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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.

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This chapter provides a methodology for evaluating plant health and leaf characteristics using spectral reflectance. It provides a step-by-step guide to using spectrometers for high-resolution point measurements of leaf spectral reflectance and multispectral imaging for capturing spatial data, emphasizing the importance of consistent measurement conditions. The chapter further explores the intricacies of multispectral imaging, including calibration, data collection, and image processing. Finally, this chapter delves into the application of various spectral indices for the quantification of key traits such as pigment content, the status of the xanthophyll cycle, water content, and how to identify spectral regions of interest for further research and development. Serving as a guide for researchers and practitioners in plant science, this chapter provides a straightforward framework for plant health assessment using spectral reflectance.

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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.

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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 (Fv/Fm) 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.

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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.

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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.

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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.

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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.

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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.

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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 ).

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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.

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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.

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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.

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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.

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The localization of carotenoids and macromolecular organization of thylakoid supercomplexes have not been reported yet in Chlamydomonas reinhardtii WT and cyclic electron transport mutants (pgrl1 and pgr5) under high light. Here, the various pigments, protein composition, and pigment-protein interactions were analyzed from the cells, thylakoids, and sucrose density gradient (SDG) fractions. Also, the supercomplexes of thylakoids were separated from BN-PAGE and SDG. The abundance of light-harvesting complex (LHC) II trimer complexes and pigment-pigment interaction were changed slightly under high light, shown by circular dichroism. However, a drastic change was seen in photosystem (PS)I-LHCI complexes than PSII complexes, especially in pgrl1 and pgr5. The lutein and ß-carotene increased under high light in LHCII trimers compared to other supercomplexes, indicating that these pigments protected the LHCII trimers against high light. However, the presence of xanthophylls, lutein, and ß-carotene was less in PSI-LHCI, indicating that pigment-protein complexes altered in high light. Even the real-time PCR data shows that the pgr5 mutant does not accumulate zeaxanthin dependent genes under high light, which shows that violaxanthin is not converting into zeaxanthin under high light. Also, the protein data confirms that the LHCSR3 expression is absent in pgr5, however it is presented in LHCII trimer in WT and pgrl1. Interestingly, some of the core proteins were aggregated in pgr5, which led to change in photosynthesis efficiency in high light.

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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.

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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.

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Background: Some Sacoglossa sea slugs steal and integrate chloroplasts derived from the algae they feed on into their cells where they continue to function photosynthetically, a process termed kleptoplasty. The stolen chloroplasts - kleptoplasts - can maintain their functionality up to several months and support animal metabolism. However, chloroplast longevity can vary depending on sea slug species and algal donor. In this study, we focused on Elysia viridis, a polyphagous species that is mostly found associated with the macroalga Codium tomentosum, but that was reported to eat other macroalgae, including Chaetomorpha sp. Methods: We have investigated the changes in E. viridis physiology when provided with the two different food sources to evaluate to which extent the photosynthetic and photoprotective mechanisms of the algae chloroplasts matched those of the plastids once in the animal cells. To perform the study, we rely on the evaluation of chlorophyll a variable fluorescence to study the photophysiological state of the integrated kleptoplasts and high-performance liquid chromatography (HPLC) to study variations in the photosynthetic pigments. Results: We observed that the photosynthetic efficiency of E. viridis is lower when fed with Chaetomorpha. Also, significant differences were observed in the non-photochemical quenching (NPQ) abilities of the sea slugs. While sea slugs fed with C. tomentosum react similarly to high-light stress as the alga, E. viridis hosting Chaetomorpha chloroplasts were unable to properly recover from photoinhibition or perform a functional xanthophyll cycle (XC). Conclusions: Our results showed that, even if the sea slugs fed with the two algae show photosynthetic activities like the respective algal donors, not all the photoprotective mechanisms present in Chaetomorpha can be maintained in E. viridis. This indicates that the functionality of the kleptoplasts does not depend solely on their origin but also on the degree of compatibility with the animal species integrating them.

Sea slugs known as Sacoglossa (also called "solar-powered sea slugs") have a fascinating ability to steal and use chloroplasts from the algae they eat. This process is called kleptoplasty. These stolen chloroplasts, also called kleptoplasts, can remain functional for several months and help the sea slugs with their metabolism by performing photosynthesis like plants. However, the time of chloroplast maintenance can vary depending on the species of sea slug and the type of algae they feed on. In this study, we focused on a species called Elysia viridis, which eats various types of algae, including Codium tomentosum and Chaetomorpha sp. These two algae have different characteristics when it comes to photosynthesis and protection against excessive light. We investigated how the physiology of E. viridis changed when it was given these two different food sources. Our results show that sea slugs had similar photosynthetic activities to their respective food alga. However, not all photoprotective mechanisms of Chaetomorpha algae could be maintained in E. viridis. This suggests that the functionality of the stolen chloroplasts depends not only on their source but also on how well they match the sea slug species that incorporate them.

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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 H2O2 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.

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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.

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