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Theory predicts that rising CO2 increases global photosynthesis, a process known as CO2 fertilization, and that this is responsible for much of the current terrestrial carbon sink. The estimated magnitude of the historic CO2 fertilization, however, differs by an order of magnitude between long-term proxies, remote sensing-based estimates and terrestrial biosphere models. Here we constrain the likely historic effect of CO2 on global photosynthesis by combining terrestrial biosphere models, ecological optimality theory, remote sensing approaches and an emergent constraint based on global carbon budget estimates. Our analysis suggests that CO2 fertilization increased global annual terrestrial photosynthesis by 13.5 ± 3.5% or 15.9 ± 2.9 PgC (mean ± s.d.) between 1981 and 2020. Our results help resolve conflicting estimates of the historic sensitivity of global terrestrial photosynthesis to CO2 and highlight the large impact anthropogenic emissions have had on ecosystems worldwide.

Terrestrial photosynthesis is the basis for vegetation growth and drives the land carbon cycle. Accurately simulating gross primary production (GPP, ecosystem-level apparent photosynthesis) is key for satellite monitoring and Earth system model predictions under climate change. While robust models exist for describing leaf-level photosynthesis, predictions diverge due to uncertain photosynthetic traits and parameters which vary on multiple spatial and temporal scales. Here, we describe and evaluate a GPP (photosynthesis per unit ground area) model, the P-model, that combines the Farquhar-von Caemmerer-Berry model for C3 photosynthesis with an optimality principle for the carbon assimilation-Transpiration trade-off, and predicts a multi-day average light use efficiency (LUE) for any climate and C3 vegetation type. The model builds on the theory developed in Prentice et al. (2014) and Wang et al. (2017a) and is extended to include low temperature effects on the intrinsic quantum yield and an empirical soil moisture stress factor. The model is forced with site-level data of the fraction of absorbed photosynthetically active radiation (fAPAR) and meteorological data and is evaluated against GPP estimates from a globally distributed network of ecosystem flux measurements. Although the P-model requires relatively few inputs, the R2 is reduced to 0.70 when not accounting for the reduction in quantum yield at low temperatures and effects of low soil moisture on LUE. The R2 for the P-model-predicted LUE is 0.32 (means by site) and 0.48 (means by vegetation type). Applying this model for global-scale simulations yields a total global GPP of 106-122 Pg C yr-1 (mean of 2001-2011), depending on the fAPAR forcing data. The P-model provides a simple but powerful method for predicting-rather than prescribing-light use efficiency and simulating terrestrial photosynthesis across a wide range of conditions. The model is available as an R package (rpmodel).

Elevated CO2 (eCO2) experiments provide critical information to quantify the effects of rising CO2 on vegetation1–6. Many eCO2 experiments suggest that nutrient limitations modulate the local magnitude of the eCO2 effect on plant biomass1,3,5, but the global extent of these limitations has not been empirically quantified, complicating projections of the capacity of plants to take up CO27,8. Here, we present a data-driven global quantification of the eCO2 effect on biomass based on 138 eCO2 experiments. The strength of CO2 fertilization is primarily driven by nitrogen (N) in ~65% of global vegetation and by phosphorus (P) in ~25% of global vegetation, with N- or P-limitation modulated by mycorrhizal association. Our approach suggests that CO2 levels expected by 2100 can potentially enhance plant biomass by 12 ± 3% above current values, equivalent to 59 ± 13 PgC. The global-scale response to eCO2 we derive from experiments is similar to past changes in greenness9 and biomass10 with rising CO2, suggesting that CO2 will continue to stimulate plant biomass in the future despite the constraining effect of soil nutrients. Our research reconciles conflicting evidence on CO2 fertilization across scales and provides an empirical estimate of the biomass sensitivity to eCO2 that may help to constrain climate projections.

Satellite retrievals of information about the Earth’s surface are widely used to monitor global terrestrial photosynthesis and primary production and to examine the ecological impacts of droughts. Methods for estimating photosynthesis from space commonly combine information on vegetation greenness, incoming radiation, temperature and atmospheric demand for water (vapour-pressure deficit), but do not account for the direct effects of low soil moisture. They instead rely on vapour-pressure deficit as a proxy for dryness, despite widespread evidence that soil moisture deficits have a direct impact on vegetation, independent of vapour-pressure deficit. Here, we use a globally distributed measurement network to assess the effect of soil moisture on photosynthesis, and identify a common bias in an ensemble of satellite-based estimates of photosynthesis that is governed by the magnitude of soil moisture effects on photosynthetic light-use efficiency. We develop methods to account for the influence of soil moisture and estimate that soil moisture effects reduce global annual photosynthesis by ~15%, increase interannual variability by more than 100% across 25% of the global vegetated land surface, and amplify the impacts of extreme events on primary production. These results demonstrate the importance of soil moisture effects for monitoring carbon-cycle variability and drought impacts on vegetation productivity from space.

Future environmental change is expected to modify the global hydrological cycle, with consequences for the regional distribution of freshwater supplies. Regional precipitation projections, however, differ largely between models, making future water resource projections highly uncertain. Using two representative concentration pathways and nine climate models, we estimate 21st century water resources across Australia, employing both a process-based dynamic vegetation model and a simple hydrological framework commonly used in water resource studies to separate the effects of climate and vegetation on water resources. We show surprisingly robust, pathway-independent regional patterns of change in water resources despite large uncertainties in precipitation projections. Increasing plant water use efficiency (due to the changing atmospheric CO2) and reduced green vegetation cover (due to the changing climate) relieve pressure on water resources for the highly populated, humid coastal regions of eastern Australia. By contrast, in semi-arid regions across Australia, runoff declines are amplified by CO2-induced greening, which leads to increased vegetation water use. These findings highlight the importance of including vegetation dynamics in future water resource projections.

Global environmental change has implications for the spatial and temporal distribution of water resources, but quantifying its effects remains a challenge. The impact of vegetation responses to increasing atmospheric CO2 concentrations on the hydrologic cycle is particularly poorly constrained. Here we combine remotely sensed normalized difference vegetation index (NDVI) data and long-term water-balance evapotranspiration (ET) measurements from 190 unimpaired river basins across Australia during 1982-2010 to show that the precipitation threshold for water limitation of vegetation cover has significantly declined during the past three decades, whereas sub-humid and semi-arid basins are not only greening but also consuming more water, leading to significant (24-28%) reductions in streamflow. In contrast, wet and arid basins show nonsignificant changes in NDVI and reductions in ET. These observations are consistent with expected effects of elevated CO2 on vegetation. They suggest that projected future decreases in precipitation are likely to be compounded by increased vegetation water use, further reducing streamflow in water-stressed regions.

