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Summary: Photoprotection in plants includes processes collectively known as nonphotochemical quenching (NPQ), which quench excess excitation‐energy in photosystem II. NPQ is triggered by acidification of the thylakoid lumen, which leads to PsbS‐protein protonation and violaxanthin de‐epoxidase activation, resulting in zeaxanthin accumulation. Despite extensive study, questions persist about the mechanisms of NPQ.We have set up a novel analytical pipeline to disentangle NPQ induction curves measured at many light intensities into a limited number of different kinetic components. To validate the method, we applied it to Chl‐fluorescence measurements, which utilised the saturating‐pulse methodology, on wild‐type (wt) and zeaxanthin‐lacking (npq1) Arabidopsis thaliana plants. NPQ induction curves in wt and npq1 can be explained by four components (α, β, γ and δ).The fastest two (β and γ) correlate with pH difference formed across the thylakoid membrane in wt and npq1. In wt, the slower component (α) appears to be due to the formation of zeaxanthin‐related quenching whilst for npq1, this component is 'replaced' by a slower component (δ), which reflects a photoinhibition‐like process that appears in the absence of zeaxanthin‐induced quenching.Expanding this approach will allow the effects of mutations and other abiotic‐stress factors to be directly probed by changes in these underlying components. [ABSTRACT FROM AUTHOR]

The article discusses the relationship between sun-induced fluorescence (SIF) and gross primary production (GPP) under heatwave conditions, challenging the notion that nonphotochemical quenching (NPQ) alters this relationship. The study suggests that environmental stresses, such as heatwaves, can disrupt the correlation between SIF and GPP due to factors like water availability and stomatal conductance. The data presented in the article highlight the complexities of energy partitioning in plants under stress conditions, emphasizing the need for further research in this area. [Extracted from the article]

SUMMARY: Algae such as diatoms and haptophytes have distinct photosynthetic pigments from plants, including a novel set of carotenoids. This includes a primary xanthophyll cycle comprised of diadinoxanthin and its de‐epoxidation product diatoxanthin that enables the switch between light harvesting and non‐photochemical quenching (NPQ)‐mediated dissipation of light energy. The enzyme responsible for the reversal of this cycle was previously unknown. Here, we identified zeaxanthin epoxidase 3 (ZEP3) from Phaeodactylum tricornutum as the candidate diatoxanthin epoxidase. Knocking out the ZEP3 gene caused a loss of rapidly reversible NPQ following saturating light exposure. This correlated with the maintenance of high concentrations of diatoxanthin during recovery in low light. Xanthophyll cycling and NPQ relaxation were restored via complementation of the wild‐type ZEP3 gene. The zep3 knockout strains showed reduced photosynthetic rates at higher light fluxes and reduced specific growth rate in variable light regimes, likely due to the mutant strains becoming locked in a light energy dissipation state. We were able to toggle the level of NPQ capacity in a time and dose dependent manner by placing the ZEP3 gene under the control of a β‐estradiol inducible promoter. Identification of this gene provides a deeper understanding of the diversification of photosynthetic control in algae compared to plants and suggests a potential target to improve the productivity of industrial‐scale cultures. Significance Statement: This work reveals the identify of the enzyme responsible for the deactivation of the xanthophyll cycle and photoprotection in diatoms. [ABSTRACT FROM AUTHOR]

SUMMARY: Photosynthetic organisms must cope with rapid fluctuations in light intensity. Nonphotochemical quenching (NPQ) enables the dissipation of excess light energy as heat under high light conditions, whereas its relaxation under low light maximizes photosynthetic productivity. We quantified variation in NPQ kinetics across a large sorghum (Sorghum bicolor) association panel in four environments, uncovering significant genetic control for NPQ. A genome‐wide association study (GWAS) confidently identified three unique regions in the sorghum genome associated with NPQ and suggestive associations in an additional 61 regions. We detected strong signals from the sorghum ortholog of Arabidopsis thaliana Suppressor Of Variegation 3 (SVR3) involved in plastid–nucleus signaling. By integrating GWAS results for NPQ across maize (Zea mays) and sorghum‐association panels, we identified a second gene, Non‐yellowing 1 (NYE1), originally studied by Gregor Mendel in pea (Pisum sativum) and involved in the degradation of photosynthetic pigments in light‐harvesting complexes. Analysis of nye1 insertion alleles in A. thaliana confirmed the effect of this gene on NPQ kinetics in eudicots. We extended our comparative genomics GWAS framework across the entire maize and sorghum genomes, identifying four additional loci involved in NPQ kinetics. These results provide a baseline for increasing the accuracy and speed of candidate gene identification for GWAS in species with high linkage disequilibrium. Significance Statement: This study identifies naturally occurring genetic variants in sorghum, which alter NPQ kinetics, of particular significance given recent evidence linking variation in NPQ kinetics to increased photosynthetic productivity and crop yield. The multi‐species GWAS approach used here—and its validation via loss of function mutant phenotype—also provides a baseline for increasing the accuracy and speed of candidate gene identification for GWAS in species with high linkage disequilibrium. [ABSTRACT FROM AUTHOR]

The article discusses the role of nonphotochemical quenching (NPQ) in the relationship between sun-induced fluorescence (SIF) and gross primary production (GPP) during heatwaves. The authors respond to criticisms by Antala et al. (2024) and present new modeling analyses to support their findings. They emphasize the importance of understanding subdiurnal dynamics and the saturation of NPQ in influencing the SIF-GPP relationship. The study highlights the need for further research to elucidate the impact of extreme environmental conditions on excitation energy distribution and carbon fluxes. [Extracted from the article]

Life on earth depends on photosynthetic primary producers that exploit sunlight to fix CO2 into biomass. Approximately half of global primary production is associated with microalgae living in aquatic environments. Microalgae also represent a promising source of biomass to complement crop cultivation, and they could contribute to the development of a more sustainable bioeconomy. Photosynthetic organisms evolved multiple mechanisms involved in the regulation of photosynthesis to respond to highly variable environmental conditions. While essential to avoid photodamage, regulation of photosynthesis results in dissipation of absorbed light energy, generating a complex trade-off between protection from stress and light-use efficiency. This work investigates the impact of the xanthophyll cycle, the light-induced reversible conversion of violaxanthin into zeaxanthin, on the protection from excess light and on biomass productivity in the marine microalgae of the genus Nannochloropsis. Zeaxanthin is shown to have an essential role in protection from excess light, contributing to the induction of nonphotochemical quenching and scavenging of reactive oxygen species. On the contrary, the overexpression of zeaxanthin epoxidase enables a faster reconversion of zeaxanthin to violaxanthin that is shown to be advantageous for biomass productivity in dense cultures in photobioreactors. These results demonstrate that zeaxanthin accumulation is critical to respond to strong illumination, but it may lead to unnecessary energy losses in light-limiting conditions and accelerating its reconversion to violaxanthin provides an advantage for biomass productivity in microalgae.

