Photosynthetic Light Use Efficiency Research Articles

Cross-species transcriptomics reveals differential regulation of essential photosynthesis genes in Hirschfeldia incana.

Photosynthesis is the only yield-related trait not yet substantially improved by plant breeding. Previously, we have established Hirschfeldia incana as the model plant for high photosynthetic light-use efficiency (LUE). Now we aim to unravel the genetic basis of this trait in H. incana, potentially contributing to the improvement of photosynthetic LUE in other species. Here, we compare its transcriptomic response to high light with that of Arabidopsis thaliana, Brassica rapa, and Brassica nigra, three fellow Brassicaceae members with lower photosynthetic LUE. We built a high-light, high-uniformity growing environment, in which the plants developed normally without signs of stress. We compared gene expression in contrasting light conditions across species, utilizing a panproteome to identify orthologous proteins. In-depth analysis of three key photosynthetic pathways showed a general trend of lower gene expression under high-light conditions for all four species. However, several photosynthesis-related genes in H. incana break this trend. We observed cases of constitutive higher expression (like antenna protein LHCB8), treatment-dependent differential expression (as for PSBE), and cumulative higher expression through simultaneous expression of multiple gene copies (like LHCA6). Thus, Hirschfeldia incana shows differential regulation of essential photosynthesis genes, with the light-harvesting complex as the first point of deviation. The effect of these expression differences on protein abundance and turnover, and ultimately the high photosynthetic LUE phenotype is relevant for further investigation. Furthermore, this transcriptomic resource of plants fully grown under, rather than briefly exposed to, a very high irradiance, will support the development of highly efficient photosynthesis in crops.

Net CO2 assimilation rate response of tomato seedlings (Solanum lycopersicum L.) to the interaction between light intensity, spectrum and ambient CO2 concentration.

Artificial lighting is complementary and single-source lighting for controlled Environment Agriculture (CEA) to increase crop productivity. Installations to control CO2 levels and luminaires with variable spectrum and intensity are becoming increasingly common. In order to see the net assimilation of CO2 based on the relationship between the three factors: intensity, spectrum and CO2 concentration, tests are proposed on tomatoes seedling with combinations of ten spectra (100B, 80B20G, 20B80G, 100G, 80G20R, 20G80R, 100R, 80R20B, 20R80B, 37R36G27B) seven light intensities (30, 90, 200, 350, 500, 700 and 1000 μmol·m-2 s-1) and nine CO2 concentrations (200, 300, 400, 500, 600, 700, 800 and 900 ppm). These tomato seedlings grew under uniform conditions with no treatments applied up to the moment of measurement by a differential gas analyzer. We have developed a model to evaluate and determine under what spectrum and intensity of light photosynthesis the Net assimilation of CO2 (An) is more significant in the leaves of tomato plants, considering the CO2 concentration as an independent variable in the model. The evaluation of the model parameters for each spectrum and intensity shows that the intensity has a more decisive influence on the maximum An rate than the spectra. For intensities lower than 350 μmol·m-2 s-1, it is observed that the spectrum has a greater influence on the variable An. The spectra with the best behaviour were 80R20B and 80B20R, which maintained An values between 2 and 4 (μmol CO2·m-2·s-1) above the spectra with the worst behaviour (100G, 80G20R, 20G80R and 37B36G27R) in practically all situations. Photosynthetic Light-Use Efficiency (PLUE) was also higher for the 80B20R and 20R80B spectra with values of 36,07 and 33,84 mmol CO2·mol photon-1, respectively, for light intensities of 200 μmol·m-2 s-1 and 400 ppm of CO2that increased to values of 49,65 and 48,38 mmol CO2·mol photon-1 for the same light intensity and concentrations of 850 ppm. The choice of spectrum is essential, as indicated by the data from this study, to optimize the photosynthesis of the plant species grown in the plant factory where light intensities are adjusted for greater profitability.

Contrasting photoprotective responses of forest trees revealed using PRI light responses sampled with airborne imaging spectrometry.

