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

Background and aims

Isoprene 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.

Methods

A 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 results

The 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.

Conclusions

A 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.

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.

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

Global terrestrial models currently predict that the Amazon rainforest will continue to act as a carbon sink in the future, primarily owing to the rising atmospheric carbon dioxide (CO2) concentration. Soil phosphorus impoverishment in parts of the Amazon basin largely controls its functioning, but the role of phosphorus availability has not been considered in global model ensembles—for example, during the Fifth Climate Model Intercomparison Project. Here we simulate the planned free-air CO2 enrichment experiment AmazonFACE with an ensemble of 14 terrestrial ecosystem models. We show that phosphorus availability reduces the projected CO2-induced biomass carbon growth by about 50% to 79 ± 63 g C m−2 yr−1 over 15 years compared to estimates from carbon and carbon–nitrogen models. Our results suggest that the resilience of the region to climate change may be much less than previously assumed. Variation in the biomass carbon response among the phosphorus-enabled models is considerable, ranging from 5 to 140 g C m−2 yr−1, owing to the contrasting plant phosphorus use and acquisition strategies considered among the models. The Amazon forest response thus depends on the interactions and relative contributions of the phosphorus acquisition and use strategies across individuals, and to what extent these processes can be upregulated under elevated CO2.

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

The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO₂ response curves, including data from 141 C₃ species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common-garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate.

Leaf-level gas exchange data support the mechanistic understanding of plant fluxes of carbon and water. These fluxes inform our understanding of ecosystem function, are an important constraint on parameterization of terrestrial biosphere models, are necessary to understand the response of plants to global environmental change, and are integral to efforts to improve crop production. Collection of these data using gas analyzers can be both technically challenging and time consuming, and individual studies generally focus on a small range of species, restricted time periods, or limited geographic regions. The high value of these data is exemplified by the many publications that reuse and synthesize gas exchange data, however the lack of metadata and data reporting conventions make full and efficient use of these data difficult. Here we propose a reporting format for leaf-level gas exchange data and metadata to provide guidance to data contributors on how to store data in repositories to maximize their discoverability, facilitate their efficient reuse, and add value to individual datasets. For data users, the reporting format will better allow data repositories to optimize data search and extraction, and more readily integrate similar data into harmonized synthesis products. The reporting format specifies data table variable naming and unit conventions, as well as metadata characterizing experimental conditions and protocols. For common data types that were the focus of this initial version of the reporting format, i.e., survey measurements, dark respiration, carbon dioxide and light response curves, and parameters derived from those measurements, we took a further step of defining required additional data and metadata that would maximize the potential reuse of those data types. To aid data contributors and the development of data ingest tools by data repositories we provided a translation table comparing the outputs of common gas exchange instruments. Extensive consultation with data collectors, data users, instrument manufacturers, and data scientists was undertaken in order to ensure that the reporting format met community needs. The reporting format presented here is intended to form a foundation for future development that will incorporate additional data types and variables as gas exchange systems and measurement approaches advance in the future. The reporting format is published in the U.S. Department of Energy's ESS-DIVE data repository, with documentation and future development efforts being maintained in a version control system.