The three major axes of terrestrial ecosystem function.

The leaf economics spectrum1,2 and the global spectrum of plant forms and functions3 revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species2. Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities4. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability4,5. Here we derive a set of ecosystem functions6 from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems7,8.

Leaf nutrient basis for the differentiation of photosynthetic traits between subtropical evergreen and deciduous trees.

Compared to evergreens, deciduous tree species usually have higher photosynthetic efficiency to complete vegetative and reproductive growth in a shorter growing season. However, the nutrient basis for the differentiation of photosynthesis functional traits between evergreen and deciduous tree species has not yet been clarified. Thirty evergreen and twenty deciduous angiosperm tree species from a subtropical common garden were compared in terms of photosynthetic traits and leaf nutrients. Generally, their differences in area-based photosynthetic capacity were uncorrelated with area-based leaf nutrient content but were caused by the fraction of nitrogen allocated to photosynthetic components. By comparison, the differences in mass-based photosynthetic capacity were more correlated with leaf nitrogen content than leaf phosphorus and potassium content. Convergence in phosphorus and potassium constraints to photosynthesis occurred in deciduous tree species but not in evergreen tree species. Furthermore, leaf C/N ratio played a more significant role than leaf mass per area in determining the differentiation of photosynthetic traits between evergreen and deciduous groups. Our findings provide insight into the nutrient basis for photosynthetic carbon gain and functional strategies across trees species.

Variable influence of photosynthetic thermal acclimation on future carbon uptake in Australian wooded ecosystems under climate change.

Climate change will impact gross primary productivity (GPP), net primary productivity (NPP), and carbon storage in wooded ecosystems. The extent of change will be influenced by thermal acclimation of photosynthesis-the ability of plants to adjust net photosynthetic rates in response to growth temperatures-yet regional differences in acclimation effects among wooded ecosystems is currently unknown. We examined the effects of changing climate on 17 Australian wooded ecosystems with and without the effects of thermal acclimation of C3 photosynthesis. Ecosystems were drawn from five ecoregions (tropical savanna, tropical forest, Mediterranean woodlands, temperate woodlands, and temperate forests) that span Australia's climatic range. We used the CABLE-POP land surface model adapted with thermal acclimation functions and forced with HadGEM2-ES climate projections from RCP8.5. For each site and ecoregion we examined (a) effects of climate change on GPP, NPP, and live tree carbon storage; and (b) impacts of thermal acclimation of photosynthesis on simulated changes. Between the end of the historical (1976-2005) and projected (2070-2099) periods simulated annual carbon uptake increased in the majority of ecosystems by 26.1%-63.3% for GPP and 15%-61.5% for NPP. Thermal acclimation of photosynthesis further increased GPP and NPP in tropical savannas by 27.2% and 22.4% and by 11% and 10.1% in tropical forests with positive effects concentrated in the wet season (tropical savannas) and the warmer months (tropical forests). We predicted minimal effects of thermal acclimation of photosynthesis on GPP, NPP, and carbon storage in Mediterranean woodlands, temperate woodlands, and temperate forests. Overall, positive effects were strongly enhanced by increasing CO2 concentrations under RCP8.5. We conclude that the direct effects of climate change will enhance carbon uptake and storage in Australian wooded ecosystems (likely due to CO2 enrichment) and that benefits of thermal acclimation of photosynthesis will be restricted to tropical ecoregions.

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.

Bridge to the future: Important lessons from 20 years of ecosystem observations made by the OzFlux network.

In 2020, the Australian and New Zealand flux research and monitoring network, OzFlux, celebrated its 20th anniversary by reflecting on the lessons learned through two decades of ecosystem studies on global change biology. OzFlux is a network not only for ecosystem researchers, but also for those 'next users' of the knowledge, information and data that such networks provide. Here, we focus on eight lessons across topics of climate change and variability, disturbance and resilience, drought and heat stress and synergies with remote sensing and modelling. In distilling the key lessons learned, we also identify where further research is needed to fill knowledge gaps and improve the utility and relevance of the outputs from OzFlux. Extreme climate variability across Australia and New Zealand (droughts and flooding rains) provides a natural laboratory for a global understanding of ecosystems in this time of accelerating climate change. As evidence of worsening global fire risk emerges, the natural ability of these ecosystems to recover from disturbances, such as fire and cyclones, provides lessons on adaptation and resilience to disturbance. Drought and heatwaves are common occurrences across large parts of the region and can tip an ecosystem's carbon budget from a net CO2 sink to a net CO2 source. Despite such responses to stress, ecosystems at OzFlux sites show their resilience to climate variability by rapidly pivoting back to a strong carbon sink upon the return of favourable conditions. Located in under-represented areas, OzFlux data have the potential for reducing uncertainties in global remote sensing products, and these data provide several opportunities to develop new theories and improve our ecosystem models. The accumulated impacts of these lessons over the last 20 years highlights the value of long-term flux observations for natural and managed systems. A future vision for OzFlux includes ongoing and newly developed synergies with ecophysiologists, ecologists, geologists, remote sensors and modellers.

