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Photosynthetic capacity, determined by light harvesting and carboxylation reactions, is a key plant trait that determines the rate of photosynthesis; however, in Earth System Models (ESMs) at a reference temperature, it is either a fixed value for a given plant functional type or derived from a linear function of leaf nitrogen content. In this study, we conducted a comprehensive analysis that considered correlations of environmental factors with photosynthetic capacity as determined by maximum carboxylation (V(cm)) rate scaled to 25 degrees C (i.e., V(c),25; μmol CO2 x m(-2)x s(-1)) and maximum electron transport rate (J(max)) scaled to 25 degrees C (i.e., J25; μmol electron x m(-2) x s(-1)) at the global scale. Our results showed that the percentage of variation in observed V(c),25 and J25 explained jointly by the environmental factors (i.e., day length, radiation, temperature, and humidity) were 2-2.5 times and 6-9 times of that explained by area-based leaf nitrogen content, respectively. Environmental factors influenced photosynthetic capacity mainly through photosynthetic nitrogen use efficiency, rather than through leaf nitrogen content. The combination of leaf nitrogen content and environmental factors was able to explain -56% and -66% of the variation in V(c),25 and J25 at the global scale, respectively. Our analyses suggest that model projections of plant photosynthetic capacity and hence land-atmosphere exchange under changing climatic conditions could be substantially improved if environmental factors are incorporated into algorithms used to parameterize photosynthetic capacity in ESMs.

The response of mature forest ecosystems to a rising atmospheric carbon dioxide concentration (Ca) is a major uncertainty in projecting the future trajectory of the Earth's climate. Although leaf-level net photosynthesis is typically stimulated by exposure to elevated Ca (eCa), it is unclear how this stimulation translates into carbon cycle responses at the ecosystem scale. Here we estimate a key component of the carbon cycle, the gross primary productivity (GPP), of a mature native eucalypt forest exposed to free-air CO2 enrichment (the EucFACE experiment). In this experiment, light-saturated leaf photosynthesis increased by 19 % in response to a 38 % increase in Ca. We used the process-based forest canopy model, MAESPA, to upscale these leaf-level measurements of photosynthesis with canopy structure to estimate the GPP and its response to eCa. We assessed the direct impact of eCa, as well as the indirect effect of photosynthetic acclimation to eCa and variability among treatment plots using different model scenarios. At the canopy scale, MAESPA estimated a GPP of 1574 g C m−2 yr−1 under ambient conditions across 4 years and a direct increase in the GPP of +11 % in response to eCa. The smaller canopy-scale response simulated by the model, as compared with the leaf-level response, could be attributed to the prevalence of RuBP regeneration limitation of leaf photosynthesis within the canopy. Photosynthetic acclimation reduced this estimated response to 10 %. After taking the baseline variability in the leaf area index across plots in account, we estimated a field GPP response to eCa of 6 % with a 95 % confidence interval (−2 %, 14 %). These findings highlight that the GPP response of mature forests to eCa is likely to be considerably lower than the response of light-saturated leaf photosynthesis. Our results provide an important context for interpreting the eCa responses of other components of the ecosystem carbon cycle.

Atmospheric carbon dioxide enrichment (eCO.sub.2) can enhance plant carbon uptake and growth.sup.1-5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO.sub.2 concentration.sup.6. Although evidence gathered from young aggrading forests has generally indicated a strong CO.sub.2 fertilization effect on biomass growth.sup.3-5, it is unclear whether mature forests respond to eCO.sub.2 in a similar way. In mature trees and forest stands.sup.7-10, photosynthetic uptake has been found to increase under eCO.sub.2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO.sub.2 unclear.sup.4,5,7-11. Here using data from the first ecosystem-scale Free-Air CO.sub.2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO.sub.2 exposure. We show that, although the eCO.sub.2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO.sub.2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO.sub.2 fertilization as a driver of increased carbon sinks in global forests. Carbon dioxide enrichment of a mature forest resulted in the emission of the excess carbon back into the atmosphere via enhanced ecosystem respiration, suggesting that mature forests may be limited in their capacity to mitigate climate change., Author(s): Mingkai Jiang [sup.1] , Belinda E. Medlyn [sup.1] , John E. Drake [sup.1] [sup.2] , Remko A. Duursma [sup.1] , Ian C. Anderson [sup.1] , Craig V. M. Barton [...]