The Photochemical Reflectance Index (PRI) provides an optical indicator of photosynthetic light-use efficiency, photoprotection, and stress in plants. Although PRI can be applied in remote sensing, its interpretation depends on irradiance, which is hard to obtain from satellite or airborne imagery. To quantify forest photoprotective responses remotely, we developed a framework for modeling and interpreting PRI-light responses of individual trees and species using airborne imaging spectrometry coupled with georeferenced forest inventory data from a temperate broad-leaved forest. We derived an irradiance proxy, used hierarchical modeling to analyze PRI-light responses, and developed a framework of physiological interpretations of model parameters as facultative and constitutive components of photoprotection. Photochemical Reflectance Index declined with illumination, and PRI-light relationships varied with landscape position and among tree crowns and species. More sun-exposed foliage had lower intercepts and slopes of the relationship, indicating greater constitutive, but less facultative, photoprotection. We show that tree photoprotective strategies can be quantified at multiple scales using airborne hyperspectral data in structurally complex forests. Our findings and approach have important implications for the remote sensing of forest stress by offering a new way to assess functional diversity through dynamic differences in photoprotection and photosynthetic downregulation and providing previsual indicators of forest stress.

Productivity, photosynthetic light-use efficiency, nitrogen metabolism and nutritional quality of C4 halophyte Portulaca oleracea L. grown indoors under different light intensities and durations.

Portulaca oleracea L. (known as purslane), is a nutritious facultative C4 halophyte. Recently, it has been successfully grown indoors under LED lightings by our team. However, basic understanding about the impacts of light on purslanes are lacking. This study aimed to investigate the effects of light intensity and duration on productivity, photosynthetic light use efficiency, nitrogen metabolism and nutritional quality of indoor grown purslanes. All plants were grown in 10% artificial seawater hydroponically under different photosynthetic photon flux densities (PPFDs) and durations and thus different daily light integrals (DLI). They are, L1 (240 µmol photon m-2 s-1, 12h, DLI = 10.368mol m-2 day-1); L2 (320 µmol photon m-2 s-1, 18h, DLI = 20.736mol m-2 day-1); L3 (240 µmol photon m-2 s-1, 24h, DLI = 20.736mol m-2 day-1); L4 (480 µmol photon m-2 s-1, 12h, DLI = 20.736mol m-2 day-1), respectively. Compared to L1, higher DLI promoted root and shoot growth and thus increased shoot productivity by 2.63-,1.96-, 3.83-folds, respectively for purslane grown under L2, L3, L4. However, under the same DLI, L3 plants (continuous light, CL) had significantly lower shoot and root productivities compared those with higher PPFDs but shorter durations (L2 and L4). While all plants had similar total chlorophyll and carotenoid concentrations, CL (L3) plants had significantly lower light use efficiency (Fv/Fm ratio), electron transport rate, effective quantum yield of PSII, photochemical- and non-photochemical quenching. Compared to L1, higher DLI with higher PPFDs (L2 and L4) increased leaf maximum nitrate reductase activity while longer durations increased leaf concentrations and total reduced nitrogen. There were no significant differences in leaf total soluble protein, total soluble sugar and total ascorbic acid concentrations in both leaf and stem regardless of light conditions. However, L2 plants had the highest leaf proline concentration but leaf total phenolic compounds concentration was higher in L3 plants instead. Generally, L2 plants had the highest dietary minerals such as K, Ca, Mg and Fe among the four different light conditions. Overall, L2 condition is the most suitable lighting strategy in enhancing productivity and nutritional quality of purslane.

The genome sequence of Hirschfeldia incana, a new Brassicaceae model to improve photosynthetic light-use efficiency.

