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The 2017–2027 National Academies' Decadal Survey, Thriving on Our Changing Planet, recommended Surface Biology and Geology (SBG) as a “Designated Targeted Observable” (DO). The SBG DO is based on the need for capabilities to acquire global, high spatial resolution, visible to shortwave infrared (VSWIR; 380–2500 nm; ~30 m pixel resolution) hyperspectral (imaging spectroscopy) and multispectral midwave and thermal infrared (MWIR: 3–5 μm; TIR: 8–12 μm; ~60 m pixel resolution) measurements with sub-monthly temporal revisits over terrestrial, freshwater, and coastal marine habitats. To address the various mission design needs, an SBG Algorithms Working Group of multidisciplinary researchers has been formed to review and evaluate the algorithms applicable to the SBG DO across a wide range of Earth science disciplines, including terrestrial and aquatic ecology, atmospheric science, geology, and hydrology. Here, we summarize current state-of-the-practice VSWIR and TIR algorithms that use airborne or orbital spectral imaging observations to address the SBG DO priorities identified by the Decadal Survey: (i) terrestrial vegetation physiology, functional traits, and health; (ii) inland and coastal aquatic ecosystems physiology, functional traits, and health; (iii) snow and ice accumulation, melting, and albedo; (iv) active surface composition (eruptions, landslides, evolving landscapes, hazard risks); (v) effects of changing land use on surface energy, water, momentum, and carbon fluxes; and (vi) managing agriculture, natural habitats, water use/quality, and urban development. We review existing algorithms in the following categories: snow/ice, aquatic environments, geology, and terrestrial vegetation, and summarize the community-state-of-practice in each category. This effort synthesizes the findings of more than 130 scientists.

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.

Plant functional traits determine vegetation responses to environmental variation, but variation in trait values is large, even within a single site. Likewise, uncertainty in how these traits map to Earth system feedbacks is large. We use a vegetation demographic model (VDM), the Functionally Assembled Terrestrial Ecosystem Simulator (FATES), to explore parameter sensitivity of model predictions, and comparison to observations, at a tropical forest site: Barro Colorado Island in Panama. We define a single 12-dimensional distribution of plant trait variation, derived primarily from observations in Panama, and define plant functional types (PFTs) as random draws from this distribution. We compare several model ensembles, where individual ensemble members vary only in the plant traits that define PFTs, and separate ensembles differ from each other based on either model structural assumptions or non-trait, ecosystem-level parameters, which include (a) the number of competing PFTs present in any simulation and (b) parameters that govern disturbance and height-based light competition. While singlePFT simulations are roughly consistent with observations of productivity at Barro Colorado Island, increasing the number of competing PFTs strongly shifts model predictions towards higher productivity and biomass forests. Different ecosystem variables show greater sensitivity than others to the number of competing PFTs, with the predictions that are most dominated by large trees, such as biomass, being the most sensitive. Changing disturbance and height-sorting parameters, i.e., the rules of competitive trait filtering, shifts regimes of dominance or coexistence between early-and late-successional PFTs in the model. Increases to the extent or severity of disturbance, or to the degree of determinism in height-based light competition, all act to shift the community towards early-successional PFTs. In turn, these shifts in competitive outcomes alter predictions of ecosystem states and fluxes, with more early-successional-dominated forests having lower biomass. It is thus crucial to differentiate between plant traits, which are under competitive pressure in VDMs, from those model parameters that are not and to better understand the relationships between these two types of model parameters to quantify sources of uncertainty in VDMs.

Vegetation plays an important role in regulating global carbon cycles and is a key component of the Earth system models (ESMs) that aim to project Earth's future climate. In the last decade, the vegetation component within ESMs has witnessed great progress from simple "big-leaf" approaches to demographically structured approaches, which have a better representation of plant size, canopy structure, and disturbances. These demographically structured vegetation models typically have a large number of input parameters, and sensitivity analysis is needed to quantify the impact of each parameter on the model outputs for a better understanding of model behavior. In this study, we conducted a comprehensive sensitivity analysis to diagnose the Community Land Model coupled to the Functionally Assembled Terrestrial Simulator, or CLM4.5(FATES). Specifically, we quantified the first- and second-order sensitivities of the model parameters to outputs that represent simulated growth and mortality as well as carbon fluxes and stocks for a tropical site with an extent of 1×1°. While the photosynthetic capacity parameter (Vc;max25) is found to be important for simulated carbon stocks and fluxes, we also show the importance of carbon storage and allometry parameters, which determine survival and growth strategies within the model. The parameter sensitivity changes with different sizes of trees and climate conditions. The results of this study highlight the importance of understanding the dynamics of the next generation of demographically enabled vegetation models within ESMs to improve model parameterization and structure for better model fidelity.

Leaf dark respiration (R dark ), an important yet rarely quantified component of carbon cycling in forest ecosystems, is often simulated from leaf traits such as the maximum carboxylation capacity (V cmax ), leaf mass per area (LMA), nitrogen (N) and phosphorus (P) concentrations, in terrestrial biosphere models. However, the validity of these relationships across forest types remains to be thoroughly assessed. Here, we analyzed R dark variability and its associations with V cmax and other leaf traits across three temperate, subtropical and tropical forests in China, evaluating the effectiveness of leaf spectroscopy as a superior monitoring alternative. We found that leaf magnesium and calcium concentrations were more significant in explaining cross-site R dark than commonly used traits like LMA, N and P concentrations, but univariate trait-R dark relationships were always weak (r 2  ≤ 0.15) and forest-specific. Although multivariate relationships of leaf traits improved the model performance, leaf spectroscopy outperformed trait-R dark relationships, accurately predicted cross-site R dark (r 2  = 0.65) and pinpointed the factors contributing to R dark variability. Our findings reveal a few novel traits with greater cross-site scalability regarding R dark , challenging the use of empirical trait-R dark relationships in process models and emphasize the potential of leaf spectroscopy as a promising alternative for estimating R dark , which could ultimately improve process modeling of terrestrial plant respiration., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

The advent of new spaceborne imaging spectrometers offers new opportunities for ecologists to map vegetation traits at global scales. However, to date most imaging spectroscopy studies exploiting satellite spectrometers have been constrained to the landscape scale. In this paper we present a new method to map vegetation traits at the landscape scale and upscale trait maps to the continental level, using historical spaceborne imaging spectroscopy (Hyperion) to derive estimates of leaf mass per area, nitrogen, and carbon concentrations of forests in Québec, Canada. We compare estimates for each species with reference field values and obtain good agreement both at the landscape and continental scales, with patterns consistent with the leaf economic spectrum. By exploiting the Hyperion satellite archive to map these traits and successfully upscale the estimates to the continental scale, we demonstrate the great potential of recent and upcoming spaceborne spectrometers to benefit plant biodiversity monitoring and conservation efforts., (© 2023. Springer Nature Limited.)

