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Heatwaves and soil droughts are increasing in frequency and intensity, leading many tree species to exceed their thermal thresholds, and driving wide‐scale forest mortality. Therefore, investigating heat tolerance and canopy temperature regulation mechanisms is essential to understanding and predicting tree vulnerability to hot droughts. We measured the diurnal and seasonal variation in leaf water potential (Ψ), gas exchange (photosynthesis Anet and stomatal conductance gs), canopy temperature (Tcan), and heat tolerance (leaf critical temperature Tcrit and thermal safety margins TSM, i.e., the difference between maximum Tcan and Tcrit) in three oak species in forests along a latitudinal gradient (Quercus petraea in Switzerland, Quercus ilex in France, and Quercus coccifera in Spain) throughout the growing season. Gas exchange and Ψ of all species were strongly reduced by increased air temperature (Tair) and soil drying, resulting in stomatal closure and inhibition of photosynthesis in Q. ilex and Q. coccifera when Tair surpassed 30°C and soil moisture dropped below 14%. Across all seasons, Tcan was mainly above Tair but increased strongly (up to 10°C > Tair) when Anet was null or negative. Although trees endured extreme Tair (up to 42°C), positive TSM were maintained during the growing season due to high Tcrit in all species (average Tcrit of 54.7°C) and possibly stomatal decoupling (i.e., Anet ≤0 while gs >0). Indeed, Q. ilex and Q. coccifera trees maintained low but positive gs (despite null Anet), decreasing Ψ passed embolism thresholds. This may have prevented Tcan from rising above Tcrit during extreme heat. Overall, our work highlighted that the mechanisms behind heat tolerance and leaf temperature regulation in oak trees include a combination of high evaporative cooling, large heat tolerance limits, and stomatal decoupling. These processes must be considered to accurately predict plant damages, survival, and mortality during extreme heatwaves. [ABSTRACT FROM AUTHOR]

Photosynthetically active radiation (PAR) is typically defined as light with a wavelength within 400–700 nm. However, ultra‐violet (UV) radiation within 280–400 nm and far‐red (FR) radiation within 700–750 nm can also excite photosystems, though not as efficiently as PAR. Vegetation and land surface models (LSMs) typically do not explicitly account for UV's contribution to energy budgets or photosynthesis, nor FR's contribution to photosynthesis. However, whether neglecting UV and FR has significant impacts remains unknown. We explored how canopy radiative transfer (RT) and photosynthesis are impacted when explicitly implementing UV in the canopy RT model and accounting for UV and FR in the photosynthesis models within a next‐generation LSM that can simulate hyperspectral canopy RT. We validated our improvements using photosynthesis measurements from plants under different light sources and intensities and surface reflection from an eddy‐covariance tower. Our model simulations suggested that at the whole plant level, after accounting for UV and FR explicitly, chlorophyll content, leaf area index (LAI), clumping index, and solar radiation all impact the modeling of gross primary productivity (GPP). At the global scale, mean annual GPP within a grid would increase by up to 7.3% and the increase is proportional to LAI; globally integrated GPP increases by 4.6 PgC year−1 (3.8% of the GPP without accounting for UV + FR). Further, using PAR to proxy UV could overestimate surface albedo by more than 0.1, particularly in the boreal forests. Our results highlight the importance of improving UV and FR in canopy RT and photosynthesis modeling and the necessity to implement hyperspectral or multispectral canopy RT schemes in future vegetation and LSMs. [ABSTRACT FROM AUTHOR]

The spatial dispersion of photoelements within a vegetation canopy, quantified by the clumping index (CI), directly regulates the within-canopy light environment and photosynthesis rate, but is not commonly implemented in terrestrial biosphere models to estimate the ecosystem carbon cycle. A few global CI products have been developed recently with remote sensing measurements, making it possible to examine the global impacts of CI. This study deployed CI in the radiative transfer scheme of the Community Land Model version 5 (CLM5) and used the revised CLM5 to quantitatively evaluate the extent to which CI can affect canopy absorbed radiation and gross primary production (GPP), and for the first time, considering the uncertainty and seasonal variation of CI with multiple remote sensing products. Compared to the results without considering the CI impact, the revised CLM5 estimated that sunlit canopy absorbed up to 9%-15% and 23%-34% less direct and diffuse radiation, respectively, while shaded canopy absorbed 3%-18% more diffuse radiation across different biome types. The CI impacts on canopy light conditions included changes in canopy light absorption, and sunlit-shaded leaf area fraction related to nitrogen distribution and thus the maximum rate of Rubisco carboxylase activity (Vcmax ), which together decreased photosynthesis in sunlit canopy by 5.9-7.2 PgC year-1 while enhanced photosynthesis by 6.9-8.2 PgC year-1 in shaded canopy. With higher light use efficiency of shaded leaves, shaded canopy increased photosynthesis compensated and exceeded the lost photosynthesis in sunlit canopy, resulting in 1.0 ± 0.12 PgC year-1 net increase in GPP. The uncertainty of GPP due to the different input CI datasets was much larger than that caused by CI seasonal variations, and was up to 50% of the magnitude of GPP interannual variations in the tropical regions. This study highlights the necessity of considering the impacts of CI and its uncertainty in terrestrial biosphere models.

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 C3 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.

Leaf coloring and fall mark the end of the growing season (EOS), playing essential roles in nutrient cycling, resource allocation, ecological interactions, and as climate change indicators. However, understanding future changes in autumn phenology is challenging due to the multitude of likely environmental cues and substantial variations in timing caused by different derivation methods. Yet, it remains unclear whether these two factors are independent or if methodological uncertainties influence the environmental cues determined. We derived start of growing season (SOS) and EOS at a mixed beech forest in Central Germany for the period 2000–2020 based on four different derivation methods using a unique long‐term data set of in‐situ data, canopy imagery, eddy covariance measurements, and satellite remote sensing data and determined their influence on a predictor analysis of leaf senescence. Both SOS and EOS exhibited substantial ranges in mean onset dates (39.5 and 28.6 days, respectively) across the different methods, although inter‐annual variations and advancing SOS trends were similar across methods. Depending on the data, EOS trends were advanced or delayed, but inter‐annual patterns correlated well (mean r =.46). Overall, warm, dry, and less photosynthetically productive growing seasons were more likely to be associated with a delayed EOS, while colder, wetter, and more photosynthetically productive vegetation periods resulted in an earlier EOS. In addition, contrary to recent results, no clear influence of pre‐solstice vegetation activity on the timing of senescence was detected. However, most notable were the large differences in sign and strength of potential drivers both in the univariate and multivariate analyses when comparing derivation methodologies. The results suggest that an ensemble analysis of all available phenological data sources and derivation methods is needed for general statements on autumn phenology and its influencing variables and correct implementation of the senescence process in ecosystem models. [ABSTRACT FROM AUTHOR]

Climate change will impact gross primary productivity (GPP), net primary productivity (NPP), and carbon storage in wooded ecosystems. The extent of change will be influenced by thermal acclimation of photosynthesis—the ability of plants to adjust net photosynthetic rates in response to growth temperatures—yet regional differences in acclimation effects among wooded ecosystems is currently unknown. We examined the effects of changing climate on 17 Australian wooded ecosystems with and without the effects of thermal acclimation of C3 photosynthesis. Ecosystems were drawn from five ecoregions (tropical savanna, tropical forest, Mediterranean woodlands, temperate woodlands, and temperate forests) that span Australia's climatic range. We used the CABLE‐POP land surface model adapted with thermal acclimation functions and forced with HadGEM2‐ES climate projections from RCP8.5. For each site and ecoregion we examined (a) effects of climate change on GPP, NPP, and live tree carbon storage; and (b) impacts of thermal acclimation of photosynthesis on simulated changes. Between the end of the historical (1976–2005) and projected (2070–2099) periods simulated annual carbon uptake increased in the majority of ecosystems by 26.1%–63.3% for GPP and 15%–61.5% for NPP. Thermal acclimation of photosynthesis further increased GPP and NPP in tropical savannas by 27.2% and 22.4% and by 11% and 10.1% in tropical forests with positive effects concentrated in the wet season (tropical savannas) and the warmer months (tropical forests). We predicted minimal effects of thermal acclimation of photosynthesis on GPP, NPP, and carbon storage in Mediterranean woodlands, temperate woodlands, and temperate forests. Overall, positive effects were strongly enhanced by increasing CO2 concentrations under RCP8.5. We conclude that the direct effects of climate change will enhance carbon uptake and storage in Australian wooded ecosystems (likely due to CO2 enrichment) and that benefits of thermal acclimation of photosynthesis will be restricted to tropical ecoregions. [ABSTRACT FROM AUTHOR]

