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Abstract Background The growth and yield of pepper, a typical temperature-loving vegetable, are limited by low-temperature environments. Using low-temperature sensitive ‘Hangjiao No. 4’ (Capsicum annuum L.) as experimental material, this study analyzed the changes in plant growth and photosynthesis under different treatments: normal control (NT), low-temperature stress alone (LT), low-temperature stress in strigolactone pretreated plants (SL_LT), and low-temperature stress in strigolactone biosynthesis inhibitor pretreated plants (Tis_LT). Results SL pretreatment increased the net photosynthetic rate (Pn) and PSII actual photochemical efficiency (φPSII), reducing the inhibition of LT on the growth of pepper by 17.44% (dry weight of shoot). Due to promoting the accumulation of carotenoids, such as lutein, and the de-epoxidation of the xanthophyll cycle [(Z + A)/(Z + A + V)] by strigolactone after long-term low-temperature stress (120 h), non-photochemical quenching (NPQ) of pepper was increased to reduce the excess excitation energy [(1-qP)/NPQ] and the photoinhibition degree (Fv/Fm) of pepper seedlings under long-term low-temperature stress was alleviated. Twelve cDNA libraries were constructed from pepper leaves by transcriptome sequencing. There were 8776 differentially expressed genes (DEGs), including 4473 (51.0%) upregulated and 4303 (49.0%) downregulated genes. Gene ontology pathway annotation showed that based on LT, the DEGs of SL_LT and Tis_LT were significantly enriched in the cellular component, which is mainly related to the photosystem and thylakoids. Further analysis of the porphyrin and chlorophyll biosynthesis, carotenoid biosynthesis, photosynthesis-antenna protein, and photosynthetic metabolic pathways and the Calvin cycle under low-temperature stress highlighted 18, 15, 21, 29, and 31 DEGs for further study, which were almost all highly expressed under SL_LT treatment and moderately expressed under LT treatment, whereas Tis_LT showed low expression. Conclusion The positive regulatory effect of SLs on the low-temperature tolerance of pepper seedlings was confirmed. This study provided new insights for the development of temperature-tolerant pepper lines through breeding programs.

Monogalactosyldiacylglycerol (MGDG) is the main lipid constituent of thylakoids and a structural component of photosystems and photosynthesis-related proteo-lipid complexes in green tissues. Previously reported changes in MGDG abundance upon stress treatments are hypothesized to reflect mobilization of MGDG-based polyunsaturated lipid intermediates to maintain extraplastidial membrane integrity. While exchange of lipid intermediates between compartmental membranes is well documented, physiological consequences of mobilizing an essential thylakoid lipid, such as MGDG, for an alternative purpose are not well understood. Arabidopsis seedlings exposed to mild (50 mM) salt treatment displayed significantly increased abundance of both MGDG and the extraplastidial lipid, phosphatidylcholine (PC). Interestingly, similar increases in MGDG and PC were observed in Arabidopsis fad3 mutant seedlings defective in endoplasmic reticulum (ER)–localized linolenic acid formation, in which compensatory plastid-to-ER-directed mobilization of linolenic acid–containing intermediates takes place. The postulated (salt) or evident (fad3) plastid–ER exchange of intermediates concurred with altered thylakoid function according to parameters of photosynthetic performance. While salt treatment of wild-type seedlings inhibited photosynthetic parameters in a dose-dependent manner, interestingly, untreated fad3 mutants did not show overall reduced photosynthetic quantum yield. By contrast, we observed a reduction specifically of non-photochemical quenching (NPQ) under high light, representing only part of observed salt effects. The decreased NPQ in the fad3 mutant was accompanied by reduced activity of the xanthophyll cycle, leading to a reduced concentration of the NPQ-effective pigment zeaxanthin. The findings suggest that altered ER-located fatty acid unsaturation and ensuing inter-organellar compensation impacts on the function of specific thylakoid enzymes, rather than globally affecting thylakoid function. [ABSTRACT FROM AUTHOR]

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

The review highlights the relationship between the molecular organization of the light-harvesting complex of photosystem II (LHCII) and sunlight utilization by higher plants. The molecular form of LHCII switches rapidly and reversibly during diurnal changes of light intensity, from low (ca. 10) to high [ca. 1,000 µmol(photon) m-2 s-1], so the sensitivity of LHCII to light may control the balance between light harvesting and photoprotection state. Our understanding and concept of this mechanism are based on the knowledge of the structure and photophysics of different LHCII molecular forms: monomer, dimer, trimer, and aggregate. It is proposed that LHCII monomers, dimers, and lateral aggregates are fundamental blocks of excess light-dissipation machinery. Trimer is exceptionally well suited to play a physiological role of an antenna complex. A correlation between the LHCII molecular form and the presence of xanthophyll cycle pigment violaxanthin and zeaxanthin in the complex structure is also shown. Moreover, the role of LHCII protein phosphorylation in thylakoid membrane architecture is also discussed. The dual function of LHCII has been studied in the natural thylakoid membranes of chloroplasts, in the artificial lipid-LHCII model membranes, and by suspension of LHCII in a detergent solution.

Conifers growing in temperate forests utilize sustained forms of thermal dissipation during winter to protect the photosynthetic apparatus from damage, which can be monitored via pronounced reductions in photochemical efficiency (F v/ F m) during winter. Eastern white pine (Pinus strobus L.) and white spruce (Picea   glauca (Moench) Voss) are known to recover from winter stress at different rates, with pine recovering more slowly than spruce, suggesting different mechanisms for sustained dissipation in these species. Our objectives were to monitor pine and spruce throughout spring recovery in order to provide insights into key mechanisms for sustained dissipation in both species. We measured chlorophyll fluorescence, pigments, and abundance and phosphorylation status of key photosynthetic proteins. We found that both species rely on two forms of sustained dissipation involving retention of high amounts of antheraxanthin (A) + zeaxanthin (Z), one that is very slowly reversible and temperature independent and one that is more dynamic and occurs only on subzero days. Differences in protein abundance suggest that spruce, but not pine, likely upregulates cyclic or alternative pathways of electron transport involving the cytochrome b6f complex and photosystem I (PSI). Both species show an increased sustained phosphorylation of the D1 protein on subzero days, and spruce additionally shows dramatic increases in the sustained phosphorylation of light-harvesting complex II (LHCII) and other PSII core proteins on subzero days only, suggesting that a mechanism of sustained dissipation that is temperature dependent requires sustained phosphorylation of photosynthetic proteins in spruce, possibly allowing for direct energy transfer from PSII to PSI as a mechanism of photoprotection. The data suggest differences in strategy among conifers in mechanisms of sustained thermal dissipation in response to winter stress. Additionally, the flexible induction of sustained A + Z and phosphorylation of photosynthetic proteins in response to subzero temperatures during spring recovery seem to be important in providing photoprotection during transitional periods with high temperature fluctuation. [ABSTRACT FROM AUTHOR]

