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Leaf nonstructural carbohydrate residence time, not concentration, correlates with leaf functional traits following the leaf economic spectrum in woody plants
Nonstructural carbohydrate (NSC) concentrations might reflect the strategies described in the leaf economic spectrum (LES) due to their dependence on photosynthesis and respiration. Here, we examined if NSC concentrations correlate with leaf structure, chemistry, and physiology traits for 114 species from 19 sites and 5 biomes around the globe. Total leaf NSC concentrations varied greatly from 16 to 199 mg g -1 dry mass and were mostly independent of leaf gas exchange and the LES traits. By contrast, leaf NSC residence time was shorter in species with higher rates of photosynthesis, following the fast-slow strategies in the LES. An average leaf held an amount of NSCs that could sustain one night of leaf respiration and could be replenished in just a few hours of photosynthesis under saturating light, indicating that most daily carbon gain is exported. Our results suggest that NSC export is clearly linked to the economics of return on resource investment.
Leaf Nitrogen, Leaf Mass Area, Leaf Water Content, Seward Peninsula, Alaska, 2017
Nitrogen and carbon content, leaf mass per area and leaf water content of leaves sampled from the NGEE Arctic Teller study site, Seward Peninsula, Alaska in 2017. Data is included for 13 species from the deciduous shrub, forb and graminoid plant functional types. See related dataset for leaf reflective spectra, sample photographs and dGPS locations. This data package includes leaf sample information and trait data (*.csv). Note that leaf nitrogen content analysis was performed on a subset of samples. Metadata files include data descriptions (_dd.csv) for tabular data and a key to species symbols used in data files. All included files are listed and described in NGA103_flmd.csv. The Next-Generation Ecosystem Experiments: Arctic (NGEE Arctic), was a research effort to reduce uncertainty in Earth System Models by developing a predictive understanding of carbon-rich Arctic ecosystems and feedbacks to climate. NGEE Arctic was supported by the Department of Energy's Office of Biological and Environmental Research. The NGEE Arctic project had two field research sites: 1) located within the Arctic polygonal tundra coastal region on the Barrow Environmental Observatory (BEO) and the North Slope near Utqiagvik (Barrow), Alaska and 2) multiple areas on the discontinuous permafrost region of the Seward Peninsula north of Nome, Alaska. Through observations, experiments, and synthesis with existing datasets, NGEE Arctic provided an enhanced knowledge base for multi-scale modeling and contributed to improved process representation at global pan-Arctic scales within the Department of Energy's Earth system Model (the Energy Exascale Earth System Model, or E3SM), and specifically within the E3SM Land Model component (ELM).
Three-dimensional estimation of deciduous forest canopy structure and leaf area using multi-directional, leaf-on and leaf-off airborne lidar data
Airborne laser scanning (ALS) has been widely used to map gap probability and leaf area index (LAI) distribution at plot and landscape scales. As an indirect measurement, most ALS methods to estimate LAI combine waveform or point density information with supporting field measurements such as the leaf angle distribution, gap probability, or direct LAI measures. The development of a more independent estimation approach would facilitate more widespread use of existing ALS data to investigate patterns of forest structure and build realistic 3-D vegetation scenes to simulate remote sensing imagery and energy balance. Here, we develop a data processing workflow (named PVlad) using ALS point cloud apparent reflectance to estimate LAI and voxel-based leaf area density (LAD), aiming to reduce the need for associated field measurements such as the gap probability. The adaptation of the path volume (PV) concept derived from apparent reflectance integrates information from multi-directional ALS pulses, and quantifies the percentage exploration of each voxel for classification and occlusion correction, such that rigorous volumetric sampling approaches can be developed to derive LAI and LAD. The PVlad workflow was applied to discrete-return lidar data (Riegl VQ480i) acquired by NASA Goddard's LiDAR, Hyperspectral and Thermal Imager (G-LiHT) Airborne Imager during leaf-on (summer) and leaf-off (spring) conditions at the Smithsonian Environmental Research Center (SERC). The estimates of LAI and LAD captured structural differences between mature, logged, and intermediate-aged stands over eight deciduous forest plots. The derived LAI values were compared to field litter collection measurements, and the derived LAD vertical distribution was compared to the output of the VoxLAD model using terrestrial laser scan (TLS) field survey data. Using voxel sizes ranging from 0.5 m to 5 m, overall LAI estimation showed linear fitting coefficient bias and for 1 and 2 m voxel sizes, and vertical LAD distribution showed strong correlation with and for 0.5 and 1m voxel sizes. For every forest stand, upper-canopy LAD had a low variance for voxel sizes of ≤ . Application of PVlad to the G-LiHT and other similar ALS data archives enables the development of fine-resolution LAI map products, including voxelization of LAD for ecosystem science and radiative transfer simulations of remote sensing imagery or surface energy balance.