Aim: The influence of soil properties on photosynthetic traits in higher plants is poorly quantified in comparison with that of climate. We address this situation by quantifying the unique and joint contributions to global leaf-trait variation from soils and climate. Location: Terrestrial ecosystems world-wide. Methods: Using a trait dataset comprising 1509 species from 288 sites, with climate and soil data derived from global datasets, we quantified the effects of 20 soil and 26 climate variables on light-saturated photosynthetic rate (Aarea), stomatal conductance (gs), leaf nitrogen and phosphorus (Narea and Parea) and specific leaf area (SLA) using mixed regression models and multivariate analyses. Results: Soil variables were stronger predictors of leaf traits than climatic variables, except for SLA. On average, Narea, Parea and Aarea increased and SLA decreased with increasing soil pH and with increasing site aridity. gs declined and Parea increased with soil available P (Pavail). Narea was unrelated to total soil N. Joint effects of soil and climate dominated over their unique effects on Narea and Parea, while unique effects of soils dominated for Aarea and gs. Path analysis indicated that variation in Aarea reflected the combined independent influences of Narea and gs, the former promoted by high pH and aridity and the latter by low Pavail. Main conclusions: Three environmental variables were key for explaining variation in leaf traits: soil pH and Pavail, and the climatic moisture index (the ratio of precipitation to potential evapotranspiration). Although the reliability of global soil datasets lags behind that of climate datasets, our results nonetheless provide compelling evidence that both can be jointly used in broad-scale analyses, and that effects uniquely attributable to soil properties are important determinants of leaf photosynthetic traits and rates. A significant future challenge is to better disentangle the covarying physiological, ecological and evolutionary mechanisms that underpin trait-environment relationships.

Efforts to control climate change require the stabilization of atmospheric CO 2 concentrations. This can only be achieved through a drastic reduction of global CO 2 emissions. Yet fossil fuel emissions increased by 29% between 2000 and 2008, in conjunction with increased contributions from emerging economies, from the production and international trade of goods and services, and from the use of coal as a fuel source. In contrast, emissions from land-use changes were nearly constant. Between 1959 and 2008, 43% of each year's CO 2 emissions remained in the atmosphere on average; the rest was absorbed by carbon sinks on land and in the oceans. In the past 50 years, the fraction of CO 2 emissions that remains in the atmosphere each year has likely increased, from about 40% to 45%, and models suggest that this trend was caused by a decrease in the uptake of CO 2 by the carbon sinks in response to climate change and variability. Changes in the CO 2 sinks are highly uncertain, but they could have a significant influence on future atmospheric CO 2 levels. It is therefore crucial to reduce the uncertainties. © 2009 Macmillan Publishers Limited. All rights reserved.

The concentration of CO2 in the atmosphere has risen from close to 280 parts per million (ppm) in 1800, at first slowly and then progressively faster to a value of 367 ppm in 1999, echoing the increasing pace of global agricultural and industrial development. This is known from numerous, well-replicated measurements of the composition of air bubbles trapped in Antarctic ice. Atmospheric CO2 concentrations have been measured directly with high precision since 1957; these measurements agree with ice-core measurements, and show a continuation of the increasing trend up to the present.

Plants acclimate to temperature by adjusting their photosynthetic capacity over weeks to months. However, most evidence for photosynthetic acclimation derives from leaf-scale experiments. Here we address the scarcity of evidence for canopy-scale photosynthetic acclimation by examining the correlation between maximum photosynthetic rates (A max,2,000 ) and growth temperature ( T air ¯ ) across a range of concurrent temperatures and canopy foliage quantity, using data from >200 eddy covariance sites. We detect widespread thermal acclimation of canopy-scale photosynthesis, demonstrated by enhanced A max,2,000 under higher T air ¯ , across flux sites with adequate water availability. A 14-day period is identified as the most relevant timescale for acclimation across all sites, with a range of 12-25 days for different plant functional types. The mean apparent thermal acclimation rate across all ecosystems is 0.41 (-0.38-1.04 for 5th-95th percentile range) µmol m -2  s -1  °C -1 , with croplands showing the largest acclimation rates and grasslands the lowest. Incorporating an optimality-based prediction of leaf photosynthetic capacities into a biochemical photosynthesis model is shown to improve the representation of thermal acclimation. Our results underscore the critical need for enhanced understanding and modelling of canopy-scale photosynthetic capacity to accurately predict plant responses to warmer growing seasons., (© 2024. The Author(s).)

Tropical forests dominate terrestrial photosynthesis, yet there are major contradictions in our understanding due to a lack of field studies, especially outside the tropical Americas. A recent field study indicated that West African forests have among the highest forests gross primary productivity (GPP) yet observed, contradicting models that rank them lower than Amazonian forests. Here, we show possible reasons for this data-model mismatch. We found that biometric GPP measurements are on average 56.3% higher than multiple global GPP products at the study sites. The underestimation of GPP largely disappears when a standard photosynthesis model is informed by local field-measured values of (a) fractional absorbed photosynthetic radiation (fAPAR), and (b) photosynthetic traits. Remote sensing products systematically underestimate fAPAR (33.9% on average at study sites) due to cloud contamination issues. The study highlights the potential widespread underestimation of tropical forests GPP and carbon cycling and hints at the ways forward for model and input data improvement., (© 2024. The Author(s).)

Interactions between carbon (C) and nitrogen (N) cycles in terrestrial ecosystems are simulated in advanced vegetation models, yet methodologies vary widely, leading to divergent simulations of past land C balance trends. This underscores the need to reassess our understanding of ecosystem processes, given recent theoretical advancements and empirical data. We review current knowledge, emphasising evidence from experiments and trait data compilations for vegetation responses to CO 2 and N input, alongside theoretical and ecological principles for modelling. N fertilisation increases leaf N content but inconsistently enhances leaf-level photosynthetic capacity. Whole-plant responses include increased leaf area and biomass, with reduced root allocation and increased aboveground biomass. Elevated atmospheric CO 2 also boosts leaf area and biomass but intensifies belowground allocation, depleting soil N and likely reducing N losses. Global leaf traits data confirm these findings, indicating that soil N availability influences leaf N content more than photosynthetic capacity. A demonstration model based on the functional balance hypothesis accurately predicts responses to N and CO 2 fertilisation on tissue allocation, growth and biomass, offering a path to reduce uncertainty in global C cycle projections., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Plant functional traits (FTs) determine growth, reproduction and survival strategies of plants adapted to their growth environment. Exploring global geographic patterns of FTs, their covariation and their relationships to climate are necessary steps towards better-founded predictions of how global environmental change will affect ecosystem composition. We compile an extensive global dataset for 16 FTs and characterise trait-trait and trait-climate relationships separately within non-woody, woody deciduous and woody evergreen plant groups, using multivariate analysis and generalised additive models (GAMs). Among the six major FTs considered, two dominant trait dimensions-representing plant size and the leaf economics spectrum (LES) respectively-are identified within all three groups. Size traits (plant height, diaspore mass) however are generally higher in warmer climates, while LES traits (leaf mass and nitrogen per area) are higher in drier climates. Larger leaves are associated principally with warmer winters in woody evergreens, but with wetter climates in non-woody plants. GAM-simulated global patterns for all 16 FTs explain up to three-quarters of global trait variation. Global maps obtained by upscaling GAMs are broadly in agreement with iNaturalist citizen-science FT data. This analysis contributes to the foundations for global trait-based ecosystem modelling by demonstrating universal relationships between FTs and climate., (© 2024. The Author(s).)

