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The ability to harvest light effectively in a changing environment is necessary to ensure efficient photosynthesis and crop growth. One mechanism, known as qE, protects photosystem II (PSII) and regulates electron transfer through the harmless dissipation of excess absorbed photons as heat. This process involves reversible clustering of the major light-harvesting complexes of PSII (LHCII) in the thylakoid membrane and relies upon the ΔpH gradient and the allosteric modulator protein PsbS. To date, the exact role of PsbS in the qE mechanism has remained elusive. Here, we show that PsbS induces hydrophobic mismatch in the thylakoid membrane through dynamic rearrangement of lipids around LHCII leading to observed membrane thinning. We found that upon illumination, the thylakoid membrane reversibly shrinks from around 4.3 to 3.2 nm, without PsbS, this response is eliminated. Furthermore, we show that the lipid digalactosyldiacylglycerol (DGDG) is repelled from the LHCII-PsbS complex due to an increase in both the p K a of lumenal residues and in the dipole moment of LHCII, which allows for further conformational change and clustering in the membrane. Our results suggest a mechanistic role for PsbS as a facilitator of a hydrophobic mismatch-mediated phase transition between LHCII-PsbS and its environment. This could act as the driving force to sort LHCII into photoprotective nanodomains in the thylakoid membrane. This work shows an example of the key role of the hydrophobic mismatch process in regulating membrane protein function in plants.

Chlorophyll a fluorescence parameters related to PSII photochemistry, photoprotection and photoinhibition were investigated in four C3 plant species growing in their natural habitat: Prosopis juliflora ; Abutilon indicum ; Salvadora persica ; and Phragmites karka . This study compared the light reaction responses of P. juliflora , an invasive species, with three native co-existing species, which adapt to varying water deficit and high salt stress. Chlorophyll a fluorescence quenching analyses revealed that P. juliflora had the highest photochemical quantum efficiency and yield, regulated by higher fraction of open reaction centres and reduced photoprotective energy dissipation without compromising the integrity of photosynthetic apparatus due to photoinhibition. Moreover, the elevated values of parameters obtained through polyphasic chlorophyll a fluorescence induction kinetics, which characterise the photochemistry of PSII and electron transport, highlighted the superior performance index of energy conservation in the transition from excitation to the reduction of intersystem electron carriers for P. juliflora compared to other species. Enhanced pigment contents and their stoichiometry in P. juliflora apparently contributed to upregulating fluxes and yields of energy absorbance, trapping and transport. This enhanced photochemistry, along with reduced non-photochemical processes, could explain the proclivity for invasion advantage in P. juliflora across diverse stress conditions.

While first-principles methods have been successfully applied to characterize individual properties of multi-principal element alloys (MPEA), their use in searching for optimal trade-offs between competing properties is hampered by high computational demands. In this work, we present a framework to explore Pareto-optimal compositions by integrating advanced ab initio-based techniques into a Bayesian multi-objective optimization workflow, complemented by a simple analytical model providing straightforward analysis of trends. We benchmark the framework by applying it to solid solution strengthening and ductility of refractory MPEAs, with the parameters of the strengthening and ductility models being efficiently computed using a combination of the coherent-potential approximation method, accounting for finite-temperature effects, and actively-learned moment-tensor potentials parameterized with ab initio data. Properties obtained from ab initio calculations are subsequently used to extend predictions of all relevant material properties to a large class of refractory alloys with the help of the analytical model validated by the data and relying on a few element-specific parameters and universal functions that describe bonding between elements. Our findings offer crucial insights into the traditional strength-vs-ductility dilemma of refractory MPEAs. The proposed framework is versatile and can be extended to other materials and properties of interest, enabling a predictive and tractable high-throughput screening of Pareto-optimal MPEAs over the entire composition space., Competing Interests: Competing interestsThe authors declare no competing interests., (© The Author(s) 2024.)

In plants, the major light-harvesting antenna complex (LHCII) is vital for both light harvesting and photoprotection in photosystem II. Previously, we proposed that the thylakoid membrane itself could switch LHCII into the photoprotective state, qE, via a process known as hydrophobic mismatch. The decrease in the membrane thickness that followed the formation of ΔpH was a key fact that prompted this idea. To test this, we made proteoliposomes from lipids with altered acyl chain length (ACL). Here, we show that ACL regulates the average chlorophyll fluorescence lifetime of LHCII. For liposomes made of lipids with an ACL of 18 carbons, the lifetime was ∼2 ns, like that for the thylakoid membrane. Furthermore, LHCII appears to be quenched in proteoliposomes with an ACL both shorter and longer than 18 carbons. The proteoliposomes made of short ACL lipids display structural heterogeneity revealing two quenched conformations of LHCII, each having characteristic 77 K fluorescence spectra. One conformation spectrally resembles isolated LHCII aggregates, whilst the other resembles LHCII immobilized in polyacrylamide gels. Overall, the decrease in the ACL appears to produce quenched conformations of LHCII, which renders plausible the idea that the trigger of qE is the hydrophobic mismatch., Competing Interests: Declaration of Competing Interest Alexander V. Ruban reports financial support was provided by Queen Mary University of London., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

