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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.

Current estimates of temperature effects on plants mostly rely on air temperature, although it can significantly deviate from leaf temperature (T leaf ). To address this, some studies have used canopy temperature (T can ). However, T can fails to capture the fine-scale variation in T leaf among leaves and species in diverse canopies. We used infrared radiometers to study T leaf and T can and how they deviate from air temperature (ΔT leaf and ΔT can ) in multispecies tropical tree plantations at three sites along an elevation and temperature gradient in Rwanda. Our results showed high T leaf (up to c. 50°C) and ΔT leaf (on average 8-10°C and up to c. 20°C) of sun-exposed leaves during 10:00 h-15:00 h, being close to or exceeding photosynthetic heat tolerance thresholds. These values greatly exceeded simultaneously measured values of T can and ΔT can , respectively, leading to strongly overestimated leaf thermal safety margins if basing those on T can data. Stomatal conductance and leaf size affected T leaf and T can in line with their expected influences on leaf energy balance. Our findings highlight the importance of leaf traits for leaf thermoregulation and show that monitoring T can is not enough to capture the peak temperatures and heat stress experienced by individual leaves of different species in tropical forest canopies., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Photosynthetic acclimation to both warming and elevated CO 2 of boreal trees remains a key uncertainty in modelling the response of photosynthesis to future climates. We investigated the impact of increased growth temperature and elevated CO 2 on photosynthetic capacity (V cmax and J max ) in mature trees of two North American boreal conifers, tamarack and black spruce. We show that V cmax and J max at a standard temperature of 25°C did not change with warming, while V cmax and J max at their thermal optima (T opt ) and growth temperature (T g ) increased. Moreover, V cmax and J max at either 25°C, T opt or T g decreased with elevated CO 2 . The J max /V cmax ratio decreased with warming when assessed at both T opt and T g but did not significantly vary at 25°C. The J max /V cmax increased with elevated CO 2 at either reference temperature. We found no significant interaction between warming and elevated CO 2 on all traits. If this lack of interaction between warming and elevated CO 2 on the V cmax , J max and J max /V cmax ratio is a general trend, it would have significant implications for improving photosynthesis representation in vegetation models. However, future research is required to investigate the widespread nature of this response in a larger number of species and biomes., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Warming shifts the thermal optimum of net photosynthesis (T optA ) to higher temperatures. However, our knowledge of this shift is mainly derived from seedlings grown in greenhouses under ambient atmospheric carbon dioxide (CO 2 ) conditions. It is unclear whether shifts in T optA of field-grown trees will keep pace with the temperatures predicted for the 21 st century under elevated atmospheric CO 2 concentrations. Here, using a whole-ecosystem warming controlled experiment under either ambient or elevated CO 2 levels, we show that T optA of mature boreal conifers increased with warming. However, shifts in T optA did not keep pace with warming as T optA only increased by 0.26-0.35 °C per 1 °C of warming. Net photosynthetic rates estimated at the mean growth temperature increased with warming in elevated CO 2 spruce, while remaining constant in ambient CO 2 spruce and in both ambient CO 2 and elevated CO 2 tamarack with warming. Although shifts in T optA of these two species are insufficient to keep pace with warming, these boreal conifers can thermally acclimate photosynthesis to maintain carbon uptake in future air temperatures., (© 2023. Springer Nature Limited.)