Vegetative transpiration (E) and photosynthetic carbon assimilation (A) are known to be seasonally dynamic, with changes in their ratio determining the marginal water use efficiency (WUE). Despite an understanding that stomata play a mechanistic role in regulating WUE, it is still unclear how stomatal and nonstomatal processes influence change in WUE over the course of the growing season. As a result, limited understanding of the primary physiological drivers of seasonal dynamics of canopy WUE remains one of the largest uncertainties in earth system model projections of carbon and water exchange in temperate deciduous forest ecosystems. We investigated seasonal patterns in leaf-level physiological, hydraulic, and anatomical properties, including the seasonal progress of the stomatal slope parameter (g 1 ; inversely proportional to WUE) and the maximum carboxylation rate (V cmax ). V cmax and g 1 were seasonally variable; however, their patterns were not temporally synchronized. g 1 generally showed an increasing trend until late in the season, while V cmax peaked during the midsummer months. Seasonal progression of V cmax was primarily driven by changes in leaf structural, and anatomical characteristics, while seasonal changes in g 1 were most strongly related to changes in V cmax and leaf hydraulics. Using a seasonally variable V cmax and g 1 to parameterize a canopy-scale gas exchange model increased seasonally aggregated A and E by 3% and 16%, respectively., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.)

Terrestrial biosphere models (TBMs) include the representation of vertical gradients in leaf traits associated with modeling photosynthesis, respiration, and stomatal conductance. However, model assumptions associated with these gradients have not been tested in complex tropical forest canopies. We compared TBM representation of the vertical gradients of key leaf traits with measurements made in a tropical forest in Panama and then quantified the impact of the observed gradients on simulated canopy-scale CO 2 and water fluxes. Comparison between observed and TBM trait gradients showed divergence that impacted canopy-scale simulations of water vapor and CO 2 exchange. Notably, the ratio between the dark respiration rate and the maximum carboxylation rate was lower near the ground than at the top-of-canopy, leaf-level water-use efficiency was markedly higher at the top-of-canopy, and the decrease in maximum carboxylation rate from the top-of-canopy to the ground was less than TBM assumptions. The representation of the gradients of leaf traits in TBMs is typically derived from measurements made within-individual plants, or, for some traits, assumed constant due to a lack of experimental data. Our work shows that these assumptions are not representative of the trait gradients observed in species-rich, complex tropical forests., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.)

Wildfires are a global crisis, but current fire models fail to capture vegetation response to changing climate. With drought and elevated temperature increasing the importance of vegetation dynamics to fire behavior, and the advent of next generation models capable of capturing increasingly complex physical processes, we provide a renewed focus on representation of woody vegetation in fire models. Currently, the most advanced representations of fire behavior and biophysical fire effects are found in distinct classes of fine-scale models and do not capture variation in live fuel (i.e. living plant) properties. We demonstrate that plant water and carbon dynamics, which influence combustion and heat transfer into the plant and often dictate plant survival, provide the mechanistic linkage between fire behavior and effects. Our conceptual framework linking remotely sensed estimates of plant water and carbon to fine-scale models of fire behavior and effects could be a critical first step toward improving the fidelity of the coarse scale models that are now relied upon for global fire forecasting. This process-based approach will be essential to capturing the influence of physiological responses to drought and warming on live fuel conditions, strengthening the science needed to guide fire managers in an uncertain future., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Within vascular plants, the partitioning of hydraulic resistance along the soil-to-leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (R leaf ) to total soil-to-leaf hydraulic resistance (R total ), or fR leaf (=R leaf /R total ), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fR leaf using new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fR leaf averaged 0.51 (95% confidence interval [CI] = 0.46-0.57) and it declined with tree height. We also used the allometric relationship between field-based measurements of soil-to-leaf hydraulic conductance and laboratory-based measurements of leaf hydraulic conductance to compute the average fR leaf for 19 tree samples, which was 0.40 (95% CI = 0.29-0.56). The in situ technique produces a more accurate descriptor of fR leaf because it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fR leaf may help stems from loss of hydraulic conductance. Thus, the decline in fR leaf with tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality., (© 2022 John Wiley & Sons Ltd.)

The representation of stomatal regulation of transpiration and CO 2 assimilation is key to forecasting terrestrial ecosystem responses to global change. Given its importance in determining the relationship between forest productivity and climate, accurate and mechanistic model representation of the relationship between stomatal conductance (g s ) and assimilation is crucial. We assess possible physiological and mechanistic controls on the estimation of the g 1 (stomatal slope, inversely proportional to water use efficiency) and g 0 (stomatal intercept) parameters, using diurnal gas exchange surveys and leaf-level response curves of six tropical broadleaf evergreen tree species. g 1 estimated from ex situ response curves averaged 50% less than g 1 estimated from survey data. While g 0 and g 1 varied between leaves of different phenological stages, the trend was not consistent among species. We identified a diurnal trend associated with g 1 and g 0 that significantly improved model projections of diurnal trends in transpiration. The accuracy of modeled g s can be improved by accounting for variation in stomatal behavior across diurnal periods, and between measurement approaches, rather than focusing on phenological variation in stomatal behavior. Additional investigation into the primary mechanisms responsible for diurnal variation in g 1 will be required to account for this phenomenon in land-surface models., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Many terrestrial biosphere models depend on an understanding of the relationship between stomatal conductance and photosynthesis. However, unlike the measurement of photosynthetic parameters, such as the maximum carboxylation capacity, where standard methods (e.g., CO2 response or ACi curves) are widely accepted, a consensus method for empirically measuring parameters representing stomatal response has not yet emerged. Most models of stomatal response to environment represent stomatal conductance as being bounded by a lower intercept parameter (g0), and linearly scaled based on a multivariate term described by the stomatal slope parameter (g1). Here we employ the widely used Unified Stomatal Optimization model, to test whether g1 and g0 parameters are impacted by the choice of measurement method, either on an intact branch or a cut branch segment stored in water. We measured paired stomatal response curves on intact and excised branches of a hybrid poplar clone (Populus deltoides Bartr. × Populus nigra L. OP367), measured twice over a diurnal period. We found that predawn branch excision did not significantly affect measured g0 and g1 when measured within 4 h of excision. Measurement in the afternoon resulted in significantly higher values of g1 and lower values of g0, with values changing by 55% and 56%, respectively. Excision combined with afternoon measurement resulted in a marked effect on parameter estimates, with g1 increasing 89% from morning to afternoon and a 25% lower g1 for cut branches than those measured in situ. We also show that in hybrid poplar the differences in parameter estimates obtained from plants measured under different conditions can directly impact models of canopy function, reducing modeled transpiration by 18% over a simulated 12.5-h period. Although these results are only for a single isohydric woody species, our findings suggest that stomatal optimality parameters may not remain constant throughout the day., (Published by Oxford University Press 2022.)

