Soil respiration-driven CO2 pulses dominate Australia's flux variability.

The Australian continent contributes substantially to the year-to-year variability of the global terrestrial carbon dioxide (CO2) sink. However, the scarcity of in situ observations in remote areas prevents the deciphering of processes that force the CO2 flux variability. In this study, by examining atmospheric CO2 measurements from satellites in the period 2009-2018, we find recurrent end-of-dry-season CO2 pulses over the Australian continent. These pulses largely control the year-to-year variability of Australia's CO2 balance. They cause two to three times larger seasonal variations compared with previous top-down inversions and bottom-up estimates. The pulses occur shortly after the onset of rainfall and are driven by enhanced soil respiration preceding photosynthetic uptake in Australia's semiarid regions. The suggested continental-scale relevance of soil-rewetting processes has substantial implications for our understanding and modeling of global climate-carbon cycle feedbacks.

On the use of Earth Observation to support estimates of national greenhouse gas emissions and sinks for the Global stocktake process: lessons learned from ESA-CCI RECCAP2.

The Global Stocktake (GST), implemented by the Paris Agreement, requires rapid developments in the capabilities to quantify annual greenhouse gas (GHG) emissions and removals consistently from the global to the national scale and improvements to national GHG inventories. In particular, new capabilities are needed for accurate attribution of sources and sinks and their trends to natural and anthropogenic processes. On the one hand, this is still a major challenge as national GHG inventories follow globally harmonized methodologies based on the guidelines established by the Intergovernmental Panel on Climate Change, but these can be implemented differently for individual countries. Moreover, in many countries the capability to systematically produce detailed and annually updated GHG inventories is still lacking. On the other hand, spatially-explicit datasets quantifying sources and sinks of carbon dioxide, methane and nitrous oxide emissions from Earth Observations (EO) are still limited by many sources of uncertainty. While national GHG inventories follow diverse methodologies depending on the availability of activity data in the different countries, the proposed comparison with EO-based estimates can help improve our understanding of the comparability of the estimates published by the different countries. Indeed, EO networks and satellite platforms have seen a massive expansion in the past decade, now covering a wide range of essential climate variables and offering high potential to improve the quantification of global and regional GHG budgets and advance process understanding. Yet, there is no EO data that quantifies greenhouse gas fluxes directly, rather there are observations of variables or proxies that can be transformed into fluxes using models. Here, we report results and lessons from the ESA-CCI RECCAP2 project, whose goal was to engage with National Inventory Agencies to improve understanding about the methods used by each community to estimate sources and sinks of GHGs and to evaluate the potential for satellite and in-situ EO to improve national GHG estimates. Based on this dialogue and recent studies, we discuss the potential of EO approaches to provide estimates of GHG budgets that can be compared with those of national GHG inventories. We outline a roadmap for implementation of an EO carbon-monitoring program that can contribute to the Paris Agreement.

Emergence of the physiological effects of elevated CO2 on land-atmosphere exchange of carbon and water.

Elevated atmospheric CO2 (eCO2 ) influences the carbon assimilation rate and stomatal conductance of plants, thereby affecting the global cycles of carbon and water. Yet, the detection of these physiological effects of eCO2 in observational data remains challenging, because natural variations and confounding factors (e.g., warming) can overshadow the eCO2 effects in observational data of real-world ecosystems. In this study, we aim at developing a method to detect the emergence of the physiological CO2 effects on various variables related to carbon and water fluxes. We mimic the observational setting in ecosystems using a comprehensive process-based land surface model QUINCY to simulate the leaf-level effects of increasing atmospheric CO2 concentrations and their century-long propagation through the terrestrial carbon and water cycles across different climate regimes and biomes. We then develop a statistical method based on the signal-to-noise ratio to detect the emergence of the eCO2 effects. The eCO2 effect on gross primary productivity (GPP) emerges at relatively low CO2 increase (∆[CO2 ] ~ 20 ppm) where the leaf area index is relatively high. Compared to GPP, the eCO2 effect causing reduced transpiration water flux (normalized to leaf area) emerges only at relatively high CO2 increase (∆[CO2 ] >> 40 ppm), due to the high sensitivity to climate variability and thus lower signal-to-noise ratio. In general, the response to eCO2 is detectable earlier for variables related to the carbon cycle than the water cycle, when plant productivity is not limited by climatic constraints, and stronger in forest-dominated rather than in grass-dominated ecosystems. Our results provide a step toward when and where we expect to detect physiological CO2 effects in in-situ flux measurements, how to detect them and encourage future efforts to improve the understanding and quantification of these effects in observations of terrestrial carbon and water dynamics.

