State of the science in reconciling top‐down and bottom‐up approaches for terrestrial CO2 budget

Robust estimates of CO₂ budget, CO₂ exchanged between the atmosphere and terrestrial biosphere, are necessary to better understand the role of the terrestrial biosphere in mitigating anthropogenic CO₂ emissions. Over the past decade, this field of research has advanced through understanding of the differences and similarities of two fundamentally different approaches: “top‐down” atmospheric inversions and “bottom‐up” biosphere models. Since the first studies were undertaken, these approaches have shown an increasing level of agreement, but disagreements in some regions still persist, in part because they do not estimate the same quantity of atmosphere–biosphere CO₂ exchange. Here, we conducted a thorough comparison of CO₂ budgets at multiple scales and from multiple methods to assess the current state of the science in estimating CO₂ budgets. Our set of atmospheric inversions and biosphere models, which were adjusted for a consistent flux definition, showed a high level of agreement for global and hemispheric CO₂ budgets in the 2000s. Regionally, improved agreement in CO₂ budgets was notable for North America and Southeast Asia. However, large gaps between the two methods remained in East Asia and South America. In other regions, Europe, boreal Asia, Africa, South Asia, and Oceania, it was difficult to determine whether those regions act as a net sink or source because of the large spread in estimates from atmospheric inversions. These results highlight two research directions to improve the robustness of CO₂ budgets: (a) to increase representation of processes in biosphere models that could contribute to fill the budget gaps, such as forest regrowth and forest degradation; and (b) to reduce sink–source compensation between regions (dipoles) in atmospheric inversion so that their estimates become more comparable. Advancements on both research areas will increase the level of agreement between the top‐down and bottom‐up approaches and yield more robust knowledge of regional CO₂ budgets.     

Interannual variability in global soil respiration, 1980–94

We used a climate‐driven regression model to develop spatially resolved estimates of soil‐CO₂ emissions from the terrestrial land surface for each month from January 1980 to December 1994, to evaluate the effects of interannual variations in climate on global soil‐to‐atmosphere CO₂ fluxes. The mean annual global soil‐CO₂ flux over this 15‐y period was estimated to be 80.4 (range 79.3–81.8) Pg C. Monthly variations in global soil‐CO₂ emissions followed closely the mean temperature cycle of the Northern Hemisphere. Globally, soil‐CO₂ emissions reached their minima in February and peaked in July and August. Tropical and subtropical evergreen broad‐leaved forests contributed more soil‐derived CO₂ to the atmosphere than did any other vegetation type (～30% of the total) and exhibited a biannual cycle in their emissions. Soil‐CO₂ emissions in other biomes exhibited a single annual cycle that paralleled the seasonal temperature cycle. Interannual variability in estimated global soil‐CO₂ production is substantially less than is variability in net carbon uptake by plants (i.e., net primary productivity). Thus, soils appear to buffer atmospheric CO₂ concentrations against far more dramatic seasonal and interannual differences in plant growth. Within seasonally dry biomes (savannas, bushlands and deserts), interannual variability in soil‐CO₂ emissions correlated significantly with interannual differences in precipitation. At the global scale, however, annual soil‐CO₂ fluxes correlated with mean annual temperature, with a slope of 3.3 Pg C y^?1 per °C. Although the distribution of precipitation influences seasonal and spatial patterns of soil‐CO₂ emissions, global warming is likely to stimulate CO₂ emissions from soils.     

Effects of 21st‐century climate,land use,and disturbances on ecosystem carbon balance in California

Terrestrial ecosystems are an important sink for atmospheric carbon dioxide (CO₂), sequestering ~30% of annual anthropogenic emissions and slowing the rise of atmospheric CO₂. However, the future direction and magnitude of the land sink is highly uncertain. We examined how historical and projected changes in climate, land use, and ecosystem disturbances affect the carbon balance of terrestrial ecosystems in California over the period 2001–2100. We modeled 32 unique scenarios, spanning 4 land use and 2 radiative forcing scenarios as simulated by four global climate models. Between 2001 and 2015, carbon storage in California's terrestrial ecosystems declined by ?188.4 Tg C, with a mean annual flux ranging from a source of ?89.8 Tg C/year to a sink of 60.1 Tg C/year. The large variability in the magnitude of the state's carbon source/sink was primarily attributable to interannual variability in weather and climate, which affected the rate of carbon uptake in vegetation and the rate of ecosystem respiration. Under nearly all future scenarios, carbon storage in terrestrial ecosystems was projected to decline, with an average loss of ?9.4% (?432.3 Tg C) by the year 2100 from current stocks. However, uncertainty in the magnitude of carbon loss was high, with individual scenario projections ranging from ?916.2 to 121.2 Tg C and was largely driven by differences in future climate conditions projected by climate models. Moving from a high to a low radiative forcing scenario reduced net ecosystem carbon loss by 21% and when combined with reductions in land‐use change (i.e., moving from a high to a low land‐use scenario), net carbon losses were reduced by 55% on average. However, reconciling large uncertainties associated with the effect of increasing atmospheric CO₂ is needed to better constrain models used to establish baseline conditions from which ecosystem‐based climate mitigation strategies can be evaluated.     