Photosynthesis is a key process in sustaining plant and human life. Improving the photosynthetic capacity of agricultural crops is an attractive means to increase their yields. While the core mechanisms of photosynthesis are highly conserved in C3 plants, these mechanisms are very flexible, allowing considerable diversity in photosynthetic properties. Among this diversity is the maintenance of high photosynthetic light-use efficiency at high irradiance as identified in a small number of exceptional C3 species. Hirschfeldia incana, a member of the Brassicaceae family, is such an exceptional species, and because it is easy to grow, it is an excellent model for studying the genetic and physiological basis of this trait. Here, we present a reference genome of H. incana and confirm its high photosynthetic light-use efficiency. While H. incana has the highest photosynthetic rates found so far in the Brassicaceae, the light-saturated assimilation rates of closely related Brassica rapa and Brassica nigra are also high. The H. incana genome has extensively diversified from that of B. rapa and B. nigra through large chromosomal rearrangements, species-specific transposon activity, and differential retention of duplicated genes. Duplicated genes in H. incana, B. rapa, and B. nigra that are involved in photosynthesis and/or photoprotection show a positive correlation between copy number and gene expression, providing leads into the mechanisms underlying the high photosynthetic efficiency of these species. Our work demonstrates that the H. incana genome serves as a valuable resource for studying the evolution of high photosynthetic light-use efficiency and enhancing photosynthetic rates in crop species.

Combining NDVI, PRI and the quantum yield of solar-induced fluorescence improves estimations of carbon fluxes in deciduous and evergreen forests

We used automated spectroradiometers to continuously monitor changes in the optical parameters of phenological and photosynthetic traits in beech and spruce forests. We examined seasonal variations in the normalized difference vegetation index (NDVI), photochemical reflectance index (PRI), and solar-induced fluorescence in the oxygen A band (SIFA) that was estimated using a 3-FLD discrimination method from radiance data. The optical parameters tracked the activation and cessation of photosynthesis in spring and autumn. Data at photon fluxes >1200 μmol m−2 s−1 during extended noon hours were used to link the seasonal PRI and SIFA variations to the dynamics of photosynthesis. Seasonal PRI was significantly correlated with photosynthetic light-use efficiency (LUE) with R2 values of 0.66 and 0.48 for the measurements in beech and spruce forests, respectively. SIFA emissions were significantly correlated with the gross primary production (GPP) of the evergreen spruce forest (R2 = 0.47), but R2 was only 0.13 when measured in the beech forest. The correlations between the optical parameters and GPP or LUE, however, tended to be lower when using a dataset with constant NDVI. Introducing an equation combining NDVI, PRI, and the quantum yield of SIFA emission increased R2 for LUE estimation to 0.77 in the spruce forest and 0.63 in the beech forest. GPP was estimated from the parametric equation with improved accuracy reaching R2 = 0.53 and RMSE = 5.95 μmol CO2 m−2 s−1 in spruce forest and R2 = 0.58 and RMSE = 5.23 μmol CO2 m−2 s−1 in beech forest. Parametric equations were more efficient in estimating photosynthesis in datasets that consisted of an entire season's data. By combining NDVI, PRI and the quantum yield of SIFA, we could thus substantially improve estimations of carbon fluxes in diverse deciduous and evergreen canopies.

EVALUATING APPROACHES RELATING ECOSYSTEM PRODUCTIVITY WITH DESIS SPECTRAL INFORMATION

Abstract. Data from the DLR Earth Sensing Imaging Spectrometer (DESIS), mounted on the International Space Station (ISS), were used to develop and test algorithms for remotely retrieving ecosystem productivity. Twenty DESIS images were used from three widely separated forested study sites representing deciduous and conifer forests. Gross primary production (GPP) values from eddy covariance flux towers at the sites were matched with DESIS spectral reflectances collected on the same days. Multiple algorithms were successful relating spectral reflectance with GPP, including: spectral vegetation indices (SVI) sensitive to chlorophyll content, SVI used in a photosynthetic light-use efficiency model framework, spectral shape characteristics through spectral derivatives and absorption feature analysis, and statistical models leading to multiband hyperspectral indices from partial least squares regression. Successful algorithms were able to achieve R2 better than 0.7 using a diverse set of observations combining data from different sites from multiple years and at multiple times during the year. The demonstrated robustness of the algorithms provides some confidence in using DESIS imagery to map spatial patterns of GPP.

Optimization of Photosynthetic Photon Flux Density and Light Quality for Increasing Radiation-Use Efficiency in Dwarf Tomato under LED Light at the Vegetative Growth Stage.