Convergence in phosphorus constraints to photosynthesis in forests around the world.

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements.

Process-oriented analysis of dominant sources of uncertainty in the land carbon sink.

The observed global net land carbon sink is captured by current land models. All models agree that atmospheric CO2 and nitrogen deposition driven gains in carbon stocks are partially offset by climate and land-use and land-cover change (LULCC) losses. However, there is a lack of consensus in the partitioning of the sink between vegetation and soil, where models do not even agree on the direction of change in carbon stocks over the past 60 years. This uncertainty is driven by plant productivity, allocation, and turnover response to atmospheric CO2 (and to a smaller extent to LULCC), and the response of soil to LULCC (and to a lesser extent climate). Overall, differences in turnover explain ~70% of model spread in both vegetation and soil carbon changes. Further analysis of internal plant and soil (individual pools) cycling is needed to reduce uncertainty in the controlling processes behind the global land carbon sink.

Contrasting anatomical and biochemical controls on mesophyll conductance across plant functional types.

Mesophyll conductance (gm ) limits photosynthesis by restricting CO2 diffusion between the substomatal cavities and chloroplasts. Although it is known that gm is determined by both leaf anatomical and biochemical traits, their relative contribution across plant functional types (PFTs) is still unclear. We compiled a dataset of gm measurements and concomitant leaf traits in unstressed plants comprising 563 studies and 617 species from all major PFTs. We investigated to what extent gm limits photosynthesis across PFTs, how gm relates to structural, anatomical, biochemical, and physiological leaf properties, and whether these relationships differ among PFTs. We found that gm imposes a significant limitation to photosynthesis in all C3 PFTs, ranging from 10-30% in most herbaceous annuals to 25-50% in woody evergreens. Anatomical leaf traits explained a significant proportion of the variation in gm (R2 > 0.3) in all PFTs except annual herbs, in which gm is more strongly related to biochemical factors associated with leaf nitrogen and potassium content. Our results underline the need to elucidate mechanisms underlying the global variability of gm . We emphasise the underestimated potential of gm for improving photosynthesis in crops and identify modifications in leaf biochemistry as the most promising pathway for increasing gm in these species.

Mismatch of N release from the permafrost and vegetative uptake opens pathways of increasing nitrous oxide emissions in the high Arctic.

Biogeochemical cycling in permafrost-affected ecosystems remains associated with large uncertainties, which could impact the Earth's greenhouse gas budget and future climate policies. In particular, increased nutrient availability following permafrost thaw could perturb the greenhouse gas exchange in these systems, an effect largely unexplored until now. Here, we enhance the terrestrial ecosystem model QUINCY (QUantifying Interactions between terrestrial Nutrient CYcles and the climate system), which simulates fully coupled carbon (C), nitrogen (N) and phosphorus (P) cycles in vegetation and soil, with processes relevant in high latitudes (e.g., soil freezing and snow dynamics). In combination with site-level and satellite-based observations, we use the model to investigate impacts of increased nutrient availability from permafrost thawing in comparison to other climate-induced effects and CO2 fertilization over 1960 to 2018 across the high Arctic. Our simulations show that enhanced availability of nutrients following permafrost thaw account for less than 15% of the total Gross primary productivity increase over the time period, despite simulated N limitation over the high Arctic scale. As an explanation for this weak fertilization effect, observational and model data indicate a mismatch between the timing of peak vegetative growth (week 26-27 of the year, corresponding to the beginning of July) and peak thaw depth (week 32-35, mid-to-late August), resulting in incomplete plant use of nutrients near the permafrost table. The resulting increasing N availability approaching the permafrost table enhances N loss pathways, which leads to rising nitrous oxide (N2 O) emissions in our model. Site-level emission trends of 2 mg N m-2  year-1 on average over the historical time period could therefore predict an emerging increasing source of N2 O emissions following future permafrost thaw in the high Arctic.

