Component Of Non-photochemical Quenching Research Articles

Gene Identification for Phototropin-dependent Photoprotection in Chlamydomonas Reinhardtii

Photosynthetic organisms convert solar energy into chemical energy through photosynthesis, generating carbohydrates and lipids that are valuable for biofuels. Algae, renowned for their rapid growth and efficiency, have developed mechanisms to adapt to varying light conditions, ensuring their survival and prosperity. Among these mechanisms, photoprotection strategies such as Non-Photochemical Quenching (NPQ) enhance their tolerance to high light intensities. Excessive light intensity can cause detrimental overexcitation of the photosystems. In Chlamydomonas, one of the proteins, LHCSR3, provides a quick protective response known as energy-dependent quenching (qE), the fastest and most important component of NPQ. Deletion of LHCSR3 leads to cell death under high light conditions. Recent research has shown that blue light, perceived by phototropin (PHOT), mediates the photoprotection of the photosynthetic machinery under high light conditions in Chlamydomonas reinhardtii. Deletion of PHOT leads to compromised expression of LHCSR3 under high light conditions, therefore leading to cell death. However, the downstream signaling components of the PHOT-LHCSR3 pathway remain largely undiscovered. The objective of this project was to identify and characterize new actors involved in PHOT-dependent photoprotection in Chlamydomonas. Using forward genetics and omics analysis, we built a mutant library that could survive under high light intensity in a phot background. We also identified 8 putative PHOT-dependent photoprotection downstream signaling components. Overall, this project brings new insights into the acclimation of photosynthesis to high light. The mutant library could be further used for additional research. Furthermore, understanding photoprotection will not only help increase microalgae biofuel production productivity but could also provide new insights into the genetic engineering of crops for high light resistance.

Chromium uptake and its impact on antioxidant level, photosynthetic machinery, and related gene expression in Brassica napus cultivars.

The development of heavy metals, particularly chromium (Cr)-tolerant crop cultivars, is hampered due to lack of understanding of the mechanisms behind Cr stress tolerance. In this study, two Brassica napus cultivars, ZS758 and ZD622, were compared for Cr stress resistance by using the chlorophyll a fluorescence technique and biochemical characteristics. In both cultivars, Cr stress dramatically decreased PSII and PSI efficiency, biomass accumulation, and antioxidant enzyme levels. Although, cultivar ZS758 showed reduction in oxidative stress by decreasing the production of reactive oxygen species (ROS) in terms of reduced H2O2 and MDA content and increased enzymatic activities of key antioxidants enzymes including SOD, APX, CAT, and POD activities that play a crucial role in the regulation of numerous transcriptional pathways involved in oxidative stress responses. Higher non-photochemical quenching (NPQ) and QY were found in tolerant ZS758 cultivar under Cr stress, indicating that tolerant cultivar had a greater capacity to preserve PSII activity under Cr stress by enhancing heat dissipation as a photo-protective component of NPQ. Lower PSI activity and electron transfer from PSII were confirmed by lower PSI efficiency and higher donor end limitation of PSI in both rapeseed cultivars. The Cr concentration was greater in the ZD622 as compared to ZS758, which affected the mineral nutrients profile and damaged the cellular ultrastructure and related gene expression levels. However, current study suggest that cultivar ZS758 is more resistant to Cr stress than ZD622 due to improved metabolism and structural integrity and Cr stress tolerance that is linked with the increased PSII activity, NPQ, and antioxidant potential; these physiological characteristics can be exploited to select cultivars for Cr stress tolerance.