Nonphotochemical quenching (NPQ) is a crucial mechanism for fine-tuning light harvesting and protecting the photosystem II (PSII) reaction centres from excess light energy in plants and algae. This process is regulated by photoprotective proteins LHCSR1, LHCSR3, and PsbS in green algae, such as Chlamydomonas reinhardtii. The det1-2 phot mutant, which overexpresses these photoprotective proteins, resulting in a significantly higher NPQ response, has been recently discovered in C. reinhardtii. Here, we analysed the physiological impact of this response on algal cells and found that det1-2 phot was capable of efficient growth under high light intensities, where wild-type (WT) cells were unable to survive. The mutant exhibited a smaller PSII cross-section in the dark and showed a detachment of the peripheral light-harvesting complex II (LHCII) antenna in the NPQ state, as suggested by a rise in the chlorophyll fluorescence parameter of photochemical quenching in the dark (qPd > 1). Furthermore, fluorescence decay-associated spectra demonstrated a decreased excitation pressure on PSII, with excess energy being directed toward PSI. The amount of LHCSR1, LHCSR3, and PsbS in the mutant correlated with the magnitude of the protective NPQ response. Overall, the study suggests the mechanism by which the overexpression of photoprotective proteins in det1-2 phot brings about an efficient and effective photoprotective response, enabling the mutant to grow and survive under high light intensities that would otherwise be lethal for WT cells., Competing Interests: Declaration of Competing Interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Jun Minagawa reports financial support was provided by Japan Society for the Promotion of Science. Eunchul Kim reports financial support was provided by Japan Society for the Promotion of Science. Eunchul Kim reports financial support was provided by Takeda Science Foundation. No additional relationships to declare., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Photosynthetic light-harvesting antennae are pigment-binding proteins that perform one of the most fundamental tasks on Earth, capturing light and transferring energy that enables life in our biosphere. Adaptation to different light environments led to the evolution of an astonishing diversity of light-harvesting systems. At the same time, several strategies have been developed to optimize the light energy input into photosynthetic membranes in response to fluctuating conditions. The basic feature of these prompt responses is the dynamic nature of antenna complexes, whose function readily adapts to the light available. High-resolution microscopy and spectroscopic studies on membrane dynamics demonstrate the crosstalk between antennae and other thylakoid membrane components. With the increased understanding of light-harvesting mechanisms and their regulation, efforts are focusing on the development of sustainable processes for effective conversion of sunlight into functional bio-products. The major challenge in this approach lies in the application of fundamental discoveries in light-harvesting systems for the improvement of plant or algal photosynthesis. Here, we underline some of the latest fundamental discoveries on the molecular mechanisms and regulation of light harvesting that can potentially be exploited for the optimization of photosynthesis., (© The Author(s) 2022. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

The major photosystem II light-harvesting antenna (LHCII) is the most abundant membrane protein in nature and plays an indispensable role in light harvesting and photoprotection in the plant thylakoid. Here, we show that "pseudothylakoid characteristics" can be observed in artificial LHCII membranes. In our proteoliposomal system, at high LHCII densities, the liposomes become stacked, mimicking the in vivo thylakoid grana membranes. Furthermore, an unexpected, unstructured emission peak at ∼730 nm appears, similar in appearance to photosystem I emission, but with a clear excimeric character that has never been previously reported. These states correlate with the increasing density of LHCII in the membrane and a decrease in its average fluorescence lifetime. The appearance of these low-energy states can also occur in natural plant membrane structures, which has unique consequences for the interpretation of the spectroscopic and physiological properties of the photosynthetic membrane.

Discovered more than 200 years ago in 1821, thermoelectricity is nowadays of global interest as it enables direct interconversion of thermal and electrical energy via the Seebeck/Peltier effect. In their seminal work, Mahan and Sofo mathematically derived the conditions for 'the best thermoelectric'-a delta-distribution-shaped electronic transport function, where charge carriers contribute to transport only in an infinitely narrow energy interval. So far, however, only approximations to this concept were expected to exist in nature. Here, we propose the Anderson transition in a narrow impurity band as a physical realisation of this seemingly unrealisable scenario. An innovative approach of continuous disorder tuning allows us to drive the Anderson transition within a single sample: variable amounts of antisite defects are introduced in a controlled fashion by thermal quenching from high temperatures. Consequently, we obtain a significant enhancement and dramatic change of the thermoelectric properties from p-type to n-type in stoichiometric Fe 2 VAl, which we assign to a narrow region of delocalised electrons in the energy spectrum near the Fermi energy. Based on our electronic transport and magnetisation experiments, supported by Monte-Carlo and density functional theory calculations, we present a novel strategy to enhance the performance of thermoelectric materials., (© 2022. The Author(s).)

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 (Ca 2+ )-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., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

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., (© 2022 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society.)

Photosystem II (PSII) uses light energy to split water into protons, electrons, and oxygen, ultimately sustaining heterotrophic life on Earth. The major light harvesting complex in plants (LHCII) is packed with chlorophylls and carotenoids and is the main supplier of excitation energy to PSII reaction centers. The protein scaffold acts as a programmed solvent for the pigments in LHCII, tuning their orientations while at the same time impeding concentration quenching to ensure efficient storage of excitation energy by chlorophylls. However, under stress, the very fuel of PSII, solar photons, can damage its delicate inner components and hamper photosynthesis. In a crucial regulatory strategy in plants, LHCII evolved a flexible design that allows it to switch between light-harvesting and dissipative conformations, thereby safely releasing the excess energy that is absorbed into heat. Several mechanisms have been proposed to explain chlorophyll de-excitation pathways in LHCII, such as chlorophyll-chlorophyll charge transfer states, resonance energy transfer from chlorophylls to a carotenoid S1 state, and chlorophyll-carotenoid reductive energy transfer. This Perspective critically assesses the listed proposals, addressing both the physical mechanism of quenching and the nature of the quenching pigment. These hypotheses are then discussed in the context of state-of-the-art biochemical, physiological, and genetic knowledge to scrutinize their likeliness to occur in the native thylakoid membranes.