Dwarf tomatoes are advantageous when cultivated in a plant factory with artificial light because they can grow well in a small volume. However, few studies have been reported on cultivation in a controlled environment for improving productivity. We performed two experiments to investigate the effects of photosynthetic photon flux density (PPFD; 300, 500, and 700 μmol m−2 s−1) with white light and light quality (white, R3B1 (red:blue = 3:1), and R9B1) with a PPFD of 300 μmol m−2 s−1 on plant growth and radiation-use efficiency (RUE) of a dwarf tomato cultivar (‘Micro-Tom’) at the vegetative growth stage. The results clearly demonstrated that higher PPFD leads to higher dry mass and lower specific leaf area, but it does not affect the stem length. Furthermore, high PPFD increased the photosynthetic rate (Pn) of individual leaves but decreased RUE. A higher blue light proportion inhibited dry mass production with the same intercepted light because the leaves under high blue light proportion had low Pn and photosynthetic light-use efficiency. In conclusion, 300 μmol m−2 s−1 PPFD and R9B1 are the recommended proper PPFD and light quality, respectively, for ‘Micro-Tom’ cultivation at the vegetative growth stage to increase the RUE.

Impacts of Reduced Nitrate Supply on Nitrogen Metabolism, Photosynthetic Light-Use Efficiency, and Nutritional Values of Edible Mesembryanthemum crystallinum.

Mesembryanthemum crystallinum (common ice plant), as a nutritious ready-to-eat salad in Singapore, has become popular in recent years. However, basic data about the impacts of NO3– supply on its NO3– accumulation and nutritional quality are lacking. In this study, all plants were first grown indoor hydroponically in 10% artificial seawater (ASW) with modified full-strength Netherlands Standard Composition nutrient solution for 11 days, before transferring them to different reduced NO3– solutions. All plants grew well and healthy after 7 days of treatment. However, plants grown with 3/4 N and 1/2 N were bigger with higher shoot and root fresh weight (FW), greater leaf number, and total leaf area (TLA) than those grown with full nitrogen (N), 1/4 N, and 0 N. Mesembryanthemum crystallinum grown with full N, 3/4 N, and 1/4 N had similar specific leaf area (SLA), while 0 N plants had significantly lower SLA. All plants had similar leaf succulence (LS). However, leaf water content (LWC) was lower, while leaf dry matter accumulation (LDMC) was higher in 0 N plants after 7 days of treatment. Compared with plants grown with full N, shoot NO3– concentrations in 3/4 N, 1/2 N, and 1/4 N plants were constant or slightly increased during the treatments. For 0 N plants, shoot NO3– concentration decreased significantly during the treatment compared with other plants. Shoot NO3– accumulation was associated with nitrate reductase activity (NRA). For instance, after 7 days of treatment, shoot NO3– concentration and NRA on a FW basis in 0 N plants were, respectively, 45 and 31% of full N plants. After transferring full N to 0 N for 7 days, all M. crystallinum had higher chlorophyll (Chl) content coupled with higher electron transport rate (ETR) and higher effective quantum yield of PSII, while full N plants had higher non-photochemical quenching (NPQ). The 0N plants had much higher concentrations of proline, total soluble sugar (TSS), and total ascorbic acid (ASC) than other plants. In conclusion, totally withdrawing NO3– from the growth media prior to harvest could be one of the strategies to reduce shoot NO3– concentration. Reduced NO3– supply further enhanced nutritional values as concentrations of proline, TSS, and ASC were enhanced markedly in M. crystallinum plants after transferring them from full N to 0 N.

Light, Not Age, Underlies the Maladaptation of Maize and Miscanthus Photosynthesis to Self-Shading.

Zea mays and Miscanthus × giganteus use NADP-ME subtype C4 photosynthesis and are important food and biomass crops, respectively. Both crops are grown in dense stands where shaded leaves can contribute a significant proportion of overall canopy productivity. This is because shaded leaves, despite intercepting little light, typically process light energy very efficiently for photosynthesis, when compared to light-saturated leaves at the top of the canopy. However, an apparently maladaptive loss in photosynthetic light-use efficiency as leaves become shaded has been shown to reduce productivity in these two species. It is unclear whether this is due to leaf aging or progressive shading from leaves forming above. This was resolved here by analysing photosynthesis in leaves of the same chronological age in the centre and exposed southern edge of field plots of these crops. Photosynthetic light-response curves were used to assess maximum quantum yield of photosynthesis; the key measure of photosynthetic capacity of a leaf in shade. Compared to the upper canopy, maximum quantum yield of photosynthesis of lower canopy leaves was significantly reduced in the plot centre; but increased slightly at the plot edge. This indicates loss of efficiency of shaded leaves is due not to aging, but to the altered light environment of the lower canopy, i.e., reduced light intensity and/or altered spectral composition. This work expands knowledge of the cause of this maladaptive shade response, which limits productivity of some of the world’s most important crops.