The carotenoid pathway in plants has been altered through metabolic engineering to enhance their nutritional value and generate keto-carotenoids, which are widely sought after in the food, feed, and human health industries. In this study, the aim was to produce keto-carotenoids by manipulating the native carotenoid pathway in tobacco plants through chloroplast engineering. Transplastomic tobacco plants were generated that express a synthetic multigene operon composed of three heterologous genes, with Intercistronic Expression Elements (IEEs) for effective mRNA splicing. The metabolic changes observed in the transplastomic plants showed a significant shift towards the xanthophyll cycle, with only a minor production of keto-lutein. The use of a ketolase gene in combination with the lycopene cyclase and hydroxylase genes was a novel approach and demonstrated a successful redirection of the carotenoid pathway towards the xanthophyll cycle and the production of keto-lutein. This study presents a scalable molecular genetic platform for the development of novel keto-carotenoids in tobacco using the Design–Build–Test–Learn (DBTL) approach. This study corroborates chloroplast metabolic engineering using a synthetic biology approach for producing novel metabolites belonging to carotenoid class in industrially important tobacco plant. The synthetic multigene construct resulted in producing a novel metabolite, keto-lutein with high accumulation of xanthophyll metabolites. This figure was drawn using BioRender (https://www.biorender.com). [ABSTRACT FROM AUTHOR]

Lesion mimic mutants (LMMs) are advantageous materials for studying programmed cell death (PCD). Although some rice LMM genes have been cloned, the diversity of functions of these genes indicates that the mechanism of cell death regulation in LMMs needs further study. In this study, we identified a rice light-dependent leaf lesion mimic mutant 4 (llm4) that showed abnormal chloroplast structure, photoinhibition, reduced photosynthetic protein levels, massive accumulation of reactive oxygen species (ROS), and PCD. Map-based cloning and complementation testing revealed that LLM4 encodes zeaxanthin epoxidase (ZEP), an enzyme involved in the xanthophyll cycle, which functions in plant photoprotection, ROS scavenging, and carotenoid and abscisic acid (ABA) biosynthesis. The ABA content was decreased, and the contents of 24 carotenoids differed between the llm4 mutant and the wild type (WT). The llm4 mutant showed reduced dormancy and greater sensitive to ABA than the WT. We concluded that the mutation of LLM4 resulted in the failure of xanthophyll cycle, in turn causing ROS accumulation. The excessive ROS accumulation damaged chloroplast structure and induced PCD, leading eventually to the formation of lesion mimics.

Abstract Background The marine alga Nannochloropsis oceanica, an emerging model belonging to Heterokont, is considered as a promising light-driven eukaryotic chassis for transforming carbon dioxide to various compounds including carotenoids. Nevertheless, the carotenogenic genes and their roles in the alga remain less understood and to be further explored. Results Here, two phylogenetically distant zeaxanthin epoxidase (ZEP) genes from N. oceanica (NoZEP1 and NoZEP2) were functionally characterized. Subcellular localization experiment demonstrated that both NoZEP1 and NoZEP2 reside in the chloroplast yet with differential distribution patterns. Overexpression of NoZEP1 or NoZEP2 led to increases of violaxanthin and its downstream carotenoids at the expense of zeaxanthin in N. oceanica, with the extent of changes mediated by NoZEP1 overexpression being greater as compared to NoZEP2 overexpression. Suppression of NoZEP1 or NoZEP2, on the other hand, caused decreases of violaxanthin and its downstream carotenoids as well as increases of zeaxanthin; similarly, the extent of changes mediated by NoZEP1 suppression was larger than that by NoZEP2 suppression. Interestingly, chlorophyll a dropped following violaxanthin decrease in a well-correlated manner in response to NoZEP suppression. The thylakoid membrane lipids including monogalactosyldiacylglycerol also correlated with the violaxanthin decreases. Accordingly, NoZEP1 suppression resulted in more attenuated algal growth than NoZEP2 suppression did under either normal light or high light stage. Conclusions The results together support that both NoZEP1 and NoZEP2, localized in the chloroplast, have overlapping roles in epoxidating zeaxanthin to violaxanthin for the light-dependent growth, yet with NoZEP1 being more functional than NoZEP2 in N. oceanica. Our study provides implications into the understanding of carotenoid biosynthesis and future manipulation of N. oceanica for carotenoid production.

The generation of violaxanthin (Vx) de-epoxidase (VDE), photosystem II subunit S (PsbS) and zeaxanthin (Zx) epoxidase (ZEP) (VPZ) lines, which simultaneously overexpress VDE, PsbS and ZEP, has been successfully used to accelerate the kinetics of the induction and relaxation of non-photochemical quenching (NPQ). Here, we studied the impact of the overexpression of VDE and ZEP on the conversion of the xanthophyll cycle pigments in VPZ lines of Arabidopsis thaliana and Nicotiana tabacum. The protein amount of both VDE and ZEP was determined to be increased to about 3- to 5-fold levels of wild-type (WT) plants for both species. Compared to WT plants, the conversion of Vx to Zx, and hence VDE activity, was only marginally accelerated in VPZ lines, whereas the conversion of Zx to Vx, and thus ZEP activity, was strongly increased in VPZ lines. This indicates that the amount of ZEP but not the amount of VDE is a critical determinant of the equilibrium of the de-epoxidation state of xanthophyll cycle pigments under saturating light conditions. Comparing the two steps of epoxidation, particularly the second step (antheraxanthin to Vx) was found to be accelerated in VPZ lines, implying that the intermediate Ax is released into the membrane during epoxidation by ZEP. [ABSTRACT FROM AUTHOR]

The photosynthetic differences between adult and juvenile Mediterranean plants were previously studied under field conditions, yet the corresponding differentiation of their photoprotective efficiency has not been sufficiently investigated. The present study aims to examine possible differences in the photoprotective potential between adults and juveniles of two native Mediterranean plants with distinct growth forms. Thus, the seasonal variations in individual carotenoids, electron transport rate (ETR), and non-photochemical quenching (NPQ) were monitored in fully exposed mature leaves from adults and juveniles of the winter deciduous tree Cercis siliquastrum L. and the evergreen sclerophyllous shrub Nerium oleander L. All plants were grown under apparently similar field conditions. In both species, juveniles displayed substantially lower ETR and increased NPQ values than adults in spring, with the differences intensifying during summer drought and diminishing in autumn. Concomitantly, juveniles showed significantly higher chlorophyll-based total carotenoids in spring and summer mainly due to the higher investment in xanthophyll cycle components (VAZ), in combination with an increased mid-day de-epoxidation state (DEPS) and partial retention of zeaxanthin in the dark. In N. oleander, although ETR was lower in juveniles during winter, NPQ was extremely low in both ages. In conclusion, juveniles exhibit enhanced photoprotection potential, especially in the summer, due to their reduced photochemical capacity. The photosynthetic superiority of adults during the favorable spring period may be attributed to the needs of the co-existing reproductive effort. [ABSTRACT FROM AUTHOR]