Plant tolerance to high light and oxidative stress is increased by overexpression of the photosynthetic enzyme Ferredoxin:NADP(H) reductase (FNR), but the specific mechanism of FNR-mediated protection remains enigmatic. It has also been reported that the localization of this enzyme within the chloroplast is related to its role in stress tolerance. Here, we dissected the impact of FNR content and location on photoinactivation of photosystem I (PSI) and photosystem II (PSII) during high light stress of Arabidopsis (Arabidopsis thaliana). The reaction center of PSII is efficiently turned over during light stress, while damage to PSI takes much longer to repair. Our results indicate a PSI sepcific effect, where efficient oxidation of the PSI primary donor (P700) upon transition from darkness to light, depends on FNR recruitment to the thylakoid membrane tether proteins: thylakoid rhodanase-like protein (TROL) and translocon at the inner envelope of chloroplasts 62 (Tic62). When these interactions were disrupted, PSI photoinactivation occurred. In contrast, there was a moderate delay in the onset of PSII damage. Based on measurements of ΔpH formation and cyclic electron flow, we propose that FNR location influences the speed at which photosynthetic control is induced, resulting in specific impact on PSI damage. Membrane tethering of FNR therefore plays a role in alleviating high light stress, by regulating electron distribution during short-term responses to light., (© The Author(s) 2021. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

Light harvesting is regulated by a process triggered by the acidification of the thylakoid lumen, known as nonphotochemical "energy-dependent quenching" (qE). In diatoms, qE is controlled by the light-harvesting complex (LHC) protein LHCX1, while the LHC stress-related (LHCSR) and photosystem II subunit S proteins are essential for green algae and plants, respectively. Here, we report a biochemical and molecular characterization of LHCX1 to investigate its role in qE. We found that, when grown under intermittent light, Phaeodactylum tricornutum forms very large qE, due to LHCX1 constitutive upregulation. This "super qE" is abolished in LHCX1 knockout mutants. Biochemical and spectroscopic analyses of LHCX1 reveal that this protein might differ in the character of binding pigments relative to the major pool of light-harvesting antenna proteins. The possibility of transient pigment binding or not binding pigments at all is discussed. Targeted mutagenesis of putative protonatable residues (D95 and E205) in transgenic P. tricornutum lines does not alter qE capacity, showing that they are not involved in sensing lumen pH, differently from residues conserved in LHCSR3. Our results suggest functional divergence between LHCX1 and LHCSR3 in qE modulation. We propose that LHCX1 evolved independently to facilitate dynamic tracking of light fluctuations in turbulent waters. The evolution of LHCX(-like) proteins in organisms with secondary red plastids, such as diatoms, might have conferred a selective advantage in the control of dynamic photoprotection, ultimately resulting in their ecological success., (© American Society of Plant Biologists 2021. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Non-photochemical chlorophyll fluorescence quenching (NPQ) remains one of the most studied topics of the 21st century in photosynthesis research. Over the past 30 years, profound knowledge has been obtained on the molecular mechanism of NPQ in higher plants. First, the largely overlooked significance of NPQ in protecting the reaction center of photosystem II (RCII) against damage, and the ways to assess its effectiveness are highlighted. Then, the key in vivo signals that can monitor the life of the major NPQ component, qE, are presented. Finally, recent knowledge on the site of qE and the possible molecular events that transmit ΔpH into the conformational change in the major LHCII [the major trimeric light harvesting complex of photosystem II (PSII)] antenna complex are discussed. Recently, number of reports on Arabidopsis mutants lacking various antenna components of PSII confirmed that the in vivo site of qE rests within the major trimeric LHCII complex. Experiments on biochemistry, spectroscopy, microscopy and molecular modeling suggest an interplay between thylakoid membrane geometry and the dynamics of LHCII, the PsbS (PSII subunit S) protein and thylakoid lipids. The molecular basis for the qE-related conformational change in the thylakoid membrane, including the possible onset of a hydrophobic mismatch between LHCII and lipids, potentiated by PsbS protein, begins to unfold., (© The Author(s) 2020. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

The proton motive force (pmf) across the thylakoid membrane couples photosynthetic electron transport and ATP synthesis. In recent years, the electrochromic carotenoid and chlorophyll absorption band shift (ECS), peaking ∼515 nm, has become a widely used probe to measure pmf in leaves. However, the use of this technique to calculate the parsing of the pmf between the proton gradient (ΔpH) and electric potential (Δψ) components remains controversial. Interpretation of the ECS signal is complicated by overlapping absorption changes associated with violaxanthin de-epoxidation to zeaxanthin (ΔA505) and energy-dependent nonphotochemical quenching (qE; ΔA535). In this study, we used Arabidopsis (Arabidopsis thaliana) plants with altered xanthophyll cycle activity and photosystem II subunit S (PsbS) content to disentangle these overlapping contributions. In plants where overlap among ΔA505, ΔA535, and ECS is diminished, such as npq4 (lacking ΔA535) and npq1npq4 (also lacking ΔA505), the parsing method implies the Δψ contribution is virtually absent and pmf is solely composed of ΔpH. Conversely, in plants where ΔA535 and ECS overlap is enhanced, such as L17 (a PsbS overexpressor) and npq1 (where ΔA535 is blue-shifted to 525 nm) the parsing method implies a dominant contribution of Δψ to the total pmf. These results demonstrate the vast majority of the pmf attributed by the ECS parsing method to Δψ is caused by ΔA505 and ΔA535 overlap, confirming pmf is dominated by ΔpH following the first 60 s of continuous illumination under both low and high light conditions. Further implications of these findings for the regulation of photosynthesis are discussed., (© American Society of Plant Biologists 2021. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Photosystems possess distinct fluorescence emissions at low (77K) temperature. PSI emits in the long-wavelength region at ~710-740 nm. In diatoms, a successful clade of marine primary producers, the contribution of PSI-associated emission (710-717 nm) has been shown to be relatively small. However, in the pennate diatom Phaeodactylum tricornutum, the source of the long-wavelength emission at ~710 nm (F710) remains controversial. Here, we addressed the origin and modulation of F710 fluorescence in this alga grown under continuous and intermittent light. The latter condition led to a strong enhancement in F710. Biochemical and spectral properties of the photosynthetic complexes isolated from thylakoid membranes were investigated for both culture conditions. F710 emission appeared to be associated with PSI regardless of light acclimation. To further assess whether PSII could also contribute to this emission, we decreased the concentration of PSII reaction centres and core antenna by growing cells with lincomycin, a chloroplast protein synthesis inhibitor. The treatment did not diminish F710 fluorescence. Our data suggest that F710 emission originates from PSI under the conditions tested and is enhanced in intermittent light-grown cells due to increased energy flow from the FCP antenna to PSI., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