The present paper aims to open discussion on the information content, physical mechanism(s), and measuring protocols to determine the partitioning of the absorbed light energy in oxygenic photosynthetic organisms. Revisiting these questions is incited by recent findings discovering that PSII, in addition to its open and closed state, assumes a light-adapted charge-separated state and that chlorophyll a fluorescence induction (ChlF), besides the photochemical activity of PSII, reflects the structural dynamics of its reaction center complex. Thus, the photochemical quantum yield of PSII cannot be determined from the conventional ChlF-based protocol. Consequently, the codependent quantity - the quantum yield of the so-called nonregulatory constitutive nonphotochemical quenching (npq) - loses its physical meaning. Processes beyond photochemistry and regulatory npq should be identified and characterized by multifaceted studies, including ChlF. Such investigations may shed light on the putative roles of dissipation and other energy-consuming events in the stress physiology of photosynthetic machinery.

Background: Glyphosate is the most widely applied herbicide in the world. Hormesis caused by low glyphosate doses has been widely documented in many plant species. However, the specific adaptative mechanism of plants responding to glyphosate hormesis stimulation remains unclear. This study focused on the biphasic relationship between glyphosate dose and tomato plant growth, and how glyphosate hormesis stimulates plant growth and enhances tolerance to environmental stress. Results: We constructed a hormesis model to describe the biphasic relationship with a maximal stimulation (MAX) of 162% above control by glyphosate at 0.063 g ha−1. Low‐dose glyphosate increased photosynthetic pigment contents and improve photosynthetic efficiency, leading to plant growth stimulation. We also found that glyphosate hormesis enhanced plant tolerance to diuron (DCMU; a representative photosynthesis inhibitor) by triggering the nonphotochemical chlorophyll fluorescence quenching (NPQ) reaction to dissipate excess energy stress from photosystem II (PSII). Transcriptomic analysis and quantitative real‐time polymerase chain reaction results revealed that the photosynthesis–antenna proteins pathway was the most sensitive to glyphosate hormesis, and PsbS (encoding photosystem II subunit S), ZEP (encoding zeaxanthin epoxidase) and VDE (encoding violaxanthin de‐epoxidase) involved in NPQ played crucial roles in the plant response to glyphosate hormesis. Conclusion: These results provide novel insights into the mechanisms of plant hormesis and is meaningful to the application of glyphosate hormesis in agriculture. © 2024 Society of Chemical Industry. [ABSTRACT FROM AUTHOR]

The present paper aims to open discussion on the information content, physical mechanism(s), and measuring protocols to determine the partitioning of the absorbed light energy in oxygenic photosynthetic organisms. Revisiting these questions is incited by recent findings discovering that PSII, in addition to its open and closed state, assumes a light-adapted charge-separated state and that chlorophyll a fluorescence induction (ChlF), besides the photochemical activity of PSII, reflects the structural dynamics of its reaction center complex. Thus, the photochemical quantum yield of PSII cannot be determined from the conventional ChlF-based protocol. Consequently, the codependent quantity – the quantum yield of the so-called nonregulatory constitutive nonphotochemical quenching (npq) – loses its physical meaning. Processes beyond photochemistry and regulatory npq should be identified and characterized by multifaceted studies, including ChlF. Such investigations may shed light on the putative roles of dissipation and other energyconsuming events in the stress physiology of photosynthetic machinery. [ABSTRACT FROM AUTHOR]

Photosynthetic organisms use sunlight as the primary energy source to fix CO2 . However, in nature, light energy is highly variable, reaching levels of saturation for periods ranging from milliseconds to hours. In the green microalga Chlamydomonas reinhardtii, safe dissipation of excess light energy by nonphotochemical quenching (NPQ) is mediated by light-harvesting complex stress-related (LHCSR) proteins and redistribution of light-harvesting antennae between the photosystems (state transition). Although each component underlying NPQ has been documented, their relative contributions to NPQ under fluctuating light conditions remain unknown. Here, by monitoring NPQ in intact cells throughout high light/dark cycles of various illumination periods, we find that the dynamics of NPQ depend on the timescales of light fluctuations. We show that LHCSRs play a major role during the light phases of light fluctuations and describe their role in growth under rapid light fluctuations. We further reveal an activation of NPQ during the dark phases of all high light/dark cycles and show that this phenomenon arises from state transition. Finally, we show that LHCSRs and state transition synergistically cooperate to enable NPQ response during light fluctuations. These results highlight the dynamic functioning of photoprotection under light fluctuations and open a new way to systematically characterize the photosynthetic response to an ever-changing light environment.

This study aimed to investigate the impact of diurnal and seasonal variations in photon flux density (PPFD) and air temperature on PSII efficiency in three sweet potato leaf-color cultivars: green (G), yellow-green (Y), and purple (P). The cultivars were exposed to full sunlight and measurements were taken from November to March. The maximal quantum yield of PSII photochemistry for the dark-adapted state (Fv/Fm) indicated Y's increased sensitivity to low temperatures at predawn, followed by G and P. Both quantum yield of PSII photochemistry for the dark and light-adapted state (ΔF/Fm') depressions were correlated with increased PPFD, with regression slopes in the order of Y > G > P. On high-light and low-temperature days, Fv/Fm values deviated below regression lines, with differences ranked as Y > G > P. These findings suggest that Y exhibits the highest sensitivity to high light and low temperatures, followed by G and then P in terms of PSII efficiency.

Woody plant areal encroachment is pervasive throughout the Southern Great Plains, USA. The ability of woody plants to dissipate excess solar radiation - dynamically over the day and sustained periods without recovery overnight -is key for maintaining photosynthetic performance during dry stretches, but our understanding of these processes remains incomplete. Photosynthetic performance and energy dissipation were assessed for co-occurring encroachers on the karst Edwards Plateau (Juniperus ashei, Prosopis glandulosa, and Quercus fusiformis) under seasonal changes in water status. Only J. ashei experienced mild photoinhibition from sustained energy dissipation overnight while experiencing the lowest photochemical yields, minimal photosynthetic rates, and the highest dynamic energy dissipation rates at midday during the dry period - indicating susceptibility to photosynthetic downregulation and increased dissipation under future drought regimes. Neither other encroacher experienced sustained energy dissipation in the dry period, though P. glandulosa did experience marked reductions in photosynthesis, photochemical yields, and increased regulatory dynamic energy dissipation.