The length of the growing season has a large influence on the carbon, water, and energy fluxes of global terrestrial ecosystems. While there has been mounting evidence of an advanced start of the growing season mostly due to elevated spring air temperatures, the mechanisms that control the end of the growing season (EOS) in most ecosystems remain relatively less well understood. Recently, a strong lagged control of EOS by growing season photosynthesis has been proposed, suggesting that more productive growing seasons lead to an earlier EOS. However, this relationship has not been extensively tested with in-situ observations across a variety of ecosystems. Here, we use observations from 40 eddy-covariance flux tower sites in temperate and boreal ecosystems in the northern hemisphere with more than 10 years of observations (594 site-years), ground observations of phenology, satellite observations from the Moderate Resolution Imaging Spectroradiometer (MODIS), and three leaf senescence models to test the extent of a relationship between growing season photosynthesis and end of season senescence. The results suggest that there is no significant negative relationship between growing season photosynthesis and observed leaf senescence, flux-inferred EOS estimates, or remotely sensed phenological metrics, in most ecosystems. On the contrary, while we found negative effects of summer air temperatures and autumn vapor pressure deficit on EOS, more productive growing seasons were typically related to a later, not earlier, EOS. Our results challenge recent reports of carry-over effects of photosynthesis on EOS timing, and suggest those results may not hold over a large range of ecosystems.

High air temperatures increase atmospheric vapor pressure deficit (VPD) and the severity of drought, threatening forests worldwide. Plants regulate stomata to maximize carbon gain and minimize water loss, resulting in a close coupling between net photosynthesis (Anet) and stomatal conductance (gs). However, evidence for decoupling of gs from Anet under extreme heat has been found. Such a response both enhances survival of leaves during heat events but also quickly depletes available water. To understand the prevalence and significance of this decoupling, we measured leaf gas exchange in 26 tree and shrub species growing in the glasshouse or at an urban site in Sydney, Australia on hot days (maximum Tair > 40°C). We hypothesized that on hot days plants with ample water access would exhibit reduced Anet and use transpirational cooling leading to stomatal decoupling, whereas plants with limited water access would rely on other mechanisms to avoid lethal temperatures. Instead, evidence for stomatal decoupling was found regardless of plant water access. Transpiration of well‐watered plants was 23% higher than model predictions during heatwaves, which effectively cooled leaves below air temperature. For hotter, droughted plants, the increase in transpiration during heatwaves was even more pronounced—gs was 77% higher than model predictions. Stomatal decoupling was found for most broadleaf evergreen and broadleaf deciduous species at the urban site, including some wilted trees with limited water access. Decoupling may simply be a passive consequence of the physical effects of high temperature on plant leaves through increased cuticular conductance of water vapor, or stomatal decoupling may be an adaptive response that is actively regulated by stomatal opening under high temperatures. This temperature response is not yet included in any land surface model, suggesting that model predictions of evapotranspiration may be underpredicted at high temperature and high VPD. [ABSTRACT FROM AUTHOR]

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.

Predicted increases in extreme droughts will likely cause major shifts in carbon sequestration and forest composition. Although growth declines during drought are widely documented, an increasing number of studies have reported both positive and negative responses to the same drought. These divergent growth patterns may reflect thresholds (i.e., nonlinear responses) promoted by changes in the dominant climatic constraints on tree growth. Here we tested whether stemwood growth exhibited linear or nonlinear responses to temperature and precipitation and whether stemwood growth thresholds co‐occurred with multiple thresholds in source and sink processes that limit tree growth. We extracted 772 tree cores, 1398 needle length records, and 1075 stable isotope samples from 27 sites across whitebark pine's (Pinus albicaulis Engelm.) climatic niche in the Sierra Nevada. Our results indicated that a temperature threshold in stemwood growth occurred at 8.4°C (7.12–9.51°C; estimated using fall‐spring maximum temperature). This threshold was significantly correlated with thresholds in foliar growth, as well as carbon (δ13C) and nitrogen (δ15N) stable isotope ratios, that emerged during drought. These co‐occurring thresholds reflected the transition between energy‐ and water‐limited tree growth (i.e., the E–W limitation threshold). This transition likely mediated carbon and nutrient cycling, as well as important differences in growth‐defense trade‐offs and drought adaptations. Furthermore, whitebark pine growing in energy‐limited regions may continue to experience elevated growth in response to climate change. The positive effect of warming, however, may be offset by growth declines in water‐limited regions, threatening the long‐term sustainability of the recently listed whitebark pine species in the Sierra Nevada. [ABSTRACT FROM AUTHOR]

Global-scale ocean acidification has spurred interest in the capacity of seagrass ecosystems to increase seawater pH within crucial shoreline habitats through photosynthetic activity. However, the dynamic variability of the coastal carbonate system has impeded generalization into whether seagrass aerobic metabolism ameliorates low pH on physiologically and ecologically relevant timescales. Here we present results of the most extensive study to date of pH modulation by seagrasses, spanning seven meadows (Zostera marina) and 1000 km of U.S. west coast over 6 years. Amelioration by seagrass ecosystems compared to non-vegetated areas occurred 65% of the time (mean increase 0.07 ± 0.008 SE). Events of continuous elevation in pH within seagrass ecosystems, indicating amelioration of low pH, were longer and of greater magnitude than opposing cases of reduced pH or exacerbation. Sustained elevations in pH of >0.1, comparable to a 30% decrease in [H+ ], were not restricted only to daylight hours but instead persisted for up to 21 days. Maximal pH elevations occurred in spring and summer during the seagrass growth season, with a tendency for stronger effects in higher latitude meadows. These results indicate that seagrass meadows can locally alleviate low pH conditions for extended periods of time with important implications for the conservation and management of coastal ecosystems.

Mechanistic photosynthesis models are at the heart of terrestrial biosphere models (TBMs) simulating the daily, monthly, annual and decadal rhythms of carbon assimilation (A). These models are founded on robust mathematical hypotheses that describe how A responds to changes in light and atmospheric CO2 concentration. Two predominant photosynthesis models are in common usage: Farquhar (FvCB) and Collatz (CBGB). However, a detailed quantitative comparison of these two models has never been undertaken. In this study, we unify the FvCB and CBGB models to a common parameter set and use novel multi-hypothesis methods (that account for both hypothesis and parameter variability) for process-level sensitivity analysis. These models represent three key biological processes: carboxylation, electron transport, triose phosphate use (TPU) and an additional model process: limiting-rate selection. Each of the four processes comprises 1-3 alternative hypotheses giving 12 possible individual models with a total of 14 parameters. To broaden inference, TBM simulations were run and novel, high-resolution photosynthesis measurements were made. We show that parameters associated with carboxylation are the most influential parameters but also reveal the surprising and marked dominance of the limiting-rate selection process (accounting for 57% of the variation in A vs. 22% for carboxylation). The limiting-rate selection assumption proposed by CBGB smooths the transition between limiting rates and always reduces A below the minimum of all potentially limiting rates, by up to 25%, effectively imposing a fourth limitation on A. Evaluation of the CBGB smoothing function in three TBMs demonstrated a reduction in global A by 4%-10%, equivalent to 50%-160% of current annual fossil fuel emissions. This analysis reveals a surprising and previously unquantified influence of a process that has been integral to many TBMs for decades, highlighting the value of multi-hypothesis methods.

