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Tree mortality during global-change-type drought is usually attributed to xylem dysfunction, but as climate change increases the frequency of extreme heat events, it is necessary to better understand the interactive role of heat stress. We hypothesized that some drought-stressed plants paradoxically open stomata in heatwaves to prevent leaves from critically overheating. We experimentally imposed heat (>40°C) and drought stress onto 20 broadleaf evergreen tree/shrub species in a glasshouse study. Most well-watered plants avoided lethal overheating, but drought exacerbated thermal damage during heatwaves. Thermal safety margins (TSM) quantifying the difference between leaf surface temperature and leaf critical temperature, where photosynthesis is disrupted, identified species vulnerability to heatwaves. Several mechanisms contributed to high heat tolerance and avoidance of damaging leaf temperatures-small leaf size, low leaf osmotic potential, high leaf mass per area (i.e., thick, dense leaves), high transpirational capacity, and access to water. Water-stressed plants had smaller TSM, greater crown dieback, and a fundamentally different stomatal heatwave response relative to well-watered plants. On average, well-watered plants closed stomata and decreased stomatal conductance (gs ) during the heatwave, but droughted plants did not. Plant species with low gs , either due to isohydric stomatal behavior under water deficit or inherently low transpirational capacity, opened stomata and increased gs under high temperatures. The current paradigm maintains that stomata close before hydraulic thresholds are surpassed, but our results suggest that isohydric species may dramatically increase gs (over sixfold increases) even past their leaf turgor loss point. By actively increasing water loss at high temperatures, plants can be driven toward mortality thresholds more rapidly than has been previously recognized. The inclusion of TSM and responses to heat stress could improve our ability to predict the vulnerability of different tree species to future droughts.

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.

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.

Drought is expected to increase in frequency and severity in many regions in the future, so it is important to improve our understanding of how drought affects plant functional traits and ecological interactions. Imposing experimental water deficits is key to gaining this understanding, but has been hindered by logistic difficulties in maintaining consistently low water availability for plants. Here, we describe a simple method for applying soil water deficits to potted plants in glasshouse experiments. We modified an existing method (the "Snow and Tingey system") in order to apply a gradual, moderate water deficit to 50 plant species of different life forms (grasses, vines, shrubs, trees). The method requires less maintenance and manual handling compared to other water deficit methods, so it can be used for extended periods of time and is relatively inexpensive to implement. With only a few modifications, it is possible to easily establish and maintain soil water deficits of differing intensity and duration, as well as to incorporate interacting stress factors. We tested this method by measuring physiological responses to an applied water deficit in a subset of 11 tree/shrub species with a wide range of drought tolerances and water-use strategies. For this subgroup of species, stomatal conductance was 2-17 times lower in droughted plants than controls, although only half of the species (5 out of 11) experienced midday leaf water potentials that exceeded their turgor loss (i.e., wilting) point. Leaf temperatures were up to 8°C higher in droughted plants than controls, indicating that droughted plants are at greater risk of thermal damage, relative to unstressed plants. The largest leaf temperature differences (between droughted and well-watered plants) were in species with high rates of water loss. Rapid osmotic adjustment was observed in leaves of five species when drought stress was combined with an experimental heatwave. These results highlight the potential value of further ecological and physiological experiments utilizing this simple water deficit method to study plant responses to drought stress.

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.

Fire-prone landscapes experience frequent fires, disrupting above-ground biomass and altering below-ground soil nutrient availability. Augmentation of leaf nutrients or leaf water balance can both reduce limitations to photosynthesis and facilitate post-fire recovery in plants. These modes of fire responses are often studied separately and hence are rarely compared. We hypothesized that under severe burning, woody plants of a coastal scrub ecosystem would have higher rates of photosynthesis (A net) than in unburned areas due to a transient release from leaf nutrient and water limitations, facilitating biomass recovery post-burn. To compare these fire recovery mechanisms in regrowing plants, we measured leaf gas exchange, leaf and soil N and P concentrations, and plant stomatal limitations in Australian native coastal scrub species across a burn sequence of sites at 1 year after severe fire, 7 years following a light controlled fire, and decades after any fire at North Head, Sydney, Australia. Recent burning stimulated increases in A net by 20% over unburned trees and across three tree species. These species showed increases in total leaf N and P as a result of burning of 28% and 50% for these macronutrients, respectively, across the three species. The boost in leaf nutrients and stimulated leaf biochemical capacity for photosynthesis, alongside species-specific stomatal conductance (g s) increases, together contributed to increased photosynthetic rates after burning compared with the long-unburned area. Photosynthetic stimulation after burning occurred due to increases in nutrient concentrations in leaves, particularly N, as well as stomatal opening for some species. The findings suggest that changes in species photosynthesis and growth with increased future fire intensity or frequency may be facilitated by changes in leaf physiology after burning. On this basis, species dominance during regrowth depends on nutrient and water availability during post-fire recovery. [ABSTRACT FROM AUTHOR]