Supplemental Light-Emitting Diode Inter-Lighting Increases Tomato Fruit Growth Through Enhanced Photosynthetic Light Use Efficiency and Modulated Root Activity.

We investigated the effect of supplemental LED inter-lighting (80% red, 20% blue; 70 W m−2; light period 04:00–22:00) on the productivity and physiological traits of tomato plants (Flavance F1) grown in an industrial greenhouse with high pressure sodium (HPS) lamps (235 W m−2, 420 µmol m−2 s−1 at canopy). Physiological trait measurements included diurnal photosynthesis and fruit relative growth rates, fruit weight at specific positions in the truss, root pressure, xylem sap hormone and ion compositions, and fruit quality. In the control treatment with HPS lamps alone, the ratio of far-red to red light (FR:R) was 1.2 at the top of the canopy and increased to 5.4 at the bottom. The supplemental LED inter-lighting decreased the FR:R ratio at the middle and low positions in the canopy and was associated with greener leaves and higher photosynthetic light use efficiency (PLUE) in the leaves in the lower canopy. The use of LED inter-lighting increased the biomass and yield by increasing the fruit weight and enhancing plant growth. The PLUE of plants receiving supplemental LED light decreased at the end of the light period, indicating that photosynthesis of the supplemented plants at the end of the day might be limited by sink capacity. The supplemental LED lighting increased the size of fruits in the middle and distal positions of the truss, resulting in a more even size for each fruit in the truss. Diurnal analysis of fruit growth showed that fruits grew more quickly during the night on the plants receiving LED light than on unsupplemented control plants. This faster fruit growth during the night was related to an increased root pressure. The LED treatment also increased the xylem levels of the phytohormone jasmonate. Supplemental LED inter-lighting increased tomato fruit weight without affecting the total soluble solid contents in fruits by increasing the total assimilates available for fruit growth and by enhancing root activity through an increase in root pressure and water supply to support fruit growth during the night.

Photosynthetic light‐use efficiency of rice leaves under fluctuating incident light

AbstractThe responsiveness of photosynthesis to continuously fluctuating light intensities can provide an estimate of the net assimilation rate (A) and photosynthetic light‐use efficiency (PLUE), which cannot be measured under steady‐state conditions. This study was conducted to investigate whether the leaves of rice (Oryza sativa L.) ‘Koshihikari’ and ‘Akitakomachi’ can efficiently utilize light under constantly fluctuating incident photosynthetic photon flux density (PPFD) and also to determine the effect of leaf N concentration on the acclimation of photosynthesis to changing PPFD. To this end, the instantaneous A and continually fluctuating incident PPFD were measured for 16 youngest fully expanded leaves of individual plants from dawn to dusk at the vegetative and reproductive phases under open‐air field conditions using two portable photosynthesis systems. The results revealed that (a) daily PLUE and calculated as the slope of the linear regression of A for values of PPFD between the light compensation point and 200 μmol m−2 s−1 can be improved with leaf N; (b) rice leaves do not utilize light efficiently under constantly fluctuating light conditions; and (c) the stomatal conductance per unit PPFD decreases under higher daily incident PPFD and diurnal fluctuation. Overall, our findings will help in refining crop photosynthesis models and would assist with the selection of high‐PLUE lines in breeding programs.