Stomata play a central role in surface-atmosphere exchange by controlling the flux of water and CO 2 between the leaf and the atmosphere. Representation of stomatal conductance (g sw ) is therefore an essential component of models that seek to simulate water and CO 2 exchange in plants and ecosystems. For given environmental conditions at the leaf surface (CO 2 concentration and vapor pressure deficit or relative humidity), models typically assume a linear relationship between g sw and photosynthetic CO 2 assimilation (A). However, measurement of leaf-level g sw 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 g sw and A to irradiance in six tropical species at different leaf phenological stages. We showed that the relationship between g sw 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 g sw 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 g 0 and g 1 . 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 g 1 and other leaf traits. We also observed nonlinear behavior between A and g sw in independent data sets that included data collected from attached and detached leaves, and from plants grown at elevated CO 2 concentration. We propose that this empirical modification of the USO model can improve the measurement of g sw parameters and the estimation of plant and ecosystem-scale water and CO 2  fluxes., (© 2022 John Wiley & Sons Ltd.)

Photosynthesis is a key target to improve crop production in many species including soybean [Glycine max (L.) Merr.]. A challenge is that phenotyping photosynthetic traits by traditional approaches is slow and destructive. There is proof-of-concept for leaf hyperspectral reflectance as a rapid method to model photosynthetic traits. However, the crucial step of demonstrating that hyperspectral approaches can be used to advance understanding of the genetic architecture of photosynthetic traits is untested. To address this challenge, we used full-range (500-2,400 nm) leaf reflectance spectroscopy to build partial least squares regression models to estimate leaf traits, including the rate-limiting processes of photosynthesis, maximum Rubisco carboxylation rate, and maximum electron transport. In total, 11 models were produced from a diverse population of soybean sampled over multiple field seasons to estimate photosynthetic parameters, chlorophyll content, leaf carbon and leaf nitrogen percentage, and specific leaf area (with R2 from 0.56 to 0.96 and root mean square error approximately <10% of the range of calibration data). We explore the utility of these models by applying them to the soybean nested association mapping population, which showed variability in photosynthetic and leaf traits. Genetic mapping provided insights into the underlying genetic architecture of photosynthetic traits and potential improvement in soybean. Notably, the maximum Rubisco carboxylation rate mapped to a region of chromosome 19 containing genes encoding multiple small subunits of Rubisco. We also mapped the maximum electron transport rate to a region of chromosome 10 containing a fructose 1,6-bisphosphatase gene, encoding an important enzyme in the regeneration of ribulose 1,5-bisphosphate and the sucrose biosynthetic pathway. The estimated rate-limiting steps of photosynthesis were low or negatively correlated with yield suggesting that these traits are not influenced by the same genetic mechanisms and are not limiting yield in the soybean NAM population. Leaf carbon percentage, leaf nitrogen percentage, and specific leaf area showed strong correlations with yield and may be of interest in breeding programs as a proxy for yield. This work is among the first to use hyperspectral reflectance to model and map the genetic architecture of the rate-limiting steps of photosynthesis., (Published by Oxford University Press on behalf of Genetics Society of America 2022. This work is written by US Government employees and is in the public domain in the US.)

As the Arctic region moves into uncharted territory under a warming climate, it is important to refine the terrestrial biosphere models (TBMs) that help us understand and predict change. One fundamental uncertainty in TBMs relates to model parameters, configuration variables internal to the model whose value can be estimated from data. We incorporate a version of the Terrestrial Ecosystem Model (TEM) developed for arctic ecosystems into the Predictive Ecosystem Analyzer (PEcAn) framework. PEcAn treats model parameters as probability distributions, estimates parameters based on a synthesis of available field data, and then quantifies both model sensitivity and uncertainty to a given parameter or suite of parameters. We examined how variation in 21 parameters in the equation for gross primary production influenced model sensitivity and uncertainty in terms of two carbon fluxes (net primary productivity and heterotrophic respiration) and two carbon (C) pools (vegetation C and soil C). We set up different parameterizations of TEM across a range of tundra types (tussock tundra, heath tundra, wet sedge tundra, and shrub tundra) in northern Alaska, along a latitudinal transect extending from the coastal plain near Utqiaġvik to the southern foothills of the Brooks Range, to the Seward Peninsula. TEM was most sensitive to parameters related to the temperature regulation of photosynthesis. Model uncertainty was mostly due to parameters related to leaf area, temperature regulation of photosynthesis, and the stomatal responses to ambient light conditions. Our analysis also showed that sensitivity and uncertainty to a given parameter varied spatially. At some sites, model sensitivity and uncertainty tended to be connected to a wider range of parameters, underlining the importance of assessing tundra community processes across environmental gradients or geographic locations. Generally, across sites, the flux of net primary productivity (NPP) and pool of vegetation C had about equal uncertainty, while heterotrophic respiration had higher uncertainty than the pool of soil C. Our study illustrates the complexity inherent in evaluating parameter uncertainty across highly heterogeneous arctic tundra plant communities. It also provides a framework for iteratively testing how newly collected field data related to key parameters may result in more effective forecasting of Arctic change., (© 2021 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

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 CO 2 . 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., (© 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 forests are one of the main carbon sinks on Earth, but the magnitude of CO2 absorbed by tropical vegetation remains uncertain. Terrestrial biosphere models (TBMs) are commonly used to estimate the CO2 absorbed by forests, but their performance is highly sensitive to the parameterization of processes that control leaf-level CO2 exchange. Direct measurements of leaf respiratory and photosynthetic traits that determine vegetation CO2 fluxes are critical, but traditional approaches are time-consuming. Reflectance spectroscopy can be a viable alternative for the estimation of these traits and, because data collection is markedly quicker than traditional gas exchange, the approach can enable the rapid assembly of large datasets. However, the application of spectroscopy to estimate photosynthetic traits across a wide range of tropical species, leaf ages and light environments has not been extensively studied. Here, we used leaf reflectance spectroscopy together with partial least-squares regression (PLSR) modeling to estimate leaf respiration (Rdark25), the maximum rate of carboxylation by the enzyme Rubisco (Vcmax25), the maximum rate of electron transport (Jmax25), and the triose phosphate utilization rate (Tp25), all normalized to 25°C. We collected data from three tropical forest sites and included leaves from fifty-three species sampled at different leaf phenological stages and different leaf light environments. Our resulting spectra-trait models validated on randomly sampled data showed good predictive performance for Vcmax25, Jmax25, Tp25 and Rdark25 (RMSE of 13, 20, 1.5 and 0.3 μmol m-2 s-1, and R2 of 0.74, 0.73, 0.64 and 0.58, respectively). The models showed similar performance when applied to leaves of species not included in the training dataset, illustrating that the approach is robust for capturing the main axes of trait variation in tropical species. We discuss the utility of the spectra-trait and traditional gas exchange approaches for enhancing tropical plant trait studies and improving the parameterization of TBMs., Competing Interests: The authors have declared that no competing interests exist.

