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The photosynthetic apparatus is a highly modular assembly of large pigment-binding proteins. Complexes called antennae can capture the sunlight and direct it from the periphery of two Photosystems (I, II) to the core reaction centers, where it is converted into chemical energy. The apparatus must cope with the natural light fluctuations that can become detrimental to the viability of the photosynthetic organism. Here we present an atomic scale view of the photoprotective mechanism that is activated on this line of defense by several photosynthetic organisms to avoid overexcitation upon excess illumination. We provide a complete macroscopic to microscopic picture with specific details on the conformations of the major antenna of Photosystem II that could be associated with the switch from the light-harvesting to the photoprotective state. This is achieved by combining insight from both experiments and all-atom simulations from our group and the literature in a perspective article. [ABSTRACT FROM AUTHOR]

Light intensity and temperature independently impact all parts of the photosynthetic machinery in plants and algae. Yet to date, the vast majority of pulse amplitude modulated (PAM) chlorophyll a fluorescence measurements have been performed at well-defined light intensities, but rarely at well-defined temperatures. In this work, we show that PAM measurements performed at various temperatures produce vastly different results in the chlorophyte Chlorella vulgaris. Using a recently developed Phenoplate technique to map quantum yield of Photosystem II (Y(II)) and non-photochemical quenching (NPQ) as a function of temperature, we show that the fast-relaxing NPQ follows an inverse normal distribution with respect to temperature and appears insensitive to previous temperature acclimation. The slow-relaxing or residual NPQ after 5 minutes of dark recovery follows a normal distribution similar to Y(II) but with a peak in the higher temperature range. Surprisingly, higher slow- and fast-relaxing NPQ values were observed in high-light relative to low-light acclimated cultures. Y(II) values peaked at the adaptation temperature regardless of temperature or light acclimation. Our novel findings show the complete temperature working spectrum of Y(II) and how excess energy quenching is managed across a wide range of temperatures in the model microalgal species C. vulgaris. Finally, we draw attention to the fact that the effect of the temperature component in PAM measurements has been wildly underestimated, and results from experiments at room temperature can be misleading. [ABSTRACT FROM AUTHOR]

Photosynthetic rates vary depending on growth conditions, even within species. Remote sensing techniques have a great potential to predict the photosynthetic rates of leaves with different characteristics. Here, we demonstrate that the photosynthetic rates of leaves acclimated to different light and nutrient conditions can be estimated based on the chlorophyll fluorescence (ChlF), the photochemical reflectance index (PRI), and a chlorophyll index. Chenopodium album plants were grown under different light and nutrient conditions. PRI, ChlF parameters, and CO2/H2O gas exchange rates of leaves were simultaneously determined under the various light and CO2 conditions. PRI was used to assess non-photochemical quenching (NPQ), but the relationship between NPQ and PRI was weakened when the data on leaves grown under different conditions were pooled, because PRI in darkness ( PRI 0 ) changed with the leaf pigment composition. Among 15 pigment indices, we found that NDVI green , a reflectance index related to the leaf chlorophyll content, had the best correlation with PRI 0 ( r 2 = 0.89 ) across the studied leaves, and the correction of PRI by NDVI green improved the predictability of NPQ ( r 2 = 0.82 ). Using the steady-state ChlF, the NPQ estimated from PRI and NDVI green , and the stomatal conductance coefficient, we calculated the CO2 assimilation rates, which were strongly correlated with the actual rates (RMSE = 4.85 μ mol m - 2 s - 1 ), irrespective of growth conditions. Our approach has the potential to contribute to a more accurate estimation of photosynthetic rates in remote sensing. However, further studies on species variations and connecting with radiative transfer models are needed to demonstrate this at the canopy scale. [ABSTRACT FROM AUTHOR]