The review highlights the relationship between the molecular organization of the light-harvesting complex of photosystem II (LHCII) and sunlight utilization by higher plants. The molecular form of LHCII switches rapidly and reversibly during diurnal changes of light intensity, from low (ca. 10) to high [ca. 1,000 µmol(photon) m-2 s-1], so the sensitivity of LHCII to light may control the balance between light harvesting and photoprotection state. Our understanding and concept of this mechanism are based on the knowledge of the structure and photophysics of different LHCII molecular forms: monomer, dimer, trimer, and aggregate. It is proposed that LHCII monomers, dimers, and lateral aggregates are fundamental blocks of excess light-dissipation machinery. Trimer is exceptionally well suited to play a physiological role of an antenna complex. A correlation between the LHCII molecular form and the presence of xanthophyll cycle pigment violaxanthin and zeaxanthin in the complex structure is also shown. Moreover, the role of LHCII protein phosphorylation in thylakoid membrane architecture is also discussed. The dual function of LHCII has been studied in the natural thylakoid membranes of chloroplasts, in the artificial lipid-LHCII model membranes, and by suspension of LHCII in a detergent solution. [ABSTRACT FROM AUTHOR]

Photosynthetic organisms have evolved to work under low and high lights in photoprotection, acting as a scavenger of reactive oxygen species. The light-dependent xanthophyll cycle involved in this process is performed by a key enzyme (present in the thylakoid lumen), Violaxanthin De-Epoxidase (VDE), in the presence of violaxanthin (Vio) and ascorbic acid substrates. Phylogenetically, VDE is found to be connected with an ancestral enzyme Chlorophycean Violaxanthin De-Epoxidase (CVDE), present in the green algae on the stromal side of the thylakoid membrane. However, the structure and functions of CVDE were not known. In search of functional similarities involving this cycle, the structure, binding conformation, stability, and interaction mechanism of CVDE are explored with the two substrates compared to VDE. The structure of CVDE was determined by homology modeling and validated. In silico docking (of first-principles optimized substrates) revealed it has a larger catalytic domain than VDE. A thorough analysis of the binding affinity and stability of four enzyme–substrate complexes is performed by computing free energies and their decomposition, the root-mean-square deviation (RMSD) and fluctuation (RMSF), the radius of gyration, salt bridge, and hydrogen bonding interactions in molecular dynamics. Based on these, violaxanthin interacts with CVDE to a similar extent as that of VDE. Hence, its role is expected to be the same for both enzymes. On the contrary, ascorbic acid has a weaker interaction with CVDE than VDE. Given these interactions drive epoxidation or de-epoxidation in the xanthophyll cycle, it immediately discerns that either ascorbic acid does not participate in de-epoxidation or a different cofactor is necessary as CVDE has a weaker interaction with ascorbic acid than VDE. [ABSTRACT FROM AUTHOR]

Background: The marine alga Nannochloropsis oceanica, an emerging model belonging to Heterokont, is considered as a promising light-driven eukaryotic chassis for transforming carbon dioxide to various compounds including carotenoids. Nevertheless, the carotenogenic genes and their roles in the alga remain less understood and to be further explored. Results: Here, two phylogenetically distant zeaxanthin epoxidase (ZEP) genes from N. oceanica (NoZEP1 and NoZEP2) were functionally characterized. Subcellular localization experiment demonstrated that both NoZEP1 and NoZEP2 reside in the chloroplast yet with differential distribution patterns. Overexpression of NoZEP1 or NoZEP2 led to increases of violaxanthin and its downstream carotenoids at the expense of zeaxanthin in N. oceanica, with the extent of changes mediated by NoZEP1 overexpression being greater as compared to NoZEP2 overexpression. Suppression of NoZEP1 or NoZEP2, on the other hand, caused decreases of violaxanthin and its downstream carotenoids as well as increases of zeaxanthin; similarly, the extent of changes mediated by NoZEP1 suppression was larger than that by NoZEP2 suppression. Interestingly, chlorophyll a dropped following violaxanthin decrease in a well-correlated manner in response to NoZEP suppression. The thylakoid membrane lipids including monogalactosyldiacylglycerol also correlated with the violaxanthin decreases. Accordingly, NoZEP1 suppression resulted in more attenuated algal growth than NoZEP2 suppression did under either normal light or high light stage. Conclusions: The results together support that both NoZEP1 and NoZEP2, localized in the chloroplast, have overlapping roles in epoxidating zeaxanthin to violaxanthin for the light-dependent growth, yet with NoZEP1 being more functional than NoZEP2 in N. oceanica. Our study provides implications into the understanding of carotenoid biosynthesis and future manipulation of N. oceanica for carotenoid production. [ABSTRACT FROM AUTHOR]

Summary: The Photochemical Reflectance Index (PRI) provides an optical indicator of photosynthetic light‐use efficiency, photoprotection, and stress in plants. Although PRI can be applied in remote sensing, its interpretation depends on irradiance, which is hard to obtain from satellite or airborne imagery.To quantify forest photoprotective responses remotely, we developed a framework for modeling and interpreting PRI‐light responses of individual trees and species using airborne imaging spectrometry coupled with georeferenced forest inventory data from a temperate broad‐leaved forest. We derived an irradiance proxy, used hierarchical modeling to analyze PRI‐light responses, and developed a framework of physiological interpretations of model parameters as facultative and constitutive components of photoprotection.Photochemical Reflectance Index declined with illumination, and PRI‐light relationships varied with landscape position and among tree crowns and species. More sun‐exposed foliage had lower intercepts and slopes of the relationship, indicating greater constitutive, but less facultative, photoprotection.We show that tree photoprotective strategies can be quantified at multiple scales using airborne hyperspectral data in structurally complex forests. Our findings and approach have important implications for the remote sensing of forest stress by offering a new way to assess functional diversity through dynamic differences in photoprotection and photosynthetic downregulation and providing previsual indicators of forest stress. [ABSTRACT FROM AUTHOR]

Ascorbic acid (AsA) plays an important role in scavenging reactive oxygen species (ROS) and reducing photoinhibition in plants, especially under stress. The function of SlGGP which encodes the key enzyme GDP-L-galactose phosphorylase in AsA synthetic pathway is relatively clear. However, there is another gene SlGGP-LIKE that encodes this enzyme in tomato, and there are few studies on it, especially under salt stress. In this study, we explored the function of this gene in tomato salt stress response using transgenic lines overexpressing SlGGP-LIKE (OE). Under normal conditions, overexpressing SlGGP-LIKE can increase the content of reduced AsA and the ratio of AsA/ DHA (dehydroascorbic acid), as well as the level of xanthophyll cycle. Under salt stress, compared with the wild-type plants (WT), the OE lines can maintain higher levels of reduced AsA. In addition, OE lines also have higher levels of reduced GSH (glutathione) and total GSH, higher ratios of AsA/DHA and GSH/oxidative GSH (GSSR), and higher level of xanthophyll cycle. Therefore, the OE lines are more tolerant to salt stress, with higher photosynthetic activity, higher antioxidative enzyme activities, higher content of D1 protein, lower production rate of ROS, and lighter membrane damage. These results indicate that overexpressing SlGGP-LIKE can enhance tomato resistance to salt stress through promoting the synthesis of AsA. [ABSTRACT FROM AUTHOR]