Data for Impact of Vertical and Seasonal Variation in Leaf Traits on Simulating Soybean Canopy Photosynthesis via 1D and 3D Modeling
Accurate modeling of photosynthesis is crucial for predicting crop productivity and quantifying the carbon cycle in agroecosystems. Leaf traits are essential inputs for modeling canopy photosynthesis. Yet, many existing models still use fixed plant functional type (PTF)-based values to parameterize leaf traits under a big-leaf or two-big-leaf assumption, neglecting their vertical profiles and seasonal changes. This simplification may introduce significant uncertainties in estimating gross primary productivity (GPP). In this study, we simulated soybean GPP and tested the effects of vertical and seasonal variation in three key leaf photosynthetic traits: the maximum carboxylation rate at 25 °C (Vcmax25), leaf chlorophyll content (LCC), and leaf mass per area (LMA) in the 1D-SCOPE and 3D-Helios models. Weekly field measurements were conducted during the growing season of 2024 to support the simulation. We designed ten leaf trait parameterization schemes by incorporating different combinations of vertical profiles and seasonal changes, while assuming homogeneous canopy architecture in both models. Our results revealed that Vcmax25 vertical and seasonal variation had the strongest influence on simulated GPP in both 1D and 3D models, while LCC and LMA effects were minimal. Particularly, the scheme with an empirically parameterized Vcmax25 profile achieved comparable performance to the scheme with the measured Vcmax25 profile. Both 1D-SCOPE and 3D-Helios accurately modeled GPP (SCOPE: R2 = 0.87, Bias = 0.55 µmol m⁻² s⁻¹; Helios: R2 = 0.9, Bias = 0.22 µmol m⁻² s⁻¹) under the most complex scheme, and their responses to vertical and seasonal variation in leaf traits were consistent, demonstrating the robustness of our findings. Based on our findings, we propose a scalable framework for parameterizing leaf traits to improve GPP simulations. This study contributes to improving the representation of leaf trait dynamics in canopy-level photosynthesis models, potentially enhancing our ability to predict crop productivity and understand agroecosystem carbon dynamics.
Optimization of leaf morphology in relation to leaf water status: A theory
The leaf economic traits such as leaf area, maximum carbon assimilation rate, and venation are all correlated and related to water availability. Furthermore, leaves are often broad and large in humid areas and narrower in arid/semiarid and hot and cold areas. We use optimization theory to explain these patterns. We have created a constrained optimization leaf model linking leaf shape to vein structure that is integrated into coupled transpiration and carbon assimilation processes. The model maximizes net leaf carbon gain (NPPleaf) over the loss of xylem water potential. Modeled relations between leaf traits are consistent with empirically observed patterns. As the results of the leaf shape-venation relation, our model further predicts that a broadleaf has overall higher NPPleaf compared to a narrowleaf. In addition, a broadleaf has a lower stomatal resistance compared to a narrowleaf under the same level of constraint. With the same leaf area, a broadleaf will have, on average, larger conduits and lower total leaf xylem resistance and thus be more efficient in water transportation but less resistant to cavitation. By linking venation structure to leaf shape and using water potential as the constraint, our model provides a physical explanation for the general pattern of the covariance of leaf traits through the safety-efficiency trade-off of leaf hydraulic design.
Leaf microscopy applications in photosynthesis research: identifying the gaps
Abstract Leaf imaging via microscopy has provided critical insights into research on photosynthesis at multiple junctures, from the early understanding of the role of stomata, through elucidating C4 photosynthesis via Kranz anatomy and chloroplast arrangement in single cells, to detailed explorations of diffusion pathways and light utilization gradients within leaves. In recent decades, the original two-dimensional (2D) explorations have begun to be visualized in three-dimensional (3D) space, revising our understanding of structure–function relationships between internal leaf anatomy and photosynthesis. In particular, advancing new technologies and analyses are providing fresh insight into the relationship between leaf cellular components and improving the ability to model net carbon fixation, water use efficiency, and metabolite turnover rate in leaves. While ground-breaking developments in imaging tools and techniques have expanded our knowledge of leaf 3D structure via high-resolution 3D and time-series images, there is a growing need for more in vivo imaging as well as metabolite imaging. However, these advances necessitate further improvement in microscopy sciences to overcome the unique challenges a green leaf poses. In this review, we discuss the available tools, techniques, challenges, and gaps for efficient in vivo leaf 3D imaging, as well as innovations to overcome these difficulties.