Leaf trait relationships are widely used to predict ecosystem function in terrestrial biosphere models (TBMs), in which leaf maximum carboxylation capacity (V c,max ), an important trait for modelling photosynthesis, can be inferred from other easier-to-measure traits. However, whether trait-V c,max relationships are robust across different forest types remains unclear. Here we used measurements of leaf traits, including one morphological trait (leaf mass per area), three biochemical traits (leaf water content, area-based leaf nitrogen content, and leaf chlorophyll content), one physiological trait (V c,max ), as well as leaf reflectance spectra, and explored their relationships within and across three contrasting forest types in China. We found weak and forest type-specific relationships between V c,max and the four morphological and biochemical traits (R 2  ≤ 0.15), indicated by significantly changing slopes and intercepts across forest types. By contrast, reflectance spectroscopy effectively collapsed the differences in the trait-V c,max relationships across three forest biomes into a single robust model for V c,max (R 2  = 0.77), and also accurately estimated the four traits (R 2  = 0.75-0.94). These findings challenge the traditional use of the empirical trait-V c,max relationships in TBMs for estimating terrestrial plant photosynthesis, but also highlight spectroscopy as an efficient alternative for characterising V c,max and multitrait variability, with critical insights into ecosystem modelling and functional trait ecology., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Drought is the most important limitation on crop yield. Understanding and detecting drought stress in crops is vital for improving water use efficiency through effective breeding and management. Leaf reflectance spectroscopy offers a rapid, non-destructive alternative to traditional techniques for measuring plant traits involved in a drought response. We measured drought stress in six glasshouse-grown agronomic species using physiological, biochemical, and spectral data. In contrast to physiological traits, leaf metabolite concentrations revealed drought stress before it was visible to the naked eye. We used full-spectrum leaf reflectance data to predict metabolite concentrations using partial least-squares regression, with validation R2 values of 0.49-0.87. We show for the first time that spectroscopy may be used for the quantitative estimation of proline and abscisic acid, demonstrating the first use of hyperspectral data to detect a phytohormone. We used linear discriminant analysis and partial least squares discriminant analysis to differentiate between watered plants and those subjected to drought based on measured traits (accuracy: 71%) and raw spectral data (66%). Finally, we validated our glasshouse-developed models in an independent field trial. We demonstrate that spectroscopy can detect drought stress via underlying biochemical changes, before visual differences occur, representing a powerful advance for measuring limitations on yield., (Published by Oxford University Press on behalf of the Society for Experimental Biology 2021.)

Partial least squares regression (PLSR) modelling is a statistical technique for correlating datasets, and involves the fitting of a linear regression between two matrices. One application of PLSR enables leaf traits to be estimated from hyperspectral optical reflectance data, facilitating rapid, high-throughput, non-destructive plant phenotyping. This technique is of interest and importance in a wide range of contexts including crop breeding and ecosystem monitoring. The lack of a consensus in the literature on how to perform PLSR means that interpreting model results can be challenging, applying existing models to novel datasets can be impossible, and unknown or undisclosed assumptions can lead to incorrect or spurious predictions. We address this lack of consensus by proposing best practices for using PLSR to predict plant traits from leaf-level hyperspectral data, including a discussion of when PLSR is applicable, and recommendations for data collection. We provide a tutorial to demonstrate how to develop a PLSR model, in the form of an R script accompanying this manuscript. This practical guide will assist all those interpreting and using PLSR models to predict leaf traits from spectral data, and advocates for a unified approach to using PLSR for predicting traits from spectra in the plant sciences., (Published by Oxford University Press on behalf of the Society for Experimental Biology 2021.)

Understanding seasonal variation in photosynthesis is important for understanding and modeling plant productivity. Here, we used shotgun sampling to examine physiological, structural and spectral leaf traits of upper canopy, sun-exposed leaves in Quercus coccinea Münchh (scarlet oak) across the growing season in order to understand seasonal trends, explore the mechanisms underpinning physiological change and investigate the impact of extrapolating measurements from a single date to the whole season. We tested the hypothesis that photosynthetic rates and capacities would peak at the summer solstice, i.e., at the time of peak photoperiod. Contrary to expectations, our results reveal a late-season peak in both photosynthetic capacity and rate before the expected sharp decrease at the start of senescence. This late-season maximum occurred after the higher summer temperatures and vapor pressure deficit and was correlated with the recovery of leaf water content and increased stomatal conductance. We modeled photosynthesis at the top of the canopy and found that the simulated results closely tracked the maximum carboxylation capacity of Rubisco. For both photosynthetic capacity and modeled top-of-canopy photosynthesis, the maximum value was therefore not observed at the summer solstice. Rather, in each case, the measurements at and around the solstice were close to the overall seasonal mean, with values later in the season leading to deviations from the mean by up to 41 and 52%, respectively. Overall, we found that the expected Gaussian pattern of photosynthesis was not observed. We conclude that an understanding of species- and environment-specific changes in photosynthesis across the season is essential for correct estimation of seasonal photosynthetic capacity., (Published by Oxford University Press 2021.)

The finely tuned balance between sources and sinks determines plant resource partitioning and regulates growth and development. Understanding and measuring metabolic indicators of source or sink limitation forms a vital part of global efforts to increase crop yield for future food security. We measured metabolic profiles of Cucurbita pepo (zucchini) grown in the field under carbon sink limitation and control conditions. We demonstrate that these profiles can be measured non-destructively using hyperspectral reflectance at both leaf and canopy scales. Total non-structural carbohydrates (TNC) increased 82% in sink-limited plants; leaf mass per unit area (LMA) increased 38% and free amino acids increased 22%. Partial least-squares regression (PLSR) models link these measured functional traits with reflectance data, enabling high-throughput estimation of traits comprising the sink limitation response. Leaf- and canopy-scale models for TNC had R 2 values of 0.93 and 0.64 and %RMSE of 13 and 38%, respectively. For LMA, R 2 values were 0.91 and 0.60 and %RMSE 7 and 14%; for free amino acids, R 2 was 0.53 and 0.21 with %RMSE 20 and 26%. Remote sensing can enable accurate, rapid detection of sink limitation in the field at the leaf and canopy scale, greatly expanding our ability to understand and measure metabolic responses to stress., (Published 2021. This article is a U.S. Government work and is in the public domain in the USA.)

Plant functional diversity is strongly connected to photosynthetic carbon assimilation in terrestrial ecosystems. However, many of the plant functional traits that regulate photosynthetic capacity, including foliar nitrogen concentration and leaf mass per area, vary significantly between and within plant functional types and vertically through forest canopies, resulting in considerable landscape-scale heterogeneity in three dimensions. Hyperspectral imagery has been used extensively to quantify functional traits across a range of ecosystems but is generally limited to providing information for top of canopy leaves only. On the other hand, lidar data can be used to retrieve the vertical structure of forest canopies. Because these data are rarely collected at the same time, there are unanswered questions about the effect of forest structure on the three -dimensional spatial patterns of functional traits across ecosystems. In the United States, the National Ecological Observatory Network's Airborne Observation Platform (NEON AOP) provides an opportunity to address this structure-function relationship by collecting lidar and hyperspectral data together across a variety of ecoregions. With a fusion of hyperspectral and lidar data from the NEON AOP and field-collected foliar trait data, we assessed the impacts of forest structure on spatial patterns of N. In addition, we examine the influence of abiotic gradients and management regimes on top-of-canopy percent N and total canopy N (i.e., the total amount of N [g/m 2 ] within a forest canopy) at a NEON site consisting of a mosaic of open longleaf pine and dense broadleaf deciduous forests. Our resulting maps suggest that, in contrast to top of canopy values, total canopy N variation is dampened across this landscape resulting in relatively homogeneous spatial patterns. At the same time, we found that leaf functional diversity and canopy structural diversity showed distinct dendritic patterns related to the spatial distribution of plant functional types., (© 2020 by the Ecological Society of America.)