High light intensities raise photosynthetic and plant growth rates but can cause damage to the photosynthetic machinery. The likelihood and severity of deleterious effects are minimised by a set of photoprotective mechanisms, one key process being the controlled dissipation of energy from chlorophyll within PSII known as non-photochemical quenching (NPQ). Although ubiquitous, the role of NPQ in plant productivity is important because it momentarily reduces the quantum efficiency of photosynthesis. Rice plants overexpressing and deficient in the gene encoding a central regulator of NPQ, the protein PsbS, were used to assess the effect of protective effectiveness of NPQ (pNPQ) at the canopy scale. Using a combination of three-dimensional reconstruction, modelling, chlorophyll fluorescence, and gas exchange, the influence of altered NPQ capacity on the distribution of pNPQ was explored. A higher phototolerance in the lower layers of a canopy was found, regardless of genotype, suggesting a mechanism for increased protection for leaves that experience relatively low light intensities interspersed with brief periods of high light. Relative to wild-type plants, psbS overexpressors have a reduced risk of photoinactivation and early growth advantage, demonstrating that manipulating photoprotective mechanisms can impact both subcellular mechanisms and whole-canopy function., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Nonphotochemical quenching (NPQ) is a mechanism of regulating light harvesting that protects the photosynthetic apparatus from photodamage by dissipating excess absorbed excitation energy as heat. In higher plants, the major light-harvesting antenna complex (LHCII) of photosystem (PS) II is directly involved in NPQ. The aggregation of LHCII is proposed to be involved in quenching. However, the lack of success in isolating native LHCII aggregates has limited the direct interrogation of this process. The isolation of LHCII in its native state from thylakoid membranes has been problematic because of the use of detergent, which tends to dissociate loosely bound proteins, and the abundance of pigment-protein complexes (e.g. PSI and PSII) embedded in the photosynthetic membrane, which hinders the preparation of aggregated LHCII. Here, we used a novel purification method employing detergent and amphipols to entrap LHCII in its natural states. To enrich the photosynthetic membrane with the major LHCII, we used Arabidopsis thaliana plants lacking the PSII minor antenna complexes (NoM), treated with lincomycin to inhibit the synthesis of PSI and PSII core proteins. Using sucrose density gradients, we succeeded in isolating the trimeric and aggregated forms of LHCII antenna. Violaxanthin- and zeaxanthin-enriched complexes were investigated in dark-adapted, NPQ, and dark recovery states. Zeaxanthin-enriched antenna complexes showed the greatest amount of aggregated LHCII. Notably, the amount of aggregated LHCII decreased upon relaxation of NPQ. Employing this novel preparative method, we obtained a direct evidence for the role of in vivo LHCII aggregation in NPQ., (Copyright © 2020 © 2020 Shukla et al. Published by Elsevier Inc. All rights reserved.)

An intriguing molecular architecture called the "semi-crystalline photosystem II (PSII) array" has been observed in the thylakoid membranes in vascular plants. It is an array of PSII-light-harvesting complex II (LHCII) supercomplexes that only appears in low light, but its functional role has not been clarified. Here, we identified PSII-LHCII supercomplexes in their monomeric and multimeric forms in low light-acclimated spinach leaves and prepared them using sucrose-density gradient ultracentrifugation in the presence of amphipol A8-35. When the leaves were acclimated to high light, only the monomeric forms were present, suggesting that the multimeric forms represent a structural adaptation to low light and that disaggregation of the PSII-LHCII supercomplex represents an adaptation to high light. Single-particle EM revealed that the multimeric PSII-LHCII supercomplexes are composed of two ("megacomplex") or three ("arraycomplex") units of PSII-LHCII supercomplexes, which likely constitute a fraction of the semi-crystalline PSII array. Further characterization with fluorescence analysis revealed that multimeric forms have a higher light-harvesting capability but a lower thermal dissipation capability than the monomeric form. These findings suggest that the configurational conversion of PSII-LHCII supercomplexes may serve as a structural basis for acclimation of plants to environmental light., Competing Interests: Conflict of interest—The authors declare that they have no conflicts of interest with the contents of this article., (© 2020 Kim et al.)