Tropical forests absorb large amounts of atmospheric CO2 through photosynthesis, but high surface temperatures suppress this absorption while promoting isoprene emissions. While mechanistic isoprene emission models predict a tight coupling to photosynthetic electron transport (ETR) as a function of temperature, direct field observations of this phenomenon are lacking in the tropics and are necessary to assess the impact of a warming climate on global isoprene emissions. Here we demonstrate that in the early successional species Vismia guianensis in the central Amazon, ETR rates increased with temperature in concert with isoprene emissions, even as stomatal conductance (gs ) and net photosynthetic carbon fixation (Pn ) declined. We observed the highest temperatures of continually increasing isoprene emissions yet reported (50°C). While Pn showed an optimum value of 32.6 ± 0.4°C, isoprene emissions, ETR, and the oxidation state of PSII reaction centers (qL ) increased with leaf temperature with strong linear correlations for ETR (ƿ = 0.98) and qL (ƿ = 0.99) with leaf isoprene emissions. In contrast, other photoprotective mechanisms, such as non-photochemical quenching, were not activated at elevated temperatures. Inhibition of isoprenoid biosynthesis repressed Pn at high temperatures through a mechanism that was independent of stomatal closure. While extreme warming will decrease gs and Pn in tropical species, our observations support a thermal tolerance mechanism where the maintenance of high photosynthetic capacity under extreme warming is assisted by the simultaneous stimulation of ETR and metabolic pathways that consume the direct products of ETR including photorespiration and the biosynthesis of thermoprotective isoprenoids. Our results confirm that models which link isoprene emissions to the rate of ETR hold true in tropical species and provide necessary "ground-truthing" for simulations of the large predicted increases in tropical isoprene emissions with climate warming.

Monitoring and estimating drought impact on plant physiological processes over large regions remains a major challenge for remote sensing and land surface modeling, with important implications for understanding plant mortality mechanisms and predicting the climate change impact on terrestrial carbon and water cycles. The Orbiting Carbon Observatory 3 (OCO‐3), with its unique diurnal observing capability, offers a new opportunity to track drought stress on plant physiology. Using radiative transfer and machine learning modeling, we derive a metric of afternoon photosynthetic depression from OCO‐3 solar‐induced chlorophyll fluorescence (SIF) as an indicator of plant physiological drought stress. This unique diurnal signal enables a spatially explicit mapping of plants' physiological response to drought. Using OCO‐3 observations, we detect a widespread increasing drought stress during the 2020 southwest US drought. Although the physiological drought stress is largely related to the vapor pressure deficit (VPD), our results suggest that plants' sensitivity to VPD increases as the drought intensifies and VPD sensitivity develops differently for shrublands and grasslands. Our findings highlight the potential of using diurnal satellite SIF observations to advance the mechanistic understanding of drought impact on terrestrial ecosystems and to improve land surface modeling. [ABSTRACT FROM AUTHOR]

Canopy greening, which is associated with significant canopy structure changes, is the most notable signal of ecosystem changes in response to anthropogenic climate change. However, our knowledge of the changing pattern of canopy development and senescence, and its endogenous and climatic drivers is still limited. Here, we used the Normalized Difference Vegetation Index (NDVI) to quantify the changes in the speed of canopy development and senescence over the Tibetan Plateau (TP) during 2000–2018, and used a solar‐induced chlorophyll fluorescence dataset as a proxy for photosynthesis, in combination with climate datasets to decipher the endogenous and climatic drivers of the interannual variation in canopy changes. We found that the canopy development during the early green‐up stage (April–May) is accelerating at a rate of 0.45–0.8 × 10−3 month−1 year−1. However, this accelerating canopy development was largely offset by a decelerating canopy development during June and July (−0.61 to −0.51 × 10−3 month−1 year−1), leading to the peak NDVI over the TP increasing at a rate of only one fifth of that in northern temperate regions, and less than one tenth of that in the Arctic and boreal regions. During the green‐down period, we observed a significant accelerating canopy senescence during October. Photosynthesis was found to be the dominant driver for canopy changes over the TP. Increasing photosynthesis stimulates canopy development during the early green‐up stage. However, slower canopy development and accelerated senescence was found with larger photosynthesis in late growth stages. This negative relationship between photosynthesis and canopy development is probably linked to the source–sink balance of plants and shifts in the allocation regime. These results suggest a sink limitation for plant growth over the TP. The impact of canopy greening on the carbon cycle may be more complicated than the source‐oriented paradigm used in current ecosystem models. [ABSTRACT FROM AUTHOR]

Solar‐induced chlorophyll fluorescence (SIF) is a remotely sensed optical signal emitted during the light reactions of photosynthesis. The past two decades have witnessed an explosion in availability of SIF data at increasingly higher spatial and temporal resolutions, sparking applications in diverse research sectors (e.g., ecology, agriculture, hydrology, climate, and socioeconomics). These applications must deal with complexities caused by tremendous variations in scale and the impacts of interacting and superimposing plant physiology and three‐dimensional vegetation structure on the emission and scattering of SIF. At present, these complexities have not been overcome. To advance future research, the two companion reviews aim to (1) develop an analytical framework for inferring terrestrial vegetation structures and function that are tied to SIF emission, (2) synthesize progress and identify challenges in SIF research via the lens of multi‐sector applications, and (3) map out actionable solutions to tackle these challenges and offer our vision for research priorities over the next 5–10 years based on the proposed analytical framework. This paper is the first of the two companion reviews, and theory oriented. It introduces a theoretically rigorous yet practically applicable analytical framework. Guided by this framework, we offer theoretical perspectives on three overarching questions: (1) The forward (mechanism) question—How are the dynamics of SIF affected by terrestrial ecosystem structure and function? (2) The inference question: What aspects of terrestrial ecosystem structure, function, and service can be reliably inferred from remotely sensed SIF and how? (3) The innovation question: What innovations are needed to realize the full potential of SIF remote sensing for real‐world applications under climate change? The analytical framework elucidates that process complexity must be appreciated in inferring ecosystem structure and function from the observed SIF; this framework can serve as a diagnosis and inference tool for versatile applications across diverse spatial and temporal scales. [ABSTRACT FROM AUTHOR]

Although our observing capabilities of solar‐induced chlorophyll fluorescence (SIF) have been growing rapidly, the quality and consistency of SIF datasets are still in an active stage of research and development. As a result, there are considerable inconsistencies among diverse SIF datasets at all scales and the widespread applications of them have led to contradictory findings. The present review is the second of the two companion reviews, and data oriented. It aims to (1) synthesize the variety, scale, and uncertainty of existing SIF datasets, (2) synthesize the diverse applications in the sector of ecology, agriculture, hydrology, climate, and socioeconomics, and (3) clarify how such data inconsistency superimposed with the theoretical complexities laid out in (Sun et al., 2023) may impact process interpretation of various applications and contribute to inconsistent findings. We emphasize that accurate interpretation of the functional relationships between SIF and other ecological indicators is contingent upon complete understanding of SIF data quality and uncertainty. Biases and uncertainties in SIF observations can significantly confound interpretation of their relationships and how such relationships respond to environmental variations. Built upon our syntheses, we summarize existing gaps and uncertainties in current SIF observations. Further, we offer our perspectives on innovations needed to help improve informing ecosystem structure, function, and service under climate change, including enhancing in‐situ SIF observing capability especially in "data desert" regions, improving cross‐instrument data standardization and network coordination, and advancing applications by fully harnessing theory and data. [ABSTRACT FROM AUTHOR]