The productivity and climate feedbacks of tropical forests depend on tree physiological responses to warmer and, over large areas, seasonally drier conditions. However, knowledge regarding such responses is limited due to data scarcity. We studied the impact of growth temperature on net photosynthesis (An), maximum rates of Rubisco carboxylation at 25 °C (Vcmax25), stomatal conductance (gs) and the slope parameter of the stomatal conductance-photosynthesis model (g1), in 10 early successional (ES) and 8 late-successional (LS) tropical tree species grown at three sites along an elevation gradient in Rwanda, differing by 6.8 °C in daytime ambient air temperature. The effect of seasonal drought on An was also investigated. We found that warm climate decreased wet-season An in LS species, but not in ES species. Values of Vcmax25 were lower at the warmest site across both successional groups, and An and Vcmax25 were higher in ES compared with LS species. Stomatal conductance exhibited no significant site differences and g1 was similar across both sites and successional groups. Drought strongly reduced An at warmer sites but not at the coolest montane site and this response was similar in both ES and LS species. Our results suggest that warming has negative effects on leaf-level photosynthesis in LS species, while both LS and ES species suffer photosynthesis declines in a warmer climate with more pronounced droughts. The contrasting responses of An between successional groups may lead to shifts in species' competitive balance in a warmer world, to the disadvantage of LS trees., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Climate warming is causing compositional changes in Andean tropical montane forests (TMFs). These shifts are hypothesised to result from differential responses to warming of cold- and warm-affiliated species, with the former experiencing mortality and the latter migrating upslope. The thermal acclimation potential of Andean TMFs remains unknown. Along a 2000 m Andean altitudinal gradient, we planted individuals of cold- and warm-affiliated species (under common soil and irrigation), exposing them to the hot and cold extremes of their thermal niches, respectively. We measured the response of net photosynthesis (A net ), photosynthetic capacity and leaf dark respiration (R dark ) to warming/cooling, 5 months after planting. In all species, A net and photosynthetic capacity at 25°C were highest when growing at growth temperatures (T g ) closest to their thermal means, declining with warming and cooling in cold-affiliated and warm-affiliated species, respectively. When expressed at T g , photosynthetic capacity and R dark remained unchanged in cold-affiliated species, but the latter decreased in warm-affiliated counterparts. R dark at 25°C increased with temperature in all species, but remained unchanged when expressed at T g . Both species groups acclimated to temperature, but only warm-affiliated species decreased R dark to photosynthetic capacity ratio at T g as temperature increased. This could confer them a competitive advantage under future warming., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (A net ) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO 2 (eCO 2 ), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO 2 on iWUE and its components A net and stomatal conductance (g s ). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller g s , but greater A net , responses to eCO 2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO 2 in all PFTs, and that increases in A net had stronger effects than reductions in g s . The USO model correctly captured stomatal behaviour with eCO 2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g 1 ) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO 2 conditions., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

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., (© 2022. The Author(s).)

The temperature sensitivity of physiological processes and growth of tropical trees remains a key uncertainty in predicting how tropical forests will adjust to future climates. In particular, our knowledge regarding warming responses of photosynthesis, and its underlying biochemical mechanisms, is very limited. We grew seedlings of two tropical montane rainforest tree species, the early-successional species Harungana montana and the late-successional species Syzygium guineense, at three different sites along an elevation gradient, differing by 6.8℃ in daytime ambient air temperature. Their physiological and growth performance was investigated at each site. The optimum temperature of net photosynthesis (T optA ) did not significantly increase in warm-grown trees in either species. Similarly, the thermal optima (T optV and T optJ ) and activation energies (E aV and E aJ ) of maximum Rubisco carboxylation capacity (V cmax ) and maximum electron transport rate (J max ) were largely unaffected by warming. However, V cmax , J max and foliar dark respiration (R d ) at 25℃ were significantly reduced by warming in both species, and this decline was partly associated with concomitant reduction in total leaf nitrogen content. The ratio of J max /V cmax decreased with increasing leaf temperature for both species, but the ratio at 25℃ was constant across sites. Furthermore, in H. montana, stomatal conductance at 25℃ remained constant across the different temperature treatments, while in S. guineense it increased with warming. Total dry biomass increased with warming in H. montana but remained constant in S. guineense. The biomass allocated to roots, stem and leaves was not affected by warming in H. montana, whereas the biomass allocated to roots significantly increased in S. guineense. Overall, our findings show that in these two tropical montane rainforest tree species, the capacity to acclimate the thermal optimum of photosynthesis is limited while warming-induced reductions in respiration and photosynthetic capacity rates are tightly coupled and linked to responses of leaf nitrogen., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Climate warming will alter photosynthesis and respiration not only via direct temperature effects on leaf biochemistry but also by increasing atmospheric dryness, thereby reducing stomatal conductance and suppressing photosynthesis. Our knowledge on how climate warming affects these processes is mainly derived from seedlings grown under highly controlled conditions. However, little is known regarding temperature responses of trees growing under field settings. We exposed mature tamarack and black spruce trees growing in a peatland ecosystem to whole-ecosystem warming of up to +9°C above ambient air temperatures in an ongoing long-term experiment (SPRUCE: Spruce and Peatland Responses Under Changing Environments). Here, we report the responses of leaf gas exchange after the first two years of warming. We show that the two species exhibit divergent stomatal responses to warming and vapor pressure deficit. Warming of up to 9°C increased leaf N in both spruce and tamarack. However, higher leaf N in the warmer plots translate into higher photosynthesis in tamarack but not in spruce, with photosynthesis being more constrained by stomatal limitations in spruce than in tamarack under warm conditions. Surprisingly, dark respiration did not acclimate to warming in spruce, and thermal acclimation of respiration was only seen in tamarack once changes in leaf N were considered. Our results highlight how warming can lead to differing stomatal responses to warming in co-occurring species, with consequent effects on both vegetation carbon and water dynamics., (© 2021 John Wiley & Sons Ltd. This article has been contributed to by US Government employees and their work is in the public domain in the USA.)