Xanthophylls in light harvesting complexes perform a number of functions ranging from structural support to light-harvesting and photoprotection. In the major light harvesting complex of photosystem II in plants (LHCII), the innermost xanthophyll binding pockets are occupied by lutein molecules. The conservation of these sites within the LHC protein family suggests their importance in LHCII functionality. In the present work, we induced the photoprotective switch in LHCII isolated from the Arabidopsis mutant npq1lut2, where the lutein molecules are exchanged with violaxanthin. Despite the differences in the energetics of the pigments and the impairment of chlorophyll fluorescence quenching in vivo, we show that isolated complexes containing violaxanthin are still able to induce the quenching switch to a similar extent to wild type LHCII monomers. Moreover, the same spectroscopic changes take place, which suggest the involvement of the terminal emitter site (L1) in energy dissipation in both complexes. These results indicate the robust nature of the L1 xanthophyll binding domain in LHCII, where protein structural cues are the major determinant of the function of the bound carotenoid.

The major light-harvesting complex of photosystem II (LHCII) is the main contributor to sunlight energy harvesting in plants. The flexible design of LHCII underlies a photoprotective mechanism whereby this complex switches to a dissipative state in response to high light stress, allowing the rapid dissipation of excess excitation energy (non-photochemical quenching, NPQ). In this work, we locked single LHCII trimers in a quenched conformation after immobilization of the complexes in polyacrylamide gels to impede protein interactions. A comparison of their pigment excited-state dynamics with quenched LHCII aggregates in buffer revealed the presence of a new spectral band at 515 nm arising after chlorophyll excitation. This is suggested to be the signature of a carotenoid excited state, linked to the quenching of chlorophyll singlet excited states. Our data highlight the marked sensitivity of pigment excited-state dynamics in LHCII to structural changes induced by the environment., Competing Interests: The authors declare no competing interests., (© 2020 The Author(s).)

Cyanobacterial thylakoid membranes represent the active sites for both photosynthetic and respiratory electron transport. We used high-resolution atomic force microscopy to visualize the native organization and interactions of photosynthetic complexes within the thylakoid membranes from the model cyanobacterium Synechococcus elongatus PCC 7942. The thylakoid membranes are heterogeneous and assemble photosynthetic complexes into functional domains to enhance their coordination and regulation. Under high light, the chlorophyll-binding proteins IsiA are strongly expressed and associate with Photosystem I (PSI), forming highly variable IsiA-PSI supercomplexes to increase the absorption cross-section of PSI. There are also tight interactions of PSI with Photosystem II (PSII), cytochrome b 6 f, ATP synthase and NAD(P)H dehydrogenase complexes. The organizational variability of these photosynthetic supercomplexes permits efficient linear and cyclic electron transport as well as bioenergetic regulation. Understanding the organizational landscape and environmental adaptation of cyanobacterial thylakoid membranes may help inform strategies for engineering efficient photosynthetic systems and photo-biofactories.

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 Q A 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., (© 2020 American Society of Plant Biologists. All Rights Reserved.)

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., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The spring ephemeral Berteroa incana is a familial relative of Arabidopsis thaliana and thrives in a diverse range of terrestrial ecosystems. Within this study, the novel chlorophyll fluorescence parameter of photochemical quenching in the dark (qPd) was used to measure the redox state of the primary quinone electron acceptor (Q A ) in order to estimate the openness of photosystem II (PSII) reaction centres (RC). From this, the early onset of photoinactivation can be sensitively quantified alongside the light tolerance of PSII and the photoprotective efficiency of nonphotochemical quenching (NPQ). This study shows that, with regards to A. thaliana, NPQ is enhanced in B. incana in both low-light (LL) and high-light (HL) acclimation states. Moreover, light tolerance is increased by up to 500%, the rate of photoinactivation is heavily diminished, and the ability to recover from light stress is enhanced in B. incana, relative to A. thaliana. This is due to faster synthesis of zeaxanthin and a larger xanthophyll cycle (XC) pool available for deepoxidation. Moreover, preferential energy transfer via CP47 around the RC further enhances efficient photoprotection. As a result, a high functional cross-section of photosystem II is maintained and is not downregulated when B. incana is acclimated to HL. A greater capacity for protective NPQ allows B. incana to maintain an enhanced light-harvesting capability when acclimated to a range of light conditions. This enhancement of flexible short-term protection saves the metabolic cost of long-term acclimatory changes., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Photoprotection refers to a set of well defined plant processes that help to prevent the deleterious effects of high and excess light on plant cells, especially within the chloroplast. Molecular components of chloroplast photoprotection are closely aligned with those of photosynthesis and together they influence productivity. Proof of principle now exists that major photoprotective processes such as non-photochemical quenching (NPQ) directly determine whole canopy photosynthesis, biomass and yield via prevention of photoinhibition and a momentary downregulation of photosynthetic quantum yield. However, this phenomenon has neither been quantified nor well characterized across different environments. Here we address this problem by assessing the existing literature with a different approach to that taken previously, beginning with our understanding of the molecular mechanism of NPQ and its regulation within dynamic environments. We then move to the leaf and the plant level, building an understanding of the circumstances (when and where) NPQ limits photosynthesis and linking to our understanding of how this might take place on a molecular and metabolic level. We argue that such approaches are needed to fine tune the relevant features necessary for improving dynamic NPQ in important crop species., (© 2019 University of Nottingham. The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

The photosynthetic apparatus of plants is a robust self-adjustable molecular system, able to function efficiently under varying environmental conditions. Under strong sunlight, it switches into photoprotective mode to avoid overexcitation by safely dissipating the excess absorbed light energy via nonphotochemical quenching (NPQ). Unfortunately, heterogeneous organization and simultaneous occurrence of multiple processes within the thylakoid membrane impede the study of natural NPQ under in vivo conditions; thus, usually artificially prepared antennae have been studied instead. However, it has never been shown directly that the origin of fluorescence quenching observed in these artificial systems underlies natural NPQ. Here we report the time-resolved fluorescence measurements of the dark-adapted and preilluminated-to induce NPQ-intact chloroplasts, performed over a broad temperature range. We show that their spectral response matches that observed in the LHCII aggregates, thus demonstrating explicitly for the first time that the latter in vitro system preserves essential properties of natural photoprotection.