Elevated temperature (Te) and drought often co-occur and interactively affect plant carbon (C) metabolism and thus the ecosystem C cycling; however, the magnitude of their interaction is unclear, making the projection of global change impacts challenging. Here, we compiled 107 journal articles in which temperature and water availability were jointly manipulated, and we performed a meta-analysis of interactive effects of Te and drought on leaf photosynthesis (Agrowth) and respiration (Rgrowth) at growth temperature, nonstructural carbohydrates and biomass of plants, and their dependencies on experimental and biological moderators (e.g., treatment intensity, plant functional type). Our results showed that, overall, there was no significant interaction of Te and drought on Agrowth. Te accelerated Rgrowth under well-watered conditions rather than under drought conditions. The Te × drought interaction on leaf soluble sugar and starch concentrations were neutral and negative, respectively. The effect of Te and drought on plant biomass displayed a negative interaction, with Te deteriorating the drought impacts. Drought induced an increase in root to shoot ratio at ambient temperature but not at Te. The magnitudes of Te and drought negatively modulated the Te × drought interactions on Agrowth. Root biomass of woody plants was more vulnerable to drought than that of herbaceous plants at ambient temperature, but this difference diminished at Te. Perennial herbs exhibited a stronger amplifying effect of Te on plant biomass in response to drought than did annual herbs. Te exacerbated the responses of Agrowth and stomatal conductance to drought for evergreen broadleaf trees rather than for deciduous broadleaf and evergreen coniferous trees. A negative Te × drought interaction on plant biomass was observed on species-level rather than on community-level. Collectively, our findings provide a mechanistic understanding of the interactive effects of Te and drought on plant C metabolism, which would improve the prediction of climate change impacts. [ABSTRACT FROM AUTHOR]

Decline in mesophyll conductance (gm) plays a key role in limiting photosynthesis in plants exposed to elevated ozone (Om). Leaf anatomical traits are known to influence gm, but the potential effects of Om-induced changes in leaf anatomy on gm have not yet been clarified. Here, two poplar clones were exposed to elevated Om. The effects of Om on the photosynthetic capacity and anatomical characteristics were assessed to investigate the leaf anatomical properties that potentially affect gm. We also conducted global meta-analysis to explore the general response patterns of gm and leaf anatomy to Om exposure. We found that the Om-induced reduction in gm was critical in limiting leaf photosynthesis. Changes in liquid-phase conductance rather than gas-phase conductance drive the decline in gm under elevated Om, and this effect was associated with thicker cell walls and smaller chloroplast sizes. The effects of Om on palisade and spongy mesophyll cell traits and their contributions to gm were highly genotype-dependent. Our results suggest that, while anatomical adjustments under elevated Om may contribute to defense against Om stress, they also cause declines in gm and photosynthesis. These results provide the first evidence of anatomical constraints on gm under elevated Om. [ABSTRACT FROM AUTHOR]

The eddy covariance (EC) technique is used to measure the net ecosystem exchange (NEE) of CO2 between ecosystems and the atmosphere, offering a unique opportunity to study ecosystem responses to climate change. NEE is the difference between the total CO2 release due to all respiration processes (RECO), and the gross carbon uptake by photosynthesis (GPP). These two gross CO2 fluxes are derived from EC measurements by applying partitioning methods that rely on physiologically based functional relationships with a limited number of environmental drivers. However, the partitioning methods applied in the global FLUXNET network of EC observations do not account for the multiple co-acting factors that modulate GPP and RECO flux dynamics. To overcome this limitation, we developed a hybrid data-driven approach based on combined neural networks (NNC-part ). NNC-part incorporates process knowledge by introducing a photosynthetic response based on the light-use efficiency (LUE) concept, and uses a comprehensive dataset of soil and micrometeorological variables as fluxes drivers. We applied the method to 36 sites from the FLUXNET2015 dataset and found a high consistency in the results with those derived from other standard partitioning methods for both GPP (R2  > .94) and RECO (R2  > .8). High consistency was also found for (a) the diurnal and seasonal patterns of fluxes and (b) the ecosystem functional responses. NNC-part performed more realistic than the traditional methods for predicting additional patterns of gross CO2 fluxes, such as: (a) the GPP response to VPD, (b) direct effects of air temperature on GPP dynamics, (c) hysteresis in the diel cycle of gross CO2 fluxes, (d) the sensitivity of LUE to the diffuse to direct radiation ratio, and (e) the post rain respiration pulse after a long dry period. In conclusion, NNC-part is a valid data-driven approach to provide GPP and RECO estimates and complementary to the existing partitioning methods.

Ozone is the most damaging air pollutant to crops, currently reducing Midwest US maize production by up to 10%, yet there has been very little effort to adapt germplasm for ozone tolerance. Ozone enters plants through stomata, reacts to form reactive oxygen species in the apoplast and ultimately decreases photosynthetic C gain. In this study, 10 diverse inbred parents were crossed in a half-diallel design to create 45 F1 hybrids, which were tested for ozone response in the field using free air concentration enrichment (FACE). Ozone stress increased the heritability of photosynthetic traits and altered genetic correlations among traits. Hybrids from parents Hp301 and NC338 showed greater sensitivity to ozone stress, and disrupted relationships among photosynthetic traits. The physiological responses underlying sensitivity to ozone differed in hybrids from the two parents, suggesting multiple mechanisms of response to oxidative stress. FACE technology was essential to this evaluation because genetic variation in photosynthesis under elevated ozone was not predictable based on performance at ambient ozone. These findings suggest that selection under elevated ozone is needed to identify deleterious alleles in the world's largest commodity crop.

Extreme climatic events, such as heat waves, cold snaps and drought spells, related to global climate change, have become more frequent and intense in recent years. Acclimation of plant physiological processes to changes in environmental conditions is a key component of plant adaptation to climate change. We assessed the temperature response of leaf photosynthetic parameters in wheat grown under contrasting water regimes and growth temperatures (Tgrowth). Two independent experiments were conducted under controlled conditions. In Experiment 1, two wheat genotypes were subjected to well‐watered or drought‐stressed treatments; in Experiment 2, the two water regimes combined with high, medium and low Tgrowth were imposed on one genotype. Parameters of a biochemical C3‐photosynthesis model were estimated at six leaf temperatures for each factor combination. Photosynthesis acclimated more to drought than to Tgrowth. Drought affected photosynthesis by lowering its optimum temperature (Topt) and the values at Topt of light‐saturated net photosynthesis, stomatal conductance, mesophyll conductance, the maximum rate of electron transport (Jmax) and the maximum rate of carboxylation by Rubisco (Vcmax). Topt for Vcmax was up to 40°C under well‐watered conditions but 24–34°C under drought. The decrease in photosynthesis under drought varied among Tgrowth but was similar between genotypes. The temperature response of photosynthetic quantum yield under drought was partly attributed to photorespiration but more to alternative electron transport. All these changes in biochemical parameters could not be fully explained by the changed leaf nitrogen content. Further model analysis showed that both diffusional and biochemical parameters of photosynthesis and their thermal sensitivity acclimate little to Tgrowth, but acclimate considerably to drought and the combination of drought and Tgrowth. The commonly used modelling approaches, which typically consider the response of diffusional parameters, but ignore acclimation responses of biochemical parameters to drought and Tgrowth, strongly overestimate leaf photosynthesis under variable temperature and drought. [ABSTRACT FROM AUTHOR]

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.

The terrestrial biosphere plays a critical role in mitigating climate change by absorbing anthropogenic CO2 emissions through photosynthesis. The rate of photosynthesis is determined jointly by environmental variables and the intrinsic photosynthetic capacity of plants (i.e. maximum carboxylation rate; Vcmax25 ). A lack of an effective means to derive spatially and temporally explicit Vcmax25 has long hampered efforts towards estimating global photosynthesis accurately. Recent work suggests that leaf chlorophyll content (Chlleaf ) is strongly related to Vcmax25 , since Chlleaf and Vcmax25 are both correlated with photosynthetic nitrogen content. We used medium resolution satellite images to derive spatially and temporally explicit Chlleaf , which we then used to parameterize Vcmax25 within a terrestrial biosphere model. Modelled photosynthesis estimates were evaluated against measured photosynthesis at 124 eddy covariance sites. The inclusion of Chlleaf in a terrestrial biosphere model improved the spatial and temporal variability of photosynthesis estimates, reducing biases at eddy covariance sites by 8% on average, with the largest improvements occurring for croplands (21% bias reduction) and deciduous forests (15% bias reduction). At the global scale, the inclusion of Chlleaf reduced terrestrial photosynthesis estimates by 9 PgC/year and improved the correlations with a reconstructed solar-induced fluorescence product and a gridded photosynthesis product upscaled from tower measurements. We found positive impacts of Chlleaf on modelled photosynthesis for deciduous forests, croplands, grasslands, savannas and wetlands, but mixed impacts for shrublands and evergreen broadleaf forests and negative impacts for evergreen needleleaf forests and mixed forests. Our results highlight the potential of Chlleaf to reduce the uncertainty of global photosynthesis but identify challenges for incorporating Chlleaf in future terrestrial biosphere models.

