Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Divergent phenological response to hydroclimate variability in forested mountain watersheds

Mountain watersheds are primary sources of freshwater, carbon sequestration, and other ecosystem services. There is significant interest in the effects of climate change and variability on these processes over short to long time scales. Much of the impact of hydroclimate variability in forest ecosystems is manifested in vegetation dynamics in space and time. In steep terrain, leaf phenology responds to topoclimate in complex ways, and can produce specific and measurable shifts in landscape forest patterns. The onset of spring is usually delayed at a specific rate with increasing elevation (often called Hopkins' Law; Hopkins, 1918), reflecting the dominant controls of temperature on greenup timing. Contrary with greenup, leaf senescence shows inconsistent trends along elevation gradients. Here, we present mechanisms and an explanation for this variability and its significance for ecosystem patterns and services in response to climate. We use moderate‐resolution imaging spectro‐radiometer (MODIS) Normalized Difference Vegetation Index (NDVI) data to derive landscape‐induced phenological patterns over topoclimate gradients in a humid temperate broadleaf forest in southern Appalachians. These phenological patterns are validated with different sets of field observations. Our data demonstrate that divergent behavior of leaf senescence with elevation is closely related to late growing season hydroclimate variability in temperature and water balance patterns. Specifically, a drier late growing season is associated with earlier leaf senescence at low elevation than at middle elevation. The effect of drought stress on vegetation senescence timing also leads to tighter coupling between growing season length and ecosystem water use estimated from observed precipitation and runoff generation. This study indicates increased late growing season drought may be leading to divergent ecosystem response between high and low elevation forests. Landscape‐induced phenological patterns are easily observed over wide areas and may be used as a unique diagnostic for sources of ecosystem vulnerability and sensitivity to hydroclimate change.     

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

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

Deriving a new phenological indicator of interannual net carbon exchange in contrasting boreal deciduous and evergreen forests

Phenology is an important variable affecting the annual net ecosystem production (NEP) of terrestrial ecosystems. A new phenological indicator was proposed based on the ratio of respiration season length and growing season length (respiration–growth length ratio, RGR). Validation of this new phenological indicator was conducted using continuous flux measurements at contrasting boreal deciduous and evergreen forests in Canada. Analyses based on yearly anomalies of both annual NEP and phenological indicators indicated that the RGR can explain more proportion of interannual NEP variability compared to existing phenological metrics, including the carbon uptake period and the autumn lag. A multivariate regression model was used to predict the respiration–growth length ratio anomaly using anomalies of spring air temperature, autumn radiation and soil water content (SWC), which serves as a prerequisite for this indicator being scaled up for regional applications where flux data were unavailable. By normalization growing season length, interannual NEP showed comparable sensitivity to RGR variations of different plant functional types, which is a great advantage over other phenological indicators. The high potential of RGR in explaining interannual NEP variability may highlight the importance of respiration process in controlling annual NEP, which has probably been overlooked or underestimated in existing phenological studies. The comparable sensitivity of RGR to annual NEP observed at different plant functional types would favor its application in tracking interannual variability of NEP regionally and complementary to existing indices to promote our understanding of carbon sequestration with future climate change.     

Modelling the interannual variability of net ecosystem CO2 exchange at a subarctic sedge fen

This paper presents an empirical model of net ecosystem CO₂ exchange (NEE) developed for a subarctic fen near Churchill, Manitoba. The model with observed data helps explain the interannual variability in growing season NEE. Five years of tower‐flux data are used to test and examine the seasonal behaviour of the model simulations. Processes controlling the observed interannual variability of CO₂ exchange at the fen are examined by exploring the sensitivity of the model to changes in air temperature, precipitation and leaf area index. Results indicate that the sensitivity of NEE to changing environmental controls is complex and varies interannually depending on the initial conditions of the wetland. Changes in air temperature and the timing of precipitation events have a strong influence on NEE, which is largely manifest in gross ecosystem photosynthesis (GEP). Climate change scenarios indicate that warmer air temperatures will increase carbon acquisition during wet years but may act to reduce wetland carbon storage in years that experience a large water deficit early in the growing season. Model simulations for this subarctic sedge fen indicate that carbon acquisition is greatest during wet and warm conditions. This suggests therefore that carbon accumulation was greatest at this subarctic fen during its early developmental stages when hydroclimatic conditions were relatively wet and warm at approximately 2500 years before present.     