Background and Aims Polyploidization can improve plant mass yield for bioenergy support, yet few studies have investigated ozone (O3) sensitivity linked to internal regulatory mechanisms at different ploidy levels. Methods Diploid and triploid Populus tomentosa plants were exposed to ambient and ambient plus 60 ppb [O3]. We explored their differences in sensitivity (leaf morphological, physiological and biochemical traits, and plant mass) as well as mechanisms of avoidance (stomatal conductance, xanthophyll cycle, thermal dissipation) and tolerance (ROS scavenging system) in response to O3 at two developmental phases. Key Results Triploid plants had the highest plant growth under ambient O3, even under O3 fumigation. However, triploid plants were the most sensitive to O3 and under elevated O3 showed the largest decreases in photosynthetic capacity and performance, as well as increased shoot:root ratio, and the highest lipid peroxidation. Thus, plant mass production could be impacted in triploid plants under long-term O3 contamination. Both diploid and triploid plants reduced stomatal aperture in response to O3, thereby reducing O3 entrance, yet only in diploid plants was reduced stomatal aperture associated with minimal (non-significant) damage to photosynthetic pigments and lower lipid peroxidation. Conclusions Tolerance mechanisms of plants of both ploidy levels mainly focused on the enzymatic reduction of hydrogen peroxide through catalase and peroxidase, yet these homeostatic regulatory mechanisms were higher in diploid plants. Our study recommends triploid white poplar as a bioenergy species only under short-term O3 contamination. Under continuously elevated O3 over the long term, diploid white poplar may perform better. [ABSTRACT FROM AUTHOR]

In the present study, low concentrations of the very mild detergent n‐dodecyl‐α‐d‐maltoside in conjunction with sucrose gradient ultracentrifugation were used to prepare fucoxanthin chlorophyll protein (FCP) complexes of the centric diatom Thalassiosira pseudonana. Two main FCP fractions were observed in the sucrose gradients, one in the upper part and one at high sucrose concentrations in the lower part of the gradient. The first fraction was dominated by the 18 kDa FCP protein band in SDS‐gels. Since this fraction also contained other protein bands, it was designated as fraction enriched in FCP‐A complex. The second fraction contained mainly the 21 kDa FCP band, which is typical for the FCP‐B complex. Determination of the lipid composition showed that both FCP fractions contained monogalactosyl diacylglycerol as the main lipid followed by the second galactolipid of the thylakoid membrane, namely digalactosyl diacylglycerol. The negatively charged lipids sulfoquinovosyl diacylglycerol and phosphatidyl glycerol were also present in both fractions in pronounced concentrations. With respect to the pigment composition, the fraction enriched in FCP‐A contained a higher amount of the xanthophyll cycle pigments diadinoxanthin (DD) and diatoxanthin (Dt), whereas the FCP‐B fraction was characterized by a lower ratio of xanthophyll cycle pigments to the light‐harvesting pigment fucoxanthin. Protein analysis by mass spectrometry revealed that in both FCP fractions the xanthophyll cycle enzyme diadinoxanthin de‐epoxidase (DDE) was present. In addition, the analysis showed an enrichment of DDE in the fraction enriched in FCP‐A but only a very low amount of DDE in the FCP‐B fraction. In‐vitro de‐epoxidation assays, employing the isolated FCP complexes, were characterized by an inefficient conversion of DD to Dt. However, in line with the heterogeneous DDE distribution, the fraction enriched in FCP‐A showed a more pronounced DD de‐epoxidation compared with the FCP‐B. [ABSTRACT FROM AUTHOR]

The Photochemical Reflectance Index (PRI) and solar induced fluorescence (SIF) provide information on plant photosynthetic activity. PRI and SIF are both strongly influenced by irradiance, but uncertainties related to the interpretation of these light responses at large spatial scales remain, partly due to a shortage of suitable data from aircraft or satellite platforms. The goal of this study was to explore interpretations of the PRI- and SIF-light responses of trees owing to species, functional types (evergreen and deciduous) and season. Using airborne hyperspectral and ultraspectral imagery in a North American urban forest, we derived PRI, SIF, and albedo (an indicator of illumination) at the 1-m pixel level. We then quantified crown-level PRI and SIF light responses of ten different tree species at three time points from late-summer to autumnal senescence using hierarchical models. Our results confirmed that both PRI and SIF were strongly influenced by illumination with PRI decreasing and SIF increasing with illumination. Both slope and intercept of the PRI-albedo relationship changed with season, but the pattern varied among species and functional types. SIF values decreased during autumnal senescence for all species, but evergreen species exhibited less seasonal decline in the slope of SIF-albedo relationship compared to deciduous species. The PRI and SIF light responses derived from the airborne imagery offer complementary information on dynamic photosynthesis responses presumably due to varying canopy structure, pigmentation and photoprotection among species and functional types. From airborne platforms, PRI- and SIF-light responses can be used to explore the contrasting physiological responses of individual tree crowns, providing a spatially and temporally explicit view of dynamic plant traits related to photoregulation and a novel view of functional diversity for entire landscapes. • PRI and SIF light responses of trees were fitted using airborne data. • Both PRI and SIF were strongly influenced by illumination. • PRI and SIF light responses varied among species, functional types and season. • PRI and SIF light responses revealed dynamic photosynthetic characteristics. [ABSTRACT FROM AUTHOR]

[Display omitted] • K. cf. nitens grows consistently at high shear stress (6600 mPa) • Fragmentation of large filaments favours vegetative multiplication of K. cf nitens. • K. cf. nitens grows in a tubular photobioreactor at high fluid velocity (70 cm s−1) • Xanthophyll cycle is activated to protect cell photosynthetic system. • Linoleic acid makes over 50 % of total fatty acids. In this study, the effects of shear stress and different culture media on the growth of the filamentous microalga Klebsormidium cf. nitens were studied. The microalga's growth, carotenoids and fatty acids were further evaluated in a pump-driven tubular photobioreactor. The results show that this microalga had the ability to withstand high shear stress and the adaptability to grow in a culture medium that lacks certain trace elements. K. cf. nitens grew consistently in the tubular photobioreactor at different average light intensities although it did not grow well in a tall bubble column. The carotenoid analysis revealed that the xanthophyll cycle was activated to protect the cell photosynthetic system. The fatty acids increased with irradiance, with linoleic acid (C18:2n6) making up over 50 % of the total fatty acids. This study supports the potential of employing pump-driven tubular photobioreactors to produce the filamentous microalga K. cf nitens at the large scale. [ABSTRACT FROM AUTHOR]

Some natural carotenoids remain unexplored in terms of their function because they occur only in trace amounts, or their stereochemistry is unclear. It is expected that scientists will discover new physiological activities that have a positive impact on human heath by analysis of such carotenoids. The purpose of this study was to analyze the carotenoids synthesized by the novel aerial microalga Coelastrella rubescens KGU–H009, and the dynamics of their accumulation. Cultivation experiments were conducted for 7 days at 25 °C, combining different light conditions with the presence or absence of nitrate ions. Short-term (2-h) cultivation was also performed, and tests were conducted with the addition of the inhibitor of photosynthesis 3-(3,4-dichlorophenyl)-1, 1-dimethylurea (DCMU). Strain KGU-H009 exhibited its highest carotenoid content (34.5 mg g−1) in the presence of nitrogen sources and strong light. Upon nitrogen source depletion, the content of zeaxanthin and β-carotene decreased, while the content of astaxanthin and adonixanthin increased. Strain KGU-H009 exhibited reversible control of the xanthophyll cycle to adapt to changes in light intensity, but the xanthophyll cycle was inhibited by the addition of DCMU. Strain KGU-H009 demonstrated the ability to synthesize carotenoids involved in both light harvesting and photoprotection, suggesting its adaptability to environmental change. [Display omitted] • The aerial microalga Coelastrella rubescens KGU-H009 was isolated. • β-Carotene, zeaxanthin, and lutein were mainly accumulated in the strain. • The carotenoid content was regulated in various light- and nitrogen-stress conditions. • The xanthophyll cycle was inhibited by photosynthetic inhibitor. [ABSTRACT FROM AUTHOR]