Rising atmospheric CO2 concentration is a key driver of enhanced global greening, thought to account for up to 70% of increased global vegetation in recent decades. CO2 fertilization effects have further profound implications for ecosystems, food security and biosphere-atmosphere feedbacks. However, it is also possible that current trends will not continue, due to ecosystem level constraints and as plants acclimate to future CO2 concentrations. Future predictions of plant response to rising [CO2 ] are often validated using single-generation short-term FACE (Free Air CO2 Enrichment) experiments but whether this accurately represents vegetation response over decades is unclear. The role of transgenerational plasticity and adaptation in the multigenerational response has yet to be elucidated. Here, we propose that naturally occurring high CO2 springs provide a proxy to quantify the multigenerational and long-term impacts of rising [CO2 ] in herbaceous and woody species respectively, such that plasticity, transgenerational effects and genetic adaptation can be quantified together in these systems. In this first meta-analysis of responses to elevated [CO2 ] at natural CO2 springs, we show that the magnitude and direction of change in eight of nine functional plant traits are consistent between spring and FACE experiments. We found increased photosynthesis (49.8% in spring experiments, comparable to 32.1% in FACE experiments) and leaf starch (58.6% spring, 84.3% FACE), decreased stomatal conductance (gs , 27.2% spring, 21.1% FACE), leaf nitrogen content (6.3% spring, 13.3% FACE) and Specific Leaf Area (SLA, 9.7% spring, 6.0% FACE). These findings not only validate the use of these sites for studying multigenerational plant response to elevated [CO2 ], but additionally suggest that long-term positive photosynthetic response to rising [CO2 ] are likely to continue as predicted by single-generation exposure FACE experiments.

Climate warming can result in both abiotic (e.g., permafrost thaw) and biotic (e.g., microbial functional genes) changes in Arctic tundra. Recent research has incorporated dynamic permafrost thaw in Earth system models (ESMs) and indicates that Arctic tundra could be a significant future carbon (C) source due to the enhanced decomposition of thawed deep soil C. However, warming-induced biotic changes may influence biologically related parameters and the consequent projections in ESMs. How model parameters associated with biotic responses will change under warming and to what extent these changes affect projected C budgets have not been carefully examined. In this study, we synthesized six data sets over 5 years from a soil warming experiment at the Eight Mile Lake, Alaska, into the Terrestrial ECOsystem (TECO) model with a probabilistic inversion approach. The TECO model used multiple soil layers to track dynamics of thawed soil under different treatments. Our results show that warming increased light use efficiency of vegetation photosynthesis but decreased baseline (i.e., environment-corrected) turnover rates of SOC in both the fast and slow pools in comparison with those under control. Moreover, the parameter changes generally amplified over time, suggesting processes of gradual physiological acclimation and functional gene shifts of both plants and microbes. The TECO model predicted that field warming from 2009 to 2013 resulted in cumulative C losses of 224 or 87 g/m2 , respectively, without or with changes in those parameters. Thus, warming-induced parameter changes reduced predicted soil C loss by 61%. Our study suggests that it is critical to incorporate biotic changes in ESMs to improve the model performance in predicting C dynamics in permafrost regions.

Recently, Yang et al. () reported a decrease in solar-induced chlorophyll fluorescence (SIF) during 2015/2016 El Niño event albeit the increase in enhanced vegetation index (EVI). They interpreted the reduced SIF as a signal of reduced ecosystem photosynthesis. However, we argue that the reduced SIF during 2015/2016 is caused by a decreasing trend of SIF due to sensor degradation and the satellite overpass time is critical for drought impact assessment.

Increases in terrestrial water‐use efficiency (WUE) have been reported in many studies, pointing to potential changes in physiological forcing of global carbon and hydrological cycles. However, gains in WUE are of uncertain magnitude over longer (i.e. >10 years) periods of time largely owing to difficulties in accounting for structural and physiological acclimation. 13C signatures (i.e. δ13C) of plant organic matter have long been used to estimate WUE at temporal scales ranging from days to centuries. Mesophyll conductance is a key uncertainty in estimated WUE owing to its influence on diffusion of CO2 to sites of carboxylation. Here we apply new knowledge of mesophyll conductance to 464 δ13C chronologies in tree‐rings of 143 species spanning global biomes. Adjusted for mesophyll conductance, gains in WUE during the 20th century (0.15 ppm year−1) were considerably smaller than those estimated from conventional modelling (0.26 ppm year−1). Across the globe, mean sensitivity of WUE to atmospheric CO2 was 0.15 ppm ppm−1. Ratios of internal‐to‐atmospheric CO2 (on a mole fraction basis; ci/ca) in leaves were mostly constant over time but differed among biomes and plant taxa—highlighting the significance of both plant structure and physiology. Together with synchronized responses in stomatal and mesophyll conductance, our results suggest that ratios of chloroplastic‐to‐atmospheric CO2 (cc/ca) are constrained over time. We conclude that forest WUE may have not increased as much as previously suggested and that projections of future climate forcing via CO2 fertilization may need to be adjusted accordingly. [ABSTRACT FROM AUTHOR]

Tropical forest photosynthesis can decline at high temperatures due to (1) biochemical responses to increasing temperature and (2) stomatal responses to increasing vapor pressure deficit (VPD), which is associated with increasing temperature. It is challenging to disentangle the influence of these two mechanisms on photosynthesis in observations, because temperature and VPD are tightly correlated in tropical forests. Nonetheless, quantifying the relative strength of these two mechanisms is essential for understanding how tropical gross primary production (GPP) will respond to climate change, because increasing atmospheric CO 2 concentration may partially offset VPD-driven stomatal responses, but is not expected to mitigate the effects of temperature-driven biochemical responses. We used two terrestrial biosphere models to quantify how physiological process assumptions (photosynthetic temperature acclimation and plant hydraulic stress) and functional traits (e.g., maximum xylem conductivity) influence the relative strength of modeled temperature versus VPD effects on light-saturated GPP at an Amazonian forest site, a seasonally dry tropical forest site, and an experimental tropical forest mesocosm. By simulating idealized climate change scenarios, we quantified the divergence in GPP predictions under model configurations with stronger VPD effects compared with stronger direct temperature effects. Assumptions consistent with stronger direct temperature effects resulted in larger GPP declines under warming, while assumptions consistent with stronger VPD effects resulted in more resilient GPP under warming. Our findings underscore the importance of quantifying the role of direct temperature and indirect VPD effects for projecting the resilience of tropical forests in the future, and demonstrate that the relative strength of temperature versus VPD effects in models is highly sensitive to plant functional parameters and structural assumptions about photosynthetic temperature acclimation and plant hydraulics., (© 2024 John Wiley & Sons Ltd.)

