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The role of spatial scale in imaging spectroscopy of plant traits and plant functional diversity
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Plant responses to rising vapor pressure deficit
Recent decades have been characterized by increasing temperatures worldwide, resulting in an exponential climb in vapor pressure deficit (VPD). VPD has been identified as an increasingly important driver of plant functioning in terrestrial biomes and has been established as a major contributor in recent drought‐induced plant mortality independent of other drivers associated with climate change. Despite this, few studies have isolated the physiological response of plant functioning to high VPD, thus limiting our understanding and ability to predict future impacts on terrestrial ecosystems. An abundance of evidence suggests that stomatal conductance declines under high VPD and transpiration increases in most species up until a given VPD threshold, leading to a cascade of subsequent impacts including reduced photosynthesis and growth, and higher risks of carbon starvation and hydraulic failure. Incorporation of photosynthetic and hydraulic traits in ‘next‐generation’ land‐surface models has the greatest potential for improved prediction of VPD responses at the plant‐ and global‐scale, and will yield more mechanistic simulations of plant responses to a changing climate. By providing a fully integrated framework and evaluation of the impacts of high VPD on plant function, improvements in forecasting and long‐term projections of climate impacts can be made.
The Impact of Alternative Trait-Scaling Hypotheses for the Maximum Photosynthetic Carboxylation Rate (V (sub cmax)) on Global Gross Primary Production
The maximum photosynthetic carboxylation rate (V (sub cmax)) is an influential plant trait that has multiple scaling hypotheses, which is a source of uncertainty in predictive understanding of global gross primary production (GPP). Four trait-scaling hypotheses (plant functional type, nutrient limitation, environmental filtering, and plant plasticity) with nine specific implementations were used to predict global V(sub cmax) distributions and their impact on global GPP in the Sheffield Dynamic Global Vegetation Model (SDGVM). Global GPP varied from 108.1 to 128.2 petagrams of Carbon (PgC) per year, 65 percent of the range of a recent model intercomparison of global GPP. The variation in GPP propagated through to a 27percent coefficient of variation in net biome productivity (NBP). All hypotheses produced global GPP that was highly correlated (r equals 0.85-0.91) with three proxies of global GPP. Plant functional type-based nutrient limitation, underpinned by a core SDGVM hypothesis that plant nitrogen (N) status is inversely related to increasing costs of N acquisition with increasing soil carbon, adequately reproduced global GPP distributions. Further improvement could be achieved with accurate representation of water sensitivity and agriculture in SDGVM. Mismatch between environmental filtering (the most data-driven hypothesis) and GPP suggested that greater effort is needed understand V(sub cmax) variation in the field, particularly in northern latitudes.
Molecular motors and their functions in plants
Molecular motors that hydrolyze ATP and use the derived energy to generate force are involved in a variety of diverse cellular functions. Genetic, biochemical, and cellular localization data have implicated motors in a variety of functions such as vesicle and organelle transport, cytoskeleton dynamics, morphogenesis, polarized growth, cell movements, spindle formation, chromosome movement, nuclear fusion, and signal transduction. In non-plant systems three families of molecular motors (kinesins, dyneins, and myosins) have been well characterized. These motors use microtubules (in the case of kinesines and dyneins) or actin filaments (in the case of myosins) as tracks to transport cargo materials intracellularly. During the last decade tremendous progress has been made in understanding the structure and function of various motors in animals. These studies are yielding interesting insights into the functions of molecular motors and the origin of different families of motors. Furthermore, the paradigm that motors bind cargo and move along cytoskeletal tracks does not explain the functions of some of the motors. Relatively little is known about the molecular motors and their roles in plants. In recent years, by using biochemical, cell biological, molecular, and genetic approaches a few molecular motors have been isolated and characterized from plants. These studies indicate that some of the motors in plants have novel features and regulatory mechanisms. The role of molecular motors in plant cell division, cell expansion, cytoplasmic streaming, cell-to-cell communication, membrane trafficking, and morphogenesis is beginning to be understood. Analyses of the Arabidopsis genome sequence database (51% of genome) with conserved motor domains of kinesin and myosin families indicates the presence of a large number (about 40) of molecular motors and the functions of many of these motors remain to be discovered. It is likely that many more motors with novel regulatory mechanisms that perform plant-specific functions are yet to be discovered. Although the identification of motors in plants, especially in Arabidopsis, is progressing at a rapid pace because of the ongoing plant genome sequencing projects, only a few plant motors have been characterized in any detail. Elucidation of function and regulation of this multitude of motors in a given species is going to be a challenging and exciting area of research in plant cell biology. Structural features of some plant motors suggest calcium, through calmodulin, is likely to play a key role in regulating the function of both microtubule- and actin-based motors in plants.