Stomatal opening in plant leaves is regulated through a balance of carbon and water exchange under different environmental conditions. Accurate estimation of stomatal regulation is crucial for understanding how plants respond to changing environmental conditions, particularly under climate change. A new generation of optimality-based modelling schemes determines instantaneous stomatal responses from a balance of trade-offs between carbon gains and hydraulic costs, but most such schemes do not account for biochemical acclimation in response to drought. Here, we compare the performance of six instantaneous stomatal optimisation models with and without accounting for photosynthetic acclimation. Using experimental data from 37 plant species, we found that accounting for photosynthetic acclimation improves the prediction of carbon assimilation in a majority of the tested models. Photosynthetic acclimation contributed significantly to the reduction of photosynthesis under drought conditions in all tested models. Drought effects on photosynthesis could not accurately be explained by the hydraulic impairment functions embedded in the stomatal models alone, indicating that photosynthetic acclimation must be considered to improve estimates of carbon assimilation during drought., (© 2024 The Authors. Plant, Cell & Environment published by John Wiley & Sons Ltd.)

During extensive periods without rain, known as dry-downs, decreasing soil moisture (SM) induces plant water stress at the point when it limits evapotranspiration, defining a critical SM threshold (θ crit ). Better quantification of θ crit is needed for improving future projections of climate and water resources, food production, and ecosystem vulnerability. Here, we combine systematic satellite observations of the diurnal amplitude of land surface temperature (dLST) and SM during dry-downs, corroborated by in-situ data from flux towers, to generate the observation-based global map of θ crit . We find an average global θ crit of 0.19 m 3 /m 3 , varying from 0.12 m 3 /m 3 in arid ecosystems to 0.26 m 3 /m 3 in humid ecosystems. θ crit simulated by Earth System Models is overestimated in dry areas and underestimated in wet areas. The global observed pattern of θ crit reflects plant adaptation to soil available water and atmospheric demand. Using explainable machine learning, we show that aridity index, leaf area and soil texture are the most influential drivers. Moreover, we show that the annual fraction of days with water stress, when SM stays below θ crit , has increased in the past four decades. Our results have important implications for understanding the inception of water stress in models and identifying SM tipping points., (© 2024. The Author(s).)

Tropical forests cover large areas of equatorial Africa and play a substantial role in the global carbon cycle. However, there has been a lack of biometric measurements to understand the forests' gross and net primary productivity (GPP, NPP) and their allocation. Here we present a detailed field assessment of the carbon budget of multiple forest sites in Africa, by monitoring 14 one-hectare plots along an aridity gradient in Ghana, West Africa. When compared with an equivalent aridity gradient in Amazonia, the studied West African forests generally had higher productivity and lower carbon use efficiency (CUE). The West African aridity gradient consistently shows the highest NPP, CUE, GPP, and autotrophic respiration at a medium-aridity site, Bobiri. Notably, NPP and GPP of the site are the highest yet reported anywhere for intact forests. Widely used data products substantially underestimate productivity when compared to biometric measurements in Amazonia and Africa. Our analysis suggests that the high productivity of the African forests is linked to their large GPP allocation to canopy and semi-deciduous characteristics., (© 2024. The Author(s).)

Leaf dark respiration (R d ) acclimates to environmental changes. However, the magnitude, controls and time scales of acclimation remain unclear and are inconsistently treated in ecosystem models. We hypothesized that R d and Rubisco carboxylation capacity (V cmax ) at 25°C (R d,25 , V cmax,25 ) are coordinated so that R d,25 variations support V cmax,25 at a level allowing full light use, with V cmax,25 reflecting daytime conditions (for photosynthesis), and R d,25 /V cmax,25 reflecting night-time conditions (for starch degradation and sucrose export). We tested this hypothesis temporally using a 5-yr warming experiment, and spatially using an extensive field-measurement data set. We compared the results to three published alternatives: R d,25 declines linearly with daily average prior temperature; R d at average prior night temperatures tends towards a constant value; and R d,25 /V cmax,25 is constant. Our hypothesis accounted for more variation in observed R d,25 over time (R 2  = 0.74) and space (R 2  = 0.68) than the alternatives. Night-time temperature dominated the seasonal time-course of R d , with an apparent response time scale of c. 2 wk. V cmax dominated the spatial patterns. Our acclimation hypothesis results in a smaller increase in global R d in response to rising CO 2 and warming than is projected by the two of three alternative hypotheses, and by current models., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

"Least-cost theory" posits that C 3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO 2 drawdown (lower ratio of leaf internal to ambient CO 2 , C i :C a ) during light-saturated photosynthesis, and at higher leaf N per area (N area ) and higher carboxylation capacity (V cmax 25 ) for a given rate of stomatal conductance to water vapour, g sw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the N area -g sw and V cmax 25 -g sw slopes, and negative effects on C i :C a . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Gross primary production (GPP) by terrestrial ecosystems is a key quantity in the global carbon cycle. The instantaneous controls of leaf-level photosynthesis are well established, but there is still no consensus on the mechanisms by which canopy-level GPP depends on spatial and temporal variation in the environment. The standard model of photosynthesis provides a robust mechanistic representation for C 3 species; however, additional assumptions are required to "scale up" from leaf to canopy. As a consequence, competing models make inconsistent predictions about how GPP will respond to continuing environmental change. This problem is addressed here by means of an empirical analysis of the light use efficiency (LUE) of GPP inferred from eddy covariance carbon dioxide flux measurements, in situ measurements of photosynthetically active radiation (PAR), and remotely sensed estimates of the fraction of PAR (fAPAR) absorbed by the vegetation canopy. Focusing on LUE allows potential drivers of GPP to be separated from its overriding dependence on light. GPP data from over 100 sites, collated over 20 years and located in a range of biomes and climate zones, were extracted from the FLUXNET2015 database and combined with remotely sensed fAPAR data to estimate daily LUE. Daytime air temperature, vapor pressure deficit, diffuse fraction of solar radiation, and soil moisture were shown to be salient predictors of LUE in a generalized linear mixed-effects model. The same model design was fitted to site-based LUE estimates generated by 16 terrestrial ecosystem models. The published models showed wide variation in the shape, the strength, and even the sign of the environmental effects on modeled LUE. These findings highlight important model deficiencies and suggest a need to progress beyond simple "goodness of fit" comparisons of inferred and predicted carbon fluxes toward an approach focused on the functional responses of the underlying dependencies., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