Tropical and subtropical evergreen broadleaved forests (TEFs) contribute more than one-third of terrestrial gross primary productivity (GPP). However, the continental-scale leaf phenology-photosynthesis nexus over TEFs is still poorly understood to date. This knowledge gap hinders most light use efficiency (LUE) models from accurately simulating the GPP seasonality in TEFs. Leaf age is the crucial plant trait to link the dynamics of leaf phenology with GPP seasonality. Thus, here we incorporated the seasonal leaf area index of different leaf age cohorts into a widely used LUE model (i.e., EC-LUE) and proposed a novel leaf age-dependent LUE model (denoted as LA-LUE model). At the site level, the LA-LUE model (average R 2  = .59, average root-mean-square error [RMSE] = 1.23 gC m -2  day -1 ) performs better than the EC-LUE model in simulating the GPP seasonality across the nine TEFs sites (average R 2  = .18; average RMSE = 1.87 gC m -2  day -1 ). At the continental scale, the monthly GPP estimates from the LA-LUE model are consistent with FLUXCOM GPP data (R 2  = .80; average RMSE = 1.74 gC m -2  day -1 ), and satellite-based GPP data retrieved from the global Orbiting Carbon Observatory-2 (OCO-2) based solar-induced chlorophyll fluorescence (SIF) product (GOSIF) (R 2  = .64; average RMSE = 1.90 gC m -2  day -1 ) and the reconstructed TROPOspheric Monitoring Instrument SIF dataset using machine learning algorithms (RTSIF) (R 2  = .78; average RMSE = 1.88 gC m -2  day -1 ). Typically, the estimated monthly GPP not only successfully represents the unimodal GPP seasonality near the Tropics of Cancer and Capricorn, but also captures well the bimodal GPP seasonality near the Equator. Overall, this study for the first time integrates the leaf age information into the satellite-based LUE model and provides a feasible implementation for mapping the continental-scale GPP seasonality over the entire TEFs., (© 2024 John Wiley & Sons Ltd.)

To bridge the knowledge gap between (a) our (instantaneous-to-seasonal-scale) process understanding of plants and water and (b) our projections of long-term coupled feedbacks between the terrestrial water and carbon cycles, we must uncover what the dominant dynamics are linking fluxes of water and carbon. This study uses the simplest empirical dynamical systems models-two-dimensional linear models-and observation-based data from satellites, eddy covariance towers, weather stations, and machine-learning-derived products to determine the dominant sub-annual timescales coupling carbon uptake and (normalized) evaporation fluxes. We find two dominant modes across the Contiguous United States: (1) a negative correlation timescale on the order of a few days during which landscapes dry after precipitation and plants increase their carbon uptake through photosynthetic upregulation. (2) A slow, seasonal-scale positive covariation through which landscape drying leads to decreased growth and carbon uptake. The slow (positively correlated) process dominates the joint distribution of local water and carbon variables, leading to similar behaviors across space, biomes, and climate regions. We propose that vegetation cover/leaf area variables link this behavior across space, leading to strong emergent spatial patterns of water/carbon coupling in the mean. The spatial pattern of local temporal dynamics-positively sloped tangent lines to a convex long-term mean-state curve-is surprisingly strong, and can serve as a benchmark for coupled Earth System Models. We show that many such models do not represent this emergent mean-state pattern, and hypothesize that this may be due to lack of water-carbon feedbacks at daily scales., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Marine heatwaves (MHWs) are increasing in frequency, duration and intensity, disrupting global marine ecosystems. While most reported impacts have been in tropical areas, New Zealand experienced its strongest and longest MHW in 2022, profoundly affecting marine sponges. Sponges are vital to rocky benthic marine communities, with their abundance influencing ecosystem functioning. This study examines the impact of this MHW on the photosynthetic sponge Cymbastella lamellata in Fiordland, New Zealand. We describe the extent, physiological responses, mortality, microbial community changes and ecological impact of this MHW on C. lamellata. The Fiordland MHW reached a maximum temperature of 4.4°C above average, lasting for 259 days. Bleaching occurred in >90% of the C. lamellata Fiordland population. The population size exceeded 66 million from 5 to 25 m, making this the largest bleaching event of its kind ever recorded. We identified the photosynthetic symbiont as a diatom, and bleached sponges had reduced photosynthetic efficiency. Post-MHW surveys in 2023 found that over 50% of sponges at sampling sites had died but that the remaining sponges had mostly recovered from earlier bleaching. Using a simulated MHW experiment, we found that temperature stress was a driver of necrosis rather than bleaching, despite necrosis only rarely being observed in the field (<2% of sponges). This suggests that bleaching may not be the cause of the mortality directly. We also identified a microbial community shift in surviving sponges, which we propose represents a microbial-mediated adaptive response to MHWs. We also found that C. lamellata are key contributors of dissolved organic carbon to the water column, with their loss likely impacting ecosystem function. We demonstrate the potential for MHWs to disrupt key marine phyla in temperate regions, highlighting how susceptible temperate sponges globally might be to MHWs., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Plants and their symbionts, such as arbuscular mycorrhizal (AM) fungi, are increasingly subjected to various environmental stressors due to climate change, including drought. As a response to drought, plants generally allocate more biomass to roots over shoots, thereby facilitating water uptake. However, whether this biomass allocation shift is modulated by AM fungi remains unknown. Based on 5691 paired observations from 154 plant species, we conducted a meta-analysis to evaluate how AM fungi modulate the responses of plant growth and biomass allocation (e.g., root-to-shoot ratio, R/S) to drought. We found that AM fungi attenuate the negative impact of drought on plant growth, including biomass production, photosynthetic performance and resource (e.g. nutrient and water) uptake. Accordingly, drought significantly increased R/S in non-inoculated plants, but not in plants symbiotic with established AM fungal symbioses. These results suggest that AM fungi promote plant growth and stabilize their R/S through facilitating nutrient and water uptake in plants under drought. Our findings highlight the crucial role of AM fungi in enhancing plant resilience to drought by optimizing resource allocation. This knowledge opens avenues for sustainable agricultural practices that leverage symbiotic relationships for climate adaptation., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Global hydroclimatic variability is increasing with more frequent extreme dry and wet years, severely destabilizing terrestrial ecosystem productivity. However, what regulates the consequence of precipitation extremes on productivity remains unclear. Based on a 9-year field manipulation experiment on the Qinghai-Tibetan Plateau, we found that the responses of gross primary productivity (GPP) to extreme drought and wetness were differentially regulated by nitrogen (N) deposition. Over increasing N deposition, extreme dry events reduced GPP more. Among the 12 biotic and abiotic factors examined, this was mostly explained by the increased plant canopy height and proportion of drought-sensitive species under N deposition, making photosynthesis more sensitive to hydraulic stress. While extreme wet events increased GPP, their effect did not shift over N deposition. These site observations were complemented by a global synthesis derived from the GOSIF GPP dataset, which showed that GPP sensitivity to extreme drought was larger in ecosystems with higher N deposition, but GPP sensitivity to extreme wetness did not change with N deposition. Our findings indicate that intensified hydroclimatic variability would lead to a greater loss of land carbon sinks in the context of increasing N deposition, due to that GPP losses during extreme dry years are more pronounced, yet without a synchronous increase in GPP gains during extreme wet years. The study implies that the conservation and management against climate extremes merit particular attention in ecosystems subject to N deposition., (© 2024 John Wiley & Sons Ltd.)

Vegetation growth is affected by past growth rates and climate variability. However, the impacts of vegetation growth carryover (VGC; biotic) and lagged climatic effects (LCE; abiotic) on tree stem radial growth may be decoupled from photosynthetic capacity, as higher photosynthesis does not always translate into greater growth. To assess the interaction of tree-species level VGC and LCE with ecosystem-scale photosynthetic processes, we utilized tree-ring width (TRW) data for three tree species: Castanopsis eyrei (CE), Castanea henryi (CH, Chinese chinquapin), and Liquidambar formosana (LF, Chinese sweet gum), along with satellite-based data on canopy greenness (EVI, enhanced vegetation index), leaf area index (LAI), and gross primary productivity (GPP). We used vector autoregressive models, impulse response functions, and forecast error variance decomposition to analyze the duration, intensity, and drivers of VGC and of LCE response to precipitation, temperature, and sunshine duration. The results showed that at the tree-species level, VGC in TRW was strongest in the first year, with an average 77% reduction in response intensity by the fourth year. VGC and LCE exhibited species-specific patterns; compared to CE and CH (diffuse-porous species), LF (ring-porous species) exhibited stronger VGC but weaker LCE. For photosynthetic capacity at the ecosystem scale (EVI, LAI, and GPP), VGC and LCE occurred within 96 days. Our study demonstrates that VGC effects play a dominant role in vegetation function and productivity, and that vegetation responses to previous growth states are decoupled from climatic variability. Additionally, we discovered the possibility for tree-ring growth to be decoupled from canopy condition. Investigating VGC and LCE of multiple indicators of vegetation growth at multiple scales has the potential to improve the accuracy of terrestrial global change models., (© 2024 John Wiley & Sons Ltd.)