Leveraging transfer learning and leaf spectroscopy for leaf trait prediction with broad spatial, species, and temporal applicability
Accurate and reliable prediction of leaf traits is crucial for understanding plant adaptations to environmental variation, monitoring terrestrial ecosystems, and enhancing comprehension of functional diversity and ecosystem functioning. Currently, various approaches (e.g., statistical, physical models) have been developed to estimate leaf traits through hyperspectral remote sensing and leaf spectroscopy. However, the absence of high-performing, transferable, and stable models across various domains of space, plant functional types (PFTs) and seasons hinder our ability to quantify and comprehend spatiotemporal variations in leaf traits. This study proposes robust and highly transferable models for better predicting leaf traits with hyperspectral reflectance. Initially, three datasets were assembled, pairing common leaf traits — chlorophyll (Chla+b), carotenoids (Ccar), leaf mass per area (LAM), equivalent water thickness (EWT) — with leaf spectra measurements collected across diverse geographic locations in the U.S. and Europe, PFTs, and seasons. Measurements were acquired using spectroradiometers (e.g., ASD FieldSpec 3/4/Pro and SVC HR-1024i) with integrating spheres, leaf clips, and contact probes. Here, we then developed transfer learning-based hybrid models that incorporated the domain knowledge of radiative transfer models (RTMs) through pretraining processes and were well-constrained by fine-tuning with field measurements. Through comparison with other state-of-the-art statistical models, including partial-least squares regression (PLSR) and Gaussian Process Regression (GPR), as well as pure physical models, we found that the proposed transfer learning models achieved better predictive performance and higher transferability. Specifically, compared to other statistical models and pure RTMs, the transfer learning model exhibited higher coefficient of determination (R 2 ) values with range of 0.01 to 0.79, lower normalized root mean square error (NRMSE) with range of 0.06 % to 33.25 % in model performance. Additionally, the models exhibited improved transferability, with higher R 2 values range from 0.04 to 0.32, lower NRMSE range from 0.08 % to 30.81 %. The findings underscore that transfer learning models through integrating domain knowledge from RTMs and limited observations, can harness the advantages of both RTMs and statistical models and serve as a promising approach for effectively predicting leaf traits.
Linking leaf dark respiration to leaf traits and reflectance spectroscopy across diverse forest types
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, in this study, 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.
Leaf angle as a leaf and canopy trait: Rejuvenating its role in ecology with new technology
Abstract Life on Earth depends on the conversion of solar energy to chemical energy by plants through photosynthesis. A fundamental challenge in optimizing photosynthesis is to adjust leaf angles to efficiently use the intercepted sunlight under the constraints of heat stress, water loss and competition. Despite the importance of leaf angle, until recently, we have lacked data and frameworks to describe and predict leaf angle dynamics and their impacts on leaves to the globe. We review the role of leaf angle in studies of ecophysiology, ecosystem ecology and earth system science, and highlight the essential yet understudied role of leaf angle as an ecological strategy to regulate plant carbon–water–energy nexus and to bridge leaf, canopy and earth system processes. Using two models, we show that leaf angle variations have significant impacts on not only canopy‐scale photosynthesis, energy balance and water use efficiency but also light competition within the forest canopy. New techniques to measure leaf angles are emerging, opening opportunities to understand the rarely‐measured intraspecific, interspecific, seasonal and interannual variations of leaf angles and their implications to plant biology and earth system science. We conclude by proposing three directions for future research.