The forest–atmosphere exchange of carbon and water is regulated by meteorological conditions as well as canopy properties such as leaf area index (LAI, m2 m−2), photosynthetic capacity (PC μmol m−2 s−1), or surface conductance in optimal conditions (Gs,opt, mmol m−2 s−1), which can vary seasonally and inter-annually. This variability is well understood for deciduous species but is poorly characterized in evergreen forests. Here, we quantify the seasonal dynamics of a temperate evergreen eucalypt forest with estimates of LAI, litterfall, carbon and water fluxes, and meteorological conditions from measurements and model simulations. We merged MODIS Enhanced Vegetation Index (EVI) values with site-based LAI measurements to establish a 17-year sequence of monthly LAI. We ran the Community Atmosphere Biosphere Land Exchange model (CABLE-POP (version r5046)) with constant and varying LAI for our site to quantify the influence of seasonal canopy dynamics on carbon and water fluxes. We observed that the peak of LAI occurred in late summer–early autumn, with a higher and earlier peak occurring in years when summer rainfall was greater. Seasonality in litterfall and allocation of net primary productivity (FNPP) to leaf growth (af, 0–1) drove this pattern, suggesting a complete renewal of the canopy before the timing of peak LAI. Litterfall peaked in spring, followed by a high af in summer, at the end of which LAI peaked, and PC and Gs,opt reached their maximum values in autumn, resulting from a combination of high LAI and efficient mature leaves. These canopy dynamics helped explain observations of maximum gross ecosystem production (FGEP) in spring and autumn and net ecosystem carbon loss in summer at our site. Inter-annual variability in LAI was positively correlated with Net Ecosystem Production (FNEP). It would be valuable to apply a similar approach to other temperate evergreen forests to identify broad patterns of seasonality in leaf growth and turnover. Because incorporating dynamic LAI was insufficient to fully capture the dynamics of FGEP, observations of seasonal variation in photosynthetic capacity, such as from solar-induced fluorescence, should be incorporated in land surface models to improve ecosystem flux estimates in evergreen forests. [ABSTRACT FROM AUTHOR]

Elevated carbon dioxide (CO2) can increase plant growth, but the magnitude of this CO2 fertilization effect is modified by soil nutrient availability. Predicting how nutrient availability affects plant responses to elevated CO2 is a key consideration for ecosystem models, and many modeling groups have moved to, or are moving towards, incorporating nutrient limitation in their models. The choice of assumptions to represent nutrient cycling processes has a major impact on model predictions, but it can be difficult to attribute outcomes to specific assumptions in complex ecosystem simulation models. Here we revisit the quasi-equilibrium analytical framework introduced by Comins and McMurtrie (1993) and explore the consequences of specific model assumptions for ecosystem net primary productivity (NPP). We review the literature applying this framework to plant–soil models and then analyze the effect of several new assumptions on predicted plant responses to elevated CO2. Examination of alternative assumptions for plant nitrogen uptake showed that a linear function of the mineral nitrogen pool or a linear function of the mineral nitrogen pool with an additional saturating function of root biomass yield similar CO2 responses at longer timescales (>5 years), suggesting that the added complexity may not be needed when these are the timescales of interest. In contrast, a saturating function of the mineral nitrogen pool with linear dependency on root biomass yields no soil nutrient feedback on the very-long-term (>500 years), near-equilibrium timescale, meaning that one should expect the model to predict a full CO2 fertilization effect on production. Secondly, we show that incorporating a priming effect on slow soil organic matter decomposition attenuates the nutrient feedback effect on production, leading to a strong medium-term (5–50 years) CO2 response. Models incorporating this priming effect should thus predict a strong and persistent CO2 fertilization effect over time. Thirdly, we demonstrate that using a “potential NPP” approach to represent nutrient limitation of growth yields a relatively small CO2 fertilization effect across all timescales. Overall, our results highlight the fact that the quasi-equilibrium analytical framework is effective for evaluating both the consequences and mechanisms through which different model assumptions affect predictions. To help constrain predictions of the future terrestrial carbon sink, we recommend the use of this framework to analyze likely outcomes of new model assumptions before introducing them to complex model structures.

It is unclear whether CO2-stimulation of photosynthesis can propagate through slower ecosystem processes and lead to long-term increases in terrestrial carbon. Here the authors show that CO2-stimulation of photosynthesis leads to a 30% increase in forest regrowth over a decade of CO2 enrichment.