Stomata play a central role in surface–atmosphere exchange by controlling the flux of water and CO2 between the leaf and the atmosphere. Representation of stomatal conductance (gsw) is therefore an essential component of models that seek to simulate water and CO2 exchange in plants and ecosystems. For given environmental conditions at the leaf surface (CO2 concentration and vapor pressure deficit or relative humidity), models typically assume a linear relationship between gsw and photosynthetic CO2 assimilation (A). However, measurement of leaf‐level gsw response curves to changes in A are rare, particularly in the tropics, resulting in only limited data to evaluate this key assumption. Here, we measured the response of gsw and A to irradiance in six tropical species at different leaf phenological stages. We showed that the relationship between gsw and A was not linear, challenging the key assumption upon which optimality theory is based—that the marginal cost of water gain is constant. Our data showed that increasing A resulted in a small increase in gsw at low irradiance, but a much larger increase at high irradiance. We reformulated the popular Unified Stomatal Optimization (USO) model to account for this phenomenon and to enable consistent estimation of the key conductance parameters g0 and g1. Our modification of the USO model improved the goodness‐of‐fit and reduced bias, enabling robust estimation of conductance parameters at any irradiance. In addition, our modification revealed previously undetectable relationships between the stomatal slope parameter g1 and other leaf traits. We also observed nonlinear behavior between A and gsw in independent data sets that included data collected from attached and detached leaves, and from plants grown at elevated CO2 concentration. We propose that this empirical modification of the USO model can improve the measurement of gsw parameters and the estimation of plant and ecosystem‐scale water and CO2 fluxes. [ABSTRACT FROM AUTHOR]

There is tremendous interspecific variability in O3 sensitivity among C3 species, but variation among C4 species has been less clearly documented. It is also unclear whether stomatal conductance and leaf structure such as leaf mass per area (LMA) determine the variation in sensitivity to O3 across species. In this study, we investigated leaf morphological, chemical, and photosynthetic responses of 22 genotypes of four C4 bioenergy species (switchgrass, sorghum, maize, and miscanthus) to elevated O3 in side‐by‐side field experiments using free‐air O3 concentration enrichment (FACE). The C4 species varied largely in leaf morphology, physiology, and nutrient composition. Elevated O3 did not alter leaf morphology, nutrient content, stomatal conductance, chlorophyll fluorescence, and respiration in most genotypes but reduced net CO2 assimilation in maize and photosynthetic capacity in sorghum and maize. Species with lower LMA and higher stomatal conductance tended to show greater losses in photosynthetic rate and capacity in elevated O3 compared with species with higher LMA and lower stomatal conductance. Stomatal conductance was the strongest determinant of leaf photosynthetic rate and capacity. The response of both area‐ and mass‐based leaf photosynthetic rate and capacity to elevated O3 were not affected by LMA directly but negatively influenced by LMA indirectly through stomatal conductance. These results demonstrate that there is significant variation in O3 sensitivity among C4 species with maize and sorghum showing greater sensitivity of photosynthesis to O3 than switchgrass and miscanthus. Interspecific variation in O3 sensitivity was determined by direct effects of stomatal conductance and indirect effects of LMA. This is the first study to provide a test of unifying theories explaining variation in O3 sensitivity in C4 bioenergy grasses. These findings advance understanding of O3 tolerance in C4 grasses and could aid in optimal placement of diverse C4 bioenergy feedstock across a polluted landscape. [ABSTRACT FROM AUTHOR]

Improving the accuracy of estimates of forest carbon exchange is a central priority for understanding ecosystem response to increased atmospheric CO2 levels and improving carbon cycle modelling. However, the spatially continuous parameterization of photosynthetic capacity (Vcmax) at global scales and appropriate temporal intervals within terrestrial biosphere models (TBMs) remains unresolved. This research investigates the use of biochemical parameters for modelling leaf photosynthetic capacity within a deciduous forest. Particular attention is given to the impacts of seasonality on both leaf biophysical variables and physiological processes, and their interdependent relationships. Four deciduous tree species were sampled across three growing seasons (2013-2015), approximately every 10 days for leaf chlorophyll content (ChlLeaf ) and canopy structure. Leaf nitrogen (NArea ) was also measured during 2014. Leaf photosynthesis was measured during 2014-2015 using a Li-6400 gas-exchange system, with A-Ci curves to model Vcmax. Results showed that seasonality and variations between species resulted in weak relationships between Vcmax normalized to 25°C (Vcmax25) and NArea (R2  = 0.62, P

Ozone (O3 ) damage to leaves can reduce plant photosynthesis, which suggests that declines in ambient O3 concentrations ([O3 ]) in the United States may have helped increase gross primary production (GPP) in recent decades. Here, we assess the effect of long-term changes in ambient [O3 ] using 20 years of observations at Harvard forest. Using artificial neural networks, we found that the effect of the inclusion of [O3 ] as a predictor was slight, and independent of O3 concentrations, which suggests limited high-frequency O3 inhibition of GPP at this site. Simulations with a terrestrial biosphere model, however, suggest an average long-term O3 inhibition of 10.4% for 1992-2011. A decline of [O3 ] over the measurement period resulted in moderate predicted GPP trends of 0.02-0.04 μmol C m-2  s-1  yr-1 , which is negligible relative to the total observed GPP trend of 0.41 μmol C m-2  s-1  yr-1 . A similar conclusion is achieved with the widely used AOT40 metric. Combined, our results suggest that ozone reductions at Harvard forest are unlikely to have had a large impact on the photosynthesis trend over the past 20 years. Such limited effects are mainly related to the slow responses of photosynthesis to changes in [O3 ]. Furthermore, we estimate that 40% of photosynthesis happens in the shade, where stomatal conductance and thus [O3 ] deposition is lower than for sunlit leaves. This portion of GPP remains unaffected by [O3 ], thus helping to buffer the changes of total photosynthesis due to varied [O3 ]. Our analyses suggest that current ozone reductions, although significant, cannot substantially alleviate the damages to forest ecosystems.

Under ongoing global climate change, drought periods are predicted to increase in frequency and intensity in the future. Under these circumstances, it is crucial for tree's survival to recover their restricted functionalities quickly after drought release. To elucidate the recovery of carbon (C) transport rates in c. 70‐year‐old Norway spruce (Picea abies [L.] KARST.) after 5 years of recurrent summer droughts, we conducted a continuous whole‐tree 13C labeling experiment in parallel with watering. We determined the arrival time of current photoassimilates in major C sinks by tracing the 13C label in stem and soil CO2 efflux, and tips of living fine roots. In the first week after watering, aboveground C transport rates (CTR) from crown to trunk base were still 50% lower in previously drought‐stressed trees (0.16 ± 0.01 m h−1) compared to controls (0.30 ± 0.06 m h−1). Conversely, CTR below ground, that is, from the trunk base to soil CO2 efflux were already similar between treatments (c. 0.03 m h−1). Two weeks after watering, aboveground C transport of previously drought‐stressed trees recovered to the level of the controls. Furthermore, regrowth of water‐absorbing fine roots upon watering was supported by faster incorporation of 13C label in previously drought‐stressed (within 12 ± 10 h upon arrival at trunk base) compared to control trees (73 ± 10 h). Thus, the whole‐tree C transport system from the crown to soil CO2 efflux fully recovered within 2 weeks after drought release, and hence showed high resilience to recurrent summer droughts in mature Norway spruce forests. This high resilience of the C transport system is an important prerequisite for the recovery of other tree functionalities and productivity. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