Diverse Photosynthetic Capacity of Global Ecosystems Mapped by Satellite Chlorophyll Fluorescence Measurements
Photosynthetic capacity is often quantified by the Rubisco-limited photosynthetic capacity (i.e. maximum carboxylation rate, V(sub cmax)). It is a key plant functional trait that is widely used in Earth System Models for simulation of the global carbon and water cycles. Measuring V(sub cmax) is time-consuming and laborious; therefore, the spatiotemporal distribution of V(sub cmax) is still poorly understood due to limited measurements of V(sub cmax). In this study, we used a data assimilation approach to map the spatial variation of V(sub cmax) for global terrestrial ecosystems from a 11-year-long satellite-observed solar-induced chlorophyll fluorescence (SIF) record. In this SIF-derived V(sub cmax) map, the mean V(sub cmax) value for each plant function type (PFT) is found to be comparable to a widely used N-derived V(sub cmax) dataset by Kattge et al. (2009). The gradient of V(sub cmax) along PFTs is clearly revealed even without land cover information as an input. Large seasonal and spatial variations of V(sub cmax) are found within each PFT, especially for diverse crop rotation systems. The distribution of major crop belts, characterized with high V(sub cmax) values, is highlighted in this V(sub cmax) map. Legume plants are characterized with high V(sub cmax) values. This V(sub cmax) map also clearly illustrates the emerging soybean revolution in South America where V(sub cmax) is the highest among the world. The gradient of V(sub cmax) in Amazon is found to follow the transition of soil types with different soil N and P contents. This study suggests that satellite-observed SIF is powerful in deriving the important plant functional trait, i.e. V(sub cmax), for global climate change studies.
Remote Sensing of Lineage Functional Types for Modeling and Monitoring Biodiversity
Hyperspectral remote sensing has the potential to continuously scale plant function and plant diversity information from landscape to global extents. Numerous studies have indicated that VSWIR (400-2500 nm) reflectance properties of vegetation capture evolutionarily conserved biochemical, structural, and other functional attributes of plant species. Spectral properties conserved in plants provide the opportunity to both 1) aggregate species into lineages with improved classification accuracy and 2) link those lineages directly to plant traits. Full realization of this goal will enable parameterization of Land Surface Models (LSMs) with remotely sensed information, e.g., canopy nitrogen, and better representations of biodiversity and functional diversity in biogeographic studies. In this study, we use hyperspectral AVIRIS data from the 2013 HyspIRI campaign over the Southern Sierra Nevada, California flight box to investigate the potential for incorporating evolutionary thinking into landcover classification. We link the airborne hyperspectral data with vegetation plot data from roughly 1372 surveys and a phylogeny representing 1361 species. We aggregate species into lineages ranging from species level groups down to similar number of Plant Functional Types as often used in LSMs. We assessed the ability of Random Forest and Partial Least Squares Discriminant Analysis to discriminate across these different phylogenetic scales and determine the optimal number of lineages to classify. Although there are some temporal and spatial differences in our training data, our best approaches achieved moderate classification accuracy (Kappa > 0.65). Given an optimal number of lineages, we explored approaches to improve classifications including machine learning and unmixing approaches. This work suggests that lineage-based methods may be a promising way to leverage the huge amounts of data that will come from high resolution and high return interval hyperspectral data planned for the Surface Biology and Geology mission with sparsely sampled existing ground-based ecological data.