Recent experimental evidence suggests that during heat extremes, wooded ecosystems may decouple photosynthesis and transpiration, reducing photosynthesis to near zero but increasing transpiration into the boundary layer. This feedback may act to dampen, rather than amplify, heat extremes in wooded ecosystems. We examined eddy covariance databases (OzFlux and FLUXNET2015) to identify whether there was field-based evidence to support these experimental findings. We focused on two types of heat extremes: (i) the 3 days leading up to a temperature extreme, defined as including a daily maximum temperature >37 ∘C (similar to the widely used TXx metric), and (ii) heatwaves, defined as 3 or more consecutive days above 35 ∘C. When focusing on (i), we found some evidence of reduced photosynthesis and sustained or increased latent heat fluxes at seven Australian evergreen wooded flux sites. However, when considering the role of vapour pressure deficit and focusing on (ii), we were unable to conclusively disentangle the decoupling between photosynthesis and latent heat flux from the effect of increasing the vapour pressure deficit. Outside of Australia, the Tier-1 FLUXNET2015 database provided limited scope to tackle this issue as it does not sample sufficient high temperature events with which to probe the physiological response of trees to extreme heat. Thus, further work is required to determine whether this photosynthetic decoupling occurs widely, ideally by matching experimental species with those found at eddy covariance tower sites. Such measurements would allow this decoupling mechanism to be probed experimentally and at the ecosystem scale. Transpiration during heatwaves remains a key issue to resolve, as no land surface model includes a decoupling mechanism, and any potential dampening of the land–atmosphere amplification is thus not included in climate model projections.

Background and aimsIsoprene is the most important volatile organic compound emitted by land plants in terms of abundance and environmental effects. Controls on isoprene emission rates include light, temperature, water supply and CO2 concentration. A need to quantify these controls has long been recognized. There are already models that give realistic results, but they are complex, highly empirical and require separate responses to different drivers. This study sets out to find a simpler, unifying principle.MethodsA simple model is presented based on the idea of balancing demands for reducing power (derived from photosynthetic electron transport) in primary metabolism versus the secondary pathway that leads to the synthesis of isoprene. This model's ability to account for key features in a variety of experimental data sets is assessed.Key resultsThe model simultaneously predicts the fundamental responses observed in short-term experiments, namely: (1) the decoupling between carbon assimilation and isoprene emission; (2) a continued increase in isoprene emission with photosynthetically active radiation (PAR) at high PAR, after carbon assimilation has saturated; (3) a maximum of isoprene emission at low internal CO2 concentration (ci) and an asymptotic decline thereafter with increasing ci; (4) maintenance of high isoprene emissions when carbon assimilation is restricted by drought; and (5) a temperature optimum higher than that of photosynthesis, but lower than that of isoprene synthase activity.ConclusionsA simple model was used to test the hypothesis that reducing power available to the synthesis pathway for isoprene varies according to the extent to which the needs of carbon assimilation are satisfied. Despite its simplicity the model explains much in terms of the observed response of isoprene to external drivers as well as the observed decoupling between carbon assimilation and isoprene emission. The concept has the potential to improve global-scale modelling of vegetation isoprene emission.

Computer models are ubiquitous tools used to represent systems across many scientific and engineering domains. For any given system, many computer models exist, each built on different assumptions and demonstrating variability in the ways in which these systems can be represented. This variability is known as epistemic uncertainty, i.e. uncertainty in our knowledge of how these systems operate. Two primary sources of epistemic uncertainty are (1) uncertain parameter values and (2) uncertain mathematical representations of the processes that comprise the system. Many formal methods exist to analyse parameter-based epistemic uncertainty, while process-representation-based epistemic uncertainty is often analysed post hoc, incompletely, informally, or is ignored. In this model description paper we present the multi-assumption architecture and testbed (MAAT v1.0) designed to formally and completely analyse process-representation-based epistemic uncertainty. MAAT is a modular modelling code that can simply and efficiently vary model structure (process representation), allowing for the generation and running of large model ensembles that vary in process representation, parameters, parameter values, and environmental conditions during a single execution of the code. MAAT v1.0 approaches epistemic uncertainty through sensitivity analysis, assigning variability in model output to processes (process representation and parameters) or to individual parameters. In this model description paper we describe MAAT and, by using a simple groundwater model example, verify that the sensitivity analysis algorithms have been correctly implemented. The main system model currently coded in MAAT is a unified, leaf-scale enzyme kinetic model of C3 photosynthesis. In the Appendix we describe the photosynthesis model and the unification of multiple representations of photosynthetic processes. The numerical solution to leaf-scale photosynthesis is verified and examples of process variability in temperature response functions are provided. For rapid application to new systems, the MAAT algorithms for efficient variation of model structure and sensitivity analysis are agnostic of the specific system model employed. Therefore MAAT provides a tool for the development of novel or toy models in many domains, i.e. not only photosynthesis, facilitating rapid informal and formal comparison of alternative modelling approaches.