Tropical climates are getting warmer, with pronounced dry periods in large areas. The productivity and climate feedbacks of future tropical forests depend on the ability of trees to acclimate their physiological processes, such as leaf dark respiration (R d ), to these new conditions. However, knowledge on this is currently limited due to data scarcity. We studied the impact of growth temperature on R d and its dependency on net photosynthesis (A n ), leaf nitrogen (N) and phosphorus (P) contents, and leaf mass per unit area (LMA) in 16 early-successional (ES) and late-successional (LS) tropical tree species in multispecies plantations along an elevation gradient (Rwanda TREE project). Moreover, we explored the effect of drought on R d in one ES and one LS species. Leaf R d at 20°C decreased at warmer sites, regardless if it was expressed per unit leaf area, mass, N or P. This acclimation resulted in an 8% and a 28% decrease in R d at prevailing nighttime temperatures in trees at the intermediate and warmest sites, respectively. Moreover, drought reduced R d , particularly in the ES species and at the coolest site. Thermal acclimation of R d is complete or overcompensatory and independent of changes in leaf nutrients or LMA in African tropical trees., (© 2020 The Authors New Phytologist © 2020 New Phytologist Foundation.)

Differences in photosynthetic capacity among tree species and tree functional types are currently assumed to be largely driven by variation in leaf nutrient content, particularly nitrogen (N). However, recent studies indicate that leaf N content is often a poor predictor of variation in photosynthetic capacity in tropical trees. In this study, we explored the relative importance of area-based total leaf N content (N tot ) and within-leaf N allocation to photosynthetic capacity versus light-harvesting in controlling the variation in photosynthetic capacity (i.e. V cmax , J max ) among mature trees of 12 species belonging to either early (ES) or late successional (LS) groups growing in a tropical montane rainforest in Rwanda, Central Africa. Photosynthetic capacity at a common leaf temperature of 25˚C (i.e. maximum rates of Rubisco carboxylation, V cmax25 and of electron transport, J max25 ) was higher in ES than in LS species (+ 58% and 68% for V cmax25 and J max25 , respectively). While N tot did not significantly differ between successional groups, the photosynthetic dependency on N tot was markedly different. In ES species, V cmax25 was strongly and positively related to N tot but this was not the case in LS species. However, there was no significant trade-off between relative leaf N investments in compounds maximizing photosynthetic capacity versus compounds maximizing light harvesting. Both leaf dark respiration at 25˚C (+ 33%) and, more surprisingly, apparent photosynthetic quantum yield (+ 35%) was higher in ES than in LS species. Moreover, R d25 was positively related to N tot for both ES and LS species. Our results imply that efforts to quantify carbon fluxes of tropical montane rainforests would be improved if they considered contrasting within-leaf N allocation and photosynthetic N tot dependencies between species with different successional strategies., (Copyright © 2020 Ziegler, Dusenge, Nyirambangutse, Zibera, Wallin and Uddling.)

