Structural and photosynthetic dynamics mediate the response of SIF to water stress in a potato crop

Solar-induced Fluorescence (SIF) has an advantage over greenness-based Vegetation Indices in detecting drought. This advantage is the mechanistic coupling between SIF and Gross Primary Productivity (GPP). Under water stress, SIF tends to decrease with photosynthesis, due to an increase in non-photochemical quenching (NPQ), resulting in rapid and/or sustained reductions in the fluorescence quantum efficiency (phi F). Water stress also affects vegetation structure via highly dynamic changes in leaf angular distributions (LAD) or slower changes in leaf area index (LAI). Critically, these responses are entangled in space and time and their relative contribution to SIF, or to the coupling between SIF and GPP, is unclear. In this study, we quantify the relative effect of structural and photosynthetic dynamics on the diurnal and spatial variation of canopy SIF in a potato crop in response to a replicated paired-plot water stress experiment. We measured SIF using two platforms: a hydraulic lift and an Unmanned Aerial Vehicle (UAV) to capture temporal and spatial variation, respectively. LAD parameters were estimated from point clouds and photographic data and used to assess structural dynamics. Leaf phi F estimated from PAM fluorescence measurements were used to represent variations in photosynthetic regulation. We also measured foliar pigments, operating quantum yield of photosystem II (PSII), photosynthetic gas exchange, stomatal conductance and LAI. We used a radiative transfer model (SCOPE) to provide a means of decoupling structural and photosynthetic factors across the diurnal and spatial domains. The results demonstrate that diurnal variation in SIF is driven by photosynthetic and structural dynamics. The influence of phi F was prominent in the diurnal SIF response to water stress, with reduced fluorescence efficiencies in stressed plants. Structural factors dominated the spatial response of SIF to water stress over and above phi F. The results showed that the relationship between SIF and GPP is maintained in response to water stress where adjustments in NPQ and leaf angle co-operate to enhance the correlation between SIF and GPP. This study points to the complexity of interpreting and modelling the spatiotemporal connection between SIF and GPP which requires simultaneous knowledge of vegetation structural and photosynthetic dynamics.Peer reviewe

Towards the quantitative and physically-based interpretation of solar-induced vegetation fluorescence retrieved from global imaging

Due to emerging high spectral resolution, remote sensing techniques and ongoing developments to retrieve the spectrally resolved vegetation fluorescence spectrum from several scales, the light reactions of photosynthesis are receiving a boost of attention for the monitoring of the Earth's carbon balance. Sensor-retrieved vegetation fluorescence (from leaf, tower, airborne or satellite scale) originating from the excited antenna chlorophyll a molecule has become a new quantitative biophysical vegetation parameter retrievable from space using global imaging techniques. However, to retrieve the actual quantum efficiencies, and hence a true photosynthetic status of the observed vegetation, all signal distortions must be accounted for, and a high-precision true vegetation reflectance must be resolved. ESA's upcoming Fluorescence Explorer aims to deliver such novel products thanks to technological and instrumental advances, and by sophisticated approaches that will enable a deeper understanding of the mechanics of energy transfer underlying the photosynthetic process in plant canopies and ecosystems

Multiangular Observation of Canopy Sun-Induced Chlorophyll Fluorescence by Combining Imaging Spectroscopy and Stereoscopy

The effect that the canopy structure and the viewing geometry have on the intensity and the spatial distribution of passively measured sun-induced chlorophyll fluorescence at canopy scale is still not well understood. These uncertainties constrain the potential use of fluorescence to quantify photosynthesis at this level. Using a novel technique, we evaluated the diurnal changes in the spatial distribution of sun-induced fluorescence at 760 nm (F760) within the canopy as a consequence of the spatial disposition of the leaves and the viewing angle of the sensor. High resolution spectral and stereo images of a full sugar beet canopy were recorded simultaneously in the field to estimate maps of F760 and the surface angle distribution, respectively. A dedicated algorithm was used to align both maps in the post-processing and its accuracy was evaluated using a sensitivity test. The relative angle between sun and the leaf surfaces primarily determined the amount of incident Photosynthetic Active Radiation (PAR), which in turn was reflected in different values of F760, with the highest values occurring in leaf surfaces that are perpendicularly oriented to the sun. The viewing angle of the sensor also had an impact in the intensity of the recorded F760. Higher viewing angles generally resulted in higher values of F760. We attribute these changes to a direct effect of the vegetation directional reflectance response on fluorescence retrieval. Consequently, at leaf surface level, the spatio-temporal variations of F760 were mainly explained by the sun–leaf–sensor geometry rather than directionality of the fluorescence emission. At canopy scale, the diurnal patterns of F760 observed on the top-of-canopy were attributed to the complex interplay between the light penetration into the canopy as a function of the display of the various leaves and the fluorescence emission of each leaf which is modulated by the exposure of the individual leaf patch to the incoming light and the functional status of photosynthesis. We expect that forward modeling can help derive analytical simplified skeleton assumptions to scale canopy measurements to the leaf functional properties