Heat stress is an abiotic factor that affects the photosynthetic parameters of plants. In this study, we examined the photosynthetic mechanisms underlying the rapid response of tobacco plants to heat stress in a controlled environment. To evaluate transient heat stress conditions, changes in photochemical, carboxylative, and fluorescence efficiencies were measured using an infrared gas analyser (IRGA Licor 6800) coupled with chlorophyll a fluorescence measurements. Our findings indicated that significant disruptions in the photosynthetic machinery occurred at 45 °C for 6 h following transient heat treatment, as explained by 76.2% in the principal component analysis. The photosynthetic mechanism analysis revealed that the dark respiration rate (Rd and Rd*CO2) increased, indicating a reduced potential for carbon fixation during plant growth and development. When the light compensation point (LCP) increased as the light saturation point (LSP) decreased, this indicated potential damage to the photosystem membrane of the thylakoids. Other photosynthetic parameters, such as AMAX, VCMAX, JMAX, and ΦCO2, also decreased, compromising both photochemical and carboxylative efficiencies in the Calvin–Benson cycle. The energy dissipation mechanism, as indicated by the NPQ, qN, and thermal values, suggested that a photoprotective strategy may have been employed. However, the observed transitory damage was a result of disruption of the electron transport rate (ETR) between the PSII and PSI photosystems, which was initially caused by high temperatures. Our study highlights the impact of rapid temperature changes on plant physiology and the potential acclimatisation mechanisms under rapid heat stress. Future research should focus on exploring the adaptive mechanisms involved in distinguishing mutants to improve crop resilience against environmental stressors. [ABSTRACT FROM AUTHOR]

This study aimed to investigate the impact of diurnal and seasonal variations in photon flux density (PPFD) and air temperature on PSII efficiency in three sweet potato leaf-color cultivars: green (G), yellow-green (Y), and purple (P). The cultivars were exposed to full sunlight and measurements were taken from November to March. The maximal quantum yield of PSII photochemistry for the dark-adapted state (Fv/Fm) indicated Y's increased sensitivity to low temperatures at predawn, followed by G and P. Both quantum yield of PSII photochemistry for the dark and light- adapted state (ΔF/Fm') depressions were correlated with increased PPFD, with regression slopes in the order of Y > G > P. On high-light and low-temperature days, Fv/Fm values deviated below regression lines, with differences ranked as Y > G > P. These findings suggest that Y exhibits the highest sensitivity to high light and low temperatures, followed by G and then P in terms of PSII efficiency. [ABSTRACT FROM AUTHOR]

High light stress decreases the photosynthetic rate in plants due to photooxidative damage to photosynthetic apparatus, photoinhibition of PSII, and/or damage to PSI. The dissipation of excess energy by nonphotochemical quenching and degradation of the D1 protein of PSII and its repair cycle help against photooxidative damage. Light stress also activates stress-responsive nuclear genes through the accumulation of phosphonucleotide-3'-phosphoadenosine- 5'-phosphate, methylerythritol cyclodiphosphate, and reactive oxygen species which comprise the chloroplast retrograde signaling pathway. Additionally, hormones, such as abscisic acid, cytokinin, brassinosteroids, and gibberellins, play a role in acclimation to light fluctuations. Several alternate electron flow mechanisms, which offset the excess of electrons, include activation of plastid or plastoquinol terminal oxidase, cytochrome b6/f complex, cyclic electron flow through PSI, Mehler ascorbate peroxidase pathway or water-water cycle, mitochondrial alternative oxidase pathway, and photorespiration. In this review, we provided insights into high light stress-mediated damage to photosynthetic apparatus and strategies to mitigate the damage by decreasing antennae size, enhancing NPQ through the introduction of mutants, expression of algal proteins to improve photosynthetic rates and engineering ATP synthase.

Summary: In natural environments, plants are exposed to rapidly changing light. Maintaining photosynthetic efficiency while avoiding photodamage requires equally rapid regulation of photoprotective mechanisms. We asked what the operation frequency range of regulation is in which plants can efficiently respond to varying light.Chlorophyll fluorescence, P700, plastocyanin, and ferredoxin responses of wild‐types Arabidopsis thaliana were measured in oscillating light of various frequencies. We also investigated the npq1 mutant lacking violaxanthin de‐epoxidase, the npq4 mutant lacking PsbS protein, and the mutants crr2‐2, and pgrl1ab impaired in different pathways of the cyclic electron transport.The fastest was the PsbS‐regulation responding to oscillation periods longer than 10 s. Processes involving violaxanthin de‐epoxidase dampened changes in chlorophyll fluorescence in oscillation periods of 2 min or longer. Knocking out the PGR5/PGRL1 pathway strongly reduced variations of all monitored parameters, probably due to congestion in the electron transport. Incapacitating the NDH‐like pathway only slightly changed the photosynthetic dynamics.Our observations are consistent with the hypothesis that nonphotochemical quenching in slow light oscillations involves violaxanthin de‐epoxidase to produce, presumably, a largely stationary level of zeaxanthin. We interpret the observed dynamics of photosystem I components as being formed in slow light oscillations partially by thylakoid remodeling that modulates the redox rates. [ABSTRACT FROM AUTHOR]

Summary: Photoprotection against excess light via nonphotochemical quenching (NPQ) is indispensable for plant survival. However, slow NPQ relaxation under low light conditions can decrease yield of field‐grown crops up to 40%.Using semi‐high‐throughput assay, we quantified the kinetics of NPQ and photosystem II operating efficiency (ΦPSII) in a replicated field trial of more than 700 maize (Zea mays) genotypes across 2 yr. Parametrized kinetics data were used to conduct genome‐wide association studies.For six candidate genes involved in NPQ and ΦPSII kinetics in maize the loss of function alleles of orthologous genes in Arabidopsis (Arabidopsis thaliana) were characterized: two thioredoxin genes, and genes encoding a transporter in the chloroplast envelope, an initiator of chloroplast movement, a putative regulator of cell elongation and stomatal patterning, and a protein involved in plant energy homeostasis.Since maize and Arabidopsis are distantly related, we propose that genes involved in photoprotection and PSII function are conserved across vascular plants. The genes and naturally occurring functional alleles identified here considerably expand the toolbox to achieving a sustainable increase in crop productivity. [ABSTRACT FROM AUTHOR]

Woody plant areal encroachment is pervasive throughout the Southern Great Plains, USA. The ability of woody plants to dissipate excess solar radiation - dynamically over the day and sustained periods without recovery overnight - is key for maintaining photosynthetic performance during dry stretches, but our understanding of these processes remains incomplete. Photosynthetic performance and energy dissipation were assessed for co-occurring encroachers on the karst Edwards Plateau (Juniperus ashei, Prosopis glandulosa, and Quercus fusiformis) under seasonal changes in water status. Only J. ashei experienced mild photoinhibition from sustained energy dissipation overnight while experiencing the lowest photochemical yields, minimal photosynthetic rates, and the highest dynamic energy dissipation rates at midday during the dry period - indicating susceptibility to photosynthetic downregulation and increased dissipation under future drought regimes. Neither other encroacher experienced sustained energy dissipation in the dry period, though P. glandulosa did experience marked reductions in photosynthesis, photochemical yields, and increased regulatory dynamic energy dissipation. [ABSTRACT FROM AUTHOR]