The life span of leaves increases with their mass per unit area (LMA). It is unclear why. Here, we show that this empirical generalization (the foundation of the worldwide leaf economics spectrum) is a consequence of natural selection, maximizing average net carbon gain over the leaf life cycle. Analyzing two large leaf trait datasets, we show that evergreen and deciduous species with diverse construction costs (assumed proportional to LMA) are selected by light, temperature, and growing-season length in different, but predictable, ways. We quantitatively explain the observed divergent latitudinal trends in evergreen and deciduous LMA and show how local distributions of LMA arise by selection under different environmental conditions acting on the species pool. These results illustrate how optimality principles can underpin a new theory for plant geography and terrestrial carbon dynamics.

Recent increases in vegetation greenness over much of the world reflect increasing CO 2 globally and warming in cold areas. However, the strength of the response to both CO 2 and warming in those areas appears to be declining for unclear reasons, contributing to large uncertainties in predicting how vegetation will respond to future global changes. Here, we investigated the changes of satellite-observed peak season absorbed photosynthetically active radiation (F max ) on the Tibetan Plateau between 1982 and 2016. Although climate trends are similar across the Plateau, we identified robust divergent responses (a greening of 0.31 ± 0.14% year -1 in drier regions and a browning of 0.12 ± 0.08% year -1 in wetter regions). Using an eco-evolutionary optimality (EEO) concept of plant acclimation/adaptation, we propose a parsimonious modelling framework that quantitatively explains these changes in terms of water and energy limitations. Our model captured the variations in F max with a correlation coefficient (r) of .76 and a root mean squared error of .12 and predicted the divergent trends of greening (0.32 ± 0.19% year -1 ) and browning (0.07 ± 0.06% year -1 ). We also predicted the observed reduced sensitivities of F max to precipitation and temperature. The model allows us to explain these changes: Enhanced growing season cumulative radiation has opposite effects on water use and energy uptake. Increased precipitation has an overwhelmingly positive effect in drier regions, whereas warming reduces F max in wetter regions by increasing the cost of building and maintaining leaf area. Rising CO 2 stimulates vegetation growth by enhancing water-use efficiency, but its effect on photosynthesis saturates. The large decrease in the sensitivity of vegetation to climate reflects a shift from water to energy limitation. Our study demonstrates the potential of EEO approaches to reveal the mechanisms underlying recent trends in vegetation greenness and provides further insight into the response of alpine ecosystems to ongoing climate change., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Plant functional traits represent adaptive strategies to the environment, linked to biophysical and biogeochemical processes and ecosystem functioning. Compilations of trait data facilitate research in multiple fields from plant ecology through to land-surface modelling. Here we present version 2 of the China Plant Trait Database, which contains information on morphometric, physical, chemical, photosynthetic and hydraulic traits from 1529 unique species in 140 sites spanning a diversity of vegetation types. Version 2 has five improvements compared to the previous version: (1) new data from a 4-km elevation transect on the edge of Tibetan Plateau, including alpine vegetation types not sampled previously; (2) inclusion of traits related to hydraulic processes, including specific sapwood conductance, the area ratio of sapwood to leaf, wood density and turgor loss point; (3) inclusion of information on soil properties to complement the existing data on climate and vegetation (4) assessments and flagging the reliability of individual trait measurements; and (5) inclusion of standardized templates for systematical field sampling and measurements., (© 2022. The Author(s).)

Here we provide the 'Global Spectrum of Plant Form and Function Dataset', containing species mean values for six vascular plant traits. Together, these traits -plant height, stem specific density, leaf area, leaf mass per area, leaf nitrogen content per dry mass, and diaspore (seed or spore) mass - define the primary axes of variation in plant form and function. The dataset is based on ca. 1 million trait records received via the TRY database (representing ca. 2,500 original publications) and additional unpublished data. It provides 92,159 species mean values for the six traits, covering 46,047 species. The data are complemented by higher-level taxonomic classification and six categorical traits (woodiness, growth form, succulence, adaptation to terrestrial or aquatic habitats, nutrition type and leaf type). Data quality management is based on a probabilistic approach combined with comprehensive validation against expert knowledge and external information. Intense data acquisition and thorough quality control produced the largest and, to our knowledge, most accurate compilation of empirically observed vascular plant species mean traits to date., (© 2022. The Author(s).)

Plant water stress occurs at the point when soil moisture (SM) limits transpiration, defining a critical SM threshold (θ crit ). Knowledge of the spatial distribution of θ crit is crucial for future projections of climate and water resources. Here, we use global eddy covariance observations to quantify θ crit and evaporative fraction (EF) regimes. Three canonical variables describe how EF is controlled by SM: the maximum EF (EF max ), θ crit , and slope (S) between EF and SM. We find systematic differences of these three variables across biomes. Variation in θ crit , S, and EF max is mostly explained by soil texture, vapor pressure deficit, and precipitation, respectively, as well as vegetation structure. Dryland ecosystems tend to operate at low θ crit and show adaptation to water deficits. The negative relationship between θ crit and S indicates that dryland ecosystems minimize θ crit through mechanisms of sustained SM extraction and transport by xylem. Our results further suggest an optimal adaptation of local EF-SM response that maximizes growing-season evapotranspiration and photosynthesis.

The global carbon and water cycles are governed by the coupling of CO 2 and water vapour exchanges through the leaves of terrestrial plants, controlled by plant adaptations to balance carbon gains and hydraulic risks. We introduce a trait-based optimality theory that unifies the treatment of stomatal responses and biochemical acclimation of plants to environments changing on multiple timescales. Tested with experimental data from 18 species, our model successfully predicts the simultaneous decline in carbon assimilation rate, stomatal conductance and photosynthetic capacity during progressive soil drought. It also correctly predicts the dependencies of gas exchange on atmospheric vapour pressure deficit, temperature and CO 2 . Model predictions are also consistent with widely observed empirical patterns, such as the distribution of hydraulic strategies. Our unified theory opens new avenues for reliably modelling the interactive effects of drying soil and rising atmospheric CO 2 on global photosynthesis and transpiration., (© 2022. The Author(s).)