Non-photochemical quenching (NPQ) in photosynthetic organisms provides the necessary photoprotection that allows them to cope with largely and quickly varying light intensities. It involves deactivation of excited states mainly at the level of the antenna complexes of photosystem II using still largely unknown molecular mechanisms. In higher plants the main contribution to NPQ is the so-called qE-quenching, which can be switched on and off in a few seconds. This quenching mechanism is affected by the low pH-induced activation of the small membrane protein PsbS which interacts with the major light-harvesting complex of photosystem II (LHCII). We are reporting here on a mechanistic study of the PsbS-induced LHCII quenching using ultrafast time-resolved chlorophyll (Chl) fluorescence. It is shown that the PsbS/LHCII interaction in reconstituted proteoliposomes induces highly effective and specific quenching of the LHCII excitation by a factor ≥ 20 via Chl–Chl charge-transfer (CT) state intermediates which are weakly fluorescent. Their characteristics are very broad fluorescence bands pronouncedly red-shifted from the typical unquenched LHCII fluorescence maximum. The observation of PsbS-induced Chl–Chl CT-state emission from LHCII in the reconstituted proteoliposomes is highly reminiscent of the in vivo quenching situation and also of LHCII quenching in vitro in aggregated LHCII, indicating a similar quenching mechanism in all those situations. The PsbS mutant lacking the two proton sensing Glu residues induced significant, but much smaller, quenching than wild type. Added zeaxanthin had only minor effects on the yield of quenching in the proteoliposomes. Overall our study shows that PsbS co-reconstituted with LHCII in liposomes represents an excellent in vitro model system with characteristics that are reflecting closely the in vivo qE-quenching situation. [ABSTRACT FROM AUTHOR]

Non-photochemical quenching, NPQ, of chlorophyll fluorescence regulates the heat dissipation of chlorophyll excited states and determines the efficiency of the oxygenic photosynthetic systems. NPQ is regulated by a pH-sensing protein, responding to the chloroplast lumen acidification induced by excess light, coupled to an actuator, a chlorophyll/xanthophyll subunit where quenching reactions are catalyzed. In plants, the sensor is PSBS, while the two pigment-binding proteins Lhcb4 (also known as CP29) and LHCII are the actuators. In algae and mosses, stress-related light-harvesting proteins (LHCSR) comprise both functions of sensor and actuator within a single subunit. Here, we report on expressing the lhcsr1 gene from the moss Physcomitrella patens into several Arabidopsis thaliana npq4 mutants lacking the pH sensing PSBS protein essential for NPQ activity. The heterologous protein LHCSR1 accumulates in thylakoids of A. thaliana and NPQ activity can be partially restored. Complementation of double mutants lacking, besides PSBS, specific xanthophylls, allowed analyzing chromophore requirement for LHCSR-dependent quenching activity. We show that the partial recovery of NPQ is mostly due to the lower levels of Zeaxanthin in A. thaliana in comparison to P. patens. Complemented npq2npq4 mutants, lacking besides PSBS, Zeaxanthin Epoxidase, showed an NPQ recovery of up to 70% in comparison to A. thaliana wild type. Furthermore, we show that Lutein is not essential for the folding nor for the quenching activity of LHCSR1. In short, we have developed a system to study the function of LHCSR proteins using heterologous expression in a variety of A. thaliana mutants. [ABSTRACT FROM AUTHOR]

Ulva lactuca is regarded as a prospective energy crop for biorefinery owing to its affluent biochemical composition and high growth rate. In fast-growing macroalgae, biomass development strictly depends on external nitrogen pools. Additionally, nitrogen uptake rates and photosynthetic pigment content vary with type of nitrogen source and light conditions. However, the combined influence of nitrogen source and light intensity on photosynthesis is not widely studied. In present study, pale green phenotype of U. lactuca was obtained under high light (HL) condition when inorganic nitrogen (nitrate) in the media was substituted with organic nitrogen (urea). Further, pale green phenotype survived the saturating light intensities in contrast to the normal pigmented control which bleached in HL. Detailed analysis of biochemical composition and photosynthesis was performed to understand functional antenna size and photoprotection in pale green phenotype. Under HL, urea-grown cultures exhibited increased growth rate, carbohydrate and lipid content while substantial reduction in protein, chlorophyll content and PSII antenna size was observed. Further, in vivo slow and polyphasic chlorophyll a (Chl a) fluorescence studies revealed reduction in excitation pressure on PSII along with low non-photochemical quenching thus, transmitting most of the absorbed energy into photochemistry. The results obtained could be correlated to previous report on cultivation of U. lactuca through saturating summer intensities (1000 µmole photons m−2 s−1) in urea based: poultry litter extract (PLE). Having proved critical role of urea in conforming photoprotection, the application PLE was authenticated for futuristic, sustainable and year-round biomass cultivation.Graphical Abstract: [ABSTRACT FROM AUTHOR]