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.

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.

Thermal optima of gross primary productivity are closely aligned with mean air temperatures across Australian wooded ecosystems.

Gross primary productivity (GPP) of wooded ecosystems (forests and savannas) is central to the global carbon cycle, comprising 67%-75% of total global terrestrial GPP. Climate change may alter this flux by increasing the frequency of temperatures beyond the thermal optimum of GPP (Topt ). We examined the relationship between GPP and air temperature (Ta) in 17 wooded ecosystems dominated by a single plant functional type (broadleaf evergreen trees) occurring over a broad climatic gradient encompassing five ecoregions across Australia ranging from tropical in the north to Mediterranean and temperate in the south. We applied a novel boundary-line analysis to eddy covariance flux observations to (a) derive ecosystem GPP-Ta relationships and Topt (including seasonal analyses for five tropical savannas); (b) quantitatively and qualitatively assess GPP-Ta relationships within and among ecoregions; (c) examine the relationship between Topt and mean daytime air temperature (MDTa) across all ecosystems; and (d) examine how down-welling short-wave radiation (Fsd) and vapour pressure deficit (VPD) influence the GPP-Ta relationship. GPP-Ta relationships were convex parabolas with narrow curves in tropical forests, tropical savannas (wet season), and temperate forests, and wider curves in temperate woodlands, Mediterranean woodlands, and tropical savannas (dry season). Ecosystem Topt ranged from 15℃ (temperate forest) to 32℃ (tropical savanna-wet and dry seasons). The shape of GPP-Ta curves was largely determined by daytime Ta range, MDTa, and maximum GPP with the upslope influenced by Fsd and the downslope influenced by VPD. Across all ecosystems, there was a strong positive linear relationship between Topt and MDTa (Adjusted R2 : 0.81; Slope: 1.08) with Topt exceeding MDTa by >1℃ at all but two sites. We conclude that ecosystem GPP has adjusted to local MDTa within Australian broadleaf evergreen forests and that GPP is buffered against small Ta increases in the majority of these ecosystems.

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.

Towards physiologically meaningful water-use efficiency estimates from eddy covariance data.

Intrinsic water-use efficiency (iWUE) characterizes the physiological control on the simultaneous exchange of water and carbon dioxide in terrestrial ecosystems. Knowledge of iWUE is commonly gained from leaf-level gas exchange measurements, which are inevitably restricted in their spatial and temporal coverage. Flux measurements based on the eddy covariance (EC) technique can overcome these limitations, as they provide continuous and long-term records of carbon and water fluxes at the ecosystem scale. However, vegetation gas exchange parameters derived from EC data are subject to scale-dependent and method-specific uncertainties that compromise their ecophysiological interpretation as well as their comparability among ecosystems and across spatial scales. Here, we use estimates of canopy conductance and gross primary productivity (GPP) derived from EC data to calculate a measure of iWUE (G1 , "stomatal slope") at the ecosystem level at six sites comprising tropical, Mediterranean, temperate, and boreal forests. We assess the following six mechanisms potentially causing discrepancies between leaf and ecosystem-level estimates of G1 : (i) non-transpirational water fluxes; (ii) aerodynamic conductance; (iii) meteorological deviations between measurement height and canopy surface; (iv) energy balance non-closure; (v) uncertainties in net ecosystem exchange partitioning; and (vi) physiological within-canopy gradients. Our results demonstrate that an unclosed energy balance caused the largest uncertainties, in particular if it was associated with erroneous latent heat flux estimates. The effect of aerodynamic conductance on G1 was sufficiently captured with a simple representation. G1 was found to be less sensitive to meteorological deviations between canopy surface and measurement height and, given that data are appropriately filtered, to non-transpirational water fluxes. Uncertainties in the derived GPP and physiological within-canopy gradients and their implications for parameter estimates at leaf and ecosystem level are discussed. Our results highlight the importance of adequately considering the sources of uncertainty outlined here when EC-derived water-use efficiency is interpreted in an ecophysiological context.