Estimating global gross primary productivity using chlorophyll fluorescence and a data assimilation system with the BETHY-SCOPE model

Abstract. This paper presents the assimilation of solar-induced chlorophyll fluorescence (SIF) into a terrestrial biosphere model to estimate the gross uptake of carbon through photosynthesis (GPP). We use the BETHY-SCOPE model to simulate both GPP and SIF using a process-based formulation, going beyond a simple linear scaling between the two. We then use satellite SIF data from the Orbiting Carbon Observatory-2 (OCO-2) for 2015 in the data assimilation system to constrain model biophysical parameters and GPP. The assimilation results in considerable improvement in the fit between model and observed SIF, despite a limited capability to fit regions with large seasonal variability in SIF. The SIF assimilation increases global GPP by 31 % to 167±5 Pg C yr−1 and shows an improvement in the global distribution of productivity relative to independent estimates, but a large difference in magnitude. This change in global GPP is driven by an overall increase in photosynthetic light-use efficiency across almost all biomes and more minor, regionally distinct changes in APAR. This process-based data assimilation opens up new pathways to the effective utilization of satellite SIF data to improve our understanding of the global carbon cycle.

Drought impacts on terrestrial primary production underestimated by satellite monitoring

Satellite retrievals of information about the Earth’s surface are widely used to monitor global terrestrial photosynthesis and primary production and to examine the ecological impacts of droughts. Methods for estimating photosynthesis from space commonly combine information on vegetation greenness, incoming radiation, temperature and atmospheric demand for water (vapour-pressure deficit), but do not account for the direct effects of low soil moisture. They instead rely on vapour-pressure deficit as a proxy for dryness, despite widespread evidence that soil moisture deficits have a direct impact on vegetation, independent of vapour-pressure deficit. Here, we use a globally distributed measurement network to assess the effect of soil moisture on photosynthesis, and identify a common bias in an ensemble of satellite-based estimates of photosynthesis that is governed by the magnitude of soil moisture effects on photosynthetic light-use efficiency. We develop methods to account for the influence of soil moisture and estimate that soil moisture effects reduce global annual photosynthesis by ~15%, increase interannual variability by more than 100% across 25% of the global vegetated land surface, and amplify the impacts of extreme events on primary production. These results demonstrate the importance of soil moisture effects for monitoring carbon-cycle variability and drought impacts on vegetation productivity from space. Soil moisture effects can substantially reduce photosynthesis and amplify the impacts of extreme events on primary production, potentially leading to biases in satellite-based estimates of photosynthesis, suggests an analysis of ground-based measurements.

Quality of supplementary LED lighting effects on growth and photosynthesis of two different Lactuca recombinant inbred lines (RILs) grown in a tropical greenhouse

LED lamps with various combinations of red (R) and blue (B) wavelengths were used to supplement sunlight for the growth of a heat-resistant (HR) and heat-sensitive (HS) recombinant inbred lines (RIL) of lettuce. The RB-LED ratios were 100R:0B (0B), 92R:8B (8B), 84R:16B (16B), and 76R:24B (24B) with an equal PPFD of 100 μmol m−2 s−1. The greatest leaf expansion rates were observed at 8B for both genotypes. All HR-RILs had similar values of growth parameters and specific leaf area (SLA). However, higher values of growth parameters were observed in HS-RIL with 0B, 8B, and 16B than that under 24B and sunlight. Furthermore, HS-RIL had higher SLA under 0B compared to other conditions. Photosynthetic light-use efficiency and maximal oxygen evolution rate were the lowest under 8B for both genotypes. The quality of LED lighting, if provided, seemed to implicate genotype dependence, probably as a result of their different sensitivities to heat stress.

Linking stand architecture with canopy reflectance to estimate vertical patterns of light-use efficiency