The photosynthetic differences between adult and juvenile Mediterranean plants were previously studied under field conditions, yet the corresponding differentiation of their photoprotective efficiency has not been sufficiently investigated. The present study aims to examine possible differences in the photoprotective potential between adults and juveniles of two native Mediterranean plants with distinct growth forms. Thus, the seasonal variations in individual carotenoids, electron transport rate (ETR), and non-photochemical quenching (NPQ) were monitored in fully exposed mature leaves from adults and juveniles of the winter deciduous tree Cercis siliquastrum L. and the evergreen sclerophyllous shrub Nerium oleander L. All plants were grown under apparently similar field conditions. In both species, juveniles displayed substantially lower ETR and increased NPQ values than adults in spring, with the differences intensifying during summer drought and diminishing in autumn. Concomitantly, juveniles showed significantly higher chlorophyll-based total carotenoids in spring and summer mainly due to the higher investment in xanthophyll cycle components (VAZ), in combination with an increased mid-day de-epoxidation state (DEPS) and partial retention of zeaxanthin in the dark. In N. oleander, although ETR was lower in juveniles during winter, NPQ was extremely low in both ages. In conclusion, juveniles exhibit enhanced photoprotection potential, especially in the summer, due to their reduced photochemical capacity. The photosynthetic superiority of adults during the favorable spring period may be attributed to the needs of the co-existing reproductive effort.

The aim of this study was to evaluate the effects of melatonin on photoinhibition under chilling stress in cucumber seedlings and to inquire into any mechanisms of mitigation. Under chilling stress, the net photosynthetic rate declined dramatically but the decline was significantly mitigated by irrigation with a melatonin solution. Possible mechanisms for this mitigation are that melatonin accelerates xanthophyll de-epoxidation by upregulating the transcription of the violaxanthin de-epoxidase gene (CsVDE) and down-regulating that of the zeaxanthin cyclase gene (CsZE) during chilling. There was also a rise in non-photochemical quenching (NPQ) if seedlings were pretreated with melatonin before chilling. The efficient operation of the xanthophyll cycle helped consume excessive excitation energy in photosystem (PS) Ⅱ and so protected the photosynthetic system. Melatonin also modulated the partitioning of absorbed excitation energy in PS Ⅱ as evidenced by alleviation of the decrease in quantum yield of photochemical energy conversion in PS Ⅱ under chilling stress, by alleviation of the rise in quantum yield of non-regulated, non-photochemical energy loss in PS II and by increasing the regulated non-photochemical energy loss in PS II. This study presents a new understanding of the mechanisms through which melatonin mitigates photoinhibition by modulating the partitioning of absorption energy in PS Ⅱ based on the xanthophyll cycle.

As a common macroalga living in the intertidal zone, Sargassum thunbergii (Sargassaceae, Phaeophyta) is often exposed to drastic changes in solar photosynthetically active radiation during a diurnal cycle; thus, the potential photosynthetic adaptation processes deserve attention. In this work, we examined the photosynthetic performance and xanthophyll cycle activity of this alga in response to high light (1,200 μmol photons m–2 s–1, the average in-situ light intensity at noon) by using chlorophyll fluorescence and high-performance liquid chromatography (HPLC). On exposure to high light, a rapid decrease in the effective quantum yield of photosystem II (PSII) (Y(II)) occurred, indicating down-regulation of PSII activity; a corresponding increase in non-photochemical quenching (NPQ) indicated the existence of energy-dissipating cycles. After turning off the light, Y(II) gradually increased to 0.7, accompanied by a decrease in NPQ. However, no complete recovery of NPQ was observed, and its value remained at ≈4, even after an 80-min dark treatment. The size of the xanthophyll cycle pigments pool was quantified using HPLC and was found to be ≈16 mol mol−1 Chl a × 100. The activity of the xanthophyll cycle, characterized by a de-epoxidation state (DPS), could reach up to ≈0.5. Such a large pigments pool and rapid accumulation of zeaxanthin may allow S. thunbergii to induce high values of NPQ (≈10). These results were further complemented by inhibitor (dithiothreitol, DTT) and pre-illumination experiments showing that (1) both NPQ and the xanthophyll cycle could be inhibited by DTT, and there was always a strong positive correlation between NPQ and DPS; (2) the previously formed antheraxanthin exhibited a long retention time and a slow epoxidation; and (3) the long retention of antheraxanthin contributed to a rapid accumulation of zeaxanthin. In conclusion, our present study demonstrated that xanthophyll cycle-induced NPQ can significantly protect S. thunbergii from light fluctuations in the intertidal zone.

5-Aminolevulinic acid (ALA), an antioxidant existing in plants, has been widely reported to participate in the process of coping with cold stress of plants. In this study, exogenous ALA promoted the growth of tomato plants and alleviated the appearance of purple tomato leaves under low-temperature stress. At the same time, exogenous ALA improved antioxidant enzyme activities, SlSOD gene expression, Fv/Fm, and proline contents and reduced H 2 O 2 contents, SlRBOH gene expression, relative electrical conductivity, and malondialdehyde contents to alleviate the damage caused by low temperature to tomato seedlings. Compared with low-temperature stress, spraying exogenous ALA before low-temperature stress could restore the indicators of photochemical quenching, actual photochemical efficiency, electron transport rate, and nonphotochemical quenching to normal. Exogenous ALA could increase the total contents of the xanthophyll cycle pool, the positive de-epoxidation rate of the xanthophyll cycle and improved the expression levels of key genes in the xanthophyll cycle under low-temperature stress. In addition, we found that exogenous ALA significantly enhanced the absorption of mineral nutrients, promoted the transfer and distribution of mineral nutrients to the leaves, and improved the expression levels of mineral nutrient absorption-related genes, which were all conducive to the improved adaptation of tomato seedlings under low-temperature stress. In summary, the application of exogenous ALA can increase tomato seedlings' tolerance to low-temperature stress by improving the xanthophyll cycle and the ability of the absorption of mineral nutrients in tomato seedlings. • Exogenous 5-aminolevulinic acid (ALA) significantly improved low temperature tolerance in tomatoes. • ALA could improve the xanthophyll cycle ability of tomato seedlings under low-temperature stress. • Exogenous ALA could improve nutrient uptake by tomato seedlings under low-temperature stress. • Exogenous ALA could promote the transformation of mineral nutrition to leaves under low-temperature stress. [ABSTRACT FROM AUTHOR]