The lack of correlation between photosynthesis and plant growth under sink-limited conditions is a long-standing puzzle in plant ecophysiology that currently severely compromises our models of vegetation responses to global change. To address this puzzle, we applied data assimilation to an experiment in which the sink strength of Eucalyptus tereticornis seedlings was manipulated by restricting root volume. Our goals were to infer which processes were affected by sink limitation and to attribute the overall reduction in growth observed in the experiment to the effects on various carbon (C) component processes. Our analysis was able to infer that, in addition to a reduction in photosynthetic rates, sink limitation reduced the rate of utilization of nonstructural carbohydrate (NSC), enhanced respiratory losses, modified C allocation and increased foliage turnover. Each of these effects was found to have a significant impact on final plant biomass accumulation. We also found that inclusion of an NSC storage pool was necessary to capture seedling growth over time, particularly for sink-limited seedlings. Our approach of applying data assimilation to infer C balance processes in a manipulative experiment enabled us to extract new information on the timing, magnitude and direction of the internal C fluxes from an existing dataset. We suggest that this approach could, if used more widely, be an invaluable tool to develop appropriate representations of sink-limited growth in terrestrial biosphere models.

The capacity of the terrestrial biosphere to sequester carbon and mitigate climate change is governed by the ability of vegetation to remove emissions of CO2 through photosynthesis. Tropospheric O3, a globally abundant and potent greenhouse gas, is, however, known to damage plants, causing reductions in primary productivity. Despite emission control policies across Europe, background concentrations of tropospheric O3 have risen significantly over the last decades due to hemispheric-scale increases in O3 and its precursors. Therefore, plants are exposed to increasing background concentrations, at levels currently causing chronic damage. Studying the impact of O3 on European vegetation at the regional scale is important for gaining greater understanding of the impact of O3 on the land carbon sink at large spatial scales. In this work we take a regional approach and update the JULES land surface model using new measurements specifically for European vegetation. Given the importance of stomatal conductance in determining the flux of O3 into plants, we implement an alternative stomatal closure parameterisation and account for diurnal variations in O3 concentration in our simulations. We conduct our analysis specifically for the European region to quantify the impact of the interactive effects of tropospheric O3 and CO2 on gross primary productivity (GPP) and land carbon storage across Europe. A factorial set of model experiments showed that tropospheric O3 can suppress terrestrial carbon uptake across Europe over the period 1901 to 2050. By 2050, simulated GPP was reduced by 4 to 9 % due to plant O3 damage and land carbon storage was reduced by 3 to 7 %. The combined physiological effects of elevated future CO2 (acting to reduce stomatal opening) and reductions in O3 concentrations resulted in reduced O3 damage in the future. This alleviation of O3 damage by CO2-induced stomatal closure was around 1 to 2 % for both land carbon and GPP, depending on plant sensitivity to O3. Reduced land carbon storage resulted from diminished soil carbon stocks consistent with the reduction in GPP. Regional variations are identified with larger impacts shown for temperate Europe (GPP reduced by 10 to 20 %) compared to boreal regions (GPP reduced by 2 to 8 %). These results highlight that O3 damage needs to be considered when predicting GPP and land carbon, and that the effects of O3 on plant physiology need to be considered in regional land carbon cycle assessments.

Predicting the seasonal dynamics of ecosystem carbon fluxes is challenging in broadleaved evergreen forests because of their moderate climates and subtle changes in canopy phenology. We assessed the climatic and biotic drivers of the seasonality of net ecosystem–atmosphere CO2 exchange (NEE) of a eucalyptus-dominated forest near Sydney, Australia, using the eddy covariance method. The climate is characterised by a mean annual precipitation of 800 mm and a mean annual temperature of 18 °C, hot summers and mild winters, with highly variable precipitation. In the 4-year study, the ecosystem was a sink each year (−225 g C m−2 yr−1 on average, with a standard deviation of 108 g C m−2 yr−1); inter-annual variations were not related to meteorological conditions. Daily net C uptake was always detected during the cooler, drier winter months (June through August), while net C loss occurred during the warmer, wetter summer months (December through February). Gross primary productivity (GPP) seasonality was low, despite longer days with higher light intensity in summer, because vapour pressure deficit (D) and air temperature (Ta) restricted surface conductance during summer while winter temperatures were still high enough to support photosynthesis. Maximum GPP during ideal environmental conditions was significantly correlated with remotely sensed enhanced vegetation index (EVI; r2 = 0.46) and with canopy leaf area index (LAI; r2 = 0.29), which increased rapidly after mid-summer rainfall events. Ecosystem respiration (ER) was highest during summer in wet soils and lowest during winter months. ER had larger seasonal amplitude compared to GPP, and therefore drove the seasonal variation of NEE. Because summer carbon uptake may become increasingly limited by atmospheric demand and high temperature, and because ecosystem respiration could be enhanced by rising temperatures, our results suggest the potential for large-scale seasonal shifts in NEE in sclerophyll vegetation under climate change.