物候变化对北美温带落叶阔叶林生态系统生产力的影响

数据源、时间范围、空间尺度等的差异导致许多物候变化对陆地生态系统碳收支影响的研究缺少可比性。该文基于4级碳通量填充数据, 采用相对阈值方法提取了两个北美典型温带阔叶林站Harvard Forest (HF)和University of Michigan Biological Station (UMBS)共20年的物候参数(返青期、枯黄期和生长季长度), 并研究了物候变化对生态系统生产力的影响。结果表明: 1)生长季长度的延长对年累积总初级生产力(GPP)有显著贡献, 但由于呼吸作用(RE)的干扰, 生长季长度变化对年净生态系统生产力(NEP)的影响并不显著; 2)返青期的提前对上半年生态系统总初级生产力的贡献最为显著, 二者的相关系数分别为0.76 (HF)和0.93 (UMBS); 3)枯黄期的延迟对生产力的影响并不显著; 4)随着春季返青期的提前或秋季枯黄期的延迟, 上、下半年GPP和RE的累积量虽均有增加趋势, 但由于各自增加的幅度不确定, 导致年NEP与二者的响应关系复杂。     

Spatial analysis of growing season length control over net ecosystem exchange

Using data from 28 flux measurement sites, we performed an analysis of the relationship between annual net ecosystem exchange (NEE) and the length of the carbon uptake period (CUP) (the number of days when the ecosystem is a net carbon sink). The observations suggest a linear correlation between the two quantities. The change in annual carbon exchange per day of the CUP differs significantly between deciduous and evergreen vegetation types. The sites containing vegetation with short‐lived foliage (less than 1 year) have higher carbon uptake and respiration rates than evergreen vegetation. The ratio between mean daily carbon exchange rates during carbon uptake and release periods is relatively invariant (2.73±1.08) across different vegetation types. This implies that a balance between carbon release and uptake periods exists despite different photosynthetic pathways, life forms, and leaf habits. The mean daily carbon sequestration rate for these ecosystems never exceeds the carbon emission rate by more than a factor of 3. Growing season lengths for the study sites were derived from the normalized difference vegetation index (NDVI) of advanced very‐high‐resolution radiometer and from the enhanced vegetation index (EVI) of VEGETATION SPOT‐4. NDVI and EVI were found to be closely related to the CUP, and consequently they also can be used to approximate annual carbon exchange of the ecosystems. This approach has potential for allowing extrapolation of NEE over large areas from remotely sensed data, given a certain amount of ancillary information. This method could complement the currently existing techniques for extrapolation, which rely upon modeling of the individual gross fluxes.     

Interannual variability in net CO2 exchange of a native tallgrass prairie

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

Maximum carbon uptake rate dominates the interannual variability of global net ecosystem exchange

Terrestrial ecosystems contribute most of the interannual variability (IAV) in atmospheric carbon dioxide (CO₂) concentrations, but processes driving the IAV of net ecosystem CO₂ exchange (NEE) remain elusive. For a predictive understanding of the global C cycle, it is imperative to identify indicators associated with ecological processes that determine the IAV of NEE. Here, we decompose the annual NEE of global terrestrial ecosystems into their phenological and physiological components, namely maximum carbon uptake (MCU) and release (MCR), the carbon uptake period (CUP), and two parameters, α and β, that describe the ratio between actual versus hypothetical maximum C sink and source, respectively. Using long‐term observed NEE from 66 eddy covariance sites and global products derived from FLUXNET observations, we found that the IAV of NEE is determined predominately by MCU at the global scale, which explains 48% of the IAV of NEE on average while α, CUP, β, and MCR explain 14%, 25%, 2%, and 8%, respectively. These patterns differ in water‐limited ecosystems versus temperature‐ and radiation‐limited ecosystems; 31% of the IAV of NEE is determined by the IAV of CUP in water‐limited ecosystems, and 60% of the IAV of NEE is determined by the IAV of MCU in temperature‐ and radiation‐limited ecosystems. The Lund‐Potsdam‐Jena (LPJ) model and the Multi‐scale Synthesis and Terrestrial Model Inter‐comparison Project (MsTMIP) models underestimate the contribution of MCU to the IAV of NEE by about 18% on average, and overestimate the contribution of CUP by about 25%. This study provides a new perspective on the proximate causes of the IAV of NEE, which suggest that capturing the variability of MCU is critical for modeling the IAV of NEE across most of the global land surface.     