Recently, we isolated a complex consisting of photosystem II (PSII) and light-harvesting complexes (LHCs) from Nannochloropsis granulata (Umetani et al. Photosynth Res 136:49-61, 2017). This complex contained stress-related protein, Lhcx, as a major component of LHC (Protein ID is Naga_100173g12.1), suggesting that non-photochemical quenching activities may be taking place in the PSII-LHC complex. In this study, we examined the energy transfer dynamics in the isolated LHCs and PSII-LHC complexes, and found substantial quenching capacity. In addition, the LHCs contained low-energy chlorophylls with fluorescence maxima at approximately 710 nm, which may enhance the quenching efficiency in the PSII-LHC. Delayed fluorescence analysis suggested that there was an approximately 50% reduction in energy trapping at the PSII reaction center in the PSII-LHC supercomplex under low-pH condition compared to neutral pH condition. Enhanced quenching may confer a survival advantage in the shallow-water habitat of Nannochloropsis. [ABSTRACT FROM AUTHOR]

Non-photochemical quenching (NPQ) is responsible for protecting the light-harvesting apparatus of plants from damage at high light conditions. Although it is agreed that the major part of NPQ, an energy-dependent quenching (qE), originates in the light-harvesting antenna, its exact mechanism is still debated. In our earlier work (Chmeliov et al. in Nat Plants 2:16045, 2016), we have analyzed the time-resolved fluorescence (TRF) from the trimers and aggregates of the major light-harvesting complexes of plants (LHCII) over a broad temperature range and came to a conclusion that three distinct states are required to describe the experimental data: two of them correspond to the emission bands centered at ~680 and ~700 nm, and the third state is responsible for the excitation quenching. This was opposite to earlier suggestions of a two-state model, where the red-shifted fluorescence and excitation quenching were assumed to be related. To examine such possibility, in the current work we repeat our analysis of the TRF data in terms of the two-state model. We find that even though it can reasonably describe the aggregate fluorescence, it fails to do so for the LHCII trimers. We conclude that the red-emitting state cannot be responsible for fluorescence quenching in the LHCII aggregates and reaffirm that the three-state model is the simplest possible description of the experimental data. [ABSTRACT FROM AUTHOR]

Cyclic electron transport around photosystem I (PSI) generates ∆pH across the thylakoid membrane without net production of NADPH. In angiosperms, two pathways of PSI cyclic electron transport operate. The main pathway depends on PGR5/PGRL1 proteins and is likely identical to the historical Arnon's pathway. The minor pathway depends on chloroplast NADH dehydrogenase-like (NDH) complex. In assays of their rates in vivo, the two independent pathways are often mixed together. Theoretically, linear electron transport from water to NADP cannot satisfy the ATP/NADPH production ratio required by the Calvin-Benson cycle and photorespiration. PGR5/PGRL1-dependent PSI cyclic electron transport contributes substantially to the supply of ATP for CO fixation, as does linear electron transport. Also, the contribution of chloroplast NDH cannot be ignored, especially at low light intensity, although the extent of the contribution depends on the plant species. An increase in proton conductivity of ATP synthase may compensate ATP synthesis to some extent in the pgr5 mutant. Combined with the decreased rate of ∆pH generation, however, this mechanism sacrifices homeostasis of the thylakoid lumen pH, seriously disturbing the pH-dependent regulation of photosynthetic electron transport, induction of qE, and downregulation of the cytochrome b f complex. PGR5/PGRL1-dependent PSI cyclic electron transport produces sufficient proton motive force for ATP synthesis and the regulation of photosynthetic electron transport. [ABSTRACT FROM AUTHOR]