Carotenoids (Cars) exhibit two functions in photosynthesis, light-harvesting and photoprotective functions, which are performed through the excited states of Cars. Therefore, increasing our knowledge on excitation relaxation dynamics of Cars is important for understanding of the functions of Cars. In light-harvesting complexes, there exist Cars functioning by converting the π-conjugation number in response to light conditions. It is well known that some microalgae have a mechanism controlling the conjugation number of Cars, called as the diadinoxanthin cycle; diadinoxanthin (10 conjugations) is accumulated under low light, whereas diatoxanthin (11 conjugations) appears under high light. However, the excitation relaxation dynamics of these two Cars have not been clarified. In the present study, we investigated excitation relaxation dynamics of diadinoxanthin and diatoxanthin in relation to their functions, by the ultrafast fluorescence spectroscopy. After an excitation to the S2 state, the intramolecular vibrational redistribution occurs, followed by the internal conversion to the S1 state. The S2 lifetimes were analyzed to be 175 fs, 155 fs, and 140 fs in diethyl ether, ethanol, and acetone, respectively, for diadinoxanthin, and 155 fs, 135 fs, and 125 fs in diethyl ether, ethanol, and acetone, respectively for diatoxanthin. By converting diadinoxanthin to diatoxanthin, the absorption spectra shift to longer wavelengths by 5–7 nm, and lifetimes of S2 and S1 states decrease by 11–13% and 52%, respectively. Differences in levels and lifetimes of excited states between diadinoxanthin and diatoxanthin are small; therefore, it is suggested that changes in the energy level of chlorophyll a are necessary to efficiently control the functions of the diadinoxanthin cycle. [ABSTRACT FROM AUTHOR]

Fucoxanthin is a major light-harvesting pigment in ecologically important algae such as diatoms, haptophytes, and brown algae (Phaeophyceae). Therefore, it is a major driver of global primary productivity. Species of these algal groups are brown colored because the high amounts of fucoxanthin bound to the proteins of their photosynthetic machineries enable efficient absorption of green light. While the structure of these fucoxanthin-chlorophyll proteins has recently been resolved, the biosynthetic pathway of fucoxanthin is still unknown. Here, we identified two enzymes central to this pathway by generating corresponding knockout mutants of the diatom Phaeodactylum tricornutum that are green due to the lack of fucoxanthin. Complementation of the mutants with the native genes or orthologs from haptophytes restored fucoxanthin biosynthesis. We propose a complete biosynthetic path to fucoxanthin in diatoms and haptophytes based on the carotenoid intermediates identified in the mutants and in vitro biochemical assays. It is substantially more complex than anticipated and reveals diadinoxanthin metabolism as the central regulatory hub connecting the photoprotective xanthophyll cycle and the formation of fucoxanthin. Moreover, our data show that the pathway evolved by repeated duplication and neofunctionalization of genes for the xanthophyll cycle enzymes violaxanthin de-epoxidase and zeaxanthin epoxidase. Brown algae lack diadinoxanthin and the genes described here and instead use an alternative pathway predicted to involve fewer enzymes. Our work represents a major step forward in elucidating the biosynthesis of fucoxanthin and understanding the evolution, biogenesis, and regulation of the photosynthetic machinery in algae. [ABSTRACT FROM AUTHOR]

Photosynthetic organisms have evolved a great variety of mechanisms to optimize their use of sunlight. Some of the clearest examples of adaptations can be seen by comparing photosynthesis in different species and in different individuals of the same species that grow under high and low light levels. While the adaptations of sun and shade higher plants have been relatively well studied, much less information is available on the photobionts of lichenized Ascomycetes. An important adaptation that can protect photosynthetic organisms from the potentially harmful effects of excess light is non-photochemical quenching (NPQ); NPQ can dissipate unused light energy as heat. Here we used chlorophyll fluorescence to compare the induction and relaxation of NPQ and the induction of electron transport (rETR) in collections of the same lichen species from exposed and from more shaded locations. All species have trebouxioid photobionts and normally grow in more exposed microhabitats but can also be readily collected from more shaded locations. Shade forms display generally higher NPQ, presumably to protect lichens from occasional rapid increases in light that occur during sunflecks. Furthermore, the NPQ of shade forms relaxes quickly when light levels are reduced, presumably to ensure efficient photosynthesis after a sunfleck has passed. The maximal relative electron transport rate is lower in shade than sun collections, probably reflecting a downregulation of photosynthetic capacity to reduce energy costs. We also compared collections of pale and melanized thalli from three species of shade lichens with Symbiochloris as their photobiont. Interestingly, NPQ in melanized thalli from slightly more exposed microhabitats induced and relaxed in a way that resembled shade rather than sun forms of the trebouxioid lichens. This might suggest that in some locations melanization induced during a temporary period of high light may be excessive and could potentially reduce photosynthesis later in the growing season. Taken together, the results suggest that lichen photobionts can flexibly adjust the amount and type of NPQ, and their levels of rETR in response to light availability. [ABSTRACT FROM AUTHOR]

Under natural conditions, photosynthesis has to be adjusted to fluctuating light intensities. Leaves exposed to high light dissipate excess light energy in form of heat at photosystem II (PSII) by a process called non‐photochemical quenching (NPQ). Upon fast transition from light to shade, plants lose light energy by a relatively slow relaxation from photoprotection. Combined overexpression of violaxanthin de‐epoxidase (VDE), PSII subunit S (PsbS) and zeaxanthin epoxidase (ZEP) in tobacco accelerates relaxation from photoprotection, and increases photosynthetic productivity. In Arabidopsis, expression of the same three genes (VPZ) resulted in a more rapid photoprotection but growth of the transgenic plants was impaired. Here we report on VPZ expressing potato plants grown under various light regimes. Similar to tobacco and Arabidopsis, induction and relaxation of NPQ was accelerated under all growth conditions tested, but did not cause an overall increased photosynthetic rate or growth of transgenic plants. Tuber yield of VPZ expressing plants was unaltered as compared to control plants under constant light conditions and even decreased under fluctuating light conditions. Under control conditions, levels of the phytohormone abscisic acid (ABA) were found to be elevated, indicating an increased violaxanthin availability in VPZ plants. However, the increased basal ABA levels did not improve drought tolerance of VPZ transgenic potato plants under greenhouse conditions. The failure to benefit from improved photoprotection is most likely caused by a reduced radiation use efficiency under high light conditions resulting from a too strong NPQ induction. Mitigating this negative effect in the future might help to improve photosynthetic performance in VPZ expressing potato plants. [ABSTRACT FROM AUTHOR]