Leaf gas exchange, spectral reflectance and leaf composition, BIONTE, Brazil, 2022
Leaf traits measured at the top of the canopy on 10 species at the BIONTE experimental plot T1B4SB3, near Manaus, Brazil, in August and September, 2022. The aim of this study was to test the relationship between leaf water use efficiency and wood density. Ten tree species located in close proximity of each other (within 50 meters) were selected and classified by wood density (low or high). Sampling was carried out on top-of-canopy leaves accessed by an articulated boom lift. Survey gas exchange measurement were performed on attached leaves at various times during the day. Following gas exchange, the leaf was harvested, and measured for spectral reflectance, leaf water content, leaf mass per area and leaf carbon content and leaf nitrogen content. Additional leaves other than those used for gas exchange were also harvested from different branches on the same tree for spectral reflectance and leaf trait measurements. Data in this data package is presented in csv files, with metadata in csv and descriptive experimental protocols as pdf. Gas exchange data and metadata meets the requirements of the ESS-DIVE reporting format for leaf gas exchange data and metadata.
The Global Spectra-Trait Initiative: A database of paired leaf spectroscopy and functional traits associated with leaf photosynthetic capacity
Accurate assessment of leaf functional traits is crucial for a diverse range of applications from crop phenotyping to parameterizing global climate models. Leaf reflectance spectroscopy offers a promising avenue to advance ecological and agricultural research by complementing traditional, time-consuming gas exchange measurements. However, the development of robust hyperspectral models for predicting leaf photosynthetic capacity and associated traits from reflectance data has been hindered by limited data availability across species and environments. Here we introduce the Global Spectra-Trait Initiative (GSTI), a collaborative repository of paired leaf hyperspectral and gas exchange measurements from diverse ecosystems. The GSTI repository currently encompasses over 7500 observations from 397 species and 41 sites gathered from 36 published and unpublished studies, thereby offering a key resource for developing and validating hyperspectral models of leaf photosynthetic capacity. The GSTI database is developed on GitHub (https://github.com/plantphys/gsti, last access: 4 January 2026) and published to ESS-DIVE https://doi.org/10.15485/2530733, Lamour et al., 2025). It includes gas exchange data, derived photosynthetic parameters, and key leaf traits often associated with traditional gas exchange measurements such as leaf mass per area and leaf elemental composition. By providing a standardized repository for data sharing and analysis, we present a critical step towards creating hyperspectral models for predicting photosynthetic traits and associated leaf traits for terrestrial plants.
Assessing soybean leaf area and leaf biomass by spectral measurements
Red and photographic infrared spectral radiances were correlated with soybean total leaf area index, green leaf area index, chlorotic leaf area index, green leaf biomass, chlorotic leaf biomass, and total biomass. The most significant correlations were found to exist between the IR/red radiance ratio data and green leaf area index and/or green leaf biomass (r squared equals 0.85 and 0.86, respectively). These findings demonstrate that remote sensing data can supply information basic to soybean canopy growth, development, and status by nondestructive determination of the green leaf area or green leaf biomass.
Spectral assessment of soybean leaf area and leaf biomass
Red and photographic infrared spectral radiance have been correlated with soybean total leaf area index, green leaf area index, chlorotic area index, green leaf biomass, chlorotic leaf biomass, and total biomass. The most significant correlations were found to exist between the spectral data and green leaf area index and/or green leaf biomass. These findings demonstrate that ground based remote sensing data can supply information basic to soybean canopy growth, development, and status by non-destructive determination of the green leaf area or green leaf biomass.
The genetic architecture of leaf stable carbon isotope composition in Zea mays and the effect of transpiration efficiency on leaf elemental accumulation
With increased demand on freshwater resources for agriculture, it is imperative that more water-use efficient crops are developed. Leaf stable carbon isotope composition, δ 13 C, is a proxy for transpiration efficiency and a possible tool for breeders, but the underlying mechanisms effecting δ 13 C in C 4 plants are not known. It has been suggested that differences in specific leaf area (SLA), which potentially reflects variation in internal CO 2 diffusion, can impact leaf δ 13 C. Furthermore, although it is known that water movement is important for elemental uptake, it is not clear how manipulation of transpiration for increased water-use efficiency may impact nutrient accumulation. Here, we characterize the genetic architecture of leaf δ 13 C and test its relationship to SLA and the ionome in five populations of maize. Five significant QTL for leaf δ 13 C were identified, including novel QTL as well as some that were identified previously in maize kernels. One of the QTL regions contains an Erecta-like gene, the ortholog of which has been shown to regulate transpiration efficiency and leaf δ 13 C in Arabidopsis. QTL for δ 13 C were located in the same general chromosome region, but slightly shifted, when comparing data from two different years. Our data does not support a relationship between δ 13 C and SLA, and of the 19 elements analyzed, only a weak correlation between molybdenum and δ 13 C was detected. Together these data add to the genetic understanding of leaf δ 13 C in maize and suggest that improvements to plant water use may be possible without significantly influencing elemental homeostasis.
