{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T04:27:03Z","timestamp":1787632023261,"version":"build-2736575974"},"reference-count":81,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T00:00:00Z","timestamp":1787097600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004731","name":"Zhejiang Provincial Natural Science Foundation","doi-asserted-by":"publisher","award":["LMS25C190005"],"award-info":[{"award-number":["LMS25C190005"]}],"id":[{"id":"10.13039\/501100004731","id-type":"DOI","asserted-by":"publisher"}]},{"award":["LMS25C190005"],"award-info":[{"award-number":["LMS25C190005"]}],"id":[{"id":"https:\/\/ror.org\/04aqat463","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32373152 and 32302228"],"award-info":[{"award-number":["32373152 and 32302228"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32373152 and 32302228"],"award-info":[{"award-number":["32373152 and 32302228"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Mar. Drugs","Marine Drugs"],"abstract":"<jats:p>This study evaluated the synergistic effects of sodium chloride (NaCl) and cytidine monophosphate (CMP) on enhancing the co-production of astaxanthin and fatty acids while suppressing secondary cell wall (SCW) formation in Haematococcus lacustris (synonym: H. pluvialis) under high-light stress. An orthogonal design identified the optimal combination (0.5 g\/L NaCl and 0.5 mM CMP), which significantly increased astaxanthin yield by over 35.6% and total fatty acid yield by 28%, while maintaining 96.8% of cells in motile state (SCW-deficient). Physiological analyses revealed elevated reactive oxygen species levels, concomitant with higher actual photochemical efficiency (Fv\u2032\/Fm\u2032) and relative electron transport rates II (rETR(II)) along with enhanced non-photochemical quenching (NPQ) capacity, and metabolic reprogramming characterized by the accumulation of lipids, sugars, and starch alongside decreased protein yield. Metabolomics indicated reduced carbon supply for SCW polysaccharide biosynthesis, coupled with decreased protein yield and altered amino acid profiles characteristic of nitrogen-limited metabolism, which collectively favored the reallocation of carbon resources toward nitrogen-free high-value products. Transcriptomics confirmed the downregulation of SCW component biosynthetic genes and the upregulation of the methylerythritol phosphate (MEP) pathway and astaxanthin biosynthetic pathway. Scale-up experiments validated this strategy for producing astaxanthin-rich motile cells, offering a promising approach for microalgal biorefinery.<\/jats:p>","DOI":"10.3390\/md24080285","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T13:14:01Z","timestamp":1787145241000},"page":"285","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Synergistic Application of Cytidine Monophosphate and Sodium Chloride for Enhanced Co-Production of Astaxanthin and Fatty Acids in Haematococcus lacustris Motile Cells Under High-Light Stress"],"prefix":"10.3390","volume":"24","author":[{"given":"Xiaoyuan","family":"Su","sequence":"first","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hailiang","family":"Xing","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai","family":"Liu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ya","family":"Zhao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-6922-8593","authenticated-orcid":false,"given":"Lijin","family":"Dong","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ziyan","family":"Zhou","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Na","family":"Zhou","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xue","family":"Sun","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liuquan","family":"Zhang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0628-4687","authenticated-orcid":false,"given":"Nianjun","family":"Xu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1177-1057","authenticated-orcid":false,"given":"Chaoyang","family":"Hu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Aquacultural Biotechnology of Ministry of Education, School of Marine Sciences, Ningbo University, Ningbo 315832, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,19]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1064","DOI":"10.1002\/jobm.202100391","article-title":"Astaxanthin: A super antioxidant from microalgae and its therapeutic potential","volume":"62","author":"Kumar","year":"2021","journal-title":"J. Basic Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Dose, J., Matsugo, S., Yokokawa, H., Koshida, Y., Okazaki, S., Seidel, U., Eggersdorfer, M., Rimbach, G., and Esatbeyoglu, T. (2016). Free radical scavenging and cellular antioxidant properties of astaxanthin. Int. J. Mol. Sci., 17.","DOI":"10.3390\/ijms17010103"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1243","DOI":"10.2174\/1386207323666200915092012","article-title":"Astaxanthin prevents lung injury due to hyperoxia and inflammation","volume":"24","author":"Akduman","year":"2021","journal-title":"Comb. Chem. High Throughput Screen."