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 CO 2 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., (© 2020 UT-Battelle, LLC. Global Change Biology published by John Wiley & Sons Ltd.)

In an era of rapid global change, our ability to understand and predict Earth's natural systems is lagging behind our ability to monitor and measure changes in the biosphere. Bottlenecks to informing models with observations have reduced our capacity to fully exploit the growing volume and variety of available data. Here, we take a critical look at the information infrastructure that connects ecosystem modeling and measurement efforts, and propose a roadmap to community cyberinfrastructure development that can reduce the divisions between empirical research and modeling and accelerate the pace of discovery. A new era of data-model integration requires investment in accessible, scalable, and transparent tools that integrate the expertise of the whole community, including both modelers and empiricists. This roadmap focuses on five key opportunities for community tools: the underlying foundations of community cyberinfrastructure; data ingest; calibration of models to data; model-data benchmarking; and data assimilation and ecological forecasting. This community-driven approach is a key to meeting the pressing needs of science and society in the 21st century., (© 2020 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Stomata regulate CO 2 uptake for photosynthesis and water loss through transpiration. The approaches used to represent stomatal conductance (g s ) in models vary. In particular, current understanding of drivers of the variation in a key parameter in those models, the slope parameter (i.e. a measure of intrinsic plant water-use-efficiency), is still limited, particularly in the tropics. Here we collected diurnal measurements of leaf gas exchange and leaf water potential (Ψ leaf ), and a suite of plant traits from the upper canopy of 15 tropical trees in two contrasting Panamanian forests throughout the dry season of the 2016 El Niño. The plant traits included wood density, leaf-mass-per-area (LMA), leaf carboxylation capacity (V c,max ), leaf water content, the degree of isohydry, and predawn Ψ leaf . We first investigated how the choice of four commonly used leaf-level g s models with and without the inclusion of Ψ leaf as an additional predictor variable influence the ability to predict g s , and then explored the abiotic (i.e. month, site-month interaction) and biotic (i.e. tree-species-specific characteristics) drivers of slope parameter variation. Our results show that the inclusion of Ψ leaf did not improve model performance and that the models that represent the response of g s to vapor pressure deficit performed better than corresponding models that respond to relative humidity. Within each g s model, we found large variation in the slope parameter, and this variation was attributable to the biotic driver, rather than abiotic drivers. We further investigated potential relationships between the slope parameter and the six available plant traits mentioned above, and found that only one trait, LMA, had a significant correlation with the slope parameter (R 2  = 0.66, n = 15), highlighting a potential path towards improved model parameterization. This study advances understanding of g s dynamics over seasonal drought, and identifies a practical, trait-based approach to improve modeling of carbon and water exchange in tropical forests., (© 2019 John Wiley & Sons Ltd.)

Leaf mass per area (LMA) is a key plant trait, reflecting tradeoffs between leaf photosynthetic function, longevity, and structural investment. Capturing spatial and temporal variability in LMA has been a long-standing goal of ecological research and is an essential component for advancing Earth system models. Despite the substantial variation in LMA within and across Earth's biomes, an efficient, globally generalizable approach to predict LMA is still lacking. We explored the capacity to predict LMA from leaf spectra across much of the global LMA trait space, with values ranging from 17 to 393 g m -2 . Our dataset contained leaves from a wide range of biomes from the high Arctic to the tropics, included broad- and needleleaf species, and upper- and lower-canopy (i.e. sun and shade) growth environments. Here we demonstrate the capacity to rapidly estimate LMA using only spectral measurements across a wide range of species, leaf age and canopy position from diverse biomes. Our model captures LMA variability with high accuracy and low error (R 2  = 0.89; root mean square error (RMSE) = 15.45 g m -2 ). Our finding highlights the fact that the leaf economics spectrum is mirrored by the leaf optical spectrum, paving the way for this technology to predict the diversity of LMA in ecosystems across global biomes., (No claim to US Government works New Phytologist © 2019 New Phytologist Trust.)

Understanding the pronounced seasonal and spatial variation in leaf carboxylation capacity (V c,max ) is critical for determining terrestrial carbon cycling in tropical forests. However, an efficient and scalable approach for predicting V c,max is still lacking. Here the ability of leaf spectroscopy for rapid estimation of V c,max was tested. V c,max was estimated using traditional gas exchange methods, and measured reflectance spectra and leaf age in leaves sampled from tropical forests in Panama and Brazil. These data were used to build a model to predict V c,max from leaf spectra. The results demonstrated that leaf spectroscopy accurately predicts V c,max of mature leaves in Panamanian tropical forests (R 2  = 0.90). However, this single-age model required recalibration when applied to broader leaf demographic classes (i.e. immature leaves). Combined use of spectroscopy models for V c,max and leaf age enabled construction of the V c,max -age relationship solely from leaf spectra, which agreed with field observations. This suggests that the spectroscopy technique can capture the seasonal variability in V c,max , assuming sufficient sampling across diverse species, leaf ages and canopy environments. This finding will aid development of remote sensing approaches that can be used to characterize V c,max in moist tropical forests and enable an efficient means to parameterize and evaluate terrestrial biosphere models., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

The maximum carboxylation capacity of Rubisco, V c,max , is an important photosynthetic parameter that is key to accurate estimation of carbon assimilation. The gold-standard technique for determining V c,max is to derive V c,max from the initial slope of an A-C i curve (the response of photosynthesis, A, to intercellular CO 2 concentration, C i ). Accurate estimates of V c,max derived from an alternative and rapid "one-point" measurement of photosynthesis could greatly accelerate data collection and model parameterization. We evaluated the practical application of the one-point method in six species measured under standard conditions (saturating irradiance and 400 μmol CO 2 mol -1 ) and under conditions that would increase the likelihood for successful estimation of V c,max : (a) ensuring Rubisco-limited A by measuring at 300 μmol CO 2 mol -1 and (b) allowing time for acclimation to saturating irradiance prior to measurement. The one-point method significantly underestimated V c,max in four of the six species, providing estimates 21%-32% below fitted values. We identified ribulose-1,5-bisphosphate-limited A, light acclimation, and the use of an assumed respiration rate as factors that limited the effective use of the one-point method to accurately estimate V c,max . We conclude that the one-point method requires a species-specific understanding of its application, is often unsuccessful, and must be used with caution., (© 2019 John Wiley & Sons Ltd.)