Remote sensing of plant carbon uptake, or gross primary production (GPP), in a repeatable and consistent manner remains a key element of a comprehensive understanding of the role of vegetation within the global carbon cycle. To further this understanding at a landscape level or global scale, accurate remote sensing of photosynthetic light-use efficiency (LUE) is required to understand photosynthetic down-regulation and environmental constraints to plant photosynthesis. The past decade has seen advances in detecting both leaf- and canopy-level physiological stress behaviours using the photochemical reflectance index (PRI), a narrow-waveband normalized difference index that relates LUE to a xanthophyll-induced absorption feature at 531nm. To date, however, much of this research has occurred using top of canopy measurements, while our understanding of the vertical distribution of LUE within the crown is limited. In this study, we demonstrate an approach which could be used to scale photosynthetic behaviour of vegetation vertically and horizontally using estimates of vertical canopy structure obtained from terrestrial Light Detection and Ranging (LiDAR) data to predict proportions of shaded and sunlit canopy which are then linked to predictions of LUE. We apply the approach over a mature Aspen study site located in central Saskatchewan, Canada utilising full-waveform LiDAR data provided by the ground-based laser scanner system and canopy spectra obtained by the AMSPEC II spectro-radiometer. Agreement between predictions of Gross Primary Productivity (GPP) using the developed approach compared to independent observations was highly significant at hourly intervals (R2=0.80, p<0.01) under clear sky conditions. A range of LUE vertical profiles for different stand structures across the growing season were developed providing estimations of how crown structure can impact LUE vertically in the crown. We conclude with a recommendation for ongoing research to verify these types of trends using concurrently acquired, independently derived leaf LUE from photosynthesis light-response curves, and forest structure variation from LiDAR, to provide a more exact quantification of these patterns.

In Memorium: Thomas Hilker

Dr. Thomas Hilker left us on 4 September 2016 following a sudden cardiac arrest. Thomas was adevoted husband to Yhasmin, and a brother and son to his family in Germany to whom we expressour deepest sympathies. Friends and colleagues of Thomas in the remote sensing and ecologicalcommunities lament this tragic loss. During his short but stellar science career, Thomas becamea world leader in the field of carbon, water and energy exchange from the land. He pioneeredstudies in the Amazonian forests, using anisotropy information acquired from satellites to describethree-dimensional structures that linked these ecosystems functionally to climatic variation.Thomas had an extreme range of interests—from the engineering of advanced spectrometers toproviding new theories and innovative methods to process remotely sensed data. Dr. Piers Sellers,Acting Director of the Earth Sciences Division at NASA/GSFC, and Deputy Director of the Sciences andExploration Directorate wrote: “Thomas Hilker was something of a renaissance man in Earth Science.He could climb towers, measure tall trees, and calculate spectral indices in his head. Working with himwas like collaborating with two or three normal people. He had some of the best and most originalideas in remote sensing that I’ve come across, but unlike most of us, he could go get the data to provehis point. And he was always the best fun. I remember him coming to a couple of parties of ours—hewas always relaxed, humorous, charming. He could make people laugh and everyone felt so goodaround him.”Thomas obtained a Bachelor of Science degree in Forestry from the University of Applied Sciences,in Goettingen, Germany in 2000, a Master in Photogrammetry and Geoinformatics from the Universityof Applied Sciences in Stuttgart in 2002 and a PhD from the University of British Columbia (UBC) inForestry in 2008. After a three year postdoctoral position at UBC (2008–2011), he worked as a ResearchAssociate at NASA’s Goddard Space Flight Center (2011–2012). From 2012 to 2016, Thomas held aposition as Assistant Professor at Oregon State University’s College of Forestry, leading the RemoteSensing Laboratory and teaching classes in Remote Sensing and Spatial Data Analysis. In 2015 and2016 he was a visiting researcher at the National Institute for Space Studies in Brazil (Instituto Nacionalde Pesquisas Espaciais, INPE). He looked forward to starting a position as an Associate Professor ofEarth System Science and Remote Sensing at the University of Southampton, UK.Thomas became fascinated by the global carbon cycle following receipt of his Master’s degree andwas keen to utilize his geospatial skills to unlock the details of the cycle. Inspired by the linksbetween canopy reflectance and photosynthesis, Thomas designed an Automated MultiangularSpectroradiometer for Estimation of Canopy reflectance [1] and improved it in subsequentiterations [2,3] to be able to continuously monitor subtle changes in the reflected spectra from

Partitioning controls on Amazon forest photosynthesis between environmental and biotic factors at hourly to interannual timescales.