Chlorophyll triplet excited states are byproducts of photosynthetic processes that can indirectly harm biological membranes by forming highly reactive oxygen species. A crucial photoprotective mechanism evolved by plants to counter this threat involves the triplet energy transfer from chlorophylls to carotenoid molecules, in which triplet states are not reactive. In the major light-harvesting complex of photosystem II (LHCII), the two central luteins play an important role in the mechanism, but it has been shown that carotenoid triplets are formed even when other carotenoids replace them in their binding sites. In this work, we have investigated carotenoid triplet formation in LHCII isolated from Arabidopsis thaliana npq1lut2 plants, in which violaxanthin replaces lutein. Although transient absorption spectroscopy showed altered singlet excited-state dynamics in the mutant LHCII without lutein, these antennae formed carotenoid triplets that were spectrally and dynamically identical to the wild-type protein. We conclude that lutein-binding sites in LHCII have conserved characteristics to ensure efficient triplet energy transfer to the carotenoid molecules that they accommodate, making the identity of the carotenoid trivial per se.

Carotenoids in photosynthetic proteins carry out the dual function of harvesting light and defending against photo-damage by quenching excess energy. The latter involves the low-lying, dark, excited state labelled S1. Here "dark" means optically-forbidden, a property that is often attributed to molecular symmetry, which leads to speculation that its optical properties may be strongly-perturbed by structural distortions. This has been both explicitly and implicitly proposed as an important feature of photo-protective energy quenching. Here we present a theoretical analysis of the relationship between structural distortions and S1 optical properties. We outline how S1 is dark not because of overall geometric symmetry but because of a topological symmetry related to bond length alternation in the conjugated backbone. Taking the carotenoid echinenone as an example and using a combination of molecular dynamics, quantum chemistry, and the theory of spectral lineshapes, we show that distortions that break this symmetry are extremely stiff. They are therefore absent in solution and only marginally present in even a very highly-distorted protein binding pocket such as in the Orange Carotenoid Protein (OCP). S1 remains resolutely optically-forbidden despite any breaking of bulk molecular symmetry by the protein environment. However, rotations of partially conjugated end-rings can result in fine tuning of the S1 transition density which may exert some influence on interactions with neighbouring chromophores.

Main Conclusion: The absence of state transitions in a Nt(Hn) cybrid is due to a cleavage of the threonine residue from the misprocessed N-terminus of the LHCII polypeptides. The cooperation between the nucleus and chloroplast genomes is essential for plant photosynthetic fitness. The rapid and specific interactions between nucleus-encoded and chloroplast-encoded proteins are under intense investigation with potential for applications in agriculture and renewable energy technology. Here, we present a novel model for photosynthesis research in which alien henbane (Hyoscyamus niger) chloroplasts function on the nuclear background of a tobacco (Nicotiana tabacum). The result of this coupling is a cytoplasmic hybrid (cybrid) with inhibited state transitions-a mechanism responsible for balancing energy absorption between photosystems. Protein analysis showed differences in the LHCII composition of the cybrid plants. SDS-PAGE analysis revealed a novel banding pattern in the cybrids with at least one additional 'LHCII' band compared to the wild-type parental species. Proteomic work suggested that the N-terminus of at least some of the cybrid Lhcb proteins was missing. These findings provide a mechanistic explanation for the lack of state transitions-the N-terminal truncation of the Lhcb proteins in the cybrid included the threonine residue that is phosphorylated/dephosphorylated in order to trigger state transitions and therefore crucial energy balancing mechanism in plants.

Plants have developed multiple self-regulatory mechanisms to efficiently function under varying sunlight conditions. At high light intensities, non-photochemical quenching (NPQ) is activated on a molecular level, safely dissipating an excess excitation as heat. The exact molecular mechanism for NPQ is still under debate, but it is widely agreed that the direct participation of the carotenoid pigments is involved, one of the proposed candidate being the zeaxanthin. In this work, we performed fluorescence measurements of violaxanthin- and zeaxanthin-enriched major light-harvesting complexes (LHCII), in ensemble and at the single pigment-protein complex level, where aggregation is prevented by immobilization of LHCIIs onto a surface. We show that a selective enrichment of LHCII with violaxanthin or zeaxanthin affects neither the ability of LHCII to switch into a dissipative conformation nor the maximal level of induced quenching. However, the kinetics of the fluorescence decrease due to aggregation on the timescale of seconds are different, prompting towards a modulatory effect of zeaxanthin in the dynamics of quenching., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Photoinhibition is the light-induced down-regulation of photosynthetic efficiency, the primary target of which is photosystem II (PSII). Currently, there is no clear consensus on the exact mechanism of this process. However, it is clear that inhibition can occur through limitations on both the acceptor- and donor side of PSII. The former mechanism is caused by electron transport limitations at the PSII acceptor side. Whilst, the latter mechanism relies on the disruption of the oxygen-evolving complex. Both of these mechanisms damage the PSII reaction centre (RC). Using a novel chlorophyll fluorescence methodology, RC photoinactivation can be sensitively measured and quantified alongside photoprotection in vivo This is achieved through estimation of the redox state of Q A , using the parameter of photochemical quenching in the dark (qPd). This study shows that through the use of PSII donor-side inhibitors, such as UV-B and Cd 2+ , there is a steeper gradient of photoinactivation in the systems with a weakened donor side, independent of the level of NPQ attained. This is coupled with a concomitant decline in the light tolerance of PSII. The native light tolerance is partially restored upon the use of 1,5-diphenylcarbazide (DPC), a PSII electron donor, allowing for the balance between the inhibitory pathways to be sensitively quantified. Thus, this study confirms that the impact of donor-side inhibition can be detected alongside acceptor-side photoinhibition using the qPd parameter and confirms qPd as a valid, sensitive and unambiguous parameter to sensitively quantify the onset of photoinhibition through both acceptor- or donor-side mechanisms., (© 2019 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society.)