Emerging satellite observations for diurnal cycling of ecosystem processes
Diurnal cycling of plant carbon uptake and water use, and their responses to water and heat stresses, provide direct insight into assessing ecosystem productivity, agricultural production and management practices, carbon and water cycles, and feedbacks to the climate. Temperature, light, atmospheric water demand, soil moisture and leaf water potential vary over the course of the day, leading to diurnal variations in stomatal conductance, photosynthesis and transpiration. Earth observations from polar-orbiting satellites are incapable of studying these diurnal variations. Here, we review the emerging satellite observations that have the potential for studying how plant functioning and ecosystem processes vary over the course of the diurnal cycle. The recently launched ECOsystem Spaceborne Thermal Radiometer Experiment on Space Station (ECOSTRESS) and Orbiting Carbon Observatory-3 (OCO-3) provide land surface temperature, evapotranspiration (ET), gross primary production (GPP) and solar-induced chlorophyll fluorescence data at different times of day. New generation operational geostationary satellites such as Himawari-8 and the GOES-R series can provide continuous, high-frequency data of land surface temperature, solar radiation, GPP and ET. Future satellite missions such as GeoCarb, TEMPO and Sentinel-4 are also planned to have diurnal sampling capability of solar-induced chlorophyll fluorescence. We explore the unprecedented opportunities for characterizing and understanding how GPP, ET and water use efficiency vary over the course of the day in response to temperature and water stresses, and management practices. We also envision that these emerging observations will revolutionize studies of plant functioning and ecosystem processes in the context of climate change and that these observations and findings can inform agricultural and forest management and lead to improvements in Earth system models and climate projections.
The Breadboard Project - A functioning CELSS plant growth system
The primary objective of the Breadboard Project for the next 3-4 years is to develop, integrate and operate a Controlled Ecological Life Support System (CELSS) at a one-person scale. The focus of this project over the past two years has been the development of the plant growth facility, the first module of the CELSS. The other major modules, food preparation, biomass processing, and resource recovery, have been researched at the laboratory scale during the past two years and facilities are currently under construction to scale-up these modules to an operational state. This paper will outline the design requirements for the Biomass Production Chamber (BPC), the plant growth facility for the project, and the control and monitoring subsystems which operate the chamber and will present results from both engineering and biological tests of the facility. Three production evaluations of wheat, conducted in the BPC during the past year, will be described and the data generated from these tests discussed.
The plasma membrane-associated NADH oxidase of spinach leaves responds to blue light
The plasma membrane-associated NADH oxidase (NOX) of spinach leaf disks is characterized by oscillations in activity with a regular period length of ca. 24 min. Within a single population of plants exposed to light at the same time, NOX activities of all plants function synchronously. Exposure of plants transferred from darkness to blue light (495 nm, 2 min, 50 micromoles m-2 s-1) resulted in a complex response pattern but with a new maximum in the rate of NOX activity 36 (24+12) min after illumination and then with maxima in the rate of NOX activity every 24 min thereafter. Transient maxima in NOX activity were observed as well after 9.3 + /- 1.4 and 20.7 +/- 2.1 min. The blue light response differed from the response to red (650 nm, 10 min, 50 micromoles m-2 s-1) or white light where activity maxima were initiated 12 min after the light exposure followed by maxima every 24 min thereafter. Green or yellow light was ineffective. The light response was independent of the time in the 24-min NOX cycle when the light was given. The net effects of blue and red light were ultimately the same with a new maximum in the rate of NOX activity at 12+24=36 min (and every 24 min thereafter), but the mechanisms appear to be distinct.
Hydraulic Diversity of Forests Regulates Ecosystem Resilience During Drought
Plants influence the atmosphere through fluxes of carbon, water and energy, and can intensify drought through land–atmosphere feedback effects. The diversity of plant functional traits in forests, especially physiological traits related to water (hydraulic) transport, may have a critical role in land–atmosphere feedback, particularly during drought. Here we combine 352 site-years of eddy covariance measurements from 40 forest sites, remote-sensing observations of plant water content and plant functional-trait data to test whether the diversity in plant traits affects the response of the ecosystem to drought. We find evidence that higher hydraulic diversity buffers variation in ecosystem flux during dry periods across temperate and boreal forests. Hydraulic traits were the predominant significant predictors of cross-site patterns in drought response. By contrast, standard leaf and wood traits, such as specific leaf area and wood density, had little explanatory power. Our results demonstrate that diversity in the hydraulic traits of trees mediates ecosystem resilience to drought and is likely to have an important role in future ecosystem–atmosphere feedback effects in a changing climate.