Long-distance electrical signals caused by the local action of stressors influence numerous physiological processes in plants including photosynthesis and increase their tolerance to the action of adverse factors. Depolarization electrical signals were mainly investigated; however, we earlier showed that hyperpolarization electrical signals (HESs) can be caused by moderate stressors (e.g., local moderate heating) and induce photosynthetic inactivation. We hypothesized that HESs are related to stressor-induced increases in the hydrostatic pressure in the zone of action of the stressor and following the propagation of a hydraulic wave. In the current work, we tested this hypothesis through the direct investigation of electrical signals induced by the local action of artificially increased pressure and an analysis of the subsequent photosynthetic changes in the nonirritated parts of plants. The electrical signals and parameters of photosynthetic light reactions were investigated in wheat plants. The local action of the increased pressure was induced by the action of weights on the wheat leaf. Extracellular electrodes were used for electrical signal measurements. Pulse–amplitude–modulation fluorescent imaging was used for measurements of the quantum yield of photosystem II and nonphotochemical quenching of chlorophyll fluorescence in wheat leaves. It was shown that the local action of pressure on wheat leaf induced electrical signals near the irritated zone: HESs were caused by low pressure (10 kPa) and depolarization signals were induced by high pressure (100 kPa). The local action of moderate pressure (50 kPa) induced weak electrical signals near the irritated zone; however, HESs were observed with increasing distance from this zone. It was also shown that the local action of this moderate pressure induced the photosynthetic inactivation (decreasing the quantum yield of photosystem II and increasing the nonphotochemical quenching of chlorophyll fluorescence) in the nonirritated parts of the wheat leaves. Thus, our results show that the local action of the increased pressure and, probably, subsequent propagation of the hydraulic wave induce electrical signals (including HESs) and photosynthetic inactivation in nonirritated parts of plants that are similar to ones caused by the local action of moderate stressors (e.g., moderate heating). This means that both HESs and depolarization electrical signals can have a hydraulic mechanism of propagation. [ABSTRACT FROM AUTHOR]

Although the importance of nonphotochemical quenching (NPQ) on photosynthetic biomass production and crop yields is well established, the in vivo operation of the individual mechanisms contributing to overall NPQ is still a matter of controversy. In order to investigate the timescale and activation dynamics of specific quenching mechanisms, we have developed a technique called snapshot transient absorption (TA) spectroscopy, which can monitor molecular species involved in the quenching response with a time resolution of 30 s. Using intact thylakoid membrane samples, we show how conventional TA kinetic and spectral analyses enable the determination of the appropriate wavelength and time delay for snapshot TA experiments. As an example, we show how the chlorophyll-carotenoid charge transfer and excitation energy transfer mechanisms can be monitored based on signals corresponding to the carotenoid (Car) radical cation and Car S1 excited state absorption, respectively. The use of snapshot TA spectroscopy together with the previously reported fluorescence lifetime snapshot technique (Sylak-Glassman et al. in Photosynth Res 127:69-76, 2016) provides valuable information such as the concurrent appearance of specific quenching species and overall quenching of excited Chl. Furthermore, we show that the snapshot TA technique can be successfully applied to completely intact photosynthetic organisms such as live cells of Nannochloropsis. This demonstrates that the snapshot TA technique is a valuable method for tracking the dynamics of intact samples that evolve over time, such as the photosynthetic system in response to high-light exposure.

Photosynthetic organisms use nonphotochemical quenching (NPQ) mechanisms to dissipate excess absorbed light energy and protect themselves from photooxidation. In the model green alga Chlamydomonas reinhardtii, the capacity for rapidly reversible NPQ (qE) is induced by high light, blue light, and UV light via increased expression of LHCSR and PSBS genes that are necessary for qE. Here, we used a forward genetics approach to identify SPA1 and CUL4, components of a putative green algal E3 ubiquitin ligase complex, as critical factors in a signaling pathway that controls light-regulated expression of the LHCSR and PSBS genes in C. reinhardtii The spa1 and cul4 mutants accumulate increased levels of LHCSR1 and PSBS proteins in high light, and unlike the wild type, they express LHCSR1 and exhibit qE capacity even when grown in low light. The spa1-1 mutation resulted in constitutively high expression of LHCSR and PSBS RNAs in both low light and high light. The qE and gene expression phenotypes of spa1-1 are blocked by mutation of CrCO, a B-box Zn-finger transcription factor that is a homolog of CONSTANS, which controls flowering time in plants. CONSTANS-like cis-regulatory sequences were identified proximal to the qE genes, consistent with CrCO acting as a direct activator of qE gene expression. We conclude that SPA1 and CUL4 are components of a conserved E3 ubiquitin ligase that acts upstream of CrCO, whose regulatory function is wired differently in C. reinhardtii to control qE capacity via cis-regulatory CrCO-binding sites at key photoprotection genes.

Energy dissipation is crucial for land and shallow-water plants exposed to direct sunlight. Almost all green plants dissipate excess excitation energy to protect the photosystem reaction centers, photosystem II (PSII) and photosystem I (PSI), and continue to grow under strong light. In our previous work, we reported that about half of the photosystem reaction centers form a PSI-PSII megacomplex in Arabidopsis thaliana, and that the excess energy was transferred from PSII to PSI fast. However, the physiological function and structure of the megacomplex remained unclear. Here, we suggest that high-light adaptable sun-plants accumulate the PSI-PSII megacomplex more than shade-plants. In addition, PSI of sun-plants has a deep trap to receive excitation energy, which is low-energy chlorophylls showing fluorescence maxima longer than 730 nm. This deep trap may increase the high-light tolerance of PSI by improving excitation energy dissipation. Electron micrographs suggest that one PSII dimer is directly sandwiched between two PSIs with 2-fold rotational symmetry in the basic form of the PSI-PSII megacomplex in green plants. This structure should enable fast energy transfer from PSII to PSI and allow energy in PSII to be dissipated via the deep trap in PSI.