Importance of recent shifts in soil thermal dynamics on growing season length, productivity, and carbon sequestration in terrestrial high-latitude ecosystems

In terrestrial high‐latitude regions, observations indicate recent changes in snow cover, permafrost, and soil freeze–thaw transitions due to climate change. These modifications may result in temporal shifts in the growing season and the associated rates of terrestrial productivity. Changes in productivity will influence the ability of these ecosystems to sequester atmospheric CO₂. We use the terrestrial ecosystem model (TEM), which simulates the soil thermal regime, in addition to terrestrial carbon (C), nitrogen and water dynamics, to explore these issues over the years 1960–2100 in extratropical regions (30–90°N). Our model simulations show decreases in snow cover and permafrost stability from 1960 to 2100. Decreases in snow cover agree well with National Oceanic and Atmospheric Administration satellite observations collected between the years 1972 and 2000, with Pearson rank correlation coefficients between 0.58 and 0.65. Model analyses also indicate a trend towards an earlier thaw date of frozen soils and the onset of the growing season in the spring by approximately 2–4 days from 1988 to 2000. Between 1988 and 2000, satellite records yield a slightly stronger trend in thaw and the onset of the growing season, averaging between 5 and 8 days earlier. In both, the TEM simulations and satellite records, trends in day of freeze in the autumn are weaker, such that overall increases in growing season length are due primarily to earlier thaw. Although regions with the longest snow cover duration displayed the greatest increase in growing season length, these regions maintained smaller increases in productivity and heterotrophic respiration than those regions with shorter duration of snow cover and less of an increase in growing season length. Concurrent with increases in growing season length, we found a reduction in soil C and increases in vegetation C, with greatest losses of soil C occurring in those areas with more vegetation, but simulations also suggest that this trend could reverse in the future. Our results reveal noteworthy changes in snow, permafrost, growing season length, productivity, and net C uptake, indicating that prediction of terrestrial C dynamics from one decade to the next will require that large‐scale models adequately take into account the corresponding changes in soil thermal regimes.     

Leaf age affects the seasonal pattern of photosynthetic capacityand net ecosystem exchange of carbon in a deciduous forest

Temporal trends in photosynthetic capacity are a critical factorin determining the seasonality and magnitude of ecosystem carbonfluxes. At a mixed deciduous forest in the south‐eastern United States (Walker Branch Watershed, Oak Ridge, TN, USA), we independently measured seasonal trends in photosynthetic capacity (using single‐leaf gas exchange techniques) and the whole‐canopycarbon flux (using the eddy covariance method). Soil respiration was also measured using chambers and an eddy covariance system beneath the canopy. These independent chamber and eddy covariance measurements, along with a biophysical model (CANOAK), areused to examine how leaf age affects the seasonal pattern of carbon uptake during the growing season. When the measured seasonality in photosynthetic capacity is representedin the CANOAK simulations, there is good agreement with the eddy covariance data on the seasonal trends in carbon uptake. Removing the temporal trends in the simulations by using the early season maximum value of photosynthetic capacity over the entire growing season over estimates the annual carbon uptake by about 300 g C m^?2 year^?1– halfthe total estimated annual net ecosystem exchange. Alternatively, use of the mean value of photosynthetic capacity incorrectly simulates the seasonality in carbon uptake by the forest. In addition to changes related to leaf development and senescence, photosynthetic capacitydecreased in the middle and late summer, even when leaf nitrogenwas essentially constant. When only these middle and late summer reductions were neglected in the model simulations, CANOAK still overestimated the carbon uptake by an amount comparable to 25% ofthe total annual net ecosystem exchange.     