Plant Responses to Rarified Atmospheres
Reduced atmospheric pressures will likely be used to minimize mass and engineering requirements for plant growth habitats used in extraterrestrial applications. This report provides a brief survey of key literature related to responses of plants to atmospheric variables and a broad rationale for designing minimal atmospheres for future plant growth structures on the Martian surface. The literature and recent work suggest that atmospheric pressure limits for normal plant function are likely to be 10 kPa or perhaps slightly lower. At Kennedy Space Center, a chamber with high vacuum capability was used to design and begin construction of a system for testing plant responses to reduced pressure atmospheres. A test rack with lighting provided by 3, high-pressure sodium vapor lamps was built to conduct measurements of short-term plant responses. Initial experiments with lettuce showed that a pressure of 10 kPa resulted in a 6.1-fold increase in the rate of water loss compared to water loss at ambient pressure (101 kPa).
Leaf-level chlorophyll fluorescence and reflectance spectra of high latitude plants
Little is known about the chlorophyll fluorescence spectra for high latitude plants. A FluoWat leaf clip was used to measure leaf-level reflectance and chlorophyll fluorescence spectra of leaves of common high latitude plants to examine general spectral characteristics of these species. Fluorescence yield (Fyield) was calculated as the ratio of the emitted fluorescence divided by the absorbed radiation for the wavelengths from 400 nm up to the wavelength of the cut-off for the FluoWat low pass filter (either 650 or 700 nm). The Fyield spectra grouped into distinctly different patterns among different plant functional types. Black spruce (Picea mariana) Fyield spectra had little red fluorescence, which was reabsorbed in the shoot, but displayed a distinct far-red peak. Quaking aspen (Populus tremuloides) had both high red and far-red Fyield peaks, as did sweet coltsfoot (Petasites frigidus). Cotton grass (Eriophorum spp.) had both red and far-red Fyield peaks, but these peaks were much lower than for aspen or coltsfoot. Sphagnum moss (Sphagnum spp.) had a distinct Fyield red peak but low far-red fluorescence. Reindeer moss lichen (Cladonia rangiferina) had very low fluorescence levels, although when damp displayed a small red Fyield peak. These high latitude vegetation samples showed wide variations in Fyield spectral shapes. The Fyield values for the individual red or far-red peaks were poorly correlated to chlorophyll content, however the ratio of far-red to red Fyield showed a strong correlation with chlorophyll content. The spectral variability of these plants may provide information for remote sensing of vegetation type but may also confound attempts to measure high latitude vegetation biophysical characteristics and function using solar induced fluorescence (SIF).
Photosynthesis-dependent Isoprene Emission from Leaf to Planet in a Global Carbon-chemistry-climate Model
We describe the implementation of a biochemical model of isoprene emission that depends on the electron requirement for isoprene synthesis into the FarquharBallBerry leaf model of photosynthesis and stomatal conductance that is embedded within a global chemistry-climate simulation framework. The isoprene production is calculated as a function of electron transport-limited photosynthesis, intercellular and atmospheric carbon dioxide concentration, and canopy temperature. The vegetation biophysics module computes the photosynthetic uptake of carbon dioxide coupled with the transpiration of water vapor and the isoprene emission rate at the 30 min physical integration time step of the global chemistry-climate model. In the model, the rate of carbon assimilation provides the dominant control on isoprene emission variability over canopy temperature. A control simulation representative of the present-day climatic state that uses 8 plant functional types (PFTs), prescribed phenology and generic PFT-specific isoprene emission potentials (fraction of electrons available for isoprene synthesis) reproduces 50 of the variability across different ecosystems and seasons in a global database of 28 measured campaign-average fluxes. Compared to time-varying isoprene flux measurements at 9 select sites, the model authentically captures the observed variability in the 30 min average diurnal cycle (R2 6496) and simulates the flux magnitude to within a factor of 2. The control run yields a global isoprene source strength of 451 TgC yr1 that increases by 30 in the artificial absence of plant water stress and by 55 for potential natural vegetation.