Leaf reflectance, leaf transmittance and sky irradiance, at four sites in Panama, 2022
Paired leaf reflectance and transmittance data measured in Panama, at the Parque Natural Metropolitano (PNM), on Barro Colorado Island (BCI), at the San Lorenzo canopy crane site (SLZ), and in Gamboa, Panama from January to April 2022. Measurements were made on leaves from 57 different tree, shrub and liana species, from sunlit canopy and understory locations. Bark reflectance was also measured on the trunks of common tree species at the PNM and SLZ sites. Also included are irradiance measurements and leaf area index (LAI) from eight vertical profiles at PNM. The aim of these data was to refine our understanding of vertical trait and environmental variation in tropical forests, and to improve models which can predict leaf traits from leaf contact spectral measurements. All spectral data and metadata are presented in .csv files and complete instrument output are included in .zip folders. Protocol details are provided as pdf documents. In addition to spectral data described here, some of these samples were also used for measurement of leaf gas exchange, carbon and nitrogen content, and leaf mass per unit leaf area (LMA) these data can be cross linked using the unique sample ID and are provided in separate related data packages.
Intra-canopy leaf trait variation facilitates high leaf area index and compensatory growth in a clonal woody encroaching shrub
Leaf trait variation enables plants to utilize large gradients of light availability that exist across canopies of high leaf area index (LAI), allowing for greater net carbon gain while reducing light availability for understory competitors. While these canopy dynamics are well understood in forest ecosystems, studies of canopy structure of woody shrubs in grasslands are lacking. To evaluate the investment strategy used by these shrubs, we investigated the vertical distribution of leaf traits and physiology across canopies of Cornus drummondii, the predominant woody encroaching shrub in the Kansas tallgrass prairie. We also examined the impact of disturbance by browsing and grazing on these factors. Our results reveal that leaf mass per area (LMA) and leaf nitrogen per area (Na) varied approximately threefold across canopies of C. drummondii, resulting in major differences in the physiological functioning of leaves. High LMA leaves had high photosynthetic capacity, while low LMA leaves had a novel strategy for maintaining light compensation points below ambient light levels. The vertical allocation of leaf traits in C. drummondii canopies was also modified in response to browsing, which increased light availability at deeper canopy depths. As a result, LMA and Na increased at lower canopy depths, leading to a greater photosynthetic capacity deeper in browsed canopies compared to control canopies. This response, along with increased light availability, facilitated greater photosynthesis and resource-use efficiency deeper in browsed canopies compared to control canopies. Furthermore, our results illustrate how C. drummondii facilitates high LAI canopies and a compensatory growth response to browsing—both of which are key factors contributing to the success of C. drummondii and other species responsible for grassland woody encroachment.
Leaf mass per area and leaf water content measurements from field survey in association with NEON AOP survey, East River, CO 2018
Leaf area, wet, and dry weights for leaf samples from sites where field sampling was conducted during the 2018 National Ecological Observatory Network (NEON) Airborne Observation Platform (AOP) imaging spectroscopy and lidar surveys in Gunnison County, Colorado. The sampling sites were located across East River, Washington Gulch, Slate River, and Coal Creek watersheds and contained a mixture of meadow, shrub, and tree sampling sites. This data package contains the leaf area and wet and dry weights for leaves from tree and shrub sampling sites, as well as samples for meadow species present within a given sampling area. Associated data packages contain species composition information at the plot level (DOI: 10.15485/1618130), as well as additional biogeochemical, microbial, and geophysical data as they become available. For full documentation, please see associated reference.This package contains zipped folders of all scans that were used for area calculations for both sampling sites (tree/shrub, site_scans.zip) as well as representative samples from a daily sampling area of meadow species that were found in the 1 m plots (meadow_area_scans.zip). It also contains the macro code that was used in ImageJ for processing of the area calculations. Finally, it includes excel files with the area (cm2), wet weight (g), dry weight (g), leaf mass per area (LMA, g/m2), and leaf water content (%) for each set of scans.