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.     

Towards a coupled paradigm of NH3‐CO2 biosphere–atmosphere exchange modelling

Stomatal conductance, one of the major plant physiological controls within NH₃ biosphere–atmosphere exchange models, is commonly estimated from semi‐empirical multiplicative schemes or simple light‐ and temperature‐response functions. However, due to their inherent parameterization on meteorological proxy variables, instead of a direct measure of stomatal opening, they are unfit for the use in climate change scenarios and of limited value for interpreting field‐scale measurements. Alternatives based on H₂O flux measurements suffer from uncertainties in the partitioning of evapotranspiration at humid sites, as well as a potential decoupling of transpiration from stomatal opening in the presence of hygroscopic particles on leaf surfaces. We argue that these problems may be avoided by directly deriving stomatal conductance from CO₂ fluxes instead. We reanalysed a data set of NH₃ flux measurements based on CO₂‐derived stomatal conductance, confirming the hypothesis that the increasing relevance of stomatal exchange with the onset of vegetation activity caused a rapid decrease of observed NH₃ deposition velocities. Finally, we argue that developing more mechanistic representations of NH₃ biosphere–atmosphere exchange can be of great benefit in many applications. These range from model‐based flux partitioning, over deposition monitoring using low‐cost samplers and inferential modelling, to a direct response of NH₃ exchange to climate change.     

Terrestrial biosphere models need better representation of vegetation phenology: results from the North American Carbon Program Site Synthesis

Phenology, by controlling the seasonal activity of vegetation on the land surface, plays a fundamental role in regulating photosynthesis and other ecosystem processes, as well as competitive interactions and feedbacks to the climate system. We conducted an analysis to evaluate the representation of phenology, and the associated seasonality of ecosystem‐scale CO₂ exchange, in 14 models participating in the North American Carbon Program Site Synthesis. Model predictions were evaluated using long‐term measurements (emphasizing the period 2000–2006) from 10 forested sites within the AmeriFlux and Fluxnet‐Canada networks. In deciduous forests, almost all models consistently predicted that the growing season started earlier, and ended later, than was actually observed; biases of 2 weeks or more were typical. For these sites, most models were also unable to explain more than a small fraction of the observed interannual variability in phenological transition dates. Finally, for deciduous forests, misrepresentation of the seasonal cycle resulted in over‐prediction of gross ecosystem photosynthesis by +160 ± 145 g C m^?2 yr^?1 during the spring transition period and +75 ± 130 g C m^?2 yr^?1 during the autumn transition period (13% and 8% annual productivity, respectively) compensating for the tendency of most models to under‐predict the magnitude of peak summertime photosynthetic rates. Models did a better job of predicting the seasonality of CO₂ exchange for evergreen forests. These results highlight the need for improved understanding of the environmental controls on vegetation phenology and incorporation of this knowledge into better phenological models. Existing models are unlikely to predict future responses of phenology to climate change accurately and therefore will misrepresent the seasonality and interannual variability of key biosphere–atmosphere feedbacks and interactions in coupled global climate models.     

