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Summary: Understanding photosynthesis in natural, dynamic light environments requires knowledge of long‐term acclimation, short‐term responses, and their mechanistic interactions. To approach the latter, we systematically determined and characterized light‐environmental effects on thylakoid ion transport‐mediated short‐term responses during light fluctuations.For this, Arabidopsis thaliana wild‐type and mutants of the Cl− channel VCCN1 and the K+ exchange antiporter KEA3 were grown under eight different light environments and characterized for photosynthesis‐associated parameters and factors in steady state and during light fluctuations. For a detailed characterization of selected light conditions, we monitored ion flux dynamics at unprecedented high temporal resolution by a modified spectroscopy approach.Our analyses reveal that daily light intensity sculpts photosynthetic capacity as a main acclimatory driver with positive and negative effects on the function of KEA3 and VCCN1 during high‐light phases, respectively. Fluctuations in light intensity boost the accumulation of the photoprotective pigment zeaxanthin (Zx). We show that KEA3 suppresses Zx accumulation during the day, which together with its direct proton transport activity accelerates photosynthetic transition to lower light intensities.In summary, both light‐environment factors, intensity and variability, modulate the function of thylakoid ion transport in dynamic photosynthesis with distinct effects on lumen pH, Zx accumulation, photoprotection, and photosynthetic efficiency. [ABSTRACT FROM AUTHOR]

In nature, soil salinity and fluctuating light (FL) often occur concomitantly. However, it is unknown whether salt stress interacts with FL on leaf photosynthesis, architecture, biochemistry, pigmentation, mineral concentrations, as well as whole-plant biomass. To elucidate this, tomato (Solanum lycopersicum) seedlings were grown under constant light (C, 200 μmol m−2 s−1) or FL (5-650 μmol m−2 s−1), in combination with no (0 mM NaCl) or moderate (80 mM NaCl) salinity, for 14 days, at identical photoperiods and daily light integrals. FL and salt stress had separate effects on leaf anatomy, biochemistry and photosynthetic capacity: FL reduced leaf thickness as well as nitrogen, chlorophyll and carotenoid contents per unit leaf area, but rarely affected steady-state and dynamic photosynthetic properties along with abundance of key proteins in the electron transport chain. Salt stress, meanwhile, mainly disorganized chloroplast grana stacking, reduced stomatal density, size and aperture as well as photosynthetic capacity. Plant biomass was affected interactively by light regime and salt stress: FL reduced biomass in salt stressed plants by 17%, but it did not affect biomass of non-stressed plants. Our results stress the importance of considering FL when inferring effects of salt-stress on photosynthesis and productivity under fluctuating light intensities. [ABSTRACT FROM AUTHOR]

The role of nitric oxide (NO) in abscisic acid (ABA)-induced stomatal closure is a matter of debate. We conducted experiments in Vicia faba leaves using NO gas and sodium nitroprusside (SNP), a NO-donor compound, and compared their effects to those of ABA. In epidermal strips, stomatal closure was induced by ABA but not by NO, casting doubt on the role of NO in ABA-mediated stomatal closure. Leaf discs and intact leaves showed a dual dose response to NO: stomatal aperture widened at low dosage and narrowed at high dosage. Overcoming stomatal resistance by means of high CO2 concentration ([CO2]) restored photosynthesis in ABA-treated leaf discs but not in those exposed to NO. NO inhibited photosynthesis immediately, causing an instantaneous increase in intercellular [CO2] (C i), followed by stomatal closure. However, lowering C i by using low ambient [CO2] showed that it was not the main factor in NO-induced stomatal closure. In intact leaves, the rate of stomatal closure in response to NO was about one order of magnitude less than after ABA application. Because of the different kinetics of photosynthesis and stomatal closure that were observed, we conclude that NO is not likely to be the key factor in ABA-induced rapid stomatal closure, but that it fine-tunes stomatal aperture via different pathways. [ABSTRACT FROM AUTHOR]

Plants are often subjected to rapidly alternating light intensity and quality. While both short‐ and long‐term changes in red and blue light affect leaf photosynthesis, their impact on dynamic photosynthesis is not well documented. It was tested how dynamic and steady‐state photosynthetic traits were affected by red/blue ratios, either during growth or during measurements, in tomato leaves. Four red/blue ratios were used: monochromatic red (R100), monochromatic blue (B100), a red/blue light ratio of 9:1 (R90B10) and a red/blue light ratio of 7:3 (R70B30). R100 grown leaves showed decreased photosynthetic capacity (maximum rates of light‐saturated photosynthesis, carboxylation, electron transport and triose phosphate use), leaf thickness and nitrogen concentrations. Acclimation to various red/blue ratios had limited effects on photosynthetic induction in dark‐adapted leaves. B100‐grown leaves had a approximately 15% larger initial NPQ transient than the other treatments, which may be beneficial for photoprotection under fluctuating light. B100‐grown leaves also showed faster stomatal closure when exposed to low light intensity, which likely resulted from smaller stomata and higher stomatal density. When measured under different red/blue ratios, stomatal opening rate and photosynthetic induction rate were hardly accelerated by increased fractions of blue light in both growth chamber‐grown leaves and greenhouse‐grown leaves. However, steady‐state photosynthesis rate 30 min after photosynthetic induction was strongly reduced in leaves exposed to B100 during the measurement. We conclude that varying red/blue light ratios during growth and measurement strongly affects steady‐state photosynthesis, but has limited effects on photosynthetic induction rate. [ABSTRACT FROM AUTHOR]