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

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

FLUXNET and modelling the global carbon cycle

Measurements of the net CO₂ flux between terrestrial ecosystems and the atmosphere using the eddy covariance technique have the potential to underpin our interpretation of regional CO₂ source–sink patterns, CO₂ flux responses to forcings, and predictions of the future terrestrial C balance. Information contained in FLUXNET eddy covariance data has multiple uses for the development and application of global carbon models, including evaluation/validation, calibration, process parameterization, and data assimilation. This paper reviews examples of these uses, compares global estimates of the dynamics of the global carbon cycle, and suggests ways of improving the utility of such data for global carbon modelling. Net ecosystem exchange of CO₂ (NEE) predicted by different terrestrial biosphere models compares favourably with FLUXNET observations at diurnal and seasonal timescales. However, complete model validation, particularly over the full annual cycle, requires information on the balance between assimilation and decomposition processes, information not readily available for most FLUXNET sites. Site history, when known, can greatly help constrain the model‐data comparison. Flux measurements made over four vegetation types were used to calibrate the land‐surface scheme of the Goddard Institute for Space Studies global climate model, significantly improving simulated climate and demonstrating the utility of diurnal FLUXNET data for climate modelling. Land‐surface temperatures in many regions cool due to higher canopy conductances and latent heat fluxes, and the spatial distribution of CO₂ uptake provides a significant additional constraint on the realism of simulated surface fluxes. FLUXNET data are used to calibrate a global production efficiency model (PEM). This model is forced by satellite‐measured absorbed radiation and suggests that global net primary production (NPP) increased 6.2% over 1982–1999. Good agreement is found between global trends in NPP estimated by the PEM and a dynamic global vegetation model (DGVM), and between the DGVM and estimates of global NEE derived from a global inversion of atmospheric CO₂ measurements. Combining the PEM, DGVM, and inversion results suggests that CO₂ fertilization is playing a major role in current increases in NPP, with lesser impacts from increasing N deposition and growing season length. Both the PEM and the inversion identify the Amazon basin as a key region for the current net terrestrial CO₂ uptake (i.e. 33% of global NEE), as well as its interannual variability. The inversion's global NEE estimate of −1.2 Pg [C] yr⁻¹ for 1982–1995 is compatible with the PEM‐ and DGVM‐predicted trends in NPP. There is, thus, a convergence in understanding derived from process‐based models, remote‐sensing‐based observations, and inversion of atmospheric data. Future advances in field measurement techniques, including eddy covariance (particularly concerning the problem of night‐time fluxes in dense canopies and of advection or flow distortion over complex terrain), will result in improved constraints on land‐atmosphere CO₂ fluxes and the rigorous attribution of mechanisms to the current terrestrial net CO₂ uptake and its spatial and temporal heterogeneity. Global ecosystem models play a fundamental role in linking information derived from FLUXNET measurements to atmospheric CO₂ variability. A number of recommendations concerning FLUXNET data are made, including a request for more comprehensive site data (particularly historical information), more measurements in undisturbed ecosystems, and the systematic provision of error estimates. The greatest value of current FLUXNET data for global carbon cycle modelling is in evaluating process representations, rather than in providing an unbiased estimate of net CO₂ exchange.     