Genotype‐dependent Strategies to “Overcome” Excessive Light: Insights into Non‐Photochemical Quenching under High Light Intensity

AbstractHigh light (HL) intensities have a significant impact on energy flux and distribution within photosynthetic apparatus. To understand the effect of high light intensity (HL) on the HL tolerance mechanisms in tomatoes, we examined the response of the photosynthesis apparatus of 12 tomato genotypes to HL. A reduced electron transfer per reaction center (ET0/RC), an increased energy dissipation (DI0/RC) and non‐photochemical quenching (NPQ), along with a reduced maximum quantum yield of photosystem II (FV/FM), and performance index per absorbed photon (PIABS) were common HL‐induced responses among genotypes; however, the magnitude of those responses was highly genotype‐dependent. Tolerant and sensitive genotypes were distinguished based on chlorophyll fluorescence and energy‐quenching responses to HL. Tolerant genotypes alleviated excess light through energy‐dependent quenching (qE), resulting in smaller photoinhibitory quenching (qI) compared to sensitive genotypes. Quantum yield components also shifted under HL, favoring the quantum yield of NPQ (ՓNPQ) and the quantum yield of basal energy loss (ՓN0), while reducing the efficient quantum yield of PSII (ՓPSII). The impact of HL on tolerant genotypes was less pronounced. While the energy partitioning ratio did not differ significantly between sensitive and tolerant genotypes, the ratio of NPQ components, especially qI, affected plant resilience against HL. These findings provide insights into different patterns of HL‐induced NPQ components in tolerant and sensitive genotypes, aiding the development of resilient crops for heterogeneous light conditions.

Combining Chlorophyll Fluorescence and Vegetation Reflectance Indices to Estimate Non-Photochemical Quenching (NPQ) of Rice at the Leaf Scale

Non-photochemical quenching (NPQ) is an indicator of crop stress. Until now, only a limited number of studies have focused on how to estimate NPQ using remote sensing technology. The main challenge is the complicated regulatory mechanism of NPQ. NPQ can be divided into energy-dependent (qE) and non-energy-dependent (non-qE) quenching. The contribution of these two components varies with environmental factors, such as light intensity and stress level due to the different response mechanisms. This study aims to explore the feasibility of estimating NPQ using photosynthesis-related vegetation parameters available from remote sensing by considering the two components of NPQ. We concurrently measured passive vegetation reflectance spectra by spectrometer, as well as active fluorescence parameters by pulse-amplitude modulated (PAM) of rice (Oryza sativa) leaves. Subsequently, we explored the ability of the selected vegetation parameters (including the photochemical reflectance index (PRI), inverted red-edge chlorophyll index (IRECI), near-infrared reflectance of vegetation (NIRv), and fluorescence quantum yield (ΦF)) to estimate NPQ. Based on different combinations of these remote sensing parameters, empirical models were established to estimate NPQ using the linear regression method. Experimental analysis shows that the contribution of qE and non-qE components varied under different illumination conditions. Under high illumination, the NPQ was attributed primarily to the qE component, while under low illumination, it was equally attributed to the qE and non-qE components. Among all tested parameters, ΦF was sensitive to the qE component variation, while IRECI and NIRv were sensitive to the non-qE component variation. Under high illumination, integrating ΦF in the regression model captured NPQ variations well (R2 > 0.74). Under low illumination, ΦF, IRECI, and NIRv explained 24%, 62%, and 65% of the variation in NPQ, respectively, while coupling IRECI or NIRv with ΦF considerably improved the accuracy of NPQ estimation (R2 > 0.9). For all the samples under both low and high illumination, the combination of ΦF with at least one of the other parameters (including IRECI, NIRv and PAR) offers a more versatile and reliable approach to estimating NPQ than using any single parameter alone. The findings of this study contribute to the further development of remote sensing methods for NPQ estimation at the canopy scale in the future.