In the present study, the high light (HL) acclimation of Chromera velia (Chromerida) was studied. HL-grown cells exhibited an increased cell volume and dry weight compared to cells grown at medium light (ML). The chlorophyll (Chl) a-specific absorption spectra ( $$a_{\text{phy}}^{*}$$ ) of the HL cells showed an increased absorption efficiency over a wavelength range from 400 to 750 nm, possibly due to differences in the packaging of Chl a molecules. In HL cells, the size of the violaxanthin ( V) cycle pigment pool was strongly increased. Despite a higher concentration of de-epoxidized V cycle pigments, non-photochemical quenching (NPQ) of the HL cells was slightly reduced compared to ML cells. The analysis of NPQ recovery during low light (LL) after a short illumination with excess light showed a fast NPQ relaxation and zeaxanthin epoxidation. Purification of the pigment-protein complexes demonstrated that the HL-synthesized V was associated with the chromera light-harvesting complex (CLH). However, the difference absorption spectrum of HL minus ML CLH, together with the 77 K fluorescence excitation spectra, suggested that the additional V was not protein bound but localized in a lipid phase associated with the CLH. The polypeptide analysis of the pigment-protein complexes showed that one out of three known LHCr proteins was associated in higher concentration with photosystem I in the HL cells, whereas in ML cells, it was enriched in the CLH fraction. In conclusion, the acclimation of C. velia to HL illumination shows features that are comparable to those of diatoms, while other characteristics more closely resemble those of higher plants and green algae. [ABSTRACT FROM AUTHOR]

In this study, we demonstrate the selective in vivo detection of diadinoxanthin (DD) and diatoxanthin (DT) in intact Cyclotella cells using resonance Raman spectroscopy. In these cells, we were able to assess both the content of DD and DT carotenoids relative to chlorophyll and their conformation. In addition, the sensitivity and selectivity of the technique allow us to discriminate between different pools of DD on a structural basis, and to follow their fate as a function of the illumination conditions. We report that the additional DD observed when cells are grown in high-light conditions adopts a more twisted conformation than the lower levels of DD present when the cells are grown in low-light (LL) conditions. Thus, we conclude that this pool of DD is more tightly bound to a protein-binding site, which must differ from the one occupied by the DD present in LL conditions. [ABSTRACT FROM AUTHOR]

Recent progress in elucidating the structure of higher plants photosynthetic membranes provides a wealth of information. It allows generation of architectural models that reveal well-organized and complex arrangements not only on whole membrane level, but also on the supramolecular level. These arrangements are not static but highly responsive to the environment. Knowledge about the interdependency between dynamic structural features of the photosynthetic machinery and the functionality of energy conversion is central to understanding the plasticity of photosynthesis in an ever-changing environment. This review summarizes the architectural switches that are realized in thylakoid membranes of green plants. [ABSTRACT FROM AUTHOR]