Reducing uncertainties in Earth System Model predictions requires carefully evaluating core model processes. Here we examined how canopy radiative transfer model (RTM) parameter uncertainties, in combination with canopy structure, affect terrestrial carbon and energy projections in a demographic land-surface model, the Ecosystem Demography model (ED2). Our analyses focused on temperate deciduous forests and tested canopies of varying structural complexity. The results showed a strong sensitivity of tree productivity, albedo, and energy balance projections to RTM parameters. Impacts of radiative parameter uncertainty on stand-level canopy net primary productivity ranged from ~2 to > 20% and was most sensitive to canopy clumping and leaf reflectance/transmittance in the visible spectrum (~400-750 nm). ED2 canopy albedo varied by ~1 to ~10% and was most sensitive to near-infrared reflectance (~800-1200 nm). Bowen ratio, in turn, was most sensitive to wood optical properties parameterization; this was much larger than expected based on literature, suggesting model instabilities. In vertically and spatially complex canopies the model response to RTM parameterization may show an apparent reduced sensitivity when compared to simpler canopies, masking much larger changes occurring within the canopy. Our findings highlight both the importance of constraining canopy RTM parameters in models and valuating how the canopy structure responds to those parameter values. Finally, we advocate for more model evaluation, similar to this study, to highlight possible issues with model behavior or process representations, particularly models with demographic representations, and identify potential ways to inform and constrain model predictions., Competing Interests: The authors have declared that no competing interests exist.

How terrestrial biosphere models (TBMs) represent leaf photosynthesis and its sensitivity to temperature are two critical components of understanding and predicting the response of the Arctic carbon cycle to global change. We measured the effect of temperature on the response of photosynthesis to irradiance in six Arctic plant species and determined the quantum yield of CO 2 fixation ( ϕ CO 2 ) and the convexity factor (θ). We also determined leaf absorptance (α) from measured reflectance to calculate ϕ CO 2 on an absorbed light basis ( ϕ CO 2 . a ) and enabled comparison with nine TBMs. The mean ϕ CO 2 . a was 0.045 mol CO 2  mol -1 absorbed quanta at 25°C and closely agreed with the mean TBM parameterisation (0.044), but as temperature decreased measured ϕ CO 2 . a diverged from TBMs. At 5°C measured ϕ CO 2 . a was markedly reduced (0.025) and 60% lower than TBM estimates. The θ also showed a significant reduction between 25°C and 5°C. At 5°C θ was 38% lower than the common model parameterisation of 0.7. These data show that TBMs are not accounting for observed reductions in ϕ CO 2 . a and θ that can occur at low temperature. Ignoring these reductions in ϕ CO 2 . a and θ could lead to a marked (45%) overestimation of CO 2 assimilation at subsaturating irradiance and low temperature., (No claim to original US government works New Phytologist © 2019 New Phytologist Trust.)

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short- and long-term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015-2016 El Niño to examine how short- and long-term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests., (© 2018 John Wiley & Sons Ltd.)

Approaches that enable high-throughput, non-destructive measurement of plant traits are essential for programs seeking to improve crop yields through physiological breeding. However, many key traits still require measurement using slow, labor-intensive, and destructive approaches. We investigated the potential to retrieve key traits associated with leaf source-sink balance and carbon-nitrogen status from leaf optical properties. Structural and biochemical traits and leaf reflectance (500-2400 nm) of eight crop species were measured and used to develop predictive 'spectra-trait' models using partial least squares regression. Independent validation data demonstrated that the models achieved very high predictive power for C, N, C:N ratio, leaf mass per area, water content, and protein content (R2>0.85), good predictive capability for starch, sucrose, glucose, and free amino acids (R2=0.58-0.80), and some predictive capability for nitrate (R2=0.51) and fructose (R2=0.44). Our spectra-trait models were developed to cover the trait space associated with food or biofuel crop plants and can therefore be applied in a broad range of phenotyping studies., (Published by Oxford University Press on behalf of the Society for Experimental Biology 2019.)

A central challenge to understanding how climate anomalies, such as drought and heatwaves, impact the terrestrial carbon cycle, is quantification and scaling of spatial and temporal variation in ecosystem gross primary productivity (GPP). Existing empirical and model-based satellite broadband spectra-based products have been shown to miss critical variation in GPP. Here, we evaluate the potential of high spectral resolution (10 nm) shortwave (400-2,500 nm) imagery to better detect spatial and temporal variations in GPP across a range of ecosystems, including forests, grassland-savannas, wetlands, and shrublands in a water-stressed region. Estimates of GPP from eddy covariance observations were compared against airborne hyperspectral imagery, collected across California during the 2013-2014 HyspIRI airborne preparatory campaign. Observations from 19 flux towers across 23 flight campaigns (102 total image-flux tower pairs) showed GPP to be strongly correlated to a suite of spectral wavelengths and band ratios associated with foliar physiology and chemistry. A partial least squares regression (PLSR) modeling approach was then used to predict GPP with higher validation accuracy (adjusted R 2  = 0.71) and low bias (0.04) compared to existing broadband approaches (e.g., adjusted R 2  = 0.68 and bias = -5.71 with the Sims et al. model). Significant wavelengths contributing to the PLSR include those previously shown to coincide with Rubisco (wavelengths 1,680, 1,740, and 2,290 nm) and V cmax (wavelengths 1,680, 1,722, 1,732, 1,760, and 2,300 nm). These results provide strong evidence that advances in satellite spectral resolution offer significant promise for improved satellite-based monitoring of GPP variability across a diverse range of terrestrial ecosystems., (© 2018 by the Ecological Society of America.)

Satellite observations of Amazon forests show seasonal and interannual variations, but the underlying biological processes remain debated. Here we combined radiative transfer models (RTMs) with field observations of Amazon forest leaf and canopy characteristics to test three hypotheses for satellite-observed canopy reflectance seasonality: seasonal changes in leaf area index, in canopy-surface leafless crown fraction and/or in leaf demography. Canopy RTMs (PROSAIL and FLiES), driven by these three factors combined, simulated satellite-observed seasonal patterns well, explaining c. 70% of the variability in a key reflectance-based vegetation index (MAIAC EVI, which removes artifacts that would otherwise arise from clouds/aerosols and sun-sensor geometry). Leaf area index, leafless crown fraction and leaf demography independently accounted for 1, 33 and 66% of FLiES-simulated EVI seasonality, respectively. These factors also strongly influenced modeled near-infrared (NIR) reflectance, explaining why both modeled and observed EVI, which is especially sensitive to NIR, captures canopy seasonal dynamics well. Our improved analysis of canopy-scale biophysics rules out satellite artifacts as significant causes of satellite-observed seasonal patterns at this site, implying that aggregated phenology explains the larger scale remotely observed patterns. This work significantly reconciles current controversies about satellite-detected Amazon phenology, and improves our use of satellite observations to study climate-phenology relationships in the tropics., (No claim to original US Government works New Phytologist © 2017 New Phytologist Trust.)