Rising atmospheric carbon dioxide (CO 2 ) concentrations may warm northern latitudes up to 8°C by the end of the century. Boreal forests play a large role in the global carbon cycle, and the responses of northern trees to climate change will thus impact the trajectory of future CO 2 increases. We grew two North American boreal tree species at a range of future climate conditions to assess how growth and carbon fluxes were altered by high CO 2 and warming. Black spruce (Picea mariana, an evergreen conifer) and tamarack (Larix laricina, a deciduous conifer) were grown under ambient (407 ppm) or elevated CO 2 (750 ppm) and either ambient temperatures, a 4°C warming, or an 8°C warming. In both species, the thermal optimum of net photosynthesis (T optA ) increased and maximum photosynthetic rates declined in warm-grown seedlings, but the strength of these changes varied between species. Photosynthetic capacity (maximum rates of Rubisco carboxylation, V cmax , and of electron transport, J max ) was reduced in warm-grown seedlings, correlating with reductions in leaf N and chlorophyll concentrations. Warming increased the activation energy for V cmax and J max (E aV and E aJ , respectively) and the thermal optimum for J max . In both species, the T optA was positively correlated with both E aV and E aJ , but negatively correlated with the ratio of J max /V cmax . Respiration acclimated to elevated temperatures, but there were no treatment effects on the Q 10 of respiration (the increase in respiration for a 10°C increase in leaf temperature). A warming of 4°C increased biomass in tamarack, while warming reduced biomass in spruce. We show that climate change is likely to negatively affect photosynthesis and growth in black spruce more than in tamarack, and that parameters used to model photosynthesis in dynamic global vegetation models (E aV and E aJ ) show no response to elevated CO 2 ., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Tropical canopies are complex, with multiple canopy layers and pronounced gap dynamics contributing to their high species diversity and productivity. An important reason for this complexity is the large variation in shade tolerance among different tree species. At present, we lack a clear understanding of which plant traits control this variation, e.g., regarding the relative contributions of whole-plant versus leaf traits or structural versus physiological traits. We investigated a broad range of traits in six tropical montane rainforest tree species with different degrees of shade tolerance, grown under three different radiation regimes (under the open sky or beneath sparse or dense canopies). The two distinct shade-tolerant species had higher fractional biomass in leaves and branches while shade-intolerant species invested more into stems, and these differences were greater under low radiation. Leaf respiration and photosynthetic light compensation point did not vary with species shade tolerance, regardless of radiation regime. Leaf temperatures in open plots were markedly higher in shade-tolerant species due to their low transpiration rates and large leaf sizes. Our results suggest that interspecific variation in shade tolerance of tropical montane trees is controlled by species differences in whole-plant biomass allocation strategy rather than by difference in physiological leaf traits determining leaf carbon balance at low radiation., (© The Author(s) 2019. Published by Oxford University Press.)