Plants in natural environments are often exposed to fluctuations in light intensity, and leaf‐level acclimation to light may be affected by those fluctuations. Concurrently, leaves acclimated to a given light climate can become progressively shaded as new leaves emerge and grow above them. Acclimation to shade alters characteristics such as photosynthetic capacity. To investigate the interaction of fluctuating light and progressive shading, we exposed three‐week old tomato (Solanum lycopersicum) plants to either lightflecks or constant light intensities. Lightflecks of 20 s length and 1000 μmol m−2 s−1 peak intensity were applied every 5 min for 16 h per day, for 3 weeks. Lightfleck and constant light treatments received identical daily light sums (15.2 mol m−2 day−1). Photosynthesis was monitored in leaves 2 and 4 (counting from the bottom) during canopy development throughout the experiment. Several dynamic and steady‐state characteristics of photosynthesis became enhanced by fluctuating light when leaves were partially shaded by the upper canopy, but much less so when they were fully exposed to lightflecks. This was the case for CO2‐saturated photosynthesis rates in leaves 2 and 4 growing under lightflecks 14 days into the treatment period. Also, leaf 2 of plants in the lightfleck treatment showed significantly faster rates of photosynthetic induction when exposed to a stepwise change in light intensity on day 15. As the plants grew larger and these leaves became increasingly shaded, acclimation of leaf‐level photosynthesis to lightflecks disappeared. These results highlight continuous acclimation of leaf photosynthesis to changing light conditions inside developing canopies. [ABSTRACT FROM AUTHOR]

Abstract: A dynamic model of leaf CO2 assimilation was developed as an extension of the canonical steady‐state model, by adding the effects of energy‐dependent non‐photochemical quenching (qE), chloroplast movement, photoinhibition, regulation of enzyme activity in the Calvin cycle, metabolite concentrations, and dynamic CO2 diffusion. The model was calibrated and tested successfully using published measurements of gas exchange and chlorophyll fluorescence on Arabidopsis thaliana ecotype Col‐0 and several photosynthetic mutants and transformants affecting the regulation of Rubisco activity (rca‐2 and rwt43), non‐photochemical quenching (npq4‐1 and npq1‐2), and sucrose synthesis (spsa1). The potential improvements on CO2 assimilation under fluctuating irradiance that can be achieved by removing the kinetic limitations on the regulation of enzyme activities, electron transport, and stomatal conductance were calculated in silico for different scenarios. The model predicted that the rates of activation of enzymes in the Calvin cycle and stomatal opening were the most limiting (up to 17% improvement) and that effects varied with the frequency of fluctuations. On the other hand, relaxation of qE and chloroplast movement had a strong effect on average low‐irradiance CO2 assimilation (up to 10% improvement). Strong synergies among processes were found, such that removing all kinetic limitations simultaneously resulted in improvements of up to 32%. [ABSTRACT FROM AUTHOR]

Leaves are often exposed to fluctuating irradiance, which limits assimilation. Elevated CO2 enhances dynamic photosynthesis (i.e. photosynthesis in fluctuating irradiance) beyond its effects on steady-state photosynthesis rates. Studying the role of CO2 in dynamic photosynthesis is important for understanding plant responses to changing atmospheric CO2 partial pressures. The rise of photosynthesis after a step-wise increase to 1000 µmol m-2 s-1, the loss of photosynthetic induction after irradiance decreases, and rates of photosynthesis during sinusoidal changes in irradiance were studied in tomato (Solanum lycopersicum L.) leaves, using three CO2 partial pressures (200, 400, and 800 µbar). Initial irradiance was set to 0, 50, 100, and 200 µmol m-2 s-1 to vary the initial induction state. Most responses at 200 µbar were not different from those at 400 µbar. In contrast, CO2 at 800 µbar increased the relative carbon gain by 12% after an increase in irradiance, decreased the loss of photosynthetic induction by 14%, and increased dynamic photosynthesis during sine waves by 17%, compared with 400 µbar. These effects were additional to steady-state effects of elevated CO2 on photosynthesis. The enhancement of dynamic photosynthesis rates by elevated CO2 may therefore additionally increase photosynthesis in a future, CO2-enriched climate. [ABSTRACT FROM AUTHOR]