Simple Summary: This study investigates the spatial analysis of morphological and chemical changes and how reflectance hyperspectroscopy and fluorescence kinetics spectroscopy can enhance our understanding of biophysical, biochemical, and photochemical changes in Codiaeum variegatum (L.) A. Juss, a plant with variegated leaves and different pigments. The analysis included pigment profiling, hyperspectral curves, chlorophyll a fluorescence induction kinetics, and multivariate analyses associated with 23 JIP test parameters and 34 vegetation indexes. The results show that the analysis of chemical composition combined with vegetation indexes, such as PRI, PSSRc, ARI1, RARS, and SIPI, are highly correlated with pigment concentration and photochemical components of photosystems in leaves. Furthermore, decreased damage to energy transfer in the electron transport chain is associated with the accumulation of carotenoids, anthocyanins, flavonoids, and phenolic compounds linked with specific wavelengths. Our results reveal the potential of optical spectroscopy techniques and multivariate data analyses to enhance the management and monitoring of the leaf color status of plants. This is the first study and report on the monitoring of nonuniform leaves, particularly in the alteration of photosystem changes in variegated leaves together with high-throughput analyses. The adjustments that occur during photosynthesis are correlated with morphological, biochemical, and photochemical changes during leaf development. Therefore, monitoring leaves, especially when pigment accumulation occurs, is crucial for monitoring organelles, cells, tissue, and whole-plant levels. However, accurately measuring these changes can be challenging. Thus, this study tests three hypotheses, whereby reflectance hyperspectroscopy and chlorophyll a fluorescence kinetics analyses can improve our understanding of the photosynthetic process in Codiaeum variegatum (L.) A. Juss, a plant with variegated leaves and different pigments. The analyses include morphological and pigment profiling, hyperspectral data, chlorophyll a fluorescence curves, and multivariate analyses using 23 JIP test parameters and 34 different vegetation indexes. The results show that photochemical reflectance index (PRI) is a useful vegetation index (VI) for monitoring biochemical and photochemical changes in leaves, as it strongly correlates with chlorophyll and nonphotochemical dissipation (Kn) parameters in chloroplasts. In addition, some vegetation indexes, such as the pigment-specific simple ratio (PSSRc), anthocyanin reflectance index (ARI1), ratio analysis of reflectance spectra (RARS), and structurally insensitive pigment index (SIPI), are highly correlated with morphological parameters and pigment levels, while PRI, moisture stress index (MSI), normalized difference photosynthetic (PVR), fluorescence ratio (FR), and normalized difference vegetation index (NDVI) are associated with photochemical components of photosynthesis. Combined with the JIP test analysis, our results showed that decreased damage to energy transfer in the electron transport chain is correlated with the accumulation of carotenoids, anthocyanins, flavonoids, and phenolic compounds in the leaves. Phenomenological energy flux modelling shows the highest changes in the photosynthetic apparatus based on PRI and SIPI when analyzed with Pearson's correlation, the hyperspectral vegetation index (HVI) algorithm, and the partial least squares (PLS) to select the most responsive wavelengths. These findings are significant for monitoring nonuniform leaves, particularly when leaves display high variation in pigment profiling in variegated and colorful leaves. This is the first study on the rapid and precise detection of morphological, biochemical, and photochemical changes combined with vegetation indexes for different optical spectroscopy techniques. [ABSTRACT FROM AUTHOR]

Heat stress is an abiotic factor that affects the photosynthetic parameters of plants. In this study, we examined the photosynthetic mechanisms underlying the rapid response of tobacco plants to heat stress in a controlled environment. To evaluate transient heat stress conditions, changes in photochemical, carboxylative, and fluorescence efficiencies were measured using an infrared gas analyser (IRGA Licor 6800) coupled with chlorophyll a fluorescence measurements. Our findings indicated that significant disruptions in the photosynthetic machinery occurred at 45 °C for 6 h following transient heat treatment, as explained by 76.2% in the principal component analysis. The photosynthetic mechanism analysis revealed that the dark respiration rate (Rd and Rd*CO2) increased, indicating a reduced potential for carbon fixation during plant growth and development. When the light compensation point (LCP) increased as the light saturation point (LSP) decreased, this indicated potential damage to the photosystem membrane of the thylakoids. Other photosynthetic parameters, such as AMAX, VCMAX, JMAX, and ΦCO2, also decreased, compromising both photochemical and carboxylative efficiencies in the Calvin–Benson cycle. The energy dissipation mechanism, as indicated by the NPQ, qN, and thermal values, suggested that a photoprotective strategy may have been employed. However, the observed transitory damage was a result of disruption of the electron transport rate (ETR) between the PSII and PSI photosystems, which was initially caused by high temperatures. Our study highlights the impact of rapid temperature changes on plant physiology and the potential acclimatisation mechanisms under rapid heat stress. Future research should focus on exploring the adaptive mechanisms involved in distinguishing mutants to improve crop resilience against environmental stressors.

Nonphotochemical quenching (NPQ) is a proxy for photoprotective thermal dissipation processes that regulate photosynthetic light harvesting. The identification of NPQ mechanisms and their molecular or physiological triggering factors under in vivo conditions is a matter of controversy. Here, to investigate chlorophyll (Chl)-zeaxanthin (Zea) excitation energy transfer (EET) and charge transfer (CT) as possible NPQ mechanisms, we performed transient absorption (TA) spectroscopy on live cells of the microalga Nannochloropsis oceanica We obtained evidence for the operation of both EET and CT quenching by observing spectral features associated with the Zea S1 and Zea●+ excited-state absorption (ESA) signals, respectively, after Chl excitation. Knockout mutants for genes encoding either violaxanthin de-epoxidase or LHCX1 proteins exhibited strongly inhibited NPQ capabilities and lacked detectable Zea S1 and Zea●+ ESA signals in vivo, which strongly suggests that the accumulation of Zea and active LHCX1 is essential for both EET and CT quenching in N. oceanica.

An important determinant of crop yields is the regulation of photosystem II (PSII) light harvesting by energy-dependent quenching (qE). However, the molecular details of excitation quenching have not been quantitatively connected to the fraction of excitations converted to chemical energy by PSII reaction centers (PSII yield), which determines flux to downstream metabolism. Here, we incorporate excitation dissipation by qE into a pigment-scale model of excitation transfer and trapping for a 200 × 200-nm patch of the grana membrane. We show that excitation transport can be rigorously coarse grained to a 2D random walk with an excitation diffusion length determined by the extent of quenching. We present an alternative method for analyzing pulse amplitude-modulated chlorophyll fluorescence measurements that incorporates the effects of a variable excitation diffusion length during qE activation.

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]

High light stress decreases the photosynthetic rate in plants due to photooxidative damage to photosynthetic apparatus, photoinhibition of PSII, and/or damage to PSI. The dissipation of excess energy by nonphotochemical quenching and degradation of the D1 protein of PSII and its repair cycle help against photooxidative damage. Light stress also activates stress-responsive nuclear genes through the accumulation of phosphonucleotide-3'-phosphoadenosine-5'-phosphate, methylerythritol cyclodiphosphate, and reactive oxygen species which comprise the chloroplast retrograde signaling pathway. Additionally, hormones, such as abscisic acid, cytokinin, brassinosteroids, and gibberellins, play a role in acclimation to light fluctuations. Several alternate electron flow mechanisms, which offset the excess of electrons, include activation of plastid or plastoquinol terminal oxidase, cytochrome b6/f complex, cyclic electron flow through PSI, Mehler ascorbate peroxidase pathway or water-water cycle, mitochondrial alternative oxidase pathway, and photorespiration. In this review, we provided insights into high light stress-mediated damage to photosynthetic apparatus and strategies to mitigate the damage by decreasing antennae size, enhancing NPQ through the introduction of mutants, expression of algal proteins to improve photosynthetic rates and engineering ATP synthase. [ABSTRACT FROM AUTHOR]