It has long been suspected that photoprotective mechanisms in green algae are similar to those in seed plants. However, exceptions have recently surfaced among aquatic and marine green algae in several taxonomic classes. Green algae are highly diverse genetically, falling into 13 named classes, and they are diverse ecologically, with many lineages including members from freshwater, marine, and terrestrial habitats. Genetically similar species living in dramatically different environments are potentially a rich source of information about variations in photoprotective function. Using aquatic and desert-derived species from three classes of green algae, we examined the induction of photoprotection under high light, exploring the relationship between nonphotochemical quenching and the xanthophyll cycle. In liquid culture, behavior of aquatic Entransia fimbriata (Klebsormidiophyceae) generally matched patterns observed in seed plants. Nonphotochemical quenching was lowest after overnight dark adaptation, increased with light intensity, and the extent of nonphotochemical quenching correlated with the extent of deepoxidation of xanthophyll cycle pigments. In contrast, overnight dark adaptation did not minimize nonphotochemical quenching in the other species studied: desert Klebsormidium sp. (Klebsormidiophyceae), desert and aquatic Cylindrocystis sp. (Zygnematophyceae), and desert Stichococcus sp. (Trebouxiophyceae). Instead, exposure to low light reduced nonphotochemical quenching below dark-adapted levels. De-epoxidation of xanthophyll cycle pigments paralleled light-induced changes in nonphotochemical quenching for species within Klebsormidiophyceae and Trebouxiophyceae, but not Zygnematophyceae. Inhibition of violaxanthin-zeaxanthin conversion by dithiothreitol reduced high-light-associated nonphotochemical quenching in all species (Zygnematophyceae the least), indicating that zeaxanthin can contribute to photoprotection as in seed plants but to different extents depending on taxon or lineage. [ABSTRACT FROM AUTHOR]

The Benson-Calvin cycle enzymes are activated in vivo when disulfide bonds are opened by reduction via the ferredoxin-thioredoxin system in chloroplasts. Iodoacetamide reacts irreversibly with free -SH groups of cysteine residues and inhibits the enzymes responsible for CO fixation. Here, we investigate the effect of iodoacetamide on electron transport, when infiltrated into spinach leaves. Using fluorescence and absorption spectroscopy, we show that (i) iodoacetamide very efficiently blocks linear electron flow upon illumination of both photosystems (decrease in the photochemical yield of photosystem II) and (ii) iodoacetamide favors cyclic electron flow upon light excitation specific to PSI. These effects account for an NPQ formation even faster in iodoacetamide under far-red illumination than in the control under saturating light. Such an increase in NPQ is dependent upon the proton gradient across the thylakoid membrane (uncoupled by nigericin addition) and PGR5 (absent in Arabidopsis pgr5 mutant). Iodoacetamide very tightly insulates the electron current at the level of the thylakoid membrane from any electron leaks toward carbon metabolism, therefore, providing choice conditions for the study of cyclic electron flow around PSI. [ABSTRACT FROM AUTHOR]

The potential of algae to acclimate to environmental stress is commonly assessed using chlorophyll a fluorescence, with changes in parameters of photosynthesis versus irradiance ( P/ E) curves measured either as rapid light curves (RLC) or steady-state light curves (LC). Here, effects of emersion on primary photosynthesis of four brown macroalgae ( Ascophyllum nodosum, Fucus serratus, Sargassum muticum, Laminaria digitata) were compared by applying both RLC and LC. When LC were used, photosynthetic performance was enhanced during emersion in A. nodosum and F. serratus as shown by increases in q, rETR and E. By contrast, emersion had no impact on photosynthetic parameters of S. muticum and L. digitata. Relative changes in the NPQ-rETR relationship were reduced in A. nodosum, F. serratus and S. muticum, but remained unaffected in L. digitata. As none of the species developed their potential NPQ, corresponding values could not be determined from RLC. Using RLC, observed photosynthetic performance of F. serratus and L. digitata was reduced upon emersion, whilst values for NPQ were enhanced. Only results derived from LC provide evidence for a potential physiological adaptation of brown macroalgae to their natural habitat; it is recommended using the LC protocol to detect environmental impacts on photosynthesis. [ABSTRACT FROM AUTHOR]