Photosynthetic organisms require rapid and reversible down-regulation of light harvesting to avoid photodamage. Response to unpredictable light fluctuations is achieved by inducing energy-dependent quenching, qE, which is the major component of the process known as non-photochemical quenching (NPQ) of chlorophyll fluorescence. qE is controlled by the operation of the xanthophyll cycle and accumulation of specific types of proteins, upon thylakoid lumen acidification. The protein cofactors so far identified to modulate qE in photosynthetic eukaryotes are the photosystem II subunit S (PsbS) and light-harvesting complex stress-related (LHCSR/LHCX) proteins. A transition from LHCSR- to PsbS-dependent qE took place during the evolution of the Viridiplantae (also known as 'green lineage' organisms), such as green algae, mosses and vascular plants. Multiple studies showed that LHCSR and PsbS proteins have distinct functions in the mechanism of qE. LHCX(-like) proteins are closely related to LHCSR proteins and found in 'red lineage' organisms that contain secondary red plastids, such as diatoms. Although LHCX proteins appear to control qE in diatoms, their role in the mechanism remains poorly understood. Here, we present the current knowledge on the functions and evolution of these crucial proteins, which evolved in photosynthetic eukaryotes to optimise light harvesting., (© 2018 The Author(s). Published by Portland Press Limited on behalf of the Biochemical Society.)

In 1991, my colleagues and I published a hypothesis article that proposed a mechanism that controls light harvesting in plants and protects them against photodamage. The major light harvesting complex, LHCII, was suggested to undergo aggregation upon exposure of the plant to damaging levels of light. Aggregated LHCII was found to be much less efficient in light harvesting, as it promptly dissipated absorbed energy into heat, possessing a very low chlorophyll fluorescence yield. Nonphotochemical quenching (NPQ) is a term coined to describe this reduction in chlorophyll fluorescence yield. This article is a story of how the hypothesis that LHCII aggregation is involved in NPQ is developed into a model that is now becoming broadly accepted by the research community., (© 2018 Federation of European Biochemical Societies.)

Non-photochemical quenching (NPQ) of chlorophyll fluorescence is the process by which excess light energy is harmlessly dissipated within the photosynthetic membrane. The fastest component of NPQ, known as energy-dependent quenching (qE), occurs within minutes, but the site and mechanism of qE remain of great debate. Here, the chlorophyll fluorescence of Arabidopsis thaliana wild type (WT) plants was compared to mutants lacking all minor antenna complexes (NoM). Upon illumination, NoM exhibits altered chlorophyll fluorescence quenching induction (i.e. from the dark-adapted state) characterised by three different stages: (i) a fast quenching component, (ii) transient fluorescence recovery and (iii) a second quenching component. The initial fast quenching component originates in light harvesting complex II (LHCII) trimers and is dependent upon PsbS and the formation of a proton gradient across the thylakoid membrane (ΔpH). Transient fluorescence recovery is likely to occur in both WT and NoM plants, but it cannot be overcome in NoM due to impaired ΔpH formation and a reduced zeaxanthin synthesis rate. Moreover, an enhanced fluorescence emission peak at ~679 nm in NoM plants indicates detachment of LHCII trimers from the bulk antenna system, which could also contribute to the transient fluorescence recovery. Finally, the second quenching component is triggered by both ΔpH and PsbS and enhanced by zeaxanthin synthesis. This study indicates that minor antenna complexes are not essential for qE, but reveals their importance in electron stransport, ΔpH formation and zeaxanthin synthesis., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Photoprotective non-photochemical quenching (NPQ) represents an effective way to dissipate the light energy absorbed in excess by most phototrophs. It is often claimed that NPQ formation/relaxation kinetics are determined by xanthophyll composition. We, however, found that, for the alveolate alga Chromera velia, this is not the case. In the present paper, we investigated the reasons for the constitutive high rate of quenching displayed by the alga by comparing its light harvesting strategies with those of a model phototroph, the land plant Spinacia oleracea. Experimental results and in silico studies support the idea that fast quenching is due not to xanthophylls, but to intrinsic properties of the Chromera light harvesting complex (CLH) protein, related to amino acid composition and protein folding. The pKa for CLH quenching was shifted by 0.5 units to a higher pH compared with higher plant antennas (light harvesting complex II; LHCII). We conclude that, whilst higher plant LHCIIs are better suited for light harvesting, CLHs are 'natural quenchers' ready to switch into a dissipative state. We propose that organisms with antenna proteins intrinsically more sensitive to protons, such as C. velia, carry a relatively high concentration of violaxanthin to improve their light harvesting. In contrast, higher plants need less violaxanthin per chlorophyll because LHCII proteins are more efficient light harvesters and instead require co-factors such as zeaxanthin and PsbS to accelerate and enhance quenching.