The ability to maintain the balance between the absorbed light energy and energy used in photosynthesis is a key factor of plant adaptation to variable environmental conditions. In this work, diurnal variations in photosynthesis, respiration, thermal energy dissipation, and the activity of the antioxidant system were studied in hoary plantain (Plantago media L.) growing on an open slope (sun plants) and under natural shading in the herbage (shade plants). The highest leaves net photosynthetic rate (Pn) was observed early in the morning and amounted to 2.6 and 9.2 μmol CO2/m2 s in shade and sun plants, respectively. In the daytime, the Pn values of sun plants decreased significantly (threefold) along with the decrease in stomatal conductance; changes of both parameters developed concurrently with the increase in insolation and air temperature. The Pn changes in leaves of shade plants were less pronounced and weakly dependent on stomatal conductance. The leaves of shade plants contained comparatively high amounts of soluble carbohydrates, whereas the sun plant leaves accumulated larger amounts of starch. In the daytime, nonphotochemical quenching (NPQ) of chlorophyll a fluorescence in photosystem II of sun plant leaves could be as large as 2.6 rel. units, which was four- to fivefold higher than NPQ in shade plants. In leaves of sun plants in the morning and evening hours, the ratio of cytochrome pathway (CP) and the alternative (AP) respiratory pathways was approximately 1.0, whereas this ratio decreased to 0.4 during the day, synchronously with an increase in NPQ. The CP/AP ratio in shade plant leaves remained constant throughout the diurnal cycle and equaled 1.4, indicating a comparatively high energy efficiency of respiration in shaded plants growing under the grass canopy. The leaves of sun plants featured an increased content of superoxide anion radical and hydrogen peroxide as well as the elevated activity of antioxidant enzymes (superoxide dismutase, guaiacol peroxidase) that control the accumulation of reactive oxygen species. The results emphasize the importance of coordinated changes in energy-dissipating processes and the activity of the antioxidant system for maintaining the energy and redox balance in phototrophic tissues during long-term adaptation of plants to environmental conditions, excessive insolation in particular. [ABSTRACT FROM AUTHOR]

Acid rain is a serious environmental problem and has obvious impacts on the growth, reproduction, and photosynthesis of terrestrial plants. Ulva prolifera, a main blooming species of green tides, was studied on its physiological response to acid rain. The photosynthetic parameters were determined under different conditions (salinity: 1, 10, 30‰; pH: 3.0, 3.5, 4.5; duration: 0.5, 1.0, 2.0 h) followed by 24-h recovering under natural conditions. Results showed 1-h treatment with pH 3.5 caused 50-70% reduction in the maximal quantum yield of PSII photochemistry (Fv/Fm) and effective quantum yield of PSII photochemistry (ФPSII) at normal salinity but when the low pH was combined with a salinity of 10‰ or lower, PSII activity was almost completely inhibited. Moreover, the low salinity (1‰ and 10‰) reduced the degree of photoprotection under low pH (3.5) conditions. Finally, we speculated if the pH of acid rain ≤ 3.5, with 1‰ salinity and 2-h rainfall time, the amount of U. prolifera and the scale of green tides would decrease.

The OJIPSMT pattern of chlorophyll (Chl) a fluorescence induction (FI) was obtained using Scenedesmus obliquus (Scenedesmus) microalgal cells exposed, after dark adaptation, to moderate and high light intensities (1200 and 1800 μmol photons m–2 s–1) in a time range from the microseconds to 10 minutes. The fast and, particularly, slow OJIPS(M)T stages of FI extending up to 100 s were quantitatively described by fitting a thylakoid membrane (TM) model. The calculations adequately reproduced the dynamics of P700 oxidoreduction transients. Parameters of TM processes were determined, and the transients of the TM model variables are shown in calculations. Dynamic stages of redox reactions in reaction centers (RCs) of photosystem II (PSII), the PQ/PQH2 quinone pool, the cytochrome b6 f complex (Cyt bf), and P700 of photosystem I (PSI) RCs were found to accompany the adaptation of the TM system to light. In silico analysis revealed the role that Cyt bf plays in regulating electron fluxes when bL/H hemes remain in a more reduced state at a high light intensity than at moderate illumination in an interval from 100 ms to 3 s. A dependence of the rate constants k42–49 on the level of reduction was revealed for non-radiative recombination of separated charges in the RC Phe–P680+ and Q P680+, together with a dependence of k42–49 on chloroplast illumination. Setting the dynamic rate constants kFNR(t) and kD-qE(t) made it possible to compare the generation ΔpHo-i(t) = pHS(t) – pHL(t), initiation of qE quenching at pHL|trigg = 6.7, activation of FNR reductase in the light, and achievement of stationary charge flows within 1–2 minutes of light induction. Fitting the model of isolated PSII to phytoplankton FI signals showed similar electron transfer parameters of PSII RCs for Scenedesmus and Chlorella monocultures and phytoplankton samples. Markers of the phytoplankton community state were an exception, differing between degrading and ecologically functional water bodies. The differences in markers included a reduced size of the antennas, an increased energy dissipation, and a decrease in pH in the lumen. [ABSTRACT FROM AUTHOR]

Photosynthetic organisms use various photoprotective mechanisms to dissipate excess photoexcitation as heat in a process called nonphotochemical quenching (NPQ). Regulation of NPQ allows for a rapid response to changes in light intensity and in vascular plants, is primarily triggered by a pH gradient across the thylakoid membrane (∆pH). The response is mediated by the PsbS protein and various xanthophylls. Time-correlated single-photon counting (TCSPC) measurements were performed on Arabidopsis thaliana to quantify the dependence of the response of NPQ to changes in light intensity on the presence and accumulation of zeaxanthin and lutein. Measurements were performed on WT and mutant plants deficient in one or both of the xanthophylls as well as a transgenic line that accumulates lutein via an engineered lutein epoxide cycle. Changes in the response of NPQ to light acclimation in WT and mutant plants were observed between two successive light acclimation cycles, suggesting that the character of the rapid and reversible response of NPQ in fully dark-acclimated plants is substantially different from in conditions plants are likely to experience caused by changes in light intensity during daylight. Mathematical models of the response of zeaxanthin- and lutein-dependent reversible NPQ were constructed that accurately describe the observed differences between the light acclimation periods. Finally, the WT response of NPQ was reconstructed from isolated components present in mutant plants with a single common scaling factor, which enabled deconvolution of the relative contributions of zeaxanthin- and lutein-dependent NPQ.