Carbon respired by terrestrial ecosystems – recent progress and challenges

Net ecosystem production is the residual of two much larger fluxes: photosynthesis and respiration. While photosynthesis is a single process with a well‐established theoretical underpinning, respiration integrates the variety of plant and microbial processes by which CO₂ returns from ecosystems to the atmosphere. Limits to current capacity for predicting ecosystem respiration fluxes across biomes or years result from the mismatch between what is usually measured – bulk CO₂ fluxes – and what process‐based models can predict – fluxes of CO₂ from plant (autotrophic) or microbial (heterotrophic) respiration. Papers in this Thematic Issue and in the recent literature, document advances in methods for separating respiration into autotrophic and heterotrophic components using three approaches: (1) continuous measurements of CO₂ fluxes and assimilation of these data into process‐based models; (2) application of isotope measurements, particularly radiocarbon; and (3) manipulation experiments. They highlight the role of allocation of C fixed by plants to respiration, storage, growth or transfer to other organisms as a control of seasonal and interannual variability in soil respiration and the oxidation state of C in the terrestrial biosphere. A second theme is the potential for comparing C isotope signatures in organic matter, CO₂ evolved in incubations and microbial biomarkers to elucidate the pathways (respiration, recycling, or transformation) of C during decomposition. Together, these factors determine the continuum of timescales over which C is returned to the atmosphere by respiration and enable testing of theories of plant and microbial respiration that go beyond empirical models and allow predictions of future respiration responses to future change in climate, pollution and land use.     

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO₂ exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO₂ exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site‐years of annual eddy covariance measurements of net and gross CO₂ exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 – 1000 mm in annual precipitation and records of 4–9 years each. In addition to evaluating spatial relationships among CO₂ and water fluxes across sites, we separately quantified site‐level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis–ET relationship was linear, suggesting ET was a better proxy for water available to drive CO₂ exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site‐level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long‐term mean CO₂ exchanges with climatic ET. Consequently, a hypothetical 100‐mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm^?2 yr^?1. Most of the unexplained NEP variability was related to persistent, site‐specific function, suggesting prioritization of research on slow‐changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site‐level responses to interannual weather can be extrapolated for prediction of CO₂ exchanges over decadal and longer timescales relevant to societal response to climate change.     

Assessing the eddy covariance technique for evaluating carbon dioxide exchange rates of ecosystems: past, present and future

The eddy covariance technique ascertains the exchange rate of CO₂ across the interface between the atmosphere and a plant canopy by measuring the covariance between fluctuations in vertical wind velocity and CO₂ mixing ratio. Two decades ago, the method was employed to study CO₂ exchange of agricultural crops under ideal conditions during short field campaigns. During the past decade the eddy covariance method has emerged as an important tool for evaluating fluxes of carbon dioxide between terrestrial ecosystems and the atmosphere over the course of a year, and more. At present, the method is being applied in a nearly continuous mode to study carbon dioxide and water vapor exchange at over a hundred and eighty field sites, worldwide. The objective of this review is to assess the eddy covariance method as it is being applied by the global change community on increasingly longer time scales and over less than ideal surfaces. The eddy covariance method is most accurate when the atmospheric conditions (wind, temperature, humidity, CO₂) are steady, the underlying vegetation is homogeneous and it is situated on flat terrain for an extended distance upwind. When the eddy covariance method is applied over natural and complex landscapes or during atmospheric conditions that vary with time, the quantification of CO₂ exchange between the biosphere and atmosphere must include measurements of atmospheric storage, flux divergence and advection. Averaging CO₂ flux measurements over long periods (days to year) reduces random sampling error to relatively small values. Unfortunately, data gaps are inevitable when constructing long data records. Data gaps are generally filled with values produced from statistical and empirical models to produce daily and annual sums of CO₂ exchange. Filling data gaps with empirical estimates do not introduce significant bias errors because the empirical algorithms are derived from large statistical populations. On the other hand, flux measurement errors can be biased at night when winds are light and intermittent. Nighttime bias errors tend to produce an underestimate in the measurement of ecosystem respiration. Despite the sources of errors associated with long‐term eddy flux measurements, many investigators are producing defensible estimates of annual carbon exchange. When measurements come from nearly ideal sites the error bound on the net annual exchange of CO₂ is less than ±50 g C m^?2 yr^?1. Additional confidence in long‐term measurements is growing because investigators are producing values of net ecosystem productivity that are converging with independent values produced by measuring changes in biomass and soil carbon, as long as the biomass inventory studies are conducted over multiple years.     

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.     