HHL1 and SOQ1 synergistically regulate nonphotochemical quenching in Arabidopsis

Nonphotochemical quenching (NPQ) is an important photoprotective mechanism that quickly dissipates excess light energy as heat. NPQ can be induced in a few seconds to several hours; most studies of this process have focused on the rapid induction of NPQ. Recently, a new, slowly induced form of NPQ, called qH, was found during the discovery of the quenching inhibitor suppressor of quenching 1 (SOQ1). However, the specific mechanism of qH remains unclear. Here, we found that hypersensitive to high light 1 (HHL1)-a damage repair factor of photosystem II-interacts with SOQ1. The enhanced NPQ phenotype of the hhl1 mutant is similar to that of the soq1 mutant, which is not related to energy-dependent quenching or other known NPQ components. Furthermore, the hhl1 soq1 double mutant showed higher NPQ than the single mutants, but its pigment content and composition were similar to those of the wildtype. Overexpressing HHL1 decreased NPQ in hhl1 to below wildtype levels, whereas NPQ in hhl1 plants overexpressing SOQ1 was lower than that in hhl1 but higher than that in the wildtype. Moreover, we found that HHL1 promotes the SOQ1-mediated inhibition of plastidial lipoprotein through its von Willebrand factor type A domain. We propose that HHL1 and SOQ1 synergistically regulate NPQ.

Sandstorms damage the photosynthetic activities of Haloxylon ammodendron seedlings

Haloxylon ammodendron is a preferred shrub species for buffering against wind and fixing sand in arid sandy areas of northwest China. To determine whether sandstorms cause damage to H. ammodendron seedlings, we investigated the effects of wind-blown sands on the photosynthetic function of H. ammodendron by simulating sand-carrying wind in the wind tunnel. The results showed that photosystem II (PSII) non-photochemical quenching (NPQ) was sensitive to wind erosion, and sustained blowing of sand-carrying winds enhanced NPQ reduction. The rapidly relaxing quenching NPQf made up the majority component of NPQ; the NPQf/NPQ ratio was approximately 64.4% in the sand-free wind group and nearly 56.2% in the sand-carrying wind group. The distribution of the quantum efficiency of the excitation energy indicated that the relative proportions of the quantum yield of PSII photochemistry ΦPSII, the quantum yield of quenching due to light-induced ΦNPQ, and non-light-induced ΦNO were influenced by both wind erosion and light intensity; the sand-carrying wind resulted in a relative decrease in ΦNPQ and a corresponding increase in ΦNO. The maximum quantum efficiency of PSII photochemistry Fv/Fm was relatively stable in the sand-free wind group, with an average value of approximately 0.81. Compared to sand-free wind, sand-carrying winds caused remarkable decreases in Fv/Fm. Light intensity was the main factor affecting ΦPSII, ΦNPQ, and relative electron transport rate (rETR); there was no interaction effect between the duration of sand-carrying wind and light intensity. Taken together, wind-blown sands cause irreversible damage to the photosynthesis of H. ammodendron seedlings, which is the main factor restricting population regeneration in shelterbelts.

Elucidating the photosynthetic responses in chlorophyll-deficient soybean (Glycine max, L.) leaf

Chlorophyll (Chl)-deficient plants can potentially increase global surface albedo of mono-cropping systems, and simultaneously maintain a similar photosynthetic efficiency by increasing light canopy penetration and thus lowering investment in pigments. However, some previous studies have shown that pale mutants might reduce productivity in field conditions. Such lower yields were suspected to be due to loss of photosynthetic efficiency at leaf level during light fluctuations as a consequence of reduced capacity and slower relaxation of non-photochemical quenching (NPQ) of Chl fluorescence. In this paper, we tested this hypothesis by comparing, CO2 assimilation (A), photosystem II (PSII) efficiency (ΦPSII), photochemical quenching and NPQ, electron transport rate (ETR) and fluorescence yield (Fyield) in a green soybean (Glycine max L.) cultivar (Eiko) and in a Chl-deficient (MinnGold) mutant under dynamically fluctuating light conditions. MinnGold had significantly slower induction of ETR and lower A and ETR than Eiko, but there was little difference in ΦPSII between the two genotypes, suggesting that the lower photosynthesis of MinnGold was mainly due to lower light energy absorption by a Chl-deficient leaf. The NPQ capacity was also smaller in MinnGold than in Eiko. As for the kinetics of the rapidly inducible component of NPQ, MinnGold showed slower induction, not relaxation, than Eiko. The combination of the effect of Chl-deficiency on lower photosynthesis, NPQ capacity and slower NPQ induction may explain the lower biomass accumulation of MinnGold in the field. Our physiological observations, combined with fluorescence kinetics, can serve as a basis to parameterize Chl content in modelling radiative transfer and photosynthesis for upscaling measures of plant and ecosystem productivity by a big leaf model.