The influence of meteorology and phenology on net ecosystem exchange in an eastern Siberian boreal larch forest

Aims Boreal forests play an important role in the global carbon cycle. Compared with the boreal forests in North America and Europe, relatively few research studies have been conducted in Siberian boreal forests. Knowledge related to the role of Siberian forests in the global carbon balance is thus essential for a full understanding of global carbon cycle.Methods This study investigated the net ecosystem exchange (NEE) during growing season (May–September) in an eastern Siberian boreal larch forest for a 3-year period in 2004–2006 with contrasting meteorological conditions.Important findings The study found that the forest served as a carbon sink during all of the 3 studied years; in addition, the meteorological conditions essentially influenced the specific annual value of the strength of the carbon sinks in each year. Although 2005 was the warmest year and much wetter than 2004, 2005 also featured the greatest amount of ecosystem respiration, which resulted in a minimum value of NEE. The study also found that the phenological changes observed during the three study years had a relatively small effect on annual NEE. Leaf expansion was 26 days earlier in 2005 than in the other 2 years, which resulted in a longer growing season in 2005. However, the NEE in 2005 was counterbalanced by the large rate of ecosystem respiration that was caused by the higher temperatures in the year. This study showed that meteorological variables had larger influences on the interannual variations in NEE for a Siberian boreal larch forest, as compared with phenological changes. The overall results of this study will improve our understanding of the carbon balance of Siberian boreal larch forests and thus can help to forecast the response of these forests to future climate change.     

Role of vegetation in determining carbon sequestration along ecological succession in the southeastern United States

Vegetation plays a central role in controlling terrestrial carbon (C) exchange, but quantifying its impacts on C cycling on time scales of ecological succession is hindered by a lack of long‐term observations. The net ecosystem exchange of carbon (NEE) was measured for several years in adjacent ecosystems that represent distinct phases of ecological succession in the southeastern USA. The experiment was designed to isolate the role of vegetation – apart from climate and soils – in controlling biosphere–atmosphere fluxes of CO₂ and water vapor. NEE was near zero over 5 years at an early successional old‐field ecosystem (OF). However, mean annual NEE was nearly equal, approximately ?450 g C m^?2 yr^?1, at an early successional planted pine forest (PP) and a late successional hardwood forest (HW) due to the sensitivity of the former to drought and ice storm damage. We hypothesize that these observations can be explained by the relationships between gross ecosystem productivity (GEP), ecosystem respiration (RE) and canopy conductance, and long‐term shifts in ecosystem physiology in response to climate to maintain near‐constant ecosystem‐level water‐use efficiency (EWUE). Data support our hypotheses, but future research should examine if GEP and RE are causally related or merely controlled by similar drivers. At successional time scales, GEP and RE observations generally followed predictions from E. P. Odum's ‘Strategy of Ecosystem Development’, with the surprising exception that the relationship between GEP and RE resulted in large NEE at the late successional HW. A practical consequence of this research suggests that plantation forestry may confer no net benefit over the conservation of mature forests for C sequestration.     

Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model

The Lund–Potsdam–Jena Dynamic Global Vegetation Model (LPJ) combines process‐based, large‐scale representations of terrestrial vegetation dynamics and land‐atmosphere carbon and water exchanges in a modular framework. Features include feedback through canopy conductance between photosynthesis and transpiration and interactive coupling between these ‘fast’ processes and other ecosystem processes including resource competition, tissue turnover, population dynamics, soil organic matter and litter dynamics and fire disturbance. Ten plants functional types (PFTs) are differentiated by physiological, morphological, phenological, bioclimatic and fire‐response attributes. Resource competition and differential responses to fire between PFTs influence their relative fractional cover from year to year. Photosynthesis, evapotranspiration and soil water dynamics are modelled on a daily time step, while vegetation structure and PFT population densities are updated annually. Simulations have been made over the industrial period both for specific sites where field measurements were available for model evaluation, and globally on a 0.5°° × 0.5°° grid. Modelled vegetation patterns are consistent with observations, including remotely sensed vegetation structure and phenology. Seasonal cycles of net ecosystem exchange and soil moisture compare well with local measurements. Global carbon exchange fields used as input to an atmospheric tracer transport model (TM2) provided a good fit to observed seasonal cycles of CO₂ concentration at all latitudes. Simulated inter‐annual variability of the global terrestrial carbon balance is in phase with and comparable in amplitude to observed variability in the growth rate of atmospheric CO₂. Global terrestrial carbon and water cycle parameters (pool sizes and fluxes) lie within their accepted ranges. The model is being used to study past, present and future terrestrial ecosystem dynamics, biochemical and biophysical interactions between ecosystems and the atmosphere, and as a component of coupled Earth system models.     