Carbon cycle responses of semi‐arid ecosystems to positive asymmetry in rainfall

Recent evidence shows that warm semi‐arid ecosystems are playing a disproportionate role in the interannual variability and greening trend of the global carbon cycle given their mean lower productivity when compared with other biomes (Ahlström et al. 2015 Science, 348, 895). Using multiple observations (land‐atmosphere fluxes, biomass, streamflow and remotely sensed vegetation cover) and two state‐of‐the‐art biospheric models, we show that climate variability and extremes lead to positive or negative responses in the biosphere, depending on vegetation type. We find Australia to be a global hot spot for variability, with semi‐arid ecosystems in that country exhibiting increased carbon uptake due to both asymmetry in the interannual distribution of rainfall (extrinsic forcing), and asymmetry in the response of gross primary production (GPP) to rainfall change (intrinsic response). The latter is attributable to the pulse‐response behaviour of the drought‐adapted biota of these systems, a response that is estimated to be as much as half of that from the CO₂ fertilization effect during 1990–2013. Mesic ecosystems, lacking drought‐adapted species, did not show an intrinsic asymmetric response. Our findings suggest that a future more variable climate will induce large but contrasting ecosystem responses, differing among biomes globally, independent of changes in mean precipitation alone. The most significant changes are occurring in the extensive arid and semi‐arid regions, and we suggest that the reported increased carbon uptake in response to asymmetric responses might be contributing to the observed greening trends there.     

Aquatic carbon fluxes dampen the overall variation of net ecosystem productivity in the Amazon basin: An analysis of the interannual variability in the boundless carbon cycle

The river–floodplain network plays an important role in the carbon (C) cycle of the Amazon basin, as it transports and processes a significant fraction of the C fixed by terrestrial vegetation, most of which evades as CO₂ from rivers and floodplains back to the atmosphere. There is empirical evidence that exceptionally dry or wet years have an impact on the net C balance in the Amazon. While seasonal and interannual variations in hydrology have a direct impact on the amounts of C transferred through the river–floodplain system, it is not known how far the variation of these fluxes affects the overall Amazon C balance. Here, we introduce a new wetland forcing file for the ORCHILEAK model, which improves the representation of floodplain dynamics and allows us to closely reproduce data‐driven estimates of net C exports through the river–floodplain network. Based on this new wetland forcing and two climate forcing datasets, we show that across the Amazon, the percentage of net primary productivity lost to the river–floodplain system is highly variable at the interannual timescale, and wet years fuel aquatic CO₂ evasion. However, at the same time overall net ecosystem productivity (NEP) and C sequestration are highest during wet years, partly due to reduced decomposition rates in water‐logged floodplain soils. It is years with the lowest discharge and floodplain inundation, often associated with El Nino events, that have the lowest NEP and the highest total (terrestrial plus aquatic) CO₂ emissions back to atmosphere. Furthermore, we find that aquatic C fluxes display greater variation than terrestrial C fluxes, and that this variation significantly dampens the interannual variability in NEP of the Amazon basin. These results call for a more integrative view of the C fluxes through the vegetation‐soil‐river‐floodplain continuum, which directly places aquatic C fluxes into the overall C budget of the Amazon basin.     

A scaling approach for quantifying the net CO2 flux of the Kuparuk River Basin,Alaska