This study aimed to evaluate the photochemical reflectance index (PRI) for assessing plant photosynthetic performance throughout the plant life cycle. The relationships between PRI, chlorophyll fluorescence parameters, and leaf pigment indices in Solanum melongena L. (aubergine; eggplant) were studied using photosynthetic induction curves both in short-term (diurnal) and long-term (seasonal) periods under different light intensities. We found good correlations between PRI/non-photochemical quenching (NPQ) and PRI/electron transport rate (ETR) in the short term at the same site of a single leaf but these relationships did not hold throughout the life of the plant. In general, changes in PRI owing to NPQ or ETR variations in the short term were <20 % of those that occurred with leaf aging. Results also showed that PRI was highly correlated to plant pigments, especially chlorophyll indices measured by spectral reflectance. Moreover, relationships of steady-state PRI/ETR and steady-state PRI/photochemical yield of photosystem II (Φ) measured at uniform light intensity at different life stages proved that overall photosynthesis capacity and steady-state PRI were better correlated through chlorophyll content than NPQ and xanthophylls. The calibrated PRI index accommodated these pigments effects and gave better correlation with NPQ and ETR than PRI. Further studies of PRI indices based on pigments other than xanthophylls, and studies on PRI mechanisms in different species are recommended. [ABSTRACT FROM AUTHOR]

During the last years significant progress was achieved in unraveling molecular characteristics of the thylakoid membrane of different diatoms. With the present review it is intended to summarize the current knowledge about the structural and functional changes within the thylakoid membrane of diatoms acclimated to different light conditions. This aspect is addressed on the level of the organization and regulation of light-harvesting proteins, the dissipation of excessively absorbed light energy by the process of non-photochemical quenching, and the lipid composition of diatom thylakoid membranes. Finally, a working hypothesis of the domain formation of the diatom thylakoid membrane is presented to highlight the most prominent differences of heterokontic thylakoids in comparison to vascular plants and green algae during the acclimation to low and high light conditions. [ABSTRACT FROM AUTHOR]

During the recent years, wide varieties of methodologies have been developed up to the level of commercial use to measure photosynthetic electron transport by modulated chlorophyll a-in vivo fluorescence. It is now widely accepted that the ratio between electron transport rates and new biomass ( P/ B) is not fixed and depends on many factors that are also taxonomically variable. In this study, the balance between photon absorption and biomass production has been measured in two phycobilin-containing phototrophs, namely, a cyanobacterium and a cryptophyte, which differ in their antenna organization. It is demonstrated that the different antenna organization exerts influence on the regulation of the primary photosynthetic reaction and the dissipation of excessively absorbed radiation. Although, growth rates and the quantum efficiency of biomass production of both phototrophs were comparable, the ratio P/ B was twice as high in the cryptophyte in comparison to the cyanobacterium. It is assumed that this discrepancy is because of differences in the metabolic regulation of cell growth. In the cryptophyte, absorbed photosynthetic energy is used to convert assimilated carbon directly into proteins and lipids, whereas in the cyanobacterium, the photosynthetic energy is preferentially stored as carbohydrates. [ABSTRACT FROM AUTHOR]

Highly time-resolved photoacclimation patterns of the chlorophyte microalga Dunaliella tertiolecta during exposure to an off-on-off (block) light pattern of saturating photon flux, and to a regime of consecutive increasing light intensities are presented. Non-photochemical quenching (NPQ) mechanisms unexpectedly responded with an initial decrease during dark-light transitions. NPQ values started to rise after light exposure of approximately 4 min. State-transitions, measured as a change of PSII:PSI fluorescence emission at 77 K, did not contribute to early NPQ oscillations. Addition of the uncoupler CCCP, however, caused a rapid increase in fluorescence and showed the significance of qE for NPQ. Partitioning of the quantum efficiencies showed that constitutive NPQ was (a) higher than qE-driven NPQ and (b) responded to light treatment within seconds, suggesting an active role of constitutive NPQ in variable energy dissipation, although it is thought to contribute statically to NPQ. The PSII connectivity parameter p correlated well with F′, F′ and NPQ during the early phase of the dark-light transients in sub-saturating light, suggesting a plastic energy distribution pattern within energetically connected PSII centres. In consecutive increasing photon flux experiments, correlations were weaker during the second light increment. Changes in connectivity can present an early photoresponse that are reflected in fluorescence signals and NPQ and might be responsive to the short-term acclimation state, and/or to the actinic photon flux. [ABSTRACT FROM AUTHOR]