Leaf dry mass per unit area (LMA), carboxylation capacity ( V cmax ) and leaf nitrogen per unit area (N area ) and mass (N mass ) are key traits for plant functional ecology and ecosystem modelling. There is however no consensus about how these traits are regulated, or how they should be modelled. Here we confirm that observed leaf nitrogen across species and sites can be estimated well from observed LMA and V cmax at 25°C ( V cmax25 ). We then test the hypothesis that global variations of both quantities depend on climate variables in specific ways that are predicted by leaf-level optimality theory, thus allowing both N area to be predicted as functions of the growth environment.A new global compilation of field measurements was used to quantify the empirical relationships of leaf N to V cmax25 and LMA. Relationships of observed V cmax25 and LMA to climate variables were estimated, and compared to independent theoretical predictions of these relationships. Soil effects were assessed by analysing biases in the theoretical predictions.LMA was the most important predictor of N area (increasing) and N mass (decreasing). About 60% of global variation across species and sites in observed N area , and 31% in N mass , could be explained by observed LMA and V cmax25 . These traits, in turn, were quantitatively related to climate variables, with significant partial relationships similar or indistinguishable from those predicted by optimality theory. Predicted trait values explained 21% of global variation in observed site-mean V cmax25 , 43% in LMA and 31% in N area . Predicted V cmax25 was biased low on clay-rich soils but predicted LMA was biased high, with compensating effects on N area . N area was overpredicted on organic soils. Synthesis . Global patterns of variation in observed site-mean N area can be explained by climate-induced variations in optimal V cmax25 and LMA. Leaf nitrogen should accordingly be modelled as a consequence (not a cause) of V cmax25 and LMA, both being optimized to the environment. Nitrogen limitation of plant growth would then be modelled principally via whole-plant carbon allocation, rather than via leaf-level traits. Further research is required to better understand and model the terrestrial nitrogen and carbon cycles and their coupling., Competing Interests: There is no conflict of interest., (© 2022 The Authors. Journal of Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Nitrogen (N) limitation has been considered as a constraint on terrestrial carbon uptake in response to rising CO 2 and climate change. By extension, it has been suggested that declining carboxylation capacity (V cmax ) and leaf N content in enhanced-CO 2 experiments and satellite records signify increasing N limitation of primary production. We predicted V cmax using the coordination hypothesis and estimated changes in leaf-level photosynthetic N for 1982-2016 assuming proportionality with leaf-level V cmax at 25°C. The whole-canopy photosynthetic N was derived using satellite-based leaf area index (LAI) data and an empirical extinction coefficient for V cmax , and converted to annual N demand using estimated leaf turnover times. The predicted spatial pattern of V cmax shares key features with an independent reconstruction from remotely sensed leaf chlorophyll content. Predicted leaf photosynthetic N declined by 0.27% yr -1 , while observed leaf (total) N declined by 0.2-0.25% yr -1 . Predicted global canopy N (and N demand) declined from 1996 onwards, despite increasing LAI. Leaf-level responses to rising CO 2 , and to a lesser extent temperature, may have reduced the canopy requirement for N by more than rising LAI has increased it. This finding provides an alternative explanation for declining leaf N that does not depend on increasing N limitation., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Aim: Biomisation has been the most widely used technique to reconstruct past regional vegetation patterns because it does not require an extensive modern pollen dataset. However, it has well-known limitations including its dependence on expert judgement for the assignment of pollen taxa to plant functional types (PFTs) and PFTs to biomes. Here we present a new method that combines the strengths of biomisation with those of the alternative dissimilarity-based techniques., Location: The Eastern Mediterranean-Black Sea Caspian Corridor (EMBSeCBIO)., Taxon: Plants., Methods: Modern pollen samples, assigned to biomes based on potential natural vegetation data, are used to characterize the within-biome means and standard deviations of the abundances of each taxon. These values are used to calculate a dissimilarity index between any pollen sample and every biome, and thus assign the sample to the most likely biome. We calculate a threshold value for each modern biome; fossil samples with scores below the threshold for all modern biomes are thus identified as non-analogue vegetation. We applied the new method to the EMBSeCBIO region to compare its performance with existing reconstructions., Results: The method captured changes in the importance of individual taxa along environmental gradients. The balanced accuracy obtained for the EMBSeCBIO region using the new method was better than obtained using biomisation (77% vs. 65%). When the method was applied to high-resolution fossil records, 70% of the entities showed more temporally stable biome assignments than obtained using biomisation. The technique also identified likely non-analogue assemblages in a synthetic modern dataset and in fossil records., Main Conclusions: The new method yields more accurate and stable reconstructions of vegetation than biomisation. It requires an extensive modern pollen dataset, but is conceptually simple, and avoids subjective choices about taxon allocations to PFTs and PFTs to biomes., Competing Interests: The authors declare no conflict of interest., (© 2022 The Authors. Journal of Biogeography published by John Wiley & Sons Ltd.)

Leaf morphological traits vary systematically along climatic gradients. However, recent studies in plant functional ecology have mainly analysed quantitative traits, while numerical models of species distributions and vegetation function have focused on traits associated with resource acquisition; both ignore the wider functional significance of leaf morphology.A dataset comprising 22 leaf morphological traits for 662 woody species from 92 sites, representing all biomes present in China, was subjected to multivariate analysis in order to identify leading dimensions of trait covariation (correspondence analysis), quantify climatic and phylogenetic contributions (canonical correspondence analysis with variation partitioning) and characterise co-occurring trait syndromes ( k -means clustering) and their climatic preferences.Three axes accounted for >20% of trait variation in both evergreen and deciduous species. Moisture index, precipitation seasonality and growing-season temperature explained 8%-10% of trait variation; family 15%-32%. Microphyll or larger, mid- to dark green leaves with drip tips in wetter climates contrasted with nanophyll or smaller glaucous leaves without drip tips in drier climates. Thick, entire leaves in less seasonal climates contrasted with thin, marginal dissected, aromatic and involute/revolute leaves in more seasonal climates. Thick, involute, hairy leaves in colder climates contrasted with thin leaves with marked surface structures (surface patterning) in warmer climates. Distinctive trait clusters were linked to the driest and most seasonal climates, for example the clustering of picophyll, fleshy and succulent leaves in the driest climates and leptophyll, linear, dissected, revolute or involute and aromatic leaves in regions with highly seasonal rainfall. Several trait clusters co-occurred in wetter climates, including clusters characterised by microphyll, moderately thick, patent and entire leaves or notophyll, waxy, dark green leaves. Synthesis . The plastic response of size, shape, colour and other leaf morphological traits to climate is muted, thus their apparent shift along climate gradients reflects plant adaptations to environment at a community level as determined by species replacement. Information on leaf morphological traits, widely available in floras, could be used to strengthen predictive models of species distribution and vegetation function., Competing Interests: The authors declare that there is no conflict of interest., (© 2022 The Authors. Journal of Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