A better understanding of how climate affects growth in tree species is essential for improved predictions of forest dynamics under climate change. Long‐term climate averages (mean climate) drive spatial variations in species' baseline growth rates, whereas deviations from these averages over time (anomalies) can create growth variation around the local baseline. However, the rarity of long‐term tree census data spanning climatic gradients has so far limited our understanding of their respective role, especially in tropical systems. Furthermore, tree growth sensitivity to climate is likely to vary widely among species, and the ecological strategies underlying these differences remain poorly understood. Here, we utilize an exceptional dataset of 49 years of growth data for 509 tree species across 23 tropical rainforest plots along a climatic gradient to examine how multiannual tree growth responds to both climate means and anomalies, and how species' functional traits mediate these growth responses to climate. We show that anomalous increases in atmospheric evaporative demand and solar radiation consistently reduced tree growth. Drier forests and fast‐growing species were more sensitive to water stress anomalies. In addition, species traits related to water use and photosynthesis partly explained differences in growth sensitivity to both climate means and anomalies. Our study demonstrates that both climate means and anomalies shape tree growth in tropical forests and that species traits can provide insights into understanding these demographic responses to climate change, offering a promising way forward to forecast tropical forest dynamics under different climate trajectories. [ABSTRACT FROM AUTHOR]

The Arctic–Boreal Region (ABR) has a large impact on global vegetation–atmosphere interactions and is experiencing markedly greater warming than the rest of the planet, a trend that is projected to continue with anticipated future emissions of CO2. The ABR is a significant source of uncertainty in estimates of carbon uptake in terrestrial biosphere models such that reducing this uncertainty is critical for more accurately estimating global carbon cycling and understanding the response of the region to global change. Process representation and parameterization associated with gross primary productivity (GPP) drives a large amount of this model uncertainty, particularly within the next 50 years, where the response of existing vegetation to climate change will dominate estimates of GPP for the region. Here we review our current understanding and model representation of GPP in northern latitudes, focusing on vegetation composition, phenology, and physiology, and consider how climate change alters these three components. We highlight challenges in the ABR for predicting GPP, but also focus on the unique opportunities for advancing knowledge and model representation, particularly through the combination of remote sensing and traditional boots‐on‐the‐ground science. [ABSTRACT FROM AUTHOR]

Changes in peak photosynthesis timing (PPT) could substantially change the seasonality of the terrestrial carbon cycle. Spring PPT in dry regions has been documented for some individual plant species on a stand scale, but both the spatio-temporal pattern of shifting PPT on a continental scale and its determinants remain unclear. Here, we use satellite measurements of vegetation greenness to find that the majority of Northern Hemisphere, mid-latitude vegetated area experienced a trend toward earlier PPT during 1982-2012, with significant trends of an average of 0.61 day yr(-1) across 19.4% of areas. These shifts correspond to increased annual accumulation of growing degree days (GDD) due to warming and are most highly concentrated in the eastern United States and Europe. Earlier mean PPT is generally a trait common among areas with summer temperatures higher than 27.6 ± 2.9 °C, summer precipitation lower than 84.2 ± 41.5 mm, and fraction of cold season precipitation greater than 89.2 ± 1.5%. The trends toward earlier PPT discovered here have co-occurred with overall increases in vegetation greenness throughout the growing season, suggesting that summer drought is not a dominant driver of these trends. These results imply that continued warming may facilitate continued shifts toward earlier PPT and cause these trends to become more pervasive, with important implications for terrestrial carbon, water, nutrient, and energy budgets.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

The integrity of coral reefs worldwide is jeopardized by ocean acidification (OA). Most studies conducted so far have focused on the vulnerability to OA of corals inhabiting shallow reefs while nothing is currently known about the response of mesophotic scleractinian corals. In this study, we assessed the susceptibility to OA of corals, together with their algal partners, inhabiting a wide depth range. We exposed fragments of the depth generalist coral Stylophora pistillata collected from either 5 or 45 m to simulated future OA conditions, and assessed key molecular, physiological and photosynthetic processes influenced by the lowered pH. Our comparative analysis reveals that mesophotic and shallow S. pistillata corals are genetically distinct and possess different symbiont types. Under the exposure to acidification conditions, we observed a 50% drop of metabolic rate in shallow corals, whereas mesophotic corals were able to maintain unaltered metabolic rates. Overall, our gene expression and physiological analyses show that mesophotic corals possess a greater capacity to cope with the effects of OA compared to their shallow counterparts. Such capability stems from physiological characteristics (i.e., biomass and lipids energetics), a greater capacity to regulate cellular acid–base parameters, and a higher baseline expression of cell adhesion and extracellular matrix genes. Moreover, our gene expression analysis suggests that the enhanced symbiont photochemical efficiency under high pCO2 levels could prevent acidosis of the host cells and it could support a greater translocation of photosynthates, increasing the energy pool available to the host. With this work, we provide new insights on the response to OA of corals living at mesophotic depths. Our investigation discloses key genetic and physiological traits underlying the potential for corals to cope with future OA conditions. [ABSTRACT FROM AUTHOR]

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 (ToptA) did not significantly increase in warm‐grown trees in either species. Similarly, the thermal optima (ToptV and ToptJ) and activation energies (EaV and EaJ) of maximum Rubisco carboxylation capacity (Vcmax) and maximum electron transport rate (Jmax) were largely unaffected by warming. However, Vcmax, Jmax and foliar dark respiration (Rd) 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 Jmax/Vcmax 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. [ABSTRACT FROM AUTHOR]

Global climate change intensifies the water cycle and makes freshest waters become fresher and vice-versa. But how this change impacts phytoplankton in coastal, particularly harmful algal blooms (HABs), remains poorly understood. Here, we monitored a coastal bay for a decade and found a significant correlation between salinity decline and the increase of Karenia mikimotoi blooms. To examine the physiological linkage between salinity decreases and K. mikimotoi blooms, we compare chemical, physiological and multi-omic profiles of this species in laboratory cultures under high (33) and low (25) salinities. Under low salinity, photosynthetic efficiency and capacity as well as growth rate and cellular protein content were significantly higher than that under high salinity. More strikingly, the omics data show that low salinity activated the glyoxylate shunt to bypass the decarboxylation reaction in the tricarboxylic acid cycle, hence redirecting carbon from CO 2 release to biosynthesis. Furthermore, the enhanced glyoxylate cycle could promote hydrogen peroxide metabolism, consistent with the detected decrease in reactive oxygen species. These findings suggest that salinity declines can reprogram metabolism to enhance cell proliferation, thus promoting bloom formation in HAB species like K. mikimotoi, which has important ecological implications for future climate-driven salinity declines in the coastal ocean with respect to HAB outbreaks., (© 2024 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. [ABSTRACT FROM AUTHOR]

In a rapidly warming world, we ask, "What limits the potential of marine diatoms to acclimate to elevated temperatures?," a group of ecologically successful unicellular eukaryotic photoautotrophs that evolved in a cooler ocean and are critical to marine food webs. To this end, we examined thermal tolerance mechanisms related to photosynthesis in the sequenced and transformable model diatom Phaeodactylum tricornutum. Data from transmission electron microscopy (TEM) and fatty acid methyl ester‐gas chromatography mass spectrometry (FAME‐GCMS) suggest that saturating thylakoid‐associated fatty acids allowed rapid (on the order of hours) thermal tolerance up to 28.5°C. Beyond this critical temperature, thylakoid ultrastructure became severely perturbed. Biophysical analyses revealed that electrochemical leakage through the thylakoid membranes was extremely sensitive to elevated temperature (Q10 of 3.5). Data suggest that the loss of the proton motive force (pmf) occurred even when heat‐labile photosystem II (PSII) was functioning, and saturation of thylakoid‐associated fatty acids was active. Indeed, growth was inhibited when leakage of pmf through thylakoid membranes was insufficiently compensated by proton input from PSII. Our findings provide a mechanistic understanding of the importance of rapid saturation of thylakoid‐associated fatty acids for ultrastructure maintenance and a generation of pmf at elevated temperatures. To the extent these experimental results apply, the ability of diatoms to generate a pmf may be a sensitive parameter for thermal sensitivity diagnosis in phytoplankton. [ABSTRACT FROM AUTHOR]