Predicting resilience through the lens of competing adjustments to vegetation function.

There is a pressing need to better understand ecosystem resilience to droughts and heatwaves. Eco-evolutionary optimization approaches have been proposed as means to build this understanding in land surface models and improve their predictive capability, but competing approaches are yet to be tested together. Here, we coupled approaches that optimize canopy gas exchange and leaf nitrogen investment, respectively, extending both approaches to account for hydraulic impairment. We assessed model predictions using observations from a native Eucalyptus woodland that experienced repeated droughts and heatwaves between 2013 and 2020, whilst exposed to an elevated [CO2 ] treatment. Our combined approaches improved predictions of transpiration and enhanced the simulated magnitude of the CO2 fertilization effect on gross primary productivity. The competing approaches also worked consistently along axes of change in soil moisture, leaf area, and [CO2 ]. Despite predictions of a significant percentage loss of hydraulic conductivity due to embolism (PLC) in 2013, 2014, 2016, and 2017 (99th percentile PLC > 45%), simulated hydraulic legacy effects were small and short-lived (2 months). Our analysis suggests that leaf shedding and/or suppressed foliage growth formed a strategy to mitigate drought risk. Accounting for foliage responses to water availability has the potential to improve model predictions of ecosystem resilience.

Long-term ecosystem nitrogen limitation from foliar δ15 N data and a land surface model.

The effect of nutrient availability on plant growth and the terrestrial carbon sink under climate change and elevated CO2 remains one of the main uncertainties of the terrestrial carbon cycle. This is partially due to the difficulty of assessing nutrient limitation at large scales over long periods of time. Consistent declines in leaf nitrogen (N) content and leaf δ15 N have been used to suggest that nitrogen limitation has increased in recent decades, most likely due to the concurrent increase in atmospheric CO2 . However, such data sets are often not straightforward to interpret due to the complex factors that contribute to the spatial and temporal variation in leaf N and isotope concentration. We use the land surface model (LSM) QUINCY, which has the unique capacity to represent N isotopic processes, in conjunction with two large data sets of foliar N and N isotope content. We run the model with different scenarios to test whether foliar δ15 N isotopic data can be used to infer large-scale N limitation and if the observed trends are caused by increasing atmospheric CO2 , changes in climate or changes in sources and magnitude of anthropogenic N deposition. We show that while the model can capture the observed change in leaf N content and predict widespread increases in N limitation, it does not capture the pronounced, but very spatially heterogeneous, decrease in foliar δ15 N observed in the data across the globe. The addition of an observation-based temporal trend in isotopic composition of N deposition leads to a more pronounced decrease in simulated leaf δ15 N. Our results show that leaf δ15 N observations cannot, on their own, be used to assess global-scale N limitation and that using such a data set in conjunction with an LSM can reveal the drivers behind the observed patterns.

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2.

Atmospheric carbon dioxide concentration ([CO2 ]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2 ] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2 ]-driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2 ] (iCO2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

A comprehensive quantification of global nitrous oxide sources and sinks.

Nitrous oxide (N2O), like carbon dioxide, is a long-lived greenhouse gas that accumulates in the atmosphere. Over the past 150 years, increasing atmospheric N2O concentrations have contributed to stratospheric ozone depletion1 and climate change2, with the current rate of increase estimated at 2 per cent per decade. Existing national inventories do not provide a full picture of N2O emissions, owing to their omission of natural sources and limitations in methodology for attributing anthropogenic sources. Here we present a global N2O inventory that incorporates both natural and anthropogenic sources and accounts for the interaction between nitrogen additions and the biochemical processes that control N2O emissions. We use bottom-up (inventory, statistical extrapolation of flux measurements, process-based land and ocean modelling) and top-down (atmospheric inversion) approaches to provide a comprehensive quantification of global N2O sources and sinks resulting from 21 natural and human sectors between 1980 and 2016. Global N2O emissions were 17.0 (minimum-maximum estimates: 12.2-23.5) teragrams of nitrogen per year (bottom-up) and 16.9 (15.9-17.7) teragrams of nitrogen per year (top-down) between 2007 and 2016. Global human-induced emissions, which are dominated by nitrogen additions to croplands, increased by 30% over the past four decades to 7.3 (4.2-11.4) teragrams of nitrogen per year. This increase was mainly responsible for the growth in the atmospheric burden. Our findings point to growing N2O emissions in emerging economies-particularly Brazil, China and India. Analysis of process-based model estimates reveals an emerging N2O-climate feedback resulting from interactions between nitrogen additions and climate change. The recent growth in N2O emissions exceeds some of the highest projected emission scenarios3,4, underscoring the urgency to mitigate N2O emissions.