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1559325820926765","DOI":"10.1177\/1559325820926765","article-title":"Effects of astaxanthin on inflammation and insulin resistance in a mouse model of gestational diabetes mellitus","volume":"18","author":"Feng","year":"2020","journal-title":"Dose-Response"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"106985","DOI":"10.1016\/j.jff.2025.106985","article-title":"Comparison of the effects and mechanisms of astaxanthin, astaxanthin esters and astacene on high-fat diet-induced NAFLD","volume":"133","author":"Gao","year":"2025","journal-title":"J. Funct. Foods"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"3760","DOI":"10.1111\/bph.15099","article-title":"Astaxanthin attenuates hepatic damage and mitochondrial dysfunction in non-alcoholic fatty liver disease by up-regulating the FGF21\/PGC-1\u03b1 pathway","volume":"177","author":"Wu","year":"2020","journal-title":"Br. J. Pharmacol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1038\/s41538-025-00654-x","article-title":"Haematococcus pluvialis ameliorates renal fibrosis by restoring mitophagy via PINK1-Parkin-p62-LC3 signaling","volume":"10","author":"Wu","year":"2025","journal-title":"npj Sci. Food"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"369","DOI":"10.5009\/gnl15208","article-title":"Astaxanthin inhibits proliferation of human gastric cancer cell lines by interrupting cell cycle progression","volume":"10","author":"Kim","year":"2016","journal-title":"Gut Liver"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Jannel, S., Caro, Y., Bermudes, M., and Petit, T. (2020). Novel insights into the biotechnological production of Haematococcus pluvialis-derived astaxanthin: Advances and key challenges to allow its industrial use as novel food ingredient. J. Mar. Sci. Eng., 8.","DOI":"10.3390\/jmse8100789"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Shah, M.M.R., Liang, Y., Cheng, J.J., and Daroch, M. (2016). Astaxanthin-producing green microalga Haematococcus pluvialis: From single cell to high value commercial products. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.00531"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Nair, A., Ahirwar, A., Singh, S., Lodhi, R., Lodhi, A., Rai, A., Jadhav, D.A., Varjani, S., and Singh, G. (2023). Astaxanthin as a king of ketocarotenoids: Structure, synthesis, accumulation, bioavailability and antioxidant properties. Mar. Drugs, 21.","DOI":"10.3390\/md21030176"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Chekanov, K. (2023). Diversity and distribution of carotenogenic algae in Europe: A review. Mar. Drugs, 21.","DOI":"10.3390\/md21020108"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Gil, T.-Y., Sim, H.-Y., Lee, H.-Y., Ryu, S., Baek, J.-S., Kim, D.G., Sim, J., and An, H.-J. (2024). Hot-melt extrusion drug delivery system-formulated Haematococcus pluvialis extracts regulate inflammation and oxidative stress in lipopolysaccharide-stimulated macrophages. Mar. Drugs, 22.","DOI":"10.3390\/md22110512"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Rudi, L., Chiriac, T., Cepoi, L., Djur, S., and Valuta, A. (2025). Stage-specific effects of TiO2, ZnO, and CuO nanoparticles on green microalga Haematococcus lacustris: Biomass and astaxanthin biosynthesis. Mar. Drugs, 23.","DOI":"10.3390\/md23050204"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1002\/bit.21391","article-title":"On the relative efficiency of two- vs. one-stage production of astaxanthin by the green alga Haematococcus pluvialis","volume":"98","author":"Aflalo","year":"2007","journal-title":"Biotechnol. Bioeng."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Li, F., Zhang, N., Zhang, Y., Lian, Q., Qin, C., Qian, Z., Wu, Y., Yang, Z., Li, C., and Huang, X. (2021). NaCl promotes the efficient formation of Haematococcus pluvialis nonmotile cells under phosphorus deficiency. Mar. Drugs, 19.","DOI":"10.3390\/md19060337"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/S0168-1656(01)00289-9","article-title":"Two-stage cultures for the production of astaxanthin from Haematococcus pluvialis","volume":"89","author":"Otero","year":"2001","journal-title":"J. Biotechnol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"101768","DOI":"10.1016\/j.algal.2019.101768","article-title":"Primary metabolism is associated with the astaxanthin biosynthesis in the green algae Haematococcus pluvialis under light stress","volume":"46","author":"Hu","year":"2020","journal-title":"Algal Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1023\/A:1018372900649","article-title":"Light-independent, astaxanthin production by the green microalga Haematococcus pluvialis under salt stress","volume":"19","author":"Kobayashi","year":"1997","journal-title":"Biotechnol. Lett."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1007\/s11738-002-0058-9","article-title":"Effect of culture conditions on growth of green alga\u2014Haematococcus pluvialis and astaxanthin production","volume":"24","author":"Tripathi","year":"2002","journal-title":"Acta Physiol. Plant."