Non-Photochemical Quenching under Drought and Fluctuating Light.

Plants grow in a variable environment in regard to soil water and light driving photochemical reactions. Light energy exceeding plant capability to use it for photochemical reactions must be dissipated by processes of non-photochemical quenching (NPQ). The aim of the study was to evaluate the impact of various components of NPQ on the response of Arabidopsis thaliana to fluctuating light and water availability. A laboratory experiment with Arabidopsis thaliana wild type (WT) and mutants npq1 and npq4 grown under optimum or reduced water availability was conducted. Dark-adapted plants were illuminated with fluctuating light (FL) of two intensities (55 and 530 μmol m−2 s−1) with each of the phases lasting for 20 s. The impact of water availability on the role of zeaxanthin and PsbS protein in NPQ induced at FL was analysed. The water deficit affected the dynamics of NPQ induced by FL. The lack of zeaxanthin or PsbS reduced plant capability to cope with FL. The synergy of both of these components was enhanced in regard to the amplitude of NPQ in the drought conditions. PsbS was shown as a component of primary importance in suiting plant response to FL under optimum and reduced water availability.

Quantifying the long-term interplay between photoprotection and repair mechanisms sustaining photosystem II activity.

The photosystem II reaction centre (RCII) protein subunit D1 is the main target of light-induced damage in the thylakoid membrane. As such, it is constantly replaced with newly synthesised proteins, in a process dubbed the 'D1 repair cycle'. The mechanism of relief of excitation energy pressure on RCII, non-photochemical quenching (NPQ), is activated to prevent damage. The contribution of the D1 repair cycle and NPQ in preserving the photochemical efficiency of RCII is currently unclear. In this work, we seek to (1) quantify the relative long-term effectiveness of photoprotection offered by NPQ and the D1 repair cycle, and (2) determine the fraction of sustained decrease in RCII activity that is due to long-term protective processes. We found that while under short-term, sunfleck-mimicking illumination, NPQ is substantially more effective in preserving RCII activity than the D1 repair cycle (Plant. Cell Environ.41, 1098-1112, 2018). Under prolonged constant illumination, its contribution is less pronounced, accounting only for up to 30% of RCII protection, while D1 repair assumes a predominant role. Exposure to a wide range of light intensities yields comparable results, highlighting the crucial role of a constant and rapid D1 turnover for the maintenance of RCII efficiency. The interplay between NPQ and D1 repair cycle is crucial to grant complete phototolerance to plants under low and moderate light intensities, and limit damage to photosystem II under high light. Additionally, we disentangled and quantified the contribution of a slowly reversible NPQ component that does not impair RCII activity, and is therefore protective.

Roles of Light-Harvesting Complex Stress-Related Proteins in the Stress Responses of <i>Chlamydomonas</i>