Temperature as a control over ecosystem CO<Subscript>2</Subscript> fluxes in a high-elevation,subalpine forest

We evaluated the hypothesis that CO(2) uptake by a subalpine, coniferous forest is limited by cool temperature during the growing season. Using the eddy covariance approach we conducted observations of net ecosystem CO(2) exchange (NEE) across two growing seasons. When pooled for the entire growing season during both years, light-saturated net ecosystem CO(2) exchange (NEE(sat)) exhibited a temperature optimum within the range 7-12 degrees C. Ecosystem respiration rate ( R(e)), calculated as the y-intercept of the NEE versus photosynthetic photon flux density (PPFD) relationship, increased with increasing temperature, causing a 15% reduction in net CO(2) uptake capacity for this ecosystem as temperatures increased from typical early season temperatures of 7 degrees C to typical mid-season temperatures of 18 degrees C. The ecosystem quantum yield and the ecosystem PPFD compensation point, which are measures of light-utilization efficiency, were highest during the cool temperatures of the early season, and decreased later in the season at higher temperatures. Branch-level measurements revealed that net photosynthesis in all three of the dominant conifer tree species exhibited a temperature optimum near 10 degrees C early in the season and 15 degrees C later in the season. Using path analysis, we statistically isolated temperature as a seasonal variable, and identified the dynamic role that temperature exhibits in controlling ecosystem fluxes early and late in the season. During the spring, an increase in temperature has a positive effect on NEE, because daytime temperatures progress from near freezing to near the photosynthetic temperature optimum, and R(e )values remain low. During the middle of the summer an increase in temperature has a negative effect on NEE, because inhibition of net photosynthesis and increases in R(e). When taken together, the results demonstrate that in this high-elevation forest ecosystem CO(2) uptake is not limited by cool-temperature constraints on photosynthetic processes during the growing-season, as suggested by some previous ecophysiological studies at the branch and needle levels. Rather, it is warm temperatures in the mid-summer, and their effect on ecosystem respiration, that cause the greatest reduction in the potential for forest carbon sequestration.     

Delayed herbivory by migratory geese increases summer‐long CO2 uptake in coastal western Alaska

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to “phenological mismatches” as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three‐year experiment to examine the consequences for CO₂ exchange of advanced spring green‐up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green‐up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO₂ uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green‐up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m^?2 s^?1 toward a greater CO₂ sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO₂ uptake. Importantly, while the direct effect of climate warming on phenology of green‐up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.     

Comparing the impact of cloudiness on carbon dioxide exchange in a grassland and a maize cropland in northwestern China

Light quantity and quality strongly influence plant growth. However, different ecosystems have different capabilities to assimilate solar radiation. In this study, the effects of cloudiness intensity on the net ecosystem exchange of carbon dioxide (NEE) were compared between an alpine grassland (with lower leaf area index) at A??Rou and an oasis maize cropland (with higher leaf area index) at Yingke, using flux data obtained during the middle of the growing season (July?CAugust) in 2008 and 2009. The results showed that the response of NEE to photosynthetically active radiation (PAR) was more negative (carbon uptake) under cloudy than under clear skies at both sites. The maximum NEE occurred when the clearness index (CI) ranged from 0.4 to 0.7 under cloudy skies. The maximum enhancements were 11.9% for solar elevation angles of 60?C65° in the grassland, and 34.9% for solar elevation angles of 60?C65° and 10.3% for angles of 35?C40° in the maize cropland before the irrigation period. The response of NEE to CI changed slightly with solar elevation angle in the grassland compared to the maize cropland. The results indicate that enhanced NEE under cloudy skies can be attributed to increasingly diffuse PAR and interactions with environmental factors (air temperature and vapor pressure deficit).     

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.