Photosystems I and II convert solar energy into the chemical energy that powers life. Chlorophyll a photochemistry, using red light (680 to 700 nm), is near universal and is considered to define the energy "red limit" of oxygenic photosynthesis. We present biophysical studies on the photosystems from a cyanobacterium grown in far-red light (750 nm). The few long-wavelength chlorophylls present are well resolved from each other and from the majority pigment, chlorophyll a. Charge separation in photosystem I and II uses chlorophyll f at 745 nm and chlorophyll f (or d) at 727 nm, respectively. Each photosystem has a few even longer-wavelength chlorophylls f that collect light and pass excitation energy uphill to the photochemically active pigments. These photosystems function beyond the red limit using far-red pigments in only a few key positions., (Copyright © 2018 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Main Conclusion: The macroalga Bryopsis corticulans relies on a sustained protective NPQ and a peculiar body architecture to efficiently adapt to the extreme light changes of intertidal shores. During low tides, intertidal algae experience prolonged high light stress. Efficient dissipation of excess light energy, measured as non-photochemical quenching (NPQ) of chlorophyll fluorescence, is therefore required to avoid photodamage. Light-harvesting regulation was studied in the intertidal macroalga Bryopsis corticulans, during high light and air exposure. Photosynthetic capacity and NPQ kinetics were assessed in different filament layers of the algal tufts and in intact chloroplasts to unravel the nature of NPQ in this siphonous green alga. We found that the morphology and pigment composition of the B. corticulans body provides functional segregation between surface sunlit filaments (protective state) and those that are underneath and undergo severe light attenuation (light-harvesting state). In the surface filaments, very high and sustained NPQ gradually formed. NPQ induction was triggered by the formation of transthylakoid proton gradient and independent of the xanthophyll cycle. PsbS and LHCSR proteins seem not to be active in the NPQ mechanism activated by this alga. Our results show that B. corticulans endures excess light energy pressure through a sustained protective NPQ, not related to photodamage, as revealed by the unusually quick restoration of photosystem II (PSII) function in the dark. This might suggest either the occurrence of transient PSII photoinactivation or a fast rate of PSII repair cycle.

The requirements for ascorbate for growth and photosynthesis were assessed under low (LL; 250 µmol m-2 s-1) or high (HL; 1600 µmol m-2 s-1) irradiance in wild-type Arabidopsis thaliana and two ascorbate synthesis mutants (vtc2-1 and vtc2-4) that have 30% wild-type ascorbate levels. The low ascorbate mutants had the same numbers of leaves but lower rosette area and biomass than the wild type under LL. Wild-type plants experiencing HL had higher leaf ascorbate, anthocyanin, and xanthophyll pigments than under LL. In contrast, leaf ascorbate levels were not increased under HL in the mutant lines. While the degree of oxidation measured using an in vivo redox reporter in the nuclei and cytosol of the leaf epidermal and stomatal cells was similar under both irradiances in all lines, anthocyanin levels were significantly lower in the low ascorbate mutants than in the wild type under HL. Differences in the photosynthetic responses of vtc2-1 and vtc2-4 mutants were observed. Unlike vtc2-1, the vtc2-4 mutants had wild-type zeaxanthin contents. While both low ascorbate mutants had lower levels of non-photochemical quenching of chlorophyll a fluorescence (NPQ) than the wild type under HL, qPd values were greater only in vtc2-1 leaves. Ascorbate is therefore essential for growth but not for photoprotection.

We present the phase diagram of Ce1-xGdxO2-x/2 (CGO), calculated by means of a combined Density Functional Theory (DFT), cluster expansion and lattice Monte Carlo approach. We show that this methodology gives reliable results for the whole range of concentrations (x ≡ xGd ≤ 1). In the thermodynamic equilibrium, we observe two transitions: the onset of oxygen-vacancy (O-Va) ordering at ca. 1200-3300 K for concentrations xGd = 0.3-1, and a phase separation into CeO2 and C-type Gd2O3 occurring below ca. 1000 K for all concentrations. We also model 'quenched' systems, with cations immobile below 1500 K, and observe that the presence of random-like cation configurations does not prevent C-type vacancy ordering. The obtained transition temperatures for Va ordering agree rather well with existing experimental data. We analyse the effect of vacancy ordering and composition on the lattice parameters and relaxation pattern of cations.

Photoinhibition is the light-induced reduction in photosynthetic efficiency and is usually associated with damage to the D1 photosystem II (PSII) reaction centre protein. This damage must either be repaired, through the PSII repair cycle, or prevented in the first place by nonphotochemical quenching (NPQ). Both NPQ and D1 repair contribute to light tolerance because they ensure the long-term maintenance of the highest quantum yield of PSII. However, the relative contribution of each of these processes is yet to be elucidated. The application of a pulse amplitude modulation fluorescence methodology, called protective NPQ, enabled us to evaluate of the protective effectiveness of the processes. Within this study, the contribution of NPQ and D1 repair to the photoprotective capacity of Arabidopsis thaliana was elucidated by using inhibitors and mutants known to affect each process. We conclude that NPQ contributes a greater amount to the maintenance of a high PSII yield than D1 repair under short periods of illumination. This research further supports the role of protective components of NPQ during light fluctuations and the value of protective NPQ and q Pd as unambiguous fluorescence parameters, as opposed to q I and F v /F m , for quantifying photoinactivation of reaction centre II and light tolerance of photosynthetic organisms., (© 2017 John Wiley & Sons Ltd.)