Although sunlight provides the energy necessary for plants to survive and grow, light can also damage reaction centers of photosystem II (PSII) and reduce photochemical efficiency. To prevent damage, plants possess photoprotective mechanisms that dissipate excess excitation. A subset of these mechanisms is collectively referred to as NPQ, or nonphotochemical quenching of chlorophyll a fluorescence. The regulation of NPQ is intrinsically linked to the cycling of xanthophylls that affects the kinetics and extent of the photoprotective response. The violaxanthin cycle (VAZ cycle) and the lutein epoxide cycle (LxL cycle) are two xanthophyll cycles found in vascular plants. The VAZ cycle has been studied extensively, owing in large part to its presence in model plant species where mutants are available to aid in its characterization. In contrast, the LxL cycle is not found in model plants, and its role in photosynthetic processes has been more difficult to define. To address this challenge, we introduced the LxL cycle into Arabidopsis thaliana and functionally isolated it from the VAZ cycle. Using these plant lines, we showed an increase in dark-acclimated PSII efficiency associated with Lx accumulation and demonstrated that violaxanthin deepoxidase is responsible for the light-driven deepoxidation of Lx. Conversion of Lx to L was reversible during periods of low light and occurred considerably faster than rates previously described in nonmodel species. Finally, we present clear evidence of the LxL cycle's role in modulating a rapid component of NPQ that is necessary to prevent photoinhibition in excess light.

Harvested mature berry clusters of two white table grape cultivars were used to study the effects of postharvest UV irradiation. One cultivar, 'Queen of Vineyard' (QV), had higher light-acclimated PSII quantum yield, higher phenolic contents, and stronger total antioxidant capacities than the other, 'White Sultana' (WS). These differences were maintained throughout the experiment. Responses of the two cultivars to a 30-min UV irradiation were also different. Antioxidant capacities and flavonol, especially quercetin-3-O-glucuronide, contents were lower 2 h after the UV treatment in both cultivars and recovered in QV but not in WS berry skins later on. Our data demonstrate that mature grapevine berries have photosynthetically active tissues capable of dynamic changes even several hours after harvest and suggest that changes in photochemistry may contribute to postharvest metabolic responses of berry skins. Results also support the potential of postharvest manipulation of fruit qualities with UV irradiation.

Long-distance electrical signals caused by the local action of stressors influence numerous physiological processes in plants including photosynthesis and increase their tolerance to the action of adverse factors. Depolarization electrical signals were mainly investigated; however, we earlier showed that hyperpolarization electrical signals (HESs) can be caused by moderate stressors (e.g., local moderate heating) and induce photosynthetic inactivation. We hypothesized that HESs are related to stressor-induced increases in the hydrostatic pressure in the zone of action of the stressor and following the propagation of a hydraulic wave. In the current work, we tested this hypothesis through the direct investigation of electrical signals induced by the local action of artificially increased pressure and an analysis of the subsequent photosynthetic changes in the nonirritated parts of plants. The electrical signals and parameters of photosynthetic light reactions were investigated in wheat plants. The local action of the increased pressure was induced by the action of weights on the wheat leaf. Extracellular electrodes were used for electrical signal measurements. Pulse–amplitude–modulation fluorescent imaging was used for measurements of the quantum yield of photosystem II and nonphotochemical quenching of chlorophyll fluorescence in wheat leaves. It was shown that the local action of pressure on wheat leaf induced electrical signals near the irritated zone: HESs were caused by low pressure (10 kPa) and depolarization signals were induced by high pressure (100 kPa). The local action of moderate pressure (50 kPa) induced weak electrical signals near the irritated zone; however, HESs were observed with increasing distance from this zone. It was also shown that the local action of this moderate pressure induced the photosynthetic inactivation (decreasing the quantum yield of photosystem II and increasing the nonphotochemical quenching of chlorophyll fluorescence) in the nonirritated parts of the wheat leaves. Thus, our results show that the local action of the increased pressure and, probably, subsequent propagation of the hydraulic wave induce electrical signals (including HESs) and photosynthetic inactivation in nonirritated parts of plants that are similar to ones caused by the local action of moderate stressors (e.g., moderate heating). This means that both HESs and depolarization electrical signals can have a hydraulic mechanism of propagation.

In vascular plants, bryophytes and algae, the photosynthetic light reaction takes place in the thylakoid membrane where two transmembrane supercomplexes PSII and PSI work together with cytochrome b6f and ATP synthase to harvest the light energy and produce ATP and NADPH. Vascular plant PSI is a 600-kDa protein--pigment supercomplex, the core complex of which is partly surrounded by peripheral light-harvesting complex I (LHCI) that captures sunlight and transfers the excitation energy to the core to be used for charge separation. PSI is unique mainly in absorption of longer-wavelengths than PSII, fast excitation energy transfer including uphill energy transfer, and an extremely high quantum efficiency. From the early 1980s, a lot of effort has been dedicated to structural and functional studies of PSI--LHCI, leading to the current understanding of how more than 200 cofactors are kept at the correct distance and geometry to facilitate fast energy transfer in this supercomplex at an atomic level. In this review, we review the history of studies on vascular plant PSI--LHCI, summarise the present research progress on its structure, and present some new and further questions to be answered in future studies. [ABSTRACT FROM AUTHOR]

When dormant, perennial plants dwelling in the regions with pronounced seasonality of climate can withstand prolonged periods of harsh environmental conditions. The period of plant dormancy is commonly divided into pre-dormancy, endodormancy, and ecodormancy. During pre-dormancy, genetic, physiological, biochemical, and morphological rearrangements increasing stress resilience of the plant organism are completed. In the course of endodormancy, meristem cells cannot resume division even under favorable conditions. Environmental stimuli trigger dormancy release and the onset of ecodormancy when plant cell division and growth are restrained only by unfavorable environmental conditions. Frequent nowadays, weather fluctuations can lead to abnormal progression of dormancy. It results in the increased risk of damage to plants, especially crop plants, by adverse climatic conditions. This situation calls for the development of methods for noninvasive express monitoring of plant dormancy. Studies of the relationships between the dormancy status of plants and the functioning of their photosynthetic apparatus made possible the development of methods for monitoring of woody plant condition by recording the variable fluorescence of chlorophyll contained either in needles or in the endoderm of the shoots. This review briefly summarizes current knowledge about the mechanism of the dormancy induction and release. The functioning and regulation of the photosynthetic apparatus during winter dormancy as well as characteristic patterns of chlorophyll fluorescence induction in this period are considered. The difficulties of interpretation of chlorophyll fluorescence signals in the context of monitoring of plant dormancy are discussed together with its potential applications. [ABSTRACT FROM AUTHOR]

In the high-light environment, young leaves accumulate anthocyanins as a photoprotection strategy. However, anthocyanin biosynthesis-related enzymes gene sequence is still unknown in the leaves of subtropical forest plants. There are thus few reports on the relationship between the expression level of these genes and photoprotection. In this study, Machilus chinensis and Castanopsis chinensis were taken as plant material in a subtropical forest. Non-full-length nucleotide sequences of chalcone synthase, dihydroflavonol 4-reductase, anthocyanin synthase, and β-tubulin gene were obtained by homologous and electronic cloning. The expression of those genes was verified and analyzed, and some physiological indicators were measured. The anthocyanin content and anthocyanin synthesis-related gene expression in the young leaves of C. chinensis were significantly higher than that of M. chinensis. Although young leaves of M. chinensis did not accumulate anthocyanins, they showed higher antioxidants and nonphotochemical quenching (NPQ). This study indicates that anthocyanins, antioxidants, and NPQ together mediate the positive effects on photoadaptation in young leaves.