SignificanceThe magnitude of the CO 2 fertilization effect on terrestrial photosynthesis is uncertain because it is not directly observed and is subject to confounding effects of climatic variability. We apply three well-established eco-evolutionary optimality theories of gas exchange and photosynthesis, constraining the main processes of CO 2 fertilization using measurable variables. Using this framework, we provide robust observationally inferred evidence that a strong CO 2 fertilization effect is detectable in globally distributed eddy covariance networks. Applying our method to upscale photosynthesis globally, we find that the magnitude of the CO 2 fertilization effect is comparable to its in situ counterpart but highlight the potential for substantial underestimation of this effect in tropical forests for many reflectance-based satellite photosynthesis products.

Accurate monitoring of vegetation stress is required for better modelling and forecasting of primary production, in a world where heatwaves and droughts are expected to become increasingly prevalent. Variability in formaldehyde (HCHO) concentrations in the troposphere is dominated by local emissions of short-lived biogenic (BVOC) and pyrogenic volatile organic compounds. BVOCs are emitted by plants in a rapid protective response to abiotic stress, mediated by the energetic status of leaves (the excess of reducing power when photosynthetic light and dark reactions are decoupled, as occurs when stomata close in response to water stress). Emissions also increase exponentially with leaf temperature. New analytical methods for the detection of spatiotemporally contiguous extremes in remote-sensing data are applied here to satellite-derived atmospheric HCHO columns. BVOC emissions are shown to play a central role in the formation of the largest positive HCHO anomalies. Although vegetation stress can be captured by various remotely sensed quantities, spaceborne HCHO emerges as the most consistent recorder of vegetation responses to the largest climate extremes, especially in forested regions., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Both low soil water content (SWC) and high atmospheric dryness (vapor pressure deficit, VPD) can negatively affect terrestrial gross primary production (GPP). The sensitivity of GPP to soil versus atmospheric dryness is difficult to disentangle, however, because of their covariation. Using global eddy-covariance observations, here we show that a decrease in SWC is not universally associated with GPP reduction. GPP increases in response to decreasing SWC when SWC is high and decreases only when SWC is below a threshold. By contrast, the sensitivity of GPP to an increase of VPD is always negative across the full SWC range. We further find canopy conductance decreases with increasing VPD (irrespective of SWC), and with decreasing SWC on drier soils. Maximum photosynthetic assimilation rate has negative sensitivity to VPD, and a positive sensitivity to decreasing SWC when SWC is high. Earth System Models underestimate the negative effect of VPD and the positive effect of SWC on GPP such that they should underestimate the GPP reduction due to increasing VPD in future climates., (© 2022. The Author(s).)

Carbon isotope discrimination (Δ 13 C) in C 3 woody plants is a key variable for the study of photosynthesis. Yet how Δ 13 C varies at decadal scales, and across regions, and how it is related to gross primary production (GPP), are still incompletely understood. Here we address these questions by implementing a new Δ 13 C modelling capability in the land-surface model JULES incorporating both photorespiratory and mesophyll-conductance fractionations. We test the ability of four leaf-internal CO 2 concentration models embedded in JULES to reproduce leaf and tree-ring (TR) carbon isotopic data. We show that all the tested models tend to overestimate average Δ 13 C values, and to underestimate interannual variability in Δ 13 C. This is likely because they ignore the effects of soil water stress on stomatal behavior. Variations in post-photosynthetic isotopic fractionations across species, sites and years, may also partly explain the discrepancies between predicted and TR-derived Δ 13 C values. Nonetheless, the "least-cost" (Prentice) model shows the lowest biases with the isotopic measurements, and lead to improved predictions of canopy-level carbon and water fluxes. Overall, modelled Δ 13 C trends vary strongly between regions during the recent (1979-2016) historical period but stay nearly constant when averaged over the globe. Photorespiratory and mesophyll effects modulate the simulated global Δ 13 C trend by 0.0015 ± 0.005‰ and -0.0006 ± 0.001‰ ppm -1 , respectively. These predictions contrast with previous findings based on atmospheric carbon isotope measurements. Predicted Δ 13 C and GPP tend to be negatively correlated in wet-humid and cold regions, and in tropical African forests, but positively related elsewhere. The negative correlation between Δ 13 C and GPP is partly due to the strong dominant influences of temperature on GPP and vapor pressure deficit on Δ 13 C in those forests. Our results demonstrate that the combined analysis of Δ 13 C and GPP can help understand the drivers of photosynthesis changes in different climatic regions., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

The global terrestrial carbon sink is increasing 1-3 , offsetting roughly a third of anthropogenic CO 2 released into the atmosphere each decade 1 , and thus serving to slow 4 the growth of atmospheric CO 2 . It has been suggested that a CO 2 -induced long-term increase in global photosynthesis, a process known as CO 2 fertilization, is responsible for a large proportion of the current terrestrial carbon sink 4-7 . The estimated magnitude of the historic increase in photosynthesis as result of increasing atmospheric CO 2 concentrations, however, differs by an order of magnitude between long-term proxies and terrestrial biosphere models 7-13 . Here we quantify the historic effect of CO 2 on global photosynthesis by identifying an emergent constraint 14-16 that combines terrestrial biosphere models with global carbon budget estimates. Our analysis suggests that CO 2 fertilization increased global annual photosynthesis by 11.85 ± 1.4%, or 13.98 ± 1.63 petagrams carbon (mean ± 95% confidence interval) between 1981 and 2020. Our results help resolve conflicting estimates of the historic sensitivity of global photosynthesis to CO 2 , and highlight the large impact anthropogenic emissions have had on ecosystems worldwide., (© 2021. The Author(s), under exclusive licence to Springer Nature Limited.)