The response of ecosystem water-use efficiency to rising atmospheric CO2 concentrations: sensitivity and large-scale biogeochemical implications.

Ecosystem water-use efficiency (WUE) is an important metric linking the global land carbon and water cycles. Eddy covariance-based estimates of WUE in temperate/boreal forests have recently been found to show a strong and unexpected increase over the 1992-2010 period, which has been attributed to the effects of rising atmospheric CO2 concentrations on plant physiology. To test this hypothesis, we forced the observed trend in the process-based land surface model JSBACH by increasing the sensitivity of stomatal conductance (gs ) to atmospheric CO2 concentration. We compared the simulated continental discharge, evapotranspiration (ET), and the seasonal CO2 exchange with observations across the extratropical northern hemisphere. The increased simulated WUE led to substantial changes in surface hydrology at the continental scale, including a significant decrease in ET and a significant increase in continental runoff, both of which are inconsistent with large-scale observations. The simulated seasonal amplitude of atmospheric CO2 decreased over time, in contrast to the observed upward trend across ground-based measurement sites. Our results provide strong indications that the recent, large-scale WUE trend is considerably smaller than that estimated for these forest ecosystems. They emphasize the decreasing CO2 sensitivity of WUE with increasing scale, which affects the physiological interpretation of changes in ecosystem WUE.

How do leaf and ecosystem measures of water-use efficiency compare?

The terrestrial carbon and water cycles are intimately linked: the carbon cycle is driven by photosynthesis, while the water balance is dominated by transpiration, and both fluxes are controlled by plant stomatal conductance. The ratio between these fluxes, the plant water-use efficiency (WUE), is a useful indicator of vegetation function. WUE can be estimated using several techniques, including leaf gas exchange, stable isotope discrimination, and eddy covariance. Here we compare global compilations of data for each of these three techniques. We show that patterns of variation in WUE across plant functional types (PFTs) are not consistent among the three datasets. Key discrepancies include the following: leaf-scale data indicate differences between needleleaf and broadleaf forests, but ecosystem-scale data do not; leaf-scale data indicate differences between C3 and C4 species, whereas at ecosystem scale there is a difference between C3 and C4 crops but not grasslands; and isotope-based estimates of WUE are higher than estimates based on gas exchange for most PFTs. Our study quantifies the uncertainty associated with different methods of measuring WUE, indicates potential for bias when using WUE measures to parameterize or validate models, and indicates key research directions needed to reconcile alternative measures of WUE.

Large potential of strengthening the land carbon sink in China through anthropogenic interventions.

The terrestrial ecosystem in China mitigates 21%-45% of the national contemporary fossil fuel CO2 emissions every year. Maintaining and strengthening the land carbon sink is essential for reaching China's target of carbon neutrality. However, this sink is subject to large uncertainties due to the joint impacts of climate change, air pollution, and human activities. Here, we explore the potential of strengthening land carbon sink in China through anthropogenic interventions, including forestation, ozone reduction, and litter removal, taking advantage of a well-validated dynamic vegetation model and meteorological forcings from 16 climate models. Without anthropogenic interventions, considering Shared Socioeconomic Pathways (SSP) scenarios, the land sink is projected to be 0.26-0.56 Pg C a-1 at 2060, to which climate change contributes 0.06-0.13 Pg C a-1 and CO2 fertilization contributes 0.08-0.44 Pg C a-1 with the stronger effects for higher emission scenarios. With anthropogenic interventions, under a close-to-neutral emission scenario (SSP1-2.6), the land sink becomes 0.47-0.57 Pg C a-1 at 2060, including the contributions of 0.12 Pg C a-1 by conservative forestation, 0.07 Pg C a-1 by ozone pollution control, and 0.06-0.16 Pg C a-1 by 20% litter removal over planted forest. This sink can mitigate 90%-110% of the residue anthropogenic carbon emissions in 2060, providing a solid foundation for the carbon neutrality in China.

Carbon-phosphorus cycle models overestimate CO2 enrichment response in a mature Eucalyptus forest.