Improving photosynthesis to raise wheat yield potential has emerged as a major target for wheat physiologists. Photosynthesis-related traits, such as nitrogen per unit leaf area (Narea) and leaf dry mass per area (LMA), require laborious, destructive, laboratory-based methods, while physiological traits underpinning photosynthetic capacity, such as maximum Rubisco activity normalized to 25 °C (Vcmax25) and electron transport rate (J), require time-consuming gas exchange measurements. The aim of this study was to assess whether hyperspectral reflectance (350-2500 nm) can be used to rapidly estimate these traits on intact wheat leaves. Predictive models were constructed using gas exchange and hyperspectral reflectance data from 76 genotypes grown in glasshouses with different nitrogen levels and/or in the field under yield potential conditions. Models were developed using half of the observed data with the remainder used for validation, yielding correlation coefficients (R2 values) of 0.62 for Vcmax25, 0.7 for J, 0.81 for SPAD, 0.89 for LMA, and 0.93 for Narea, with bias <0.7%. The models were tested on elite lines and landraces that had not been used to create the models. The bias varied between -2.3% and -5.5% while relative error of prediction was similar for SPAD but slightly greater for LMA and Narea., (© The Author(s) 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Numerous current efforts seek to improve the representation of ecosystem ecology and vegetation demographic processes within Earth System Models (ESMs). These developments are widely viewed as an important step in developing greater realism in predictions of future ecosystem states and fluxes. Increased realism, however, leads to increased model complexity, with new features raising a suite of ecological questions that require empirical constraints. Here, we review the developments that permit the representation of plant demographics in ESMs, and identify issues raised by these developments that highlight important gaps in ecological understanding. These issues inevitably translate into uncertainty in model projections but also allow models to be applied to new processes and questions concerning the dynamics of real-world ecosystems. We argue that stronger and more innovative connections to data, across the range of scales considered, are required to address these gaps in understanding. The development of first-generation land surface models as a unifying framework for ecophysiological understanding stimulated much research into plant physiological traits and gas exchange. Constraining predictions at ecologically relevant spatial and temporal scales will require a similar investment of effort and intensified inter-disciplinary communication., (© 2017 John Wiley & Sons Ltd.)

Terrestrial biosphere models (TBMs) are highly sensitive to model representation of photosynthesis, in particular the parameters maximum carboxylation rate and maximum electron transport rate at 25°C (V c,max.25 and J max.25 , respectively). Many TBMs do not include representation of Arctic plants, and those that do rely on understanding and parameterization from temperate species. We measured photosynthetic CO 2 response curves and leaf nitrogen (N) content in species representing the dominant vascular plant functional types found on the coastal tundra near Barrow, Alaska. The activation energies associated with the temperature response functions of V c,max and J max were 17% lower than commonly used values. When scaled to 25°C, V c,max.25 and J max.25 were two- to five-fold higher than the values used to parameterize current TBMs. This high photosynthetic capacity was attributable to a high leaf N content and the high fraction of N invested in Rubisco. Leaf-level modeling demonstrated that current parameterization of TBMs resulted in a two-fold underestimation of the capacity for leaf-level CO 2 assimilation in Arctic vegetation. This study highlights the poor representation of Arctic photosynthesis in TBMs, and provides the critical data necessary to improve our ability to project the response of the Arctic to global environmental change., (No claim to original US Government works. New Phytologist © 2017 New Phytologist Trust.)

Leaf quantity (i.e., canopy leaf area index, LAI), quality (i.e., per-area photosynthetic capacity), and longevity all influence the photosynthetic seasonality of tropical evergreen forests. However, these components of tropical leaf phenology are poorly represented in most terrestrial biosphere models (TBMs). Here, we explored alternative options for the representation of leaf phenology effects in TBMs that employ the Farquahar, von Caemmerer & Berry (FvCB) representation of CO 2 assimilation. We developed a two-fraction leaf (sun and shade), two-layer canopy (upper and lower) photosynthesis model to evaluate different modeling approaches and assessed three components of phenological variations (i.e., leaf quantity, quality, and within-canopy variation in leaf longevity). Our model was driven by the prescribed seasonality of leaf quantity and quality derived from ground-based measurements within an Amazonian evergreen forest. Modeled photosynthetic seasonality was not sensitive to leaf quantity, but was highly sensitive to leaf quality and its vertical distribution within the canopy, with markedly more sensitivity to upper canopy leaf quality. This is because light absorption in tropical canopies is near maximal for the entire year, implying that seasonal changes in LAI have little impact on total canopy light absorption; and because leaf quality has a greater effect on photosynthesis of sunlit leaves than light limited, shade leaves and sunlit foliage are more abundant in the upper canopy. Our two-fraction leaf, two-layer canopy model, which accounted for all three phenological components, was able to simulate photosynthetic seasonality, explaining ~90% of the average seasonal variation in eddy covariance-derived CO 2 assimilation. This work identifies a parsimonious approach for representing tropical evergreen forest photosynthetic seasonality in TBMs that utilize the FvCB model of CO 2 assimilation and highlights the importance of incorporating more realistic phenological mechanisms in models that seek to improve the projection of future carbon dynamics in tropical evergreen forests., (© 2017 John Wiley & Sons Ltd.)

Leaf age structures the phenology and development of plants, as well as the evolution of leaf traits over life histories. However, a general method for efficiently estimating leaf age across forests and canopy environments is lacking. Here, we explored the potential for a statistical model, previously developed for Peruvian sunlit leaves, to consistently predict leaf ages from leaf reflectance spectra across two contrasting forests in Peru and Brazil and across diverse canopy environments. The model performed well for independent Brazilian sunlit and shade canopy leaves (R 2  = 0.75-0.78), suggesting that canopy leaves (and their associated spectra) follow constrained developmental trajectories even in contrasting forests. The model did not perform as well for mid-canopy and understory leaves (R 2  = 0.27-0.29), because leaves in different environments have distinct traits and trait developmental trajectories. When we accounted for distinct environment-trait linkages - either by explicitly including traits and environments in the model, or, even better, by re-parameterizing the spectra-only model to implicitly capture distinct trait-trajectories in different environments - we achieved a more general model that well-predicted leaf age across forests and environments (R 2  = 0.79). Fundamental rules, linked to leaf environments, constrain the development of leaf traits and allow for general prediction of leaf age from spectra across species, sites and canopy environments., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Accurate representation of photosynthesis in terrestrial biosphere models (TBMs) is essential for robust projections of global change. However, current representations vary markedly between TBMs, contributing uncertainty to projections of global carbon fluxes. Here we compared the representation of photosynthesis in seven TBMs by examining leaf and canopy level responses of photosynthetic CO 2 assimilation (A) to key environmental variables: light, temperature, CO 2 concentration, vapor pressure deficit and soil water content. We identified research areas where limited process knowledge prevents inclusion of physiological phenomena in current TBMs and research areas where data are urgently needed for model parameterization or evaluation. We provide a roadmap for new science needed to improve the representation of photosynthesis in the next generation of terrestrial biosphere and Earth system models., (No claim to original US Government works New Phytologist © 2016 New Phytologist Trust.)