Low phosphorus supply constrains plant responses to elevated CO2 : A meta-analysis.

Phosphorus (P) is an essential macro-nutrient required for plant metabolism and growth. Low P availability could potentially limit plant responses to elevated carbon dioxide (eCO2 ), but consensus has yet to be reached on the extent of this limitation. Here, based on data from experiments that manipulated both CO2 and P for young individuals of woody and non-woody species, we present a meta-analysis of P limitation impacts on plant growth, physiological, and morphological response to eCO2 . We show that low P availability attenuated plant photosynthetic response to eCO2 by approximately one-quarter, leading to a reduced, but still positive photosynthetic response to eCO2 compared to those under high P availability. Furthermore, low P limited plant aboveground, belowground, and total biomass responses to eCO2 , by 14.7%, 14.3%, and 12.4%, respectively, equivalent to an approximate halving of the eCO2 responses observed under high P availability. In comparison, low P availability did not significantly alter the eCO2 -induced changes in plant tissue nutrient concentration, suggesting tissue nutrient flexibility is an important mechanism allowing biomass response to eCO2 under low P availability. Low P significantly reduced the eCO2 -induced increase in leaf area by 14.3%, mirroring the aboveground biomass response, but low P did not affect the eCO2 -induced increase in root length. Woody plants exhibited stronger attenuation effect of low P on aboveground biomass response to eCO2 than non-woody plants, while plants with different mycorrhizal associations showed similar responses to low P and eCO2 interaction. This meta-analysis highlights crucial data gaps in capturing plant responses to eCO2 and low P availability. Field-based experiments with longer-term exposure of both CO2 and P manipulations are critically needed to provide ecosystem-scale understanding. Taken together, our results provide a quantitative baseline to constrain model-based hypotheses of plant responses to eCO2 under P limitation, thereby improving projections of future global change impacts.

Organizing principles for vegetation dynamics.

Plants and vegetation play a critical-but largely unpredictable-role in global environmental changes due to the multitude of contributing processes at widely different spatial and temporal scales. In this Perspective, we explore approaches to master this complexity and improve our ability to predict vegetation dynamics by explicitly taking account of principles that constrain plant and ecosystem behaviour: natural selection, self-organization and entropy maximization. These ideas are increasingly being used in vegetation models, but we argue that their full potential has yet to be realized. We demonstrate the power of natural selection-based optimality principles to predict photosynthetic and carbon allocation responses to multiple environmental drivers, as well as how individual plasticity leads to the predictable self-organization of forest canopies. We show how models of natural selection acting on a few key traits can generate realistic plant communities and how entropy maximization can identify the most probable outcomes of community dynamics in space- and time-varying environments. Finally, we present a roadmap indicating how these principles could be combined in a new generation of models with stronger theoretical foundations and an improved capacity to predict complex vegetation responses to environmental change.

The fate of carbon in a mature forest under carbon dioxide enrichment.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1-5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3-5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7-10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7-11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests.

Ensemble projections elucidate effects of uncertainty in terrestrial nitrogen limitation on future carbon uptake.