Gross ecosystem productivity (GEP) in tropical forests varies both with the environment and with biotic changes in photosynthetic infrastructure, but our understanding of the relative effects of these factors across timescales is limited. Here, we used a statistical model to partition the variability of seven years of eddy covariance-derived GEP in a central Amazon evergreen forest into two main causes: variation in environmental drivers (solar radiation, diffuse light fraction, and vapor pressure deficit) that interact with model parameters that govern photosynthesis and biotic variation in canopy photosynthetic light-use efficiency associated with changes in the parameters themselves. Our fitted model was able to explain most of the variability in GEP at hourly (R2 =0.77) to interannual (R2 =0.80) timescales. At hourly timescales, we found that 75% of observed GEP variability could be attributed to environmental variability. When aggregating GEP to the longer timescales (daily, monthly, and yearly), however, environmental variation explained progressively less GEP variability: At monthly timescales, it explained only 3%, much less than biotic variation in canopy photosynthetic light-use efficiency, which accounted for 63%. These results challenge modeling approaches that assume GEP is primarily controlled by the environment at both short and long timescales. Our approach distinguishing biotic from environmental variability can help to resolve debates about environmental limitations to tropical forest photosynthesis. For example, we found that biotically regulated canopy photosynthetic light-use efficiency (associated with leaf phenology) increased with sunlight during dry seasons (consistent with light but not water limitation of canopy development) but that realized GEP was nonetheless lower relative to its potential efficiency during dry than wet seasons (consistent with water limitation of photosynthesis in given assemblages of leaves). This work highlights the importance of accounting for differential regulation of GEP at different timescales and of identifying the underlying feedbacks and adaptive mechanisms.

The influence of leaf anatomy on the internal light environment and photosynthetic electron transport rate: exploration with a new leaf ray tracing model.

Leaf photosynthesis is determined by biochemical properties and anatomical features. Here we developed a three-dimensional leaf model that can be used to evaluate the internal light environment of a leaf and its implications for whole-leaf electron transport rates (J). This model includes (i) the basic components of a leaf, such as the epidermis, palisade and spongy tissues, as well as the physical dimensions and arrangements of cell walls, vacuoles and chloroplasts; and (ii) an efficient forward ray-tracing algorithm, predicting the internal light environment for light of wavelengths between 400 and 2500nm. We studied the influence of leaf anatomy and ambient light on internal light conditions and J The results show that (i) different chloroplasts can experience drastically different light conditions, even when they are located at the same distance from the leaf surface; (ii) bundle sheath extensions, which are strips of parenchyma, collenchyma or sclerenchyma cells connecting the vascular bundles with the epidermis, can influence photosynthetic light-use efficiency of leaves; and (iii) chloroplast positioning can also influence the light-use efficiency of leaves. Mechanisms underlying leaf internal light heterogeneity and implications of the heterogeneity for photoprotection and for the convexity of the light response curves are discussed.

Seasonality of temperate forest photosynthesis and daytime respiration.

Terrestrial ecosystems currently offset one-quarter of anthropogenic carbon dioxide (CO2) emissions because of a slight imbalance between global terrestrial photosynthesis and respiration. Understanding what controls these two biological fluxes is therefore crucial to predicting climate change. Yet there is no way of directly measuring the photosynthesis or daytime respiration of a whole ecosystem of interacting organisms; instead, these fluxes are generally inferred from measurements of net ecosystem-atmosphere CO2 exchange (NEE), in a way that is based on assumed ecosystem-scale responses to the environment. The consequent view of temperate deciduous forests (an important CO2 sink) is that, first, ecosystem respiration is greater during the day than at night; and second, ecosystem photosynthetic light-use efficiency peaks after leaf expansion in spring and then declines, presumably because of leaf ageing or water stress. This view has underlain the development of terrestrial biosphere models used in climate prediction and of remote sensing indices of global biosphere productivity. Here, we use new isotopic instrumentation to determine ecosystem photosynthesis and daytime respiration in a temperate deciduous forest over a three-year period. We find that ecosystem respiration is lower during the day than at night-the first robust evidence of the inhibition of leaf respiration by light at the ecosystem scale. Because they do not capture this effect, standard approaches overestimate ecosystem photosynthesis and daytime respiration in the first half of the growing season at our site, and inaccurately portray ecosystem photosynthetic light-use efficiency. These findings revise our understanding of forest-atmosphere carbon exchange, and provide a basis for investigating how leaf-level physiological dynamics manifest at the canopy scale in other ecosystems.