Simulations of photosynthesis by terrestrial biosphere models typically need a specification of the maximum carboxylation rate (Vcmax ). Estimating this parameter using A-Ci curves (net photosynthesis, A, vs intercellular CO2 concentration, Ci ) is laborious, which limits availability of Vcmax data. However, many multispecies field datasets include net photosynthetic rate at saturating irradiance and at ambient atmospheric CO2 concentration (Asat ) measurements, from which Vcmax can be extracted using a 'one-point method'. We used a global dataset of A-Ci curves (564 species from 46 field sites, covering a range of plant functional types) to test the validity of an alternative approach to estimate Vcmax from Asat via this 'one-point method'. If leaf respiration during the day (Rday ) is known exactly, Vcmax can be estimated with an r(2) value of 0.98 and a root-mean-squared error (RMSE) of 8.19 μmol m(-2) s(-1) . However, Rday typically must be estimated. Estimating Rday as 1.5% of Vcmax, we found that Vcmax could be estimated with an r(2) of 0.95 and an RMSE of 17.1 μmol m(-2) s(-1) . The one-point method provides a robust means to expand current databases of field-measured Vcmax , giving new potential to improve vegetation models and quantify the environmental drivers of Vcmax variation., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

The soybean aphid, Aphis glycines Matsumura, an exotic species in North America that has been detected in 21 U.S. states and Canada, is a major pest for soybean that can reduce maximum photosynthetic capacity and yields. Our existing knowledge is based on relatively few studies that do not span a wide variety of environmental conditions, and often focus on relatively high and damaging population pressure. We examined the effects of varied populations and duration of soybean aphids on soybean photosynthetic rates and yield in two experiments. In a 2011 field study, we found that plants with low cumulative aphid days (CAD, less than 2,300) had higher yields than plants not experiencing significant aphid pressure, suggesting a compensatory growth response to low aphid pressure. This response did not hold at higher CAD, and yields declined. In a 2013 controlled-environment greenhouse study, soybean plants were well-watered and fertilized with nitrogen (N), and aphid populations were manipulated to reach moderate to high levels (8,000-50,000 CAD). Plants tolerated these population levels when aphids were introduced during the vegetative or reproductive phenological stages of the plant, showing no significant reduction in yield. Leaf N concentration and CAD were positively and significantly correlated with increasing ambient photosynthetic rates. Our findings suggest that, given the right environmental conditions, modern soybean plants can withstand higher aphid pressure than previously assumed. Moreover, soybean plants also responded positively through a compensatory photosynthetic effect to moderate population pressure, contributing to stable or increased yield., (© The Authors 2016. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The morphological and biochemical properties of plant canopies are strong predictors of photosynthetic capacity and nutrient cycling. Remote sensing research at the leaf and canopy scales has demonstrated the ability to characterize the biochemical status of vegetation canopies using reflectance spectroscopy, including at the leaf level and canopy level from air- and spaceborne imaging spectrometers. We developed a set of accurate and precise spectroscopic calibrations for the determination of leaf chemistry (contents of nitrogen, carbon, and fiber constituents), morphology (leaf mass per area, Marea), and isotopic composition (δ15N) of temperate and boreal tree species using spectra of dried and ground leaf material. The data set consisted of leaves from both broadleaf and needle-leaf conifer species and displayed a wide range in values, determined with standard analytical approaches: 0.7–4.4% for nitrogen (Nmass), 42–54% for carbon (Cmass), 17–58% for fiber (acid-digestible fiber, ADF), 7–44% for lignin (acid-digestible lignin, ADL), 3–31% for cellulose, 17–265 g/m2 for Marea, and −9.4‰ to 0.8‰ for δ15N. The calibrations were developed using a partial least-squares regression (PLSR) modeling approach combined with a novel uncertainty analysis. Our PLSR models yielded model calibration (independent validation shown in parentheses) R2 and the root mean square error (RMSE) values, respectively, of 0.98 (0.97) and 0.10% (0.13%) for Nmass, R2 = 0.77 (0.73) and RMSE = 0.88% (0.95%) for Cmass, R2 = 0.89 (0.84) and RMSE = 2.8% (3.4%) for ADF, R2 = 0.77 (0.69) and RMSE = 2.4% (3.9%) for ADL, R2 = 0.77 (0.72) and RMSE = 1.4% (1.9%) for leaf cellulose, R2 = 0.62 (0.60) and RMSE = 0.91‰ (1.5‰) for δ15N, and R2 = 0.88 (0.87) with RMSE = 17.2 g/m2 (22.8 g/m2) for Marea. This study demonstrates the potential for rapid and accurate estimation of key foliar traits of forest canopies that are important for ecological research and modeling activities, with a single calibration equation valid over a wide range of northern temperate and boreal species and leaf physiognomies. The results provide the basis to characterize important variability between and within species, and across ecological gradients using a rapid, cost-effective, easily replicated method.

A major goal of remote sensing is the development of generalizable algorithms to repeatedly and accurately map ecosystem properties across space and time. Imaging spectroscopy has great potential to map vegetation traits that cannot be retrieved from broadband spectral data, but rarely have such methods been tested across broad regions. Here we illustrate a general approach for estimating key foliar chemical and morphological traits through space and time using NASA's Airborne Visible/Infrared Imaging Spectrometer (AVIRIS-Classic). We apply partial least squares regression (PLSR) to data from 237 field plots within 51 images acquired between 2008 and 2011. Using a series of 500 randomized 50/50 subsets of the original data, we generated spatially explicit maps of seven traits (leaf mass per area (M(area)), percentage nitrogen, carbon, fiber, lignin, and cellulose, and isotopic nitrogen concentration, δ15N) as well as pixel-wise uncertainties in their estimates based on error propagation in the analytical methods. Both M(area) and %N PLSR models had a R2 > 0.85. Root mean square errors (RMSEs) for both variables were less than 9% of the range of data. Fiber and lignin were predicted with R2 > 0.65 and carbon and cellulose with R2 > 0.45. Although R2 of %C and cellulose were lower than M(area) and %N, the measured variability of these constituents (especially %C) was also lower, and their RMSE values were beneath 12% of the range in overall variability. Model performance for δ15N was the lowest (R2 = 0.48, RMSE = 0.95 per thousand), but within 15% of the observed range. The resulting maps of chemical and morphological traits, together with their overall uncertainties, represent a first-of-its-kind approach for examining the spatiotemporal patterns of forest functioning and nutrient cycling across a broad range of temperate and sub-boreal ecosystems. These results offer an alternative to categorical maps of functional or physiognomic types by providing non-discrete maps (i.e., on a continuum) of traits that define those functional types. A key contribution of this work is the ability to assign retrieval uncertainties by pixel, a requirement to enable assimilation of these data products into ecosystem modeling frameworks to constrain carbon and nutrient cycling projections.