Close coupling between water loss and carbon dioxide uptake requires coordination of plant hydraulics and photosynthesis. However, there is still limited information on the quantitative relationships between hydraulic and photosynthetic traits. We propose a basis for these relationships based on optimality theory, and test its predictions by analysis of measurements on 107 species from 11 sites, distributed along a nearly 3000-m elevation gradient. Hydraulic and leaf economic traits were less plastic, and more closely associated with phylogeny, than photosynthetic traits. The two sets of traits were linked by the sapwood to leaf area ratio (Huber value, v H ). The observed coordination between v H and sapwood hydraulic conductivity (K S ) and photosynthetic capacity (V cmax ) conformed to the proposed quantitative theory. Substantial hydraulic diversity was related to the trade-off between K S and v H . Leaf drought tolerance (inferred from turgor loss point, -Ψ tlp ) increased with wood density, but the trade-off between hydraulic efficiency (K S ) and -Ψ tlp was weak. Plant trait effects on v H were dominated by variation in K S , while effects of environment were dominated by variation in temperature. This research unifies hydraulics, photosynthesis and the leaf economics spectrum in a common theoretical framework, and suggests a route towards the integration of photosynthesis and hydraulics in land-surface models., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

We introduce the AusTraits database - a compilation of values of plant traits for taxa in the Australian flora (hereafter AusTraits). AusTraits synthesises data on 448 traits across 28,640 taxa from field campaigns, published literature, taxonomic monographs, and individual taxon descriptions. Traits vary in scope from physiological measures of performance (e.g. photosynthetic gas exchange, water-use efficiency) to morphological attributes (e.g. leaf area, seed mass, plant height) which link to aspects of ecological variation. AusTraits contains curated and harmonised individual- and species-level measurements coupled to, where available, contextual information on site properties and experimental conditions. This article provides information on version 3.0.2 of AusTraits which contains data for 997,808 trait-by-taxon combinations. We envision AusTraits as an ongoing collaborative initiative for easily archiving and sharing trait data, which also provides a template for other national or regional initiatives globally to fill persistent gaps in trait knowledge., (© 2021. The Author(s).)

Global vegetation and land-surface models embody interdisciplinary scientific understanding of the behaviour of plants and ecosystems, and are indispensable to project the impacts of environmental change on vegetation and the interactions between vegetation and climate. However, systematic errors and persistently large differences among carbon and water cycle projections by different models highlight the limitations of current process formulations. In this review, focusing on core plant functions in the terrestrial carbon and water cycles, we show how unifying hypotheses derived from eco-evolutionary optimality (EEO) principles can provide novel, parameter-sparse representations of plant and vegetation processes. We present case studies that demonstrate how EEO generates parsimonious representations of core, leaf-level processes that are individually testable and supported by evidence. EEO approaches to photosynthesis and primary production, dark respiration and stomatal behaviour are ripe for implementation in global models. EEO approaches to other important traits, including the leaf economics spectrum and applications of EEO at the community level are active research areas. Independently tested modules emerging from EEO studies could profitably be integrated into modelling frameworks that account for the multiple time scales on which plants and plant communities adjust to environmental change., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

Leaf mass per area (Ma), nitrogen content per unit leaf area (Narea), maximum carboxylation capacity (Vcmax) and the ratio of leaf-internal to ambient CO2 partial pressure (χ) are important traits related to photosynthetic function, and they show systematic variation along climatic and elevational gradients. Separating the effects of air pressure and climate along elevational gradients is challenging due to the covariation of elevation, pressure and climate. However, recently developed models based on optimality theory offer an independent way to predict leaf traits and thus to separate the contributions of different controls. We apply optimality theory to predict variation in leaf traits across 18 sites in the Gongga Mountain region. We show that the models explain 59% of trait variability on average, without site- or region-specific calibration. Temperature, photosynthetically active radiation, vapor pressure deficit, soil moisture and growing season length are all necessary to explain the observed patterns. The direct effect of air pressure is shown to have a relatively minor impact. These findings contribute to a growing body of research indicating that leaf-level traits vary with the physical environment in predictable ways, suggesting a promising direction for the improvement of terrestrial ecosystem models., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Atmospheric aridity and drought both influence physiological function in plant leaves, but their relative contributions to changes in the ratio of leaf internal to ambient partial pressure of CO 2 (χ) - an index of adjustments in both stomatal conductance and photosynthetic rate to environmental conditions - are difficult to disentangle. Many stomatal models predicting χ include the influence of only one of these drivers. In particular, the least-cost optimality hypothesis considers the effect of atmospheric demand for water on χ but does not predict how soils with reduced water further influence χ, potentially leading to an overestimation of χ under dry conditions. Here, we use a large network of stable carbon isotope measurements in C 3 woody plants to examine the acclimated response of χ to soil water stress. We estimate the ratio of cost factors for carboxylation and transpiration (β) expected from the theory to explain the variance in the data, and investigate the responses of β (and thus χ) to soil water content and suction across seed plant groups, leaf phenological types and regions. Overall, β decreases linearly with soil drying, implying that the cost of water transport along the soil-plant-atmosphere continuum increases as water available in the soil decreases. However, despite contrasting hydraulic strategies, the stomatal responses of angiosperms and gymnosperms to soil water tend to converge, consistent with the optimality theory. The prediction of β as a simple, empirical function of soil water significantly improves χ predictions by up to 6.3 ± 2.3% (mean ± SD of adjusted-R 2 ) over 1980-2018 and results in a reduction of around 2% of mean χ values across the globe. Our results highlight the importance of soil water status on stomatal functions and plant water-use efficiency, and suggest the implementation of trait-based hydraulic functions into the model to account for soil water stress., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Quantitative reconstructions of past climates are an important resource for evaluating how well climate models reproduce climate changes. One widely used statistical approach for making such reconstructions from fossil biotic assemblages is weighted averaging partial least-squares regression (WA-PLS). There is however a known tendency for WA-PLS to yield reconstructions compressed towards the centre of the climate range used for calibration, potentially biasing the reconstructed past climates. We present an improvement of WA-PLS by assuming that: (i) the theoretical abundance of each taxon is unimodal with respect to the climate variable considered; (ii) observed taxon abundances follow a multinomial distribution in which the total abundance of a sample is climatically uninformative; and (iii) the estimate of the climate value at a given site and time makes the observation most probable, i.e. it maximizes the log-likelihood function. This climate estimate is approximated by weighting taxon abundances in WA-PLS by the inverse square of their climate tolerances. We further improve the approach by considering the frequency (  fx ) of the climate variable in the training dataset. Tolerance-weighted WA-PLS with fx correction greatly reduces the compression bias, compared with WA-PLS, and improves model performance in reconstructions based on an extensive modern pollen dataset., Competing Interests: We declare we have no competing interests., (© 2020 The Author(s).)