Canopy height and climate dryness parsimoniously explain spatial variation of unstressed stomatal conductance
The spatio-temporal variation of stomatal conductance directly regulates photosynthesis, water partitioning, and biosphere-atmosphere interactions. While many studies have focused on stomatal response to stresses, the spatial variation of unstressed stomatal conductance remains poorly determined, and is usually characterized in land surface models (LSMs) simply based on plant functional type (PFT). Here, we derived unstressed stomatal conductance at the ecosystem-scale using observations from 115 global FLUXNET sites. When aggregated by PFTs, the across-PFT pattern was highly consistent with the parameterizations of LSMs. However, PFTs alone captured only 17% of the variation in unstressed stomatal conductance across sites. Within the same PFT, unstressed stomatal conductance was negatively related to climate dryness and canopy height, which explained 45% of the total spatial variation. Our results highlight the importance of plant environment interactions in shaping stomatal traits. The trait-environment relationship established here provides an empirical approach for improved parameterizations of stomatal conductance in LSMs.
The Flora Mission for Ecosystem Composition, Disturbance and Productivity
Global land use and climate variability alter ecosystem conditions - including structure, function, and biological diversity - at a pace that requires unambiguous observations from satellite vantage points. Current global measurements are limited to general land cover, some disturbances, vegetation leaf area index, and canopy energy absorption. Flora is a pathfinding mission that provides new measurements of ecosystem structure, function, and diversity to understand the spatial and temporal dynamics of human and natural disturbances, and the biogeochemical and physiological responses of ecosystems to disturbance. The mission relies upon high-fidelity imaging spectroscopy to deliver full optical spectrum measurements (400-2500 nm) of the global land surface on a monthly time step at 45 meter spatial resolution for three years. The Flora measurement objectives are: (i) fractional cover of biological materials, (ii) canopy water content, (iii) vegetation pigments and light-use efficiency, (iv) plant functional types, (v) fire fuel load and fuel moisture content, and (vi) disturbance occurrence, type and intensity. These measurements are made using a multi-parameter, spectroscopic analysis approach afforded by observation of the full optical spectrum. Combining these measurements, along with additional observations from multispectral sensors, Flora will far advance global studies and models of ecosystem dynamics and change.
Structure and Parameter Uncertainty in Centennial Projections of Forest Community Structure and Carbon Cycling
Secondary forest regrowth shapes community succession and biogeochemistry for decades, including in the Upper Great Lakes region. Vegetation models encapsulate our understanding of forest function, and whether models can reproduce multi‐decadal succession patterns is an indication of our ability to predict forest responses to future change. We test the ability of a vegetation model to simulate C cycling and community composition during 100 years of forest regrowth following stand‐replacing disturbance, asking (a) Which processes and parameters are most important to accurately model Upper Midwest forest succession? (b) What is the relative importance of model structure versus parameter values to these predictions? We ran ensembles of the Ecosystem Demography model v2.2 with different representations of processes important to competition for light. We compared the magnitude of structural and parameter uncertainty and assessed which sub‐model–parameter combinations best reproduced observed C fluxes and community composition. On average, our simulations underestimated observed net primary productivity (NPP) and leaf area index (LAI) after 100 years and predicted complete dominance by a single plant functional type (PFT). Out of 4,000 simulations, only nine fell within the observed range of both NPP and LAI, but these predicted unrealistically complete dominance by either early hardwood or pine PFTs. A different set of seven simulations were ecologically plausible but under‐predicted observed NPP and LAI. Parameter uncertainty was large; NPP and LAI ranged from ~0% to >200% of their mean value, and any PFT could become dominant. The two parameters that contributed most to uncertainty in predicted NPP were plant–soil water conductance and growth respiration, both unobservable empirical coefficients. We conclude that (a) parameter uncertainty is more important than structural uncertainty, at least for ED‐2.2 in Upper Midwest forests and (b) simulating both productivity and plant community composition accurately without physically unrealistic parameters remains challenging for demographic vegetation models.