Patient‐derived cancer organoids (PDOs) hold considerable promise for personalizing therapy selection and improving patient outcomes. However, it is challenging to generate PDOs in sufficient numbers to test therapies in standard culture platforms. This challenge is particularly acute for pancreatic ductal adenocarcinoma (PDAC) where most patients are diagnosed at an advanced stage with non‐resectable tumors and where patient tissue is in the form of needle biopsies. Here the development and characterization of microfluidic devices for testing therapies using a limited amount of tissue or PDOs available from PDAC biopsies is described. It is demonstrated that microfluidic PDOs are phenotypically and genotypically similar to the gold‐standard Matrigel organoids with the advantages of 1) spheroid uniformity, 2) minimal cell number requirement, and 3) not relying on Matrigel. The utility of microfluidic PDOs is proven by testing PDO responses to several chemotherapies, including an inhibitor of glycogen synthase kinase (GSKI). In addition, microfluidic organoid cultures are used to test effectiveness of immunotherapy comprised of NK cells in combination with a novel biologic. In summary, our microfluidic device offers considerable benefits for personalizing oncology based on cancer biopsies and may, in the future, be developed into a companion diagnostic for chemotherapy or immunotherapy treatments. [ABSTRACT FROM AUTHOR]

Severe droughts have caused widespread tree mortality across many forest biomes with profound effects on the function of ecosystems and carbon balance. Climate change is expected to intensify regional-scale droughts, focusing attention on the physiological basis of drought-induced tree mortality. Recent work has shown that catastrophic failure of the plant hydraulic system is a principal mechanism involved in extensive crown death and tree mortality during drought, but the multi-dimensional response of trees to desiccation is complex. Here we focus on the current understanding of tree hydraulic performance under drought, the identification of physiological thresholds that precipitate mortality and the mechanisms of recovery after drought. Building on this, we discuss the potential application of hydraulic thresholds to process-based models that predict mortality. Because climate change is expected to intensify regional-scale droughts, it is important to identify the physiological thresholds that precipitate the mortality of trees and the mechanisms of recovery after drought., Author(s): Brendan Choat [sup.1] , Timothy J. Brodribb [sup.2] , Craig R. Brodersen [sup.3] , Remko A. Duursma [sup.1] , Rosana López [sup.1] [sup.4] , Belinda E. Medlyn [sup.1] Author [...]

Understanding the sensitivity of transpiration to stomatal conductance is critical to simulating the water cycle. This sensitivity is a function of the degree of coupling between the vegetation and the atmosphere and is commonly expressed by the decoupling factor. The degree of coupling assumed by models varies considerably and has previously been shown to be a major cause of model disagreement when simulating changes in transpiration in response to elevated CO2. The degree of coupling also offers us insight into how different vegetation types control transpiration fluxes, which is fundamental to our understanding of land–atmosphere interactions. To explore this issue, we combined an extensive literature summary from 41 studies with estimates of the decoupling coefficient estimated from FLUXNET data. We found some notable departures from the values previously reported in single-site studies. There was large variability in estimated decoupling coefficients (range 0.05–0.51) for evergreen needleleaf forests. This is a result that was broadly supported by our literature review but contrasts with the early literature which suggests that evergreen needleleaf forests are generally well coupled. Estimates from FLUXNET indicated that evergreen broadleaved forests were the most tightly coupled, differing from our literature review and instead suggesting that it was evergreen needleleaf forests. We also found that the assumption that grasses would be strongly decoupled (due to vegetation stature) was only true for high precipitation sites. These results were robust to assumptions about aerodynamic conductance and, to a lesser extent, energy balance closure. Thus, these data form a benchmarking metric against which to test model assumptions about coupling. Our results identify a clear need to improve the quantification of the processes involved in scaling from the leaf to the whole ecosystem. Progress could be made with targeted measurement campaigns at flux sites and greater site characteristic information across the FLUXNET network.