Forest production efficiency (FPE) metric describes how efficiently the assimilated carbon is partitioned into plants organs (biomass production, BP) or-more generally-for the production of organic matter (net primary production, NPP). We present a global analysis of the relationship of FPE to stand-age and climate, based on a large compilation of data on gross primary production and either BP or NPP. FPE is important for both forest production and atmospheric carbon dioxide uptake. We find that FPE increases with absolute latitude, precipitation and (all else equal) with temperature. Earlier findings-FPE declining with age-are also supported by this analysis. However, the temperature effect is opposite to what would be expected based on the short-term physiological response of respiration rates to temperature, implying a top-down regulation of carbon loss, perhaps reflecting the higher carbon costs of nutrient acquisition in colder climates. Current ecosystem models do not reproduce this phenomenon. They consistently predict lower FPE in warmer climates, and are therefore likely to overestimate carbon losses in a warming climate.

Leaf area (LA), mass per area (LMA), nitrogen per unit area (N area ) and the leaf-internal to ambient CO 2 ratio (χ) are fundamental traits for plant functional ecology and vegetation modelling. Here we aimed to assess how their variation, within and between species, tracks environmental gradients. Measurements were made on 705 species from 116 sites within a broad north-south transect from tropical to temperate Australia. Trait responses to environment were quantified using multiple regression; within- and between-species responses were compared using analysis of covariance and trait-gradient analysis. Leaf area, the leaf economics spectrum (indexed by LMA and N area ) and χ (from stable carbon isotope ratios) varied almost independently among species. Across sites, however, χ and LA increased with mean growing-season temperature (mGDD 0 ) and decreased with vapour pressure deficit (mVPD 0 ) and soil pH. LMA and N area showed the reverse pattern. Climate responses agreed with expectations based on optimality principles. Within-species variability contributed < 10% to geographical variation in LA but > 90% for χ, with LMA and N area intermediate. These findings support the hypothesis that acclimation within individuals, adaptation within species and selection among species combine to create predictable relationships between traits and environment. However, the contribution of acclimation/adaptation vs species selection differs among traits., (© 2020 The Authors. New Phytologist © 2020 New Phytologist Trust.)

Photosynthetic 'least-cost' theory posits that the optimal trait combination for a given environment is that where the summed costs of photosynthetic water and nutrient acquisition/use are minimised. The effects of soil water and nutrient availability on photosynthesis should be stronger as climate-related costs for both resources increase. Two independent datasets of photosynthetic traits, Globamax (1509 species, 288 sites) and Glob13C (3645 species, 594 sites), were used to quantify biophysical and biochemical limitations of photosynthesis and the key variable C i /C a (CO 2 drawdown during photosynthesis). Climate and soil variables were associated with both datasets. The biochemical photosynthetic capacity was higher on alkaline soils. This effect was strongest at more arid sites, where water unit-costs are presumably higher. Higher values of soil silt and depth increased C i /C a , likely by providing greater H 2 O supply, alleviating biophysical photosynthetic limitation when soil water is scarce. Climate is important in controlling the optimal balance of H 2 O and N costs for photosynthesis, but soil properties change these costs, both directly and indirectly. In total, soil properties modify the climate-demand driven predictions of C i /C a by up to 30% at a global scale., (© 2020 The Authors New Phytologist © 2020 New Phytologist Trust.)

Pollen data are widely used to reconstruct past climate changes, using relationships between modern pollen abundance in surface samples and climate at the surface-sample sites as a calibration. Visualization of modern pollen data in multidimensional climate space provides a way to establish that taxon abundances are well behaved before using them in climate reconstructions. Visualization is also helpful for ecological interpretation of variations in pollen abundance in space and time. Here, we present Generalized Additive Models for the distribution of 195 European pollen and pteridophyte spore taxa in a bioclimate space defined by seasonal temperatures (as mean temperature of the coldest month and annual growing degree days) and an annual moisture index. These models can be used to explore the realized climate niche of pollen taxa and to build statistical models for palaeoclimate reconstruction. The data set is released under a Creative Commons BY license. When using the data set, we kindly request that you cite this article., (© 2020 by the Ecological Society of America.)

Several publications have examined leaf-trait and carbon-cycling shifts along an Amazon-Andes transect spanning 3.5 km in elevation and 16°C in mean annual temperature. Photosynthetic capacity was previously shown to increase as temperature declines with increasing elevation, counteracting enzyme-kinetic effects. Primary production declines, nonetheless, due to decreasing light availability. We aimed to predict leaf-trait and production gradients from first principles, using published data to test an emerging theory whereby photosynthetic traits and primary production depend on optimal acclimation and/or adaptation to environment. We re-analysed published data for 210 species at 25 sites, fitting linear relationships to elevation for both predicted and observed photosynthetic traits and primary production. Declining leaf-internal/ambient CO 2 ratio (χ) and increasing carboxylation (V cmax ) and electron-transport (J max ) capacities with increasing elevation were predicted. Increases in leaf nitrogen content with elevation were explained by increasing V cmax and leaf mass-per-area. Leaf and soil phosphorus covaried, but after controlling for elevation, no nutrient metric accounted for any additional variance in photosynthetic traits. Primary production was predicted to decline with elevation. This analysis unifies leaf and ecosystem observations in a common theoretical framework. The insensitivity of primary production to temperature is shown to emerge as a consequence of the optimisation of photosynthetic traits., (© 2020 The Authors. New Phytologist © 2020 New Phytologist Trust.)

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.

Plant respiration is an important contributor to the proposed positive global carbon-cycle feedback to climate change. However, as a major component, leaf mitochondrial ('dark') respiration (R d ) differs among species adapted to contrasting environments and is known to acclimate to sustained changes in temperature. No accepted theory explains these phenomena or predicts its magnitude. Here we propose that the acclimation of R d follows an optimal behaviour related to the need to maintain long-term average photosynthetic capacity (V cmax ) so that available environmental resources can be most efficiently used for photosynthesis. To test this hypothesis, we extend photosynthetic co-ordination theory to predict the acclimation of R d to growth temperature via a link to V cmax , and compare predictions to a global set of measurements from 112 sites spanning all terrestrial biomes. This extended co-ordination theory predicts that field-measured R d and V cmax accessed at growth temperature (R d,tg and V cmax,tg ) should increase by 3.7% and 5.5% per degree increase in growth temperature. These acclimated responses to growth temperature are less steep than the corresponding instantaneous responses, which increase 8.1% and 9.9% per degree of measurement temperature for R d and V cmax respectively. Data-fitted responses proof indistinguishable from the values predicted by our theory, and smaller than the instantaneous responses. Theory and data are also shown to agree that the basal rates of both R d and V cmax assessed at 25°C (R d,25 and V cmax,25 ) decline by ~4.4% per degree increase in growth temperature. These results provide a parsimonious general theory for R d acclimation to temperature that is simpler-and potentially more reliable-than the plant functional type-based leaf respiration schemes currently employed in most ecosystem and land-surface models., (© 2019 John Wiley & Sons Ltd.)