Net CO₂ flux measurements conducted during the summer and winter of 1994–96 were scaled in space and time to provide estimates of net CO₂ exchange during the 1995–96 (9 May 1995–8 May 1996) annual cycle for the Kuparuk River Basin, a 9200 km² watershed located in NE Alaska. Net CO₂ flux was measured using dynamic chambers and eddy covariance in moist‐acidic, nonacidic, wet‐sedge, and shrub tundra, which comprise 95% of the terrestrial landscape of the Kuparuk Basin. CO₂ flux data were used as input to multivariate models that calculated instantaneous and daily rates of gross primary production (GPP) and whole‐ecosystem respiration (R) as a function of meteorology and ecosystem development. Net CO₂ flux was scaled up to the Kuparuk Basin using a geographical information system (GIS) consisting of a vegetation map, digital terrain map, dynamic temperature and radiation fields, and the models of GPP and R. Basin‐wide estimates of net CO₂ exchange for the summer growing season (9 May?5 September 1995) indicate that nonacidic tundra was a net sink of ?31.7 ± 21.3 GgC (1 Gg = 10⁹ g), while shrub tundra lost 32.5 ± 6.3 GgC to the atmosphere (negative values denote net ecosystem CO₂ uptake). Acidic and wet sedge tundra were in balance, and when integrated for the entire Kuparuk River Basin (including aquatic surfaces), whole basin summer net CO₂ exchange was estimated to be in balance (?0.9 ± 50.3 GgC). Autumn to winter (6 September 1995–8 May 1996) estimates of net CO₂ flux indicate that acidic, nonacidic, and shrub tundra landforms were all large sources of CO₂ to the atmosphere (75.5 ± 8.3, 96.4 ± 11.4, and 43.3 ± 4.7 GgC for acidic, nonacidic, and shrub tundra, respectively). CO₂ loss from wet sedge surfaces was not substantially different from zero, but the large losses from the other terrestrial landforms resulted in a whole basin net CO₂ loss of 217.2 ± 24.1 GgC during the 1995–96 cold season. When integrated for the 1995–96 annual cycle, acidic (66.4 + 25.25 GgC), nonacidic (64.7 ± 29.2 GgC), and shrub tundra (75.8 ± 8.4 GgC) were substantial net sources of CO₂ to the atmosphere, while wet sedge tundra was in balance (0.4 + 0.8 GgC). The Kuparuk River Basin as a whole was estimated to be a net CO₂ source of 218.1 ± 60.6 GgC over the 1995–96 annual cycle. Compared to direct measurements of regional net CO₂ flux obtained from aircraft‐based eddy covariance, the scaling procedure provided realistic estimates of CO₂ exchange during the summer growing season. Although winter estimates could not be assessed directly using aircraft measurements of net CO₂ exchange, the estimates reported here are comparable to measured values reported in the literature. Thus, we have high confidence in the summer estimates of net CO₂ exchange and reasonable confidence in the winter net CO₂ flux estimates for terrestrial landforms of the Kuparuk river basin. Although there is larger uncertainty in the aquatic estimates, the small surface area of aquatic surfaces in the Kuparuk river basin (≈ 5%) presumably reduces the potential for this uncertainty to result in large errors in basin‐wide CO₂ flux estimates.     

Temperature and precipitation drive temporal variability in aquatic carbon and GHG concentrations and fluxes in a peatland catchment

The aquatic pathway is increasingly being recognized as an important component of catchment carbon and greenhouse gas (GHG) budgets, particularly in peatland systems due to their large carbon store and strong hydrological connectivity. In this study, we present a complete 5‐year data set of all aquatic carbon and GHG species from an ombrotrophic Scottish peatland. Measured species include particulate and dissolved forms of organic carbon (POC, DOC), dissolved inorganic carbon (DIC), CO₂, CH₄ and N₂O. We show that short‐term variability in concentrations exists across all species and this is strongly linked to discharge. Seasonal cyclicity was only evident in DOC, CO₂ and CH₄ concentration; however, temperature correlated with monthly means in all species except DIC. Although the temperature correlation with monthly DOC and POC concentrations appeared to be related to biological productivity in the terrestrial system, we suggest the temperature correlation with CO₂ and CH₄ was primarily due to in‐stream temperature‐dependent solubility. Interannual variability in total aquatic carbon concentration was strongly correlated with catchment gross primary productivity (GPP) indicating a strong potential terrestrial aquatic linkage. DOC represented the largest aquatic carbon flux term (19.3 ± 4.59 g C m^?2 yr^?1), followed by CO₂ evasion (10.0 g C m^?2 yr^?1). Despite an estimated contribution to the total aquatic carbon flux of between 8 and 48%, evasion estimates had the greatest uncertainty. Interannual variability in total aquatic carbon export was low in comparison with variability in terrestrial biosphere–atmosphere exchange, and could be explained primarily by temperature and precipitation. Our results therefore suggest that climatic change is likely to have a significant impact on annual carbon losses through the aquatic pathway, and as such, aquatic exports are fundamental to the understanding of whole catchment responses to climate change.     

Observed and modelled historical trends in the water‐use efficiency of plants and ecosystems