Light is very important for photosynthesis in plants. However, excess light can result in photodamage to the photosynthetic apparatus. Via nonphotochemical quenching (NPQ), the oxidative stress caused by excess light energy can be counteracted by photoprotective mechanisms that evolve photosynthetic/oxygenic organisms. Energy-dependent quenching (qE), as the major NPQ component, relies on the accumulation of specific proteins that are termed light-harvesting complex stress-related (LHCSR) proteins in microalgae and mosses. LHCSRs have been reported to participate in adaptation to diverse environmental stresses, including excess light. In this review, we discuss the identification of LHCSRs in <i>Chlamydomonas</i> and the basic biochemical properties and functions of LHCSRs in acclimation to environmental stresses such as excess light and salt stress. We further review the potential interactive factors and upstream regulators of LHCSRs in <i>Chlamydomonas</i>, aiming to explore the underlying mechanism of <i>LHCSRs </i>in adaptation to multiple environmental stresses. We also discuss the evolution of LHCSRs in green algae and mosses and tentatively speculate about their participation in the adaptation to environmental change of the Earth. Work on <i>Chlamydomonas</i> LHCSR could provide clues to analyze the roles of LHCSR in both green algae and mosses. Thus, we offer an overview of current knowledge on the characteristics and functions of <i>Chlamydomonas</i> LHCSRs, which could shed new light on their detailed studies in both green algae and moss in the future.

The role of photoprotection in defence of two wheat genotypes against Zymoseptoria tritici

AbstractThis study provides new insights into the role of photoprotection in preformed and induced defence of two wheat genotypes with contrasting phenotypes to infection by Zymoseptoria tritici. We investigated the mechanisms of the photoprotective response during early infection, including nonphotochemical quenching (NPQ), β‐carotene‐derived xanthophylls, reactive oxygen species, and the phytohormones abscisic acid (ABA), jasmonic acid (JA), and salicylic acid (SA). Furthermore, we quantified the effects of pathogenesis on photosynthesis, stomatal control, and expression of plant defence molecular markers. The photoprotective mechanism of successful defence involved the qI component of NPQ leading to rapid down‐regulation of photosystem II quantum yield and chlorophyll a:b, increased biosynthesis of the xanthophyll neoxanthin and ABA, and the expression of chloroplast‐specific enzymes to engage in scavenging of O2●−. Elevated ABA in the resistant genotype correlated with preformed leaf defence traits including low stomatal density, increased expression of wax biosynthesis, and lignification. Z. tritici exhibited reduced germination and branching on the resistant host genotype and hijacked stomatal control in both genotypes by enhancing stomatal sensitivity to light. Increased biosynthesis of JA and anthocyanins, in contrast to SA, were quantified in the incompatible interaction. Our results indicate that ABA and JA in antagonistic action to SA were associated with defence in the resistant genotype, Cougar, against Z. tritici.

An underlying mechanism of qE deficiency in marine angiosperm Zostera marina.

Non-photochemical quenching (NPQ) of photosystem II (PSII) fluorescence is one of the most important protective mechanisms enabling the survival of phototropic organisms under high-light conditions. A low-efficiency NPQ, characterized by weak NPQ induction capacity and a low level of protective NPQ, was observed in the marine angiosperm Zostera marina, which inhabits the shallow water regions. Furthermore, chlorophyll fluorescence and Western blot analysis verified that the fast-inducted component of NPQ, i.e., the energy-dependent quenching (qE), was not present in this species. In contrast with the lack of PSII antenna quenching sites for qE induction in brown algae and the lack of functional XC in Ulvophyceae belonging to green algae, all the antenna proteins and the functional XC are present in Z. marina. A novel underlying mechanism was observed that the limited construction of the trans-thylakoid proton gradient (ΔpH) caused by photoinactivation of the oxygen evolving complex (OEC) did not induce protonation of PsbS, thus explaining the inability to form quenching sites for qE induction. Although the ΔpH established under light exposure activated violaxanthin (V) de-epoxidase enzyme to catalyze conversion of V via antheraxanthin (A) and then to zeaxanthin (Z), the quenching capacity of de-epoxidized pigment was weak in Z. marina. We suggest that the low-efficiency NPQ was conducive to efficiently utilize the limited electrons to perform photosynthesis, resisting the adverse effect of OEC photoinactivation on the photosynthetic rate.

Diurnal dynamics of nonphotochemical quenching in Arabidopsis npq mutants assessed by solar-induced fluorescence and reflectance measurements in the field.