Life on earth is dependent on the photosynthetic conversion of light energy into chemical energy. However, absorption of excess sunlight can damage the photosynthetic machinery and limit photosynthetic activity, thereby affecting growth and productivity. Photosynthetic light harvesting can be down-regulated by nonphotochemical quenching (NPQ). A major component of NPQ is qE (energy-dependent nonphotochemical quenching), which allows dissipation of light energy as heat. Photodamage peaks in the UV-B part of the spectrum, but whether and how UV-B induces qE are unknown. Plants are responsive to UV-B via the UVR8 photoreceptor. Here, we report in the green alga Chlamydomonas reinhardtii that UVR8 induces accumulation of specific members of the light-harvesting complex (LHC) superfamily that contribute to qE, in particular LHC Stress-Related 1 (LHCSR1) and Photosystem II Subunit S (PSBS). The capacity for qE is strongly induced by UV-B, although the patterns of qE-related proteins accumulating in response to UV-B or to high light are clearly different. The competence for qE induced by acclimation to UV-B markedly contributes to photoprotection upon subsequent exposure to high light. Our study reveals an anterograde link between photoreceptor-mediated signaling in the nucleocytosolic compartment and the photoprotective regulation of photosynthetic activity in the chloroplast.

Summary: Diatoms are successful phytoplankton clades able to acclimate to changing environmental conditions, including e.g. variable light intensity. Diatoms are outstanding at dissipating light energy exceeding the maximum photosynthetic electron transfer (PET) capacity via the nonphotochemical quenching (NPQ) process. While the molecular effectors of NPQ as well as the involvement of the proton motive force (PMF) in its regulation are known, the regulators of the PET/PMF relationship remain unidentified in diatoms.We generated mutants of the H+/K+ antiporter KEA3 in the model diatom Phaeodactylum tricornutum.Loss of KEA3 activity affects the PET/PMF coupling and NPQ responses at the onset of illumination, during transients and in steady‐state conditions. Thus, this antiporter is a main regulator of the PET/PMF coupling. Consistent with this conclusion, a parsimonious model including only two free components, KEA3 and the diadinoxanthin de‐epoxidase, describes most of the feedback loops between PET and NPQ.This simple regulatory system allows for efficient responses to fast (minutes) or slow (e.g. diel) changes in light environment, thanks to the presence of a regulatory calcium ion (Ca2+)‐binding domain in KEA3 modulating its activity. This circuit is likely tuned by the NPQ‐effector proteins, LHCXs, providing diatoms with the required flexibility to thrive in different ocean provinces. [ABSTRACT FROM AUTHOR]

Photosynthetic pigments are an integral and vital part of all photosynthetic machinery and are present in different types and abundances throughout the photosynthetic apparatus. Chlorophyll, carotenoids and phycobilins are the prime photosynthetic pigments which facilitate efficient light absorption in plants, algae, and cyanobacteria. The chlorophyll family plays a vital role in light harvesting by absorbing light at different wavelengths and allowing photosynthetic organisms to adapt to different environments, either in the long-term or during transient changes in light. Carotenoids play diverse roles in photosynthesis, including light capture and as crucial antioxidants to reduce photodamage and photoinhibition. In the marine habitat, phycobilins capture a wide spectrum of light and have allowed cyanobacteria and red algae to colonise deep waters where other frequencies of light are attenuated by the water column. In this review, we discuss the potential strategies that photosynthetic pigments provide, coupled with development of molecular biological techniques, to improve crop yields through enhanced light harvesting, increased photoprotection and improved photosynthetic efficiency. [ABSTRACT FROM AUTHOR]

The orange carotenoid protein (OCP) is responsible for nonphotochemical quenching (NPQ) in cyanobacteria, a defense mechanism against potentially damaging effects of excess light conditions. This soluble two‐domain protein undergoes profound conformational changes upon photoactivation, involving translocation of the ketocarotenoid inside the cavity followed by domain separation. Domain separation is a critical step in the photocycle of OCP because it exposes the N‐terminal domain (NTD) to perform quenching of the phycobilisomes. Many details regarding the mechanism and energetics of OCP domain separation remain unknown. In this work, we apply metadynamics to elucidate the protein rearrangements that lead to the active, domain‐separated, form of OCP. We find that translocation of the ketocarotenoid canthaxanthin has a profound effect on the energetic landscape and that domain separation only becomes favorable following translocation. We further explore, characterize, and validate the free energy surface (FES) using equilibrium simulations initiated from different states on the FES. Through pathway optimization methods, we characterize the most probable path to domain separation and reveal the barriers along that pathway. We find that the free energy barriers are relatively small (<5 kcal/mol), but the overall estimated kinetic rate is consistent with experimental measurements (>1 ms). Overall, our results provide detailed information on the requirement for canthaxanthin translocation to precede domain separation and an energetically feasible pathway to dissociation. [ABSTRACT FROM AUTHOR]

Summary: Sun‐induced fluorescence in the far‐red region (SIF) is increasingly used as a remote and proximal‐sensing tool capable of tracking vegetation gross primary production (GPP). However, the use of SIF to probe changes in GPP is challenged during extreme climatic events, such as heatwaves.Here, we examined how the 2018 European heatwave (HW) affected the GPP–SIF relationship in evergreen broadleaved trees with a relatively invariant canopy structure. To do so, we combined canopy‐scale SIF measurements, GPP estimated from an eddy covariance tower, and active pulse amplitude modulation fluorescence.The HW caused an inversion of the photosynthesis–fluorescence relationship at both the canopy and leaf scales. The highly nonlinear relationship was strongly shaped by nonphotochemical quenching (NPQ), that is, a dissipation mechanism to protect from the adverse effects of high light intensity. During the extreme heat stress, plants experienced a saturation of NPQ, causing a change in the allocation of energy dissipation pathways towards SIF.Our results show the complex modulation of the NPQ–SIF–GPP relationship at an extreme level of heat stress, which is not completely represented in state‐of‐the‐art coupled radiative transfer and photosynthesis models. [ABSTRACT FROM AUTHOR]

Barriers to continuous cropping of poplar plantations are closely related to the accumulation of phenolic acids in the soil. The purpose of this study was to explore the mechanism through which phenolic acid stress affects poplar productivity. The results showed that phenolic acids had a significant inhibitory effect on the photosynthesis of poplar. The inhibition of photosynthesis due to phenolic acids occurred mainly because, during electron transfer at the PSII reaction center, the primary quinone acceptor of PSII (QA) in the oxidized state was reduced, and excess light energy was lost in the form of heat dissipation; thus, poplar productivity decreased. At low phenolic acid concentrations (0X-1.5X), the activity of the PSII reaction center was temporarily inactivated, mainly because of stomatal limiting factors. At high phenolic acid concentrations (1.5X-3.0X), the PSII reaction centers were damaged, and photoinhibition occurred, mainly because of nonstomatal limiting factors.