Contents Summary 32 I. The importance of plant carbon metabolism for climate change 32 II. Rising atmospheric CO2 and carbon metabolism 33 III. Rising temperatures and carbon metabolism 37 IV. Thermal acclimation responses of carbon metabolic processes can be best understood when studied together 38 V. Will elevated CO2 offset warming-induced changes in carbon metabolism? 40 VI. No plant is an island: water and nutrient limitations define plant responses to climate drivers 41 VII. Conclusions 42 Acknowledgements 42 References 42 Appendix A1 48 SUMMARY: Plant carbon metabolism is impacted by rising CO 2 concentrations and temperatures, but also feeds back onto the climate system to help determine the trajectory of future climate change. Here we review how photosynthesis, photorespiration and respiration are affected by increasing atmospheric CO 2 concentrations and climate warming, both separately and in combination. We also compile data from the literature on plants grown at multiple temperatures, focusing on net CO 2 assimilation rates and leaf dark respiration rates measured at the growth temperature (A growth and R growth , respectively). Our analyses show that the ratio of A growth to R growth is generally homeostatic across a wide range of species and growth temperatures, and that species that have reduced A growth at higher growth temperatures also tend to have reduced R growth , while species that show stimulations in A growth under warming tend to have higher R growth in the hotter environment. These results highlight the need to study these physiological processes together to better predict how vegetation carbon metabolism will respond to climate change., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Stomatal CO 2 responsiveness and photosynthetic capacity vary greatly among plant species, but the factors controlling these physiological leaf traits are often poorly understood. To explore if these traits are linked to taxonomic group identity and/or to other plant functional traits, we investigated the short-term stomatal CO 2 responses and the maximum rates of photosynthetic carboxylation (V cmax ) and electron transport (J max ) in an evolutionary broad range of tropical woody plant species. The study included 21 species representing four major seed plant taxa: gymnosperms, monocots, rosids and asterids. We found that stomatal closure responses to increased CO 2 were stronger in angiosperms than in gymnosperms, and in monocots compared to dicots. Stomatal CO 2 responsiveness was not significantly related to any of the other functional traits investigated, while a parameter describing the relationship between photosynthesis and stomatal conductance in combined leaf gas exchange models (g 1 ) was related to leaf area-specific plant hydraulic conductance. For photosynthesis, we found that the interspecific variation in V cmax and J max was related to within leaf nitrogen (N) allocation rather than to area-based total leaf N content. Within-leaf N allocation and water use were strongly co-ordinated (r 2  = 0.67), such that species with high fractional N investments into compounds maximizing photosynthetic capacity also had high stomatal conductance. We conclude that while stomatal CO 2 responsiveness of tropical woody species seems poorly related to other plant functional traits, photosynthetic capacity is linked to fractional within-leaf N allocation rather than total leaf N content and is closely co-ordinated with leaf water use.

The sensitivity of photosynthetic metabolism to temperature has been identified as a key uncertainty for projecting the magnitude of the terrestrial feedback on future climate change. While temperature responses of photosynthetic capacities have been comparatively well investigated in temperate species, the responses of tropical tree species remain unexplored. We compared the responses of seedlings of native cold-adapted tropical montane rainforest tree species with those of exotic warm-adapted plantation species, all growing in an intermediate temperature common garden in Rwanda. Leaf gas exchange responses to carbon dioxide (CO2 ) at different temperatures (20-40°C) were used to assess the temperature responses of biochemical photosynthetic capacities. Analyses revealed a lower optimum temperature for photosynthetic electron transport rates than for Rubisco carboxylation rates, along with lower electron transport optima in the native cold-adapted than in the exotic warm-adapted species. The photosynthetic optimum temperatures were generally exceeded by daytime peak leaf temperatures, in particular in the native montane rainforest climax species. This study thus provides evidence of pronounced negative effects of high temperature in tropical trees and indicates high susceptibility of montane rainforest climax species to future global warming., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Photosynthetic capacity of tree leaves is typically positively related to nutrient content and little affected by changes in growth temperature. These relationships are, however, often poorly supported for tropical trees, for which interspecific differences may be more strongly controlled by within-leaf nutrient allocation than by absolute leaf nutrient content, and little is known regarding photosynthetic acclimation to temperature. To explore the influence of leaf nutrient status, successional strategy and growth temperature on the photosynthetic capacity of tropical trees, we collected data on photosynthetic, chemical and morphological leaf traits of ten tree species in Rwanda. Seven species were studied in a forest plantation at mid-altitude (~1,700 m), whereas six species were studied in a cooler montane rainforest at higher altitude (~2,500 m). Three species were common to both sites, and, in the montane rainforest, three pioneer species and three climax species were investigated. Across species, interspecific variation in photosynthetic capacity was not related to leaf nutrient content. Instead, this variation was related to differences in within-leaf nitrogen allocation, with a tradeoff between investments into compounds related to photosynthetic capacity (higher in pioneer species) versus light-harvesting compounds (higher in climax species). Photosynthetic capacity was significantly lower at the warmer site at 1,700 m altitude. We conclude that (1) within-leaf nutrient allocation is more important than leaf nutrient content per se in controlling interspecific variation in photosynthetic capacity among tree species in tropical Rwanda, and that (2) tropical montane rainforest species exhibit decreased photosynthetic capacity when grown in a warmer environment.