Evaluating solar-induced fluorescence across spatial and temporal scales to monitor primary productivity

Solar-induced chlorophyll fluorescence (SIF) has been widely cited in carbon cycling studies as a proxy for photosynthesis, and SIF data are commonly incorporated into terrestrial primary productivity models. Though satellite-based SIF products show close relationships with gross primary productivity (GPP), this is not universally true at intermediate scales. A meta-analysis of the tower-based and airborne SIF literature revealed that mean SIF retrievals from unstressed vegetation span three orders of magnitude. While reporting on spectrometer calibration procedures, hardware characterizations, and associated corrections is inconsistent, laboratory and field experiments show that these factors may contribute to significant uncertainty in SIF retrievals. Additionally, there remain ongoing questions regarding the interpretation of SIF data made across spatial scales and the link between satellite SIF retrievals and primary productivity on the ground. Chlorophyll fluorescence originates from dynamic energy partitioning at the leaf level and does not exhibit a uniformly linear relationship with photosynthesis at finer scales. As a standalone metric, SIF measured at the tower scale was not found to track changes in carbon assimilation following stomatal closure induced in deciduous woody tree branches. This lack of relationship may be explained by alternative energy partitioning pathways, such as thermal energy dissipation mediated by xanthophyll cycle pigments; the activity of these pigments can be tracked using the photochemical reflectance index (PRI). Gradual, phenological changes in energy partitioning are observed as changes in the slope of the SIF-PRI relationship over the course of a season. Along with high frequency effects such as wind-mediated changes in leaf orientation and reflectance, and rapid changes in sky condition due to clouds, PRI offers crucial insights needed to link SIF to leaf physiology. While SIF offers tremendous promise for improving the characterization of terrestrial carbon exchange, and a fuller understanding of the boundaries on its utility and interpretation as a biophysical phenomenon will help to create more reliable models of global productivity

Evaluating wheat for genetic variation in radiation use efficiency: scaling traits from leaves to canopies

Wheat yields are stagnating or declining in many regions of the planet, requiring efforts to improve the light conversion efficiency, i.e., radiation use efficiency (RUE). RUE is a key trait in plant physiology because it links light capture and primary metabolism with biomass accumulation and yield. High-throughput phenotyping (HTP) was used among a population of field grown wheat with variation in RUE and photosynthetic traits to build predictive models of RUE, biomass and intercepted photosynthetically active radiation (IPAR). The use of remote sensing models predicted RUE with up to 70% accuracy compared to ground truth data. Wheat yield can be defined as the product of solar radiation intercepted throughout the crop cycle, radiation use efficiency and harvest index. Photosynthesis is a central component of RUE but normally measured in the upper layers of the canopy where light conditions are saturating. Significant relationships were found between light saturated photosynthetic rates measured at initiation of booting in the top, middle and bottom layers of the canopy and yield. These findings indicate that there is an opportunity for yield improvement if we consider the requirements of photosynthesis in the middle and bottom layers of wheat canopies where conditions are not light saturating. The study of photosynthesis in the field is constrained by low throughput and lack of integrative measurements at canopy level. Partial least squares regression (PLSR) modelling was used to build predictive models of photosynthetic, biophysical and biochemical traits at the top, middle and bottom layers of wheat canopies. The combined layer model predictions performed better than individual layer predictions. Using HTP allowed us to increase phenotyping capacity 30-fold compared to conventional phenotyping methods and our models can be used to screen varieties for high and low RUE. There is clear consensus in the physiological and breeding communities that improving RUE will be key to boost wheat yield. In the previous years of RUE research little has been explored on the role of root biomass accumulation and its interaction with aboveground biomass accumulation, RUE and yield. Strong positive associations were found between above and belowground biomass accumulation with RUE and root biomass during the vegetative period, and negative associations between yield components and root biomass accumulation, suggesting there is a coordination between roots and shoot in the vegetative period to maximize growth. However, if too much energy is invested in root biomass this will have an effect in decreasing aboveground biomass during grain filling. More research will be needed to explore new hypothesis in the field that accounts the effect of root biomass in canopy RUE and yield