The magnitude of the nitrogen (N) limitation of terrestrial carbon (C) storage over the 21st century is highly uncertain because of the complex interactions between the terrestrial C and N cycles. We use an ensemble approach to quantify and attribute process-level uncertainty in C-cycle projections by analysing a 30-member ensemble representing published alternative representations of key N cycle processes (stoichiometry, biological nitrogen fixation (BNF) and ecosystem N losses) within the framework of one terrestrial biosphere model. Despite large differences in the simulated present-day N cycle, primarily affecting simulated productivity north of 40°N, ensemble members generally conform with global C-cycle benchmarks for present-day conditions. Ensemble projections for two representative concentration pathways (RCP 2.6 and RCP 8.5) show that the increase in land C storage due to CO2 fertilization is reduced by 24 ± 15% due to N constraints, whereas terrestrial C losses associated with climate change are attenuated by 19 ± 20%. As a result, N cycling reduces projected land C uptake for the years 2006-2099 by 19% (37% decrease to 3% increase) for RCP 2.6, and by 21% (40% decrease to 9% increase) for RCP 8.5. Most of the ensemble spread results from uncertainty in temperate and boreal forests, and is dominated by uncertainty in BNF (10% decrease to 50% increase for RCP 2.6, 5% decrease to 100% increase for RCP 8.5). However, choices about the flexibility of ecosystem C:N ratios and processes controlling ecosystem N losses regionally also play important roles. The findings of this study demonstrate clearly the need for an ensemble approach to quantify likely future terrestrial C-N cycle trajectories. Present-day C-cycle observations only weakly constrain the future ensemble spread, highlighting the need for better observational constraints on large-scale N cycling, and N cycle process responses to global change.

Enhanced regional terrestrial carbon uptake over Korea revealed by atmospheric CO2 measurements from 1999 to 2017.

Understanding changes in terrestrial carbon balance is important to improve our knowledge of the regional carbon cycle and climate change. However, evaluating regional changes in the terrestrial carbon balance is challenging due to the lack of surface flux measurements. This study reveals that the terrestrial carbon uptake over the Republic of Korea has been enhanced from 1999 to 2017 by analyzing long-term atmospheric CO2 concentration measurements at the Anmyeondo Station (36.53°N, 126.32°E) located in the western coast. The influence of terrestrial carbon flux on atmospheric CO2 concentrations (ΔCO2 ) is estimated from the difference of CO2 concentrations that were influenced by the land sector (through easterly winds) and the Yellow Sea sector (through westerly winds). We find a significant trend in ΔCO2 of -4.75 ppm per decade (p < .05) during the vegetation growing season (May through October), suggesting that the regional terrestrial carbon uptake has increased relative to the surrounding ocean areas. Combined analysis with satellite measured normalized difference vegetation index and gross primary production shows that the enhanced carbon uptake is associated with significant nationwide increases in vegetation and its production. Process-based terrestrial model and inverse model simulations estimate that regional terrestrial carbon uptake increases by up to 18.9 and 8.0 Tg C for the study period, accounting for 13.4% and 5.7% of the average annual domestic carbon emissions, respectively. Atmospheric chemical transport model simulations indicate that the enhanced terrestrial carbon sink is the primary reason for the observed ΔCO2 trend rather than anthropogenic emissions and atmospheric circulation changes. Our results highlight the fact that atmospheric CO2 measurements could open up the possibility of detecting regional changes in the terrestrial carbon cycle even where anthropogenic emissions are not negligible.

Mesophyll conductance in land surface models: effects on photosynthesis and transpiration.

The CO2 transfer conductance within plant leaves (mesophyll conductance, gm ) is currently not considered explicitly in most land surface models (LSMs), but instead treated implicitly as an intrinsic property of the photosynthetic machinery. Here, we review approaches to overcome this model deficiency by explicitly accounting for gm , which comprises the re-adjustment of photosynthetic parameters and a model describing the variation of gm in dependence of environmental conditions. An explicit representation of gm causes changes in the response of photosynthesis to environmental factors, foremost leaf temperature, and ambient CO2 concentration, which are most pronounced when gm is small. These changes in leaf-level photosynthesis translate into a stronger climate and CO2 response of gross primary productivity (GPP) and transpiration at the global scale. The results from two independent studies show consistent latitudinal patterns of these effects with biggest differences in GPP in the boreal zone (up to ~15%). Transpiration and evapotranspiration show spatially similar, but attenuated, changes compared with GPP. These changes are indirect effects of gm caused by the assumed strong coupling between stomatal conductance and photosynthesis in current LSMs. Key uncertainties in these simulations are the variation of gm with light and the robustness of its temperature response across plant types and growth conditions. Future research activities focusing on the response of gm to environmental factors and its relation to other plant traits have the potential to improve the representation of photosynthesis in LSMs and to better understand its present and future role in the Earth system.