Terrestrial ecosystems have absorbed roughly 30 % of anthropogenic CO2 emissions over the past decades, but it is unclear whether this carbon (C) sink will endure into the future. Despite extensive modeling and experimental and observational studies, what fundamentally determines transient dynamics of terrestrial C storage under global change is still not very clear. Here we develop a new framework for understanding transient dynamics of terrestrial C storage through mathematical analysis and numerical experiments. Our analysis indicates that the ultimate force driving ecosystem C storage change is the C storage capacity, which is jointly determined by ecosystem C input (e.g., net primary production, NPP) and residence time. Since both C input and residence time vary with time, the C storage capacity is time-dependent and acts as a moving attractor that actual C storage chases. The rate of change in C storage is proportional to the C storage potential, which is the difference between the current storage and the storage capacity. The C storage capacity represents instantaneous responses of the land C cycle to external forcing, whereas the C storage potential represents the internal capability of the land C cycle to influence the C change trajectory in the next time step. The influence happens through redistribution of net C pool changes in a network of pools with different residence times. Moreover, this and our other studies have demonstrated that one matrix equation can replicate simulations of most land C cycle models (i.e., physical emulators). As a result, simulation outputs of those models can be placed into a three-dimensional (3-D) parameter space to measure their differences. The latter can be decomposed into traceable components to track the origins of model uncertainty. In addition, the physical emulators make data assimilation computationally feasible so that both C flux- and pool-related datasets can be used to better constrain model predictions of land C sequestration. Overall, this new mathematical framework offers new approaches to understanding, evaluating, diagnosing, and improving land C cycle models.

Elevated CO2 affects C cycling processes which in turn can influence the nitrogen (N) and phosphorus (P) concentrations of plant tissues. Given differences in how N and P are used by plants, we asked if their stoichiometry in leaves and wood was maintained or altered in a long-term elevated CO2 experiment in a mature Eucalyptus forest on a low P soil (EucFACE). We measured N and P concentrations in green leaves at different ages at the top of mature trees across 6 years including 5 years in elevated CO2. N and P concentrations in green and senesced leaves and wood were determined to evaluate both spatial and temporal variation of leaf N and P concentrations, including the N and P retranslocation in leaves and wood. Leaf P concentrations were 32% lower in old mature leaves compared to newly flushed leaves with no effect of elevated CO2 on leaf P. By contrast, elevated CO2 significantly decreased leaf N concentrations in newly flushed leaves but this effect disappeared as leaves matured. As such, newly flushed leaves had 9% lower N:P ratios in elevated CO2 and N:P ratios were not different in mature green leaves (CO2 by Age effect, P = 0.02). Over time, leaf N and P concentrations in the upper canopy slightly declined in both CO2 treatments compared to before the start of the experiment. P retranslocation in leaves was 50%, almost double that of N retranslocation (29%), indicating that this site was P-limited and that P retranslocation was an important mechanism in this ecosystem to retain P in plants. As P-limited trees tend to store relatively more N than P, we found an increased N:P ratio in sapwood in response to elevated CO2 (P < 0.01), implying N accumulation in live wood. The flexible stoichiometric ratios we observed can have important implications for how plants adjust to variable environmental conditions including climate change. Hence, variable nutrient stoichiometry should be accounted for in large-scale Earth Systems models invoking biogeochemical processes.

Global climate change is expected to increase drought duration and intensity in certain regions while increasing rainfall in others. The quantitative consequences of increased drought for ecosystems are not easy to predict. Process-based models must be informed by experiments to determine the resilience of plants and ecosystems from different climates. Here, we demonstrate what and how experimentally derived quantitative information can improve the representation of stomatal and non-stomatal photosynthetic responses to drought in large-scale vegetation models. In particular, we review literature on the answers to four key questions: (1) Which photosynthetic processes are affected under short-term drought? (2) How do the stomatal and non-stomatal responses to short-term drought vary among species originating from different hydro-climates? (3) Do plants acclimate to prolonged water stress, and do mesic and xeric species differ in their degree of acclimation? (4) Does inclusion of experimentally based plant functional type specific stomatal and non-stomatal response functions to drought help Land Surface Models to reproduce key features of ecosystem responses to drought? We highlighted the need for evaluating model representations of the fundamental eco-physiological processes under drought. Taking differential drought sensitivity of different vegetation into account is necessary for Land Surface Models to accurately model drought responses, or the drought impacts on vegetation in drier environments may be over-estimated.