Solar-induced fluorescence (SIF) is highly relevant in mapping photosynthesis from remote-sensing platforms. This requires linking SIF to photosynthesis and understanding the role of nonphotochemical quenching (NPQ) mechanisms under field conditions. Hence, active and passive fluorescence were measured in Arabidopsis with altered NPQ in outdoor conditions. Plants with mutations in either violaxanthin de-epoxidase (npq1) or PsbS protein (npq4) exhibited reduced NPQ capacity. Parallel measurements of NPQ, photosystem II efficiency, SIF and spectral reflectance (ρ) were conducted diurnally on one sunny summer day and two consecutive days during a simulated cold spell. Results showed that both npq mutants exhibited higher levels of SIF compared to wild-type plants. Changes in reflectance were related to changes in the violaxanthin-antheraxanthin-zeaxanthin cycle and not to PsbS-mediated conformational changes. When plants were exposed to cold temperatures, rapid onset of photoinhibition strongly quenched SIF in all lines. Using well-characterized Arabidopsis npq mutants, we showed for the first time the quantitative link between SIF, photosynthetic efficiency, NPQ components and leaf reflectance. We discuss the functional potential and limitations of SIF and reflectance measurements for estimating photosynthetic efficiency and NPQ in the field.

Rethinking the Influence of Chloroplast Movements on Non-photochemical Quenching and Photoprotection.

Under blue light, plant chloroplasts relocate to different areas of the cell. The photoreceptor phototropin2 (phot2) mediates the chloroplast movement mechanism under excess blue light alongside the chloroplast unusual positioning1 (chup1) protein. Recently, it has been proposed that leaf transmittance changes associated with chloroplast relocation affect measurements of nonphotochemical quenching (NPQ), resulting in kinetic differences due to these movements (termed "qM"). We evaluated these claims using Arabidopsis (Arabidopsis thaliana) knock-out mutants lacking either phot2 or chup1 and analyzed the kinetics of both the onset and recovery of NPQ under equivalent intensities of both red and blue light. We also evaluated the photoprotective ability of chloroplast movements both during the early onset of photoinhibition and under sustained excess light. We monitored photoinhibition using the chlorophyll fluorescence parameter of photochemical quenching in the dark, which measures the redox state of QA within PSII without any of the complications of traditional F v /F m measurements. While there were noticeable differences between the responses under red and blue light, the chloroplast movement mechanism had no effect on the rate or amplitude of NPQ onset or recovery. Therefore, we were unable to replicate the "qM" component and its corresponding influence on the kinetics of NPQ in Arabidopsis grown under "shade" conditions. Furthermore, chloroplast relocation had no effect on the high-light tolerance of these plants. These data cast doubt upon the existence of a chloroplast movement-dependent component of NPQ Therefore, the influence of chloroplast movements on photoprotection should be thoroughly reevaluated.

Rapid regulation of photosynthetic light harvesting in the absence of minor antenna and reaction centre complexes.

Plants are subject to dramatic fluctuations in the intensity of sunlight throughout the day. When the photosynthetic machinery is exposed to high light, photons are absorbed in excess, potentially leading to oxidative damage of its delicate membrane components. A photoprotective molecular process called non-photochemical quenching (NPQ) is the fastest response carried out in the thylakoid membranes to harmlessly dissipate excess light energy. Despite having been intensely studied, the site and mechanism of this essential regulatory process are still debated. Here, we show that the main NPQ component called energy-dependent quenching (qE) is present in plants with photosynthetic membranes largely enriched in the major trimeric light-harvesting complex (LHC) II, while being deprived of all minor LHCs and most photosystem core proteins. This fast and reversible quenching depends upon thylakoid lumen acidification (ΔpH). Enhancing ΔpH amplifies the extent of the quenching and restores qE in the membranes lacking PSII subunit S protein (PsbS), whereas the carotenoid zeaxanthin modulates the kinetics and amplitude of the quenching. These findings highlight the self-regulatory properties of the photosynthetic light-harvesting membranes in vivo, where the ability to switch reversibly between the harvesting and dissipative states is an intrinsic property of the major LHCII.