Whole-plant optimality predicts changes in leaf nitrogen under variable CO2 and nutrient availability.

Vegetation nutrient limitation is essential for understanding ecosystem responses to global change. In particular, leaf nitrogen (N) is known to be plastic under changed nutrient limitation. However, models can often not capture these observed changes, leading to erroneous predictions of whole-ecosystem stocks and fluxes. We hypothesise that an optimality approach can improve representation of leaf N content compared to existing empirical approaches. Unlike previous optimality-based approaches, which adjust foliar N concentrations based on canopy carbon export, we use a maximisation criterion based on whole-plant growth, and allow for a lagged response of foliar N to this maximisation criterion to account for the limited plasticity of this plant trait. We test these model variants at a range of Free-Air CO2 Enrichment and N fertilisation experimental sites. We show that a model based solely on canopy carbon export fails to reproduce observed patterns and predicts decreasing leaf N content with increased N availability. However, an optimal model which maximises total plant growth can correctly reproduce the observed patterns. The optimality model we present here is a whole-plant approach which reproduces biologically realistic changes in leaf N and can thereby improve ecosystem-level predictions under transient conditions.

Effects of mesophyll conductance on vegetation responses to elevated CO2 concentrations in a land surface model.

Mesophyll conductance (gm ) is known to affect plant photosynthesis. However, gm is rarely explicitly considered in land surface models (LSMs), with the consequence that its role in ecosystem and large-scale carbon and water fluxes is poorly understood. In particular, the different magnitudes of gm across plant functional types (PFTs) are expected to cause spatially divergent vegetation responses to elevated CO2 concentrations. Here, an extensive literature compilation of gm across major vegetation types is used to parameterize an empirical model of gm in the LSM JSBACH and to adjust photosynthetic parameters based on simulated An  - Ci curves. We demonstrate that an explicit representation of gm changes the response of photosynthesis to environmental factors, which cannot be entirely compensated by adjusting photosynthetic parameters. These altered responses lead to changes in the photosynthetic sensitivity to atmospheric CO2 concentrations which depend both on the magnitude of gm and the climatic conditions, particularly temperature. We then conducted simulations under ambient and elevated (ambient + 200 µmol/mol) CO2 concentrations for contrasting ecosystems and for historical and anticipated future climate conditions (representative concentration pathways; RCPs) globally. The gm -explicit simulations using the RCP8.5 scenario resulted in significantly higher increases in gross primary productivity (GPP) in high latitudes (+10% to + 25%), intermediate increases in temperate regions (+5% to + 15%), and slightly lower to moderately higher responses in tropical regions (-2% to +5%), which summed up to moderate GPP increases globally. Similar patterns were found for transpiration, but with a lower magnitude. Our results suggest that the effect of an explicit representation of gm is most important for simulated carbon and water fluxes in the boreal zone, where a cold climate coincides with evergreen vegetation.

Decadal biomass increment in early secondary succession woody ecosystems is increased by CO2 enrichment.

Increasing atmospheric CO2 stimulates photosynthesis which can increase net primary production (NPP), but at longer timescales may not necessarily increase plant biomass. Here we analyse the four decade-long CO2-enrichment experiments in woody ecosystems that measured total NPP and biomass. CO2 enrichment increased biomass increment by 1.05 ± 0.26 kg C m-2 over a full decade, a 29.1 ± 11.7% stimulation of biomass gain in these early-secondary-succession temperate ecosystems. This response is predictable by combining the CO2 response of NPP (0.16 ± 0.03 kg C m-2 y-1) and the CO2-independent, linear slope between biomass increment and cumulative NPP (0.55 ± 0.17). An ensemble of terrestrial ecosystem models fail to predict both terms correctly. Allocation to wood was a driver of across-site, and across-model, response variability and together with CO2-independence of biomass retention highlights the value of understanding drivers of wood allocation under ambient conditions to correctly interpret and predict CO2 responses.