Background and Aims Plants depend on photosynthesis for growth. In nature, factors such as temperature, humidity, CO2 partial pressure, and spectrum and intensity of irradiance often fluctuate. Whereas irradiance intensity is most influential and has been studied in detail, understanding of interactions with other factors is lacking. Methods We tested how photosynthetic induction after dark-light transitions was affected by CO2 partial pressure (20, 40, 80 Pa), leaf temperatures (15·5, 22·8, 30·5 °C), leaf-to-air vapour pressure deficits (VPDleaf-air; 0·5, 0·8, 1·6, 2·3 kPa) and blue irradiance (0-20 %) in tomato leaves (Solanum lycopersicum). Key Results Rates of photosynthetic induction strongly increased with CO2partial pressure, due to increased apparent Rubisco activation rates and reduced diffusional limitations. High leaf temperature produced slightly higher induction rates, and increased intrinsic water use efficiency and diffusional limitation. High VPDleaf-air slowed down induction rates and apparent Rubisco activation and (at 2·3 kPa) induced damped stomatal oscillations. Blue irradiance had no effect. Slower apparent Rubisco activation in elevated VPDleaf-air may be explained by low leaf internal CO2partial pressure at the beginning of induction. Conclusions The environmental factors CO2 partial pressure, temperature and VPDleaf-air had significant impacts on rates of photosynthetic induction, as well as on underlying diffusional, carboxylation and electron transport processes. Furthermore, maximizing Rubisco activation rates would increase photosynthesis by at most 6-8 % in ambient CO2 partial pressure (across temperatures and humidities), while maximizing rates of stomatal opening would increase photosynthesis by at most 1-3 %. [ABSTRACT FROM AUTHOR]

The stimulating effect of diffuse light on radiation use efficiency (RUE) of crops is often explained by the more homogeneous spatial light distribution, while rarely considering differences in temporal light distribution at leaf level. This study investigated whether diffuse light effects on crop RUE can be explained by dynamic responses of leaf photosynthesis to temporal changes of photosynthetic photon flux density (PPFD). Two Anthurium andreanum cultivars ('Pink Champion' and 'Royal Champion') were grown in two glasshouses covered by clear (control) and diffuse glass, with similar light transmission. On clear days, diffusing the light resulted in less temporal fluctuations of PPFD. Stomatal conductance (gs) varied strongly in response to transient PPFD in 'Royal Champion,' whereas it remained relatively constant in 'Pink Champion.' Instantaneous net leaf photosynthesis (Pn) in both cultivars approached steady state Pn in diffuse light treatment. In control treatment this only occurred in 'Pink Champion.' These cultivar differences were reflected by a higher RUE (8%) in 'Royal Champion' in diffuse light treatment compared with control, whereas no effect on RUE was observed in 'Pink Champion.' We conclude that the stimulating effect of diffuse light on RUE depends on the stomatal response to temporal PPFD fluctuations, which response is cultivar dependent. [ABSTRACT FROM AUTHOR]

Understanding photosynthetic acclimation to elevated CO 2 (eCO 2 ) is important for predicting plant physiology and optimizing management decisions under global climate change, but is underexplored in important horticultural crops. We grew three crops differing in stomatal density-namely chrysanthemum, tomato, and cucumber-at near-ambient CO 2 (450 μmol mol -1 ) and eCO 2 (900 μmol mol -1 ) for 6 weeks. Steady-state and dynamic photosynthetic and stomatal conductance (g s ) responses were quantified by gas exchange measurements. Opening and closure of individual stomata were imaged in situ, using a novel custom-made microscope. The three crop species acclimated to eCO 2 with very different strategies: Cucumber (with the highest stomatal density) acclimated to eCO 2 mostly via dynamic g s responses, whereas chrysanthemum (with the lowest stomatal density) acclimated to eCO 2 mostly via photosynthetic biochemistry. Tomato exhibited acclimation in both photosynthesis and g s kinetics. eCO 2 acclimation in individual stomatal pore movement increased rates of pore aperture changes in chrysanthemum, but such acclimation responses resulted in no changes in g s responses. Although eCO 2 acclimation occurred in all three crops, photosynthesis under fluctuating irradiance was hardly affected. Our study stresses the importance of quantifying eCO 2 acclimatory responses at different integration levels to understand photosynthetic performance under future eCO 2 environments., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Image-based high-throughput phenotyping promises the rapid determination of functional traits in large plant populations. However, interpretation of some traits - such as those related to photosynthesis or transpiration rates - is only meaningful if the irradiance absorbed by the measured leaves is known, which can differ greatly between different parts of the same plant and within canopies. No feasible method currently exists to rapidly measure absorbed irradiance in three-dimensional plants and canopies. We developed a method and protocols to derive absorbed irradiance at any visible part of a canopy with a thermal camera, by fitting a leaf energy balance model to transient changes in leaf temperature. Leaves were exposed to short light pulses (30 s) that were not long enough to trigger stomatal opening but strong enough to induce transient changes in leaf temperature that was proportional to the absorbed irradiance. The method was successfully validated against point measurements of absorbed irradiance in plant species with relatively simple architecture (sweet pepper, cucumber, tomato, and lettuce). Once calibrated, the model was used to produce absorbed irradiance maps from thermograms. Our method opens new avenues for the interpretation of plant responses derived from imaging techniques and can be adapted to existing high-throughput phenotyping platforms., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)