Ascorbate Deficiency Does Not Limit Nonphotochemical Quenching in Chlamydomonas reinhardtii.

Ascorbate (Asc; vitamin C) plays essential roles in development, signaling, hormone biosynthesis, regulation of gene expression, stress resistance, and photoprotection. In vascular plants, violaxanthin de-epoxidase requires Asc as a reductant; thereby, Asc is required for the energy-dependent component of nonphotochemical quenching (NPQ). To assess the role of Asc in NPQ in green algae, which are known to contain low amounts of Asc, we searched for an insertional Chlamydomonas reinhardtii mutant affected in theVTC2 gene encoding GDP-l-Gal phosphorylase, which catalyzes the first committed step in the biosynthesis of Asc. The Crvtc2-1 knockout mutant was viable and, depending on the growth conditions, contained 10% to 20% Asc relative to its wild type. When C. reinhardtii was grown photomixotrophically at moderate light, the zeaxanthin-dependent component of NPQ emerged upon strong red illumination both in the Crvtc2-1 mutant and in its wild type. Deepoxidation was unaffected by Asc deficiency, demonstrating that the Chlorophycean violaxanthin de-epoxidase found in C. reinhardtii does not require Asc as a reductant. The rapidly induced, energy-dependent NPQ component characteristic of photoautotrophic C. reinhardtii cultures grown at high light was not limited by Asc deficiency either. On the other hand, a reactive oxygen species-induced photoinhibitory NPQ component was greatly enhanced upon Asc deficiency, both under photomixotrophic and photoautotrophic conditions. These results demonstrate that Asc has distinct roles in NPQ formation in C. reinhardtii as compared to vascular plants.

The rise and fall of Light-Harvesting Complex Stress-Related proteins as photoprotection agents during evolution.

Photosynthesis depends on light. However, excess light can be harmful for the photosynthetic apparatus because it produces reactive oxygen species (ROS) that cause photoinhibition. Oxygenic organisms evolved photoprotection mechanisms to counteract light-dependent ROS production, including preventive dissipation of excited states of chlorophyll (1Chl*) into heat in the process termed non-photochemical quenching (NPQ). This consists in the activation of 1Chl* quenching reactions when the thylakoid luminal pH drops below 5.2. In turn, acidification occurs when the rate of the CO2 reducing cycle is saturated and cannot regenerate ADP+Pi, thus inhibiting ATPase activity and the return of protons (H+) to the stromal compartment. The major and fastest component of NPQ is energy quenching, qE, which in algae depends on the Light-Harvesting Complex Stress-Related (LHCSR) proteins. In mosses, LHCSR proteins have remained the major catalysts of energy dissipation, but a minor contribution also occurs via a homologous protein, Photosystem II Subunit S (PSBS). In vascular plants, however, LHCSR has disappeared and PSBS is the only pH-sensitive trigger of qE. Why did PSBS replace LHCSR in the later stages of land colonization? Both PSBS and LHCSR belong to the Light Harvesting Complex superfamily (LHC) and share properties such as harboring protonatable residues that are exposed to the chloroplast lumen, which is essential for pH sensing. However, there are also conspicuous differences: LHCSR binds chlorophylls and xanthophylls while PSBS does not, implying that the former may well catalyse quenching reactions while the latter needs pigment-binding partners for its quenching function. Here, the evolution of quenching mechanisms for excess light is reviewed with a focus on the role of LHCSR versus PSBS, and the reasons for the redundancy of LHCSR in vascular plants as PSBS became established.

