and has fundamental implications to scale leaf level responses to ecosystem dynamics.