Induction of metabolite biosynthesis and accumulation is one of the most prominent UV-mediated changes in plants, whether during eustress (positive response) or distress (negative response). However, despite evidence suggesting multiple linkages between UV exposure and carotenoid induction in plants, there is no consensus in the literature concerning the direction and/or amplitude of these effects. Here, we compiled publications that characterised the relative impact of UV on the content of individual carotenoids and subjected the created database to a meta-analysis in order to acquire new, fundamental insights in responses of the carotenoid pool to UV exposure. Overall, it was found that violaxanthin was the only carotenoid compound that was significantly and consistently induced as a result of UV exposure. Violaxanthin accumulation was accompanied by a UV dose dependent decrease in antheraxanthin and zeaxanthin. The resulting shift in the state of the xanthophyll cycle would normally occur when plants are exposed to low light and this is associated with increased susceptibility to photoinhibition. Although UV induced violaxanthin accumulation is positively linked to the daily UV dose, the current dataset is too small to establish a link with plant stress, or even experimental growth conditions. In summary, the effects of UV radiation on carotenoids are multifaceted and compound-specific, and there is a need for a systematic analysis of dose-response and wavelength dependencies, as well as of interactive effects with further environmental parameters. • No consensus exists in the literature concerning effects of UV exposure on carotenoids in plants. • Studies that characterised the relative impact of UV on the content of individual carotenoids were compiled. • UV induces violaxanthin accumulation in plants. • A degree of interplay exists between UV effects on individual carotenoids. [ABSTRACT FROM AUTHOR]

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

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

• D1 turnover plays a more important role than xanthophyll cycle in photoprotection under sub-high temperature and high light (HH) conditions in tomato. • D1 protein turnover and xanthophyll cycle avoided the accumulation of excess energy and photooxidative damage of photosystem through maintaining photosynthetic pigments content and inducing changes in NPQ core components. • D1 protein turnover and xanthophyll cycle can also avoided accumulation of ROS and membrane lipid peroxidation by maintaining the activity of the ROS scavenging system. D1 protein turnover and the xanthophyll cycle (XC) are important photoprotective mechanisms in plants that operate under adverse conditions. Here, streptomycin sulfate (SM) and dithiothreitol (DTT) were used in tomato plants as inhibitors of D1 protein turnover and XC to elucidate their photoprotective impacts under sub-high temperature and high light conditions (HH, 35°C, 1000 µmol·m-2·s-1). SM and DTT treatments significantly reduced the net photosynthetic rate, apparent quantum efficiency, maximum photochemical efficiency, and potential activity of photosystem II, leading to photoinhibition and a decline in plant biomass under HH. The increase in reactive oxygen species levels resulted in thylakoid membrane lipid peroxidation. In addition, there were increased non-photochemical quenching and decreased chlorophyll pigments in SM and DTT application, causing an inhibition of D1 protein production at both transcriptional and translational levels. Overall, inhibition of D1 turnover caused greater photoinhibition than XC inhibition. Additionally, the recovery levels of most photosynthesis indicators in DTT-treated plants were higher than in SM-treated plants. These findings support the view that D1 turnover has a more important role than XC in photoprotection in tomato under HH conditions.

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

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

High light exposure rapidly activates non-photochemical quenching (NPQ), protecting plants from photooxidative damage. Contrarily, its relaxation upon transition to normal light occurs quite slowly, limiting photosynthetic efficiency. De Souza et al. demonstrated that by overexpressing NPQ-related genes, faster NPQ relaxation and enhanced photosynthesis can be achieved under fluctuating light conditions. [ABSTRACT FROM AUTHOR]

Plants must switch rapidly between light harvesting and photoprotection in response to environmental fluctuations in light intensity. This switch can lead to losses in absorbed energy usage, as photoprotective energy dissipation mechanisms can take minutes to hours to fully relax. One possible way to improve photosynthesis is to engineer these energy dissipation mechanisms (measured as non-photochemical quenching of chlorophyll a fluorescence, NPQ) to induce and relax more quickly, resulting in smaller losses under dynamic light conditions. Previous studies aimed at understanding the enzymes involved in the regulation of NPQ have relied primarily on labor-intensive and time-consuming generation of stable transgenic lines and mutant populations - approaches limited to organisms amenable to genetic manipulation and mapping. To enable rapid functional testing of NPQ-related genes from diverse organisms, we performed Agrobacterium tumefaciens-mediated transient expression assays in Nicotiana benthamiana to test if NPQ kinetics could be modified in fully expanded leaves. By expressing Arabidopsis thaliana genes known to be involved in NPQ, we confirmed the viability of this method for studying dynamic photosynthetic processes. Subsequently, we used naturally occurring variation in photosystem II subunit S, a modulator of NPQ in plants, to explore how differences in amino acid sequence affect NPQ capacity and kinetics. Finally, we functionally characterized four predicted carotenoid biosynthesis genes from the marine algae Nannochloropsis oceanica and Thalassiosira pseudonana and examined the effect of their expression on NPQ in N. benthamiana. This method offers a powerful alternative to traditional gene characterization methods by providing a fast and easy platform for assessing gene function in planta.

Evergreen conifers growing in high-latitude regions must endure prolonged winters that are characterized by sub-zero temperatures combined with light, conditions that can cause significant photooxidative stress. Understanding overwintering mechanisms is crucial for addressing winter adversity in temperate forest ecosystems and enhancing the ability of conifers to adapt to climate change. This review synthesizes the current understanding of the photoprotective mechanisms that conifers employ to mitigate photooxidative stress, particularly non-photochemical "sustained quenching", the mechanism of which is hypothesized to be a recombination or deformation of the original mechanism employed by conifers in response to short-term low temperature and intense light stress in the past. Based on this hypothesis, scattered studies in this field are assembled and integrated into a complete mechanism of sustained quenching embedded in the adaptation process of plant physiology. It also reveals which parts of the whole system have been verified in conifers and which have only been verified in non-conifers, and proposes specific directions for future research. The functional implications of studies of non-coniferous plant species for the study of coniferous trees are also considered, as a wide range of plant responses lead to sustained quenching, even among different conifer species. In addition, the review highlights the challenges of measuring sustained quenching and discusses the application of ultrafast-time-resolved fluorescence and decay-associated spectra for the elucidation of photosynthetic principles. • The sustained quenching mechanism involves recombination or deformation of the original mechanism. • Existing scattered studies are consolidated into a complete sustained quenching system. • It clearly identifies verified parts of the system in conifers and those only verified in non-conifers. • Discussing the application of ultrafast time-resolved fluorescence and DAS to elucidate photosynthetic principles. [ABSTRACT FROM AUTHOR]

Most C4 plants that naturally occur in tropical or subtropical climates, in high light environments, had to evolve a series of adaptations of photosynthesis that allowed them to grow under these conditions. In this review, we summarize mechanisms that ensure the balancing of energy distribution, counteract photoinhibition, and allow the dissipation of excess light energy. They secure effective electron transport in light reactions of photosynthesis, which will lead to the production of NADPH and ATP. Furthermore, a higher content of the cyclic electron transport components and an increase in ATP production are observed, which is necessary for the metabolism of C4 for effective assimilation of CO2. Most of the data are provided by studies of the genus Flaveria, where species belonging to different metabolic subtypes and intermediate forms between C3 and C4 are present. All described mechanisms that function in mesophyll and bundle sheath chloroplasts, into which photosynthetic reactions are divided, may differ in metabolic subtypes as a result of the different organization of thylakoid membranes, as well as the different demand for ATP and NADPH. This indicates that C4 plants have plasticity in the utilization of pathways in which efficient use and dissipation of excitation energy are realized. [ABSTRACT FROM AUTHOR]