The need to incorporate fast and slow relaxation kinetic parameters into photosynthesis-measuring systems

Abstract Increasing global population demands the doubling of crop production by 2050. Advancements in agricultural techniques make it possible to increase the rate of crop improvement, though utilization of molecular tools to identify traits related to yield and stress tolerance is poor. The development of effective physiological selection criteria that are fast and non-destructive, particularly those related to yield and stress tolerance, will help in developing improved crop cultivars. In plant stress physiology, established chlorophyll fluorescence parameters, such as the efficiency of photosystem II (ФPSII), photosystem I electron transport rate (PSI ETR), and non-photochemical quenching (NPQ), have received attention. The photoprotective processes conferred by NPQ serve fundamental roles in maintaining plant fitness and sustainable yields where their importance in predicting crop performance and yield under normal or stressful environments has been widely recognized. In this review, we focus on two informative parameters, - q E and q I ,- which are components of NPQ that have been ignored by most photosynthesis-measuring systems. We argue for adding these parameters to photosynthesis machines as calculating them later can be cumbersome and their inclusion will help plant breeders and non-specialists in phenotyping using chlorophyll fluorescence.

Plant biodiversity and regulation of photosynthesis in the natural environment.

Investigation of photosynthesis regulation in different plant groups exposed to variable conditions showed that all species have similar photosynthetic electron transport modulation while excess energy dissipation is species specific. Photosynthesis is regulated in response to dynamic environmental conditions to satisfy plant metabolic demands while also avoiding possible over-excitation of the electron transport chain and the generation of harmful reactive oxygen species. Photosynthetic organisms evolved several mechanisms to modulate light harvesting and electron transport efficiency to respond to conditions changing at different timescales, going from fast sun flecks to slow seasonal variations. These regulatory mechanisms changed during evolution of photosynthetic organisms, also adapting to various ecological niches, making the investigation of plant biodiversity highly valuable to uncover conserved traits and plasticity of photosynthetic regulation and complement studies on model species. In this work, a set of plants belonging to different genera of angiosperms, gymnosperms, ferns and lycophytes were investigated by monitoring their photosynthetic parameters in different seasons looking for common trends and differences. In all plants, analysed photosynthetic electron transport rate was found to be modulated by growth light intensity, ensuring a balance between available energy and photochemical capacity. Growth light also influenced the threshold where heat dissipation of excitation energy, a mechanism called non-photochemical quenching (NPQ), was activated. On the contrary, NPQ amplitude did not correlate with light intensity experienced by the plants but was a species-specific feature. The zeaxanthin-dependent component of NPQ, qZ, was found to be the most variable in different plants and its modulation influenced the intensity and the kinetic properties of the response.

Changes in the photosynthesis properties and photoprotection capacity in rice (Oryza sativa) grown under red, blue, or white light.

Non-photochemical quenching (NPQ) of the excited state of chlorophyll a is a major photoprotective mechanism plants utilize to survive under high light. Here, we report the impact of long-term light quality treatment on photosynthetic properties, especially NPQ in rice. We used three LED-based light regimes, i.e., red (648-672nm), blue (438-460nm), and "warm" white light (529-624nm), with the incident photon flux density of 300µmolphotonsm-2s-1, the difference in the absorbed photon flux densities by leaves grown under different light quality being less than 7%. Our results show that blue light, as compared to white light, induced a significant decrease in Fv/Fm, a decreased rate of reduction of P700+ after P700 was completely oxidized; furthermore, blue light also induced higher NPQ with an increased initial speed of NPQ induction, which corresponds to the qE component of NPQ, and a lower maximum quantum yield of PSII, i.e., Y(II). In contrast, rice grown under long-term red light showed decreased Y(II) and increased NPQ, but with no change in Fv/Fm. Furthermore, we found that rice grown under either blue or red light showed decreased transcript abundance of both catalase and ascorbate peroxidase, together with an increased H2O2 content, as compared to rice grown under white light. All these data suggest that even under a moderate incident light level, rice grown under blue or red light led to compromised antioxidant system, which contributed to decreased quantum yield of photosystem II and increased NPQ.