The xanthophyll cycle represents one of the important photoprotection mechanisms in plant cells. In the present review, we summarize current knowledge about the violaxanthin cycle of vascular plants, green and brown algae, and the diadinoxanthin cycle of the algal classes Bacillariophyceae, Xanthophyceae, Haptophyceae, and Dinophyceae. We address the biochemistry of the xanthophyll cycle enzymes with a special focus on protein structure, co-substrate requirements and regulation of enzyme activity. We present recent ideas regarding the structural basis of xanthophyll cycle-dependent photoprotection, including different models for the mechanism of non-photochemical quenching of chlorophyll a fluorescence. In a dedicated chapter, we also describe the unique violaxanthin antheraxanthin cycle of the Prasinophyceae, together with its implication for the mechanism of xanthophyll cycle-dependent heat dissipation. The interaction between the diadinoxanthin cycle and alternative electron flow pathways in the chloroplasts of diatoms is an additional topic of this review, and in the last chapter we cover aspects of the importance of xanthophyll cycle-dependent photoprotection for different algal species in their natural environments. [ABSTRACT FROM AUTHOR]

Photoprotection of the chloroplast is an important component of abiotic stress resistance in plants. Carotenoids have a central role in photoprotection. We review here the recent evidence, derived mainly from in vitro reconstitution of recombinant Lhc proteins with different carotenoids and from carotenoid biosynthesis mutants, for the existence of different mechanisms of photoprotection and regulation based on xanthophyll binding to Lhc proteins into multiple sites and the exchange of chromophores between different Lhc proteins during exposure of plants to high light stress and the operation of the xanthophyll cycle. The use of recombinant Lhc proteins has revealed up to four binding sites in members of Lhc families with distinct selectivity for xanthophyll species which are here hypothesised to have different functions. Site L1 is selective for lutein and is here proposed to be essential for catalysing the protection from singlet oxygen by quenching chlorophyll triplets. Site L2 and N1 are here proposed to act as allosteric sites involved in the regulation of chlorophyll singlet excited states by exchanging ligand during the operation of the xanthophyll cycle. Site V1 of the major antenna complex LHC II is here hypothesised to be a deposit for readily available substrate for violaxanthin de-epoxidase rather than a light harvesting pigment. Moreover, xanthophylls bound to Lhc proteins can be released into the lipid bilayer where they contribute to the scavenging of reactive oxygen species produced in excess light. [ABSTRACT FROM AUTHOR]

Photosynthetic rates vary depending on growth conditions, even within species. Remote sensing techniques have a great potential to predict the photosynthetic rates of leaves with different characteristics. Here, we demonstrate that the photosynthetic rates of leaves acclimated to different light and nutrient conditions can be estimated based on the chlorophyll fluorescence (ChlF), the photochemical reflectance index (PRI), and a chlorophyll index. Chenopodium album plants were grown under different light and nutrient conditions. PRI, ChlF parameters, and CO 2 /H 2 O gas exchange rates of leaves were simultaneously determined under the various light and CO 2 conditions. PRI was used to assess non-photochemical quenching (NPQ), but the relationship between NPQ and PRI was weakened when the data on leaves grown under different conditions were pooled, because PRI in darkness ([Formula: see text]) changed with the leaf pigment composition. Among 15 pigment indices, we found that [Formula: see text], a reflectance index related to the leaf chlorophyll content, had the best correlation with [Formula: see text] ([Formula: see text]) across the studied leaves, and the correction of PRI by [Formula: see text] improved the predictability of NPQ ([Formula: see text]). Using the steady-state ChlF, the NPQ estimated from PRI and [Formula: see text], and the stomatal conductance coefficient, we calculated the CO 2 assimilation rates, which were strongly correlated with the actual rates (RMSE = 4.85 [Formula: see text]mol m[Formula: see text] s[Formula: see text]), irrespective of growth conditions. Our approach has the potential to contribute to a more accurate estimation of photosynthetic rates in remote sensing. However, further studies on species variations and connecting with radiative transfer models are needed to demonstrate this at the canopy scale.