Plant water‐use efficiency (WUE, the carbon gained through photosynthesis per unit of water lost through transpiration) is a tracer of the plant physiological controls on the exchange of water and carbon dioxide between terrestrial ecosystems and the atmosphere. At the leaf level, rising CO₂ concentrations tend to increase carbon uptake (in the absence of other limitations) and to reduce stomatal conductance, both effects leading to an increase in leaf WUE. At the ecosystem level, indirect effects (e.g. increased leaf area index, soil water savings) may amplify or dampen the direct effect of CO₂. Thus, the extent to which changes in leaf WUE translate to changes at the ecosystem scale remains unclear. The differences in the magnitude of increase in leaf versus ecosystem WUE as reported by several studies are much larger than would be expected with current understanding of tree physiology and scaling, indicating unresolved issues. Moreover, current vegetation models produce inconsistent and often unrealistic magnitudes and patterns of variability in leaf and ecosystem WUE, calling for a better assessment of the underlying approaches. Here, we review the causes of variations in observed and modelled historical trends in WUE over the continuum of scales from leaf to ecosystem, including methodological issues, with the aim of elucidating the reasons for discrepancies observed within and across spatial scales. We emphasize that even though physiological responses to changing environmental drivers should be interpreted differently depending on the observational scale, there are large uncertainties in each data set which are often underestimated. Assumptions made by the vegetation models about the main processes influencing WUE strongly impact the modelled historical trends. We provide recommendations for improving long‐term observation‐based estimates of WUE that will better inform the representation of WUE in vegetation models.     

Evolutionary trends in RuBisCO kinetics and their co‐evolution with CO2 concentrating mechanisms

RuBisCO‐catalyzed CO₂ fixation is the main source of organic carbon in the biosphere. This enzyme is present in all domains of life in different forms (III, II, and I) and its origin goes back to 3500 Mya, when the atmosphere was anoxygenic. However, the RuBisCO active site also catalyzes oxygenation of ribulose 1,5‐bisphosphate, therefore, the development of oxygenic photosynthesis and the subsequent oxygen‐rich atmosphere promoted the appearance of CO₂ concentrating mechanisms (CCMs) and/or the evolution of a more CO₂‐specific RuBisCO enzyme. The wide variability in RuBisCO kinetic traits of extant organisms reveals a history of adaptation to the prevailing CO₂/O₂ concentrations and the thermal environment throughout evolution. Notable differences in the kinetic parameters are found among the different forms of RuBisCO, but the differences are also associated with the presence and type of CCMs within each form, indicative of co‐evolution of RuBisCO and CCMs. Trade‐offs between RuBisCO kinetic traits vary among the RuBisCO forms and also among phylogenetic groups within the same form. These results suggest that different biochemical and structural constraints have operated on each type of RuBisCO during evolution, probably reflecting different environmental selective pressures. In a similar way, variations in carbon isotopic fractionation of the enzyme point to significant differences in its relationship to the CO₂ specificity among different RuBisCO forms. A deeper knowledge of the natural variability of RuBisCO catalytic traits and the chemical mechanism of RuBisCO carboxylation and oxygenation reactions raises the possibility of finding unrevealed landscapes in RuBisCO evolution.     

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.     

Regional atmospheric CO2 inversion reveals seasonal and geographic differences in Amazon net biome exchange

Understanding tropical rainforest carbon exchange and its response to heat and drought is critical for quantifying the effects of climate change on tropical ecosystems, including global climate–carbon feedbacks. Of particular importance for the global carbon budget is net biome exchange of CO₂ with the atmosphere (NBE), which represents nonfire carbon fluxes into and out of biomass and soils. Subannual and sub‐Basin Amazon NBE estimates have relied heavily on process‐based biosphere models, despite lack of model agreement with plot‐scale observations. We present a new analysis of airborne measurements that reveals monthly, regional‐scale (~1–8 × 10⁶ km²) NBE variations. We develop a regional atmospheric CO₂ inversion that provides the first analysis of geographic and temporal variability in Amazon biosphere–atmosphere carbon exchange and that is minimally influenced by biosphere model‐based first guesses of seasonal and annual mean fluxes. We find little evidence for a clear seasonal cycle in Amazon NBE but do find NBE sensitivity to aberrations from long‐term mean climate. In particular, we observe increased NBE (more carbon emitted to the atmosphere) associated with heat and drought in 2010, and correlations between wet season NBE and precipitation (negative correlation) and temperature (positive correlation). In the eastern Amazon, pulses of increased NBE persisted through 2011, suggesting legacy effects of 2010 heat and drought. We also identify regional differences in postdrought NBE that appear related to long‐term water availability. We examine satellite proxies and find evidence for higher gross primary productivity (GPP) during a pulse of increased carbon uptake in 2011, and lower GPP during a period of increased NBE in the 2010 dry season drought, but links between GPP and NBE changes are not conclusive. These results provide novel evidence of NBE sensitivity to short‐term temperature and moisture extremes in the Amazon, where monthly and sub‐Basin estimates have not been previously available.