The importance of phosphorus (P) in regulating ecosystem responses to climate change has fostered P-cycle implementation in land surface models, but their CO2 effects predictions have not been evaluated against measurements. Here, we perform a data-driven model evaluation where simulations of eight widely used P-enabled models were confronted with observations from a long-term free-air CO2 enrichment experiment in a mature, P-limited Eucalyptus forest. We show that most models predicted the correct sign and magnitude of the CO2 effect on ecosystem carbon (C) sequestration, but they generally overestimated the effects on plant C uptake and growth. We identify leaf-to-canopy scaling of photosynthesis, plant tissue stoichiometry, plant belowground C allocation, and the subsequent consequences for plant-microbial interaction as key areas in which models of ecosystem C-P interaction can be improved. Together, this data-model intercomparison reveals data-driven insights into the performance and functionality of P-enabled models and adds to the existing evidence that the global CO2-driven carbon sink is overestimated by models.

Higher global gross primary productivity under future climate with more advanced representations of photosynthesis.

Gross primary productivity (GPP) is the key determinant of land carbon uptake, but its representation in terrestrial biosphere models (TBMs) does not reflect our latest physiological understanding. We implemented three empirically well supported but often omitted mechanisms into the TBM CABLE-POP: photosynthetic temperature acclimation, explicit mesophyll conductance, and photosynthetic optimization through redistribution of leaf nitrogen. We used the RCP8.5 climate scenario to conduct factorial model simulations characterizing the individual and combined effects of the three mechanisms on projections of GPP. Simulated global GPP increased more strongly (up to 20% by 2070-2099) in more comprehensive representations of photosynthesis compared to the model lacking the three mechanisms. The experiments revealed non-additive interactions among the mechanisms as combined effects were stronger than the sum of the individual effects. The modeled responses are explained by changes in the photosynthetic sensitivity to temperature and CO2 caused by the added mechanisms. Our results suggest that current TBMs underestimate GPP responses to future CO2 and climate conditions.

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.

Multi-decadal increase of forest burned area in Australia is linked to climate change.

Fire activity in Australia is strongly affected by high inter-annual climate variability and extremes. Through changes in the climate, anthropogenic climate change has the potential to alter fire dynamics. Here we compile satellite (19 and 32 years) and ground-based (90 years) burned area datasets, climate and weather observations, and simulated fuel loads for Australian forests. Burned area in Australia's forests shows a linear positive annual trend but an exponential increase during autumn and winter. The mean number of years since the last fire has decreased consecutively in each of the past four decades, while the frequency of forest megafire years (>1 Mha burned) has markedly increased since 2000. The increase in forest burned area is consistent with increasingly more dangerous fire weather conditions, increased risk factors associated with pyroconvection, including fire-generated thunderstorms, and increased ignitions from dry lightning, all associated to varying degrees with anthropogenic climate change.

Bigleaf-An R package for the calculation of physical and physiological ecosystem properties from eddy covariance data.

We present the R package bigleaf (version 0.6.5), an open source toolset for the derivation of meteorological, aerodynamic, and physiological ecosystem properties from eddy covariance (EC) flux observations and concurrent meteorological measurements. A 'big-leaf' framework, in which vegetation is represented as a single, uniform layer, is employed to infer bulk ecosystem characteristics top-down from the measured fluxes. Central to the package is the calculation of a bulk surface/canopy conductance (Gs/Gc) and a bulk aerodynamic conductance (Ga), with the latter including formulations for the turbulent and canopy boundary layer components. The derivation of physical land surface characteristics such as surface roughness parameters, wind profile, aerodynamic and radiometric surface temperature, surface vapor pressure deficit (VPD), potential evapotranspiration (ET), imposed and equilibrium ET, as well as vegetation-atmosphere decoupling coefficients, is described. The package further provides calculation routines for physiological ecosytem properties (stomatal slope parameters, stomatal sensitivity to VPD, bulk intercellular CO2 concentration, canopy photosynthetic capacity), energy balance characteristics (closure, biochemical energy), ancillary meteorological variables (psychrometric constant, saturation vapor pressure, air density, etc.), customary unit interconversions and data filtering. The target variables can be calculated with a different degree of complexity, depending on the amount of available site-specific information. The utilities of the package are demonstrated for three single-level (above-canopy) eddy covariance sites representing a temperate grassland, a temperate needle-leaf forest, and a Mediterranean evergreen broadleaf forest. The routines are further tested for a two-level EC site (tree and grass layer) located in a Mediterranean oak savanna. The limitations and the ecophysiological interpretation of the derived ecosystem properties are discussed and practical guidelines are given. The package provides the basis for a consistent, physically sound, and reproducible characterization of biometeorological conditions and ecosystem physiology, and is applicable to EC sites across vegetation types and climatic conditions with minimal ancillary data requirements.