In recent years, with the global climate change, the intensity, frequency and duration of drought have increased significantly, which has become the main limiting factor for agricultural development in many areas. Glycine betaine (GB) is an effective stress-resistant substance. In this experiment, the effects of the genetic engineering of GB synthesis on photosynthetic apparatus of tobacco under drought stress were studied using transgenic tobacco (T) accumulating GB and wild-type tobacco (K326, WT). Potted tobaccos were subjected to drought stress (controlled irrigation, 25 °C ± 1 °C, a relative humidity: 75–80%) for 9 days, photosynthetic gas exchange parameters, chlorophyll a fluorescence, structure of chloroplast and thylakoid membrane, and protein function of thylakoid membrane were examined under different drought stress time (days). The results showed that T tobacco could accumulate GB and the accumulated GB improved the resistance of the photosynthetic apparatus to drought stress. Under drought stress, the damage of chloroplast and thylakoid lamellae in T tobacco was less than that in WT tobacco, the accumulation of GB in T tobacco could maintain the stability of thylakoid membrane, improved the unsaturated fatty acid index (IUFA) of thylakoid membrane lipid, increased the contents of digalactosyl diacylglycerol (DGDG) and phosphatidylglycerol (PG), and decreased the ratio of monogalactosyl diaylglycerol (MGDG) to DGDG. In addition, under drought stress, the accumulation of GB in T tobacco alleviated the photo-inhibition of PSII, and the increase of xanthophyll cycle de-epoxidation may be one of the reasons for the enhancement of PSII function. [ABSTRACT FROM AUTHOR]

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

Xanthophyll cycles (XC) have proven to be major contributors to photoacclimation for many organisms. This work describes a light-driven XC operating in the chlorophyte Chlamydomonas reinhardtii and involving the xanthophylls Lutein (L) and Loroxanthin (Lo). Pigments were quantified during a switch from high to low light (LL) and at different time points from cells grown in Day/Night cycle. Trimeric LHCII was purified from cells acclimated to high or LL and their pigment content and spectroscopic properties were characterized. The Lo/(L + Lo) ratio in the cells varies by a factor of 10 between cells grown in low or high light (HL) leading to a change in the Lo/(L + Lo) ratio in trimeric LHCII from.5 in low light to.07 in HL. Trimeric LhcbMs binding Loroxanthin have 5 ± 1% higher excitation energy (EE) transfer (EET) from carotenoid to Chlorophyll as well as higher thermo- and photostability than trimeric LhcbMs that only bind Lutein. The Loroxanthin cycle operates on long time scales (hours to days) and likely evolved as a shade adaptation. It has many similarities with the Lutein-epoxide – Lutein cycle (LLx) of plants. [ABSTRACT FROM AUTHOR]

Xanthophyll cycles (XC) have proven to be major contributors to photoacclimation for many organisms. This work describes a light-driven XC operating in the chlorophyte Chlamydomonas reinhardtii and involving the xanthophylls Lutein (L) and Loroxanthin (Lo). Pigments were quantified during a switch from high to low light (LL) and at different time points from cells grown in Day/Night cycle. Trimeric LHCII was purified from cells acclimated to high or LL and their pigment content and spectroscopic properties were characterized. The Lo/(L + Lo) ratio in the cells varies by a factor of 10 between cells grown in low or high light (HL) leading to a change in the Lo/(L + Lo) ratio in trimeric LHCII from .5 in low light to .07 in HL. Trimeric LhcbMs binding Loroxanthin have 5 ± 1% higher excitation energy (EE) transfer (EET) from carotenoid to Chlorophyll as well as higher thermo- and photostability than trimeric LhcbMs that only bind Lutein. The Loroxanthin cycle operates on long time scales (hours to days) and likely evolved as a shade adaptation. It has many similarities with the Lutein-epoxide – Lutein cycle (LLx) of plants.

Summary: Photosynthetic organisms in nature often experience light fluctuations. While low light conditions limit the energy uptake by algae, light absorption exceeding the maximal rate of photosynthesis may go along with enhanced formation of potentially toxic reactive oxygen species. To preempt high light‐induced photodamage, photosynthetic organisms evolved numerous photoprotective mechanisms. Among these, energy‐dependent fluorescence quenching (qE) provides a rapid mechanism to dissipate thermally the excessively absorbed energy. Diatoms thrive in all aquatic environments and thus belong to the most important primary producers on earth. qE in diatoms is provided by a concerted action of Lhcx proteins and the xanthophyll cycle pigment diatoxanthin. While the exact Lhcx activation mechanism of diatom qE is unknown, two lumen‐exposed acidic amino acids within Lhcx proteins were proposed to function as regulatory switches upon light‐induced lumenal acidification. By introducing a modified Lhcx1 lacking these amino acids into a Phaeodactylum tricornutum Lhcx1‐null qE knockout line, we demonstrate that qE is unaffected by these two amino acids. Based on sequence comparisons with Lhcx4, being incapable of providing qE, we perform domain swap experiments of Lhcx4 with Lhcx1 and identify two peptide motifs involved in conferring qE. Within one of these motifs, we identify a tryptophan residue with a major influence on qE establishment. This tryptophan residue is located in close proximity to the diadinoxanthin/diatoxanthin‐binding site based on the recently revealed diatom Lhc crystal structure. Our findings provide a structural explanation for the intimate link of Lhcx and diatoxanthin in providing qE in diatoms. Significance Statement: Together with a functional xanthophyll cycle, Lhcx proteins confer rapid photoprotection (energy‐dependent fluorescence quenching) in diatoms in response to excess light. By domain swap experiments, we identify essential parts of the Lhcx proteins that are mandatory to fulfill this role and identify a putative link between Lhcx proteins and the xanthophyll cycle in triggering energy‐dependent fluorescence quenching. [ABSTRACT FROM AUTHOR]

Low temperature in winter depresses rates of photosynthesis, which, in evergreen plants, can exacerbate imbalances between light absorption and photochemical light use. Damage that could result from increased excess light absorption is minimized by the conversion of excitation energy to heat in a process known as energy dissipation, which involves the de-epoxidized carotenoids of the xanthophyll cycle. Overwintering evergreens employ sustained forms of energy dissipation observable even after lengthy periods of dark acclimation. Whereas most studies of photoprotective energy dissipation examine one or a small number of species; here, we measured the levels of sustained thermal energy dissipation of seventy conifer taxa growing outdoors under common-garden conditions at the Red Butte Garden in Salt Lake City, Utah, U.S.A. (forty nine taxa were also sampled for needle pigment content). We observed an extremely wide range of wintertime engagement of sustained energy dissipation; the percentage decrease in dark-acclimated photosystem II quantum efficiency from summer to winter ranged from 6 to 95%. Of the many pigment-based parameters measured, the magnitude of the seasonal decrease in quantum efficiency was most closely associated with the seasonal increase in zeaxanthin content expressed on a total chlorophyll basis, which explained only slightly more than one-third of the variation. We did not find evidence for a consistent wintertime decrease in needle chlorophyll content. Thus, the prevailing mechanism for winter decreases in solar-induced fluorescence emitted by evergreen forests may be decreases in fluorescence quantum yield, and wintertime deployment of sustained energy dissipation likely underlies this effect. [ABSTRACT FROM AUTHOR]