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"6468","DOI":"10.1016\/j.biortech.2010.03.072","article-title":"Methyl jasmonate- or gibberellins A3-induced astaxanthin accumulation is associated with up-regulation of transcription of \u03b2-carotene ketolase genes (bkts) in microalga Haematococcus pluvialis","volume":"101","author":"Lu","year":"2010","journal-title":"Bioresour. Technol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.enzmictec.2012.07.001","article-title":"Induction of salicylic acid (SA) on transcriptional expression of eight carotenoid genes and astaxanthin accumulation in Haematococcus pluvialis","volume":"51","author":"Gao","year":"2012","journal-title":"Enzym. Microb. Technol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"735757","DOI":"10.1016\/j.aquaculture.2020.735757","article-title":"Transcriptome analysis reveals pathways responsible for the promoting effect of sucrose on astaxanthin accumulation in Haematococcus pluvialis under high light condition","volume":"530","author":"Du","year":"2021","journal-title":"Aquaculture"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1007\/s10811-018-1622-z","article-title":"Exogenous sodium acetate enhances astaxanthin accumulation and photoprotection in Haematococcus pluvialis at the non-motile stage","volume":"31","author":"Zhang","year":"2018","journal-title":"J. Appl. Phycol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"130001","DOI":"10.1016\/j.biortech.2023.130001","article-title":"Exogenous arginine promotes the coproduction of biomass and astaxanthin under high-light conditions in Haematococcus pluvialis","volume":"393","author":"Acheampong","year":"2024","journal-title":"Bioresour. Technol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"128488","DOI":"10.1016\/j.biortech.2022.128488","article-title":"Exogenous proline boosts the co-accumulation of astaxanthin and biomass in stress-induced Haematococcus pluvialis","volume":"369","author":"Xing","year":"2023","journal-title":"Bioresour. Technol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"129802","DOI":"10.1016\/j.biortech.2023.129802","article-title":"Enhancement of astaxanthin production, recovery, and bio-accessibility in Haematococcus pluvialis through taurine-mediated inhibition of secondary cell wall formation under high light conditions","volume":"389","author":"Zhang","year":"2023","journal-title":"Bioresour. Technol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1007\/s1012601-0051-0","article-title":"Presence of a nonhydrolyzable biopolymer in the cell wall of vegetative cells and astaxanthin-rich cysts of Haematococcus pluvialis (Chlorophyceae)","volume":"3","author":"Montsant","year":"2001","journal-title":"Mar. Biotechnol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"5725","DOI":"10.1007\/s00253-020-10648-2","article-title":"Microbial astaxanthin biosynthesis: Recent achievements, challenges, and commercialization outlook","volume":"104","author":"Zhang","year":"2020","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1111\/ijfs.14030","article-title":"Ionic liquid as an effective solvent for cell wall deconstructing through astaxanthin extraction from Haematococcus pluvialis","volume":"54","author":"Liu","year":"2018","journal-title":"Int. J. Food Sci. Technol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"5023","DOI":"10.1021\/acssuschemeng.2c06726","article-title":"One-pot, water-based disruption of cell walls and astaxanthin extraction from Haematococcus pluvialis by mechanochemistry","volume":"11","author":"Su","year":"2023","journal-title":"ACS Sustain. Chem. Eng."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"129041","DOI":"10.1016\/j.biortech.2023.129041","article-title":"Cell wall breaking of Haematococcus pluvialis biomass facilitated by Baijiu jiuqu fermentation with simultaneously production of beverages","volume":"379","author":"Zhang","year":"2023","journal-title":"Bioresour. Technol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"128067","DOI":"10.1016\/j.biortech.2022.128067","article-title":"Increasing production and bio-accessibility of natural astaxanthin in Haematococcus pluvialis by screening and culturing red motile cells under high light condition","volume":"364","author":"Chen","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"127783","DOI":"10.1016\/j.biortech.2022.127783","article-title":"A strategy for interfering with the formation of thick cell walls in Haematococcus pluvialis by down-regulating the mannan synthesis pathway","volume":"362","author":"Xu","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"130150","DOI":"10.1016\/j.biortech.2023.130150","article-title":"The combination of uridine and nitrogen-deprivation promotes the efficient formation of astaxanthin-rich motile cells in Haematococcus pluvialis","volume":"393","author":"Xing","year":"2024","journal-title":"Bioresour. Technol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"742156","DOI":"10.1016\/j.aquaculture.2025.742156","article-title":"Xanthosine-mediated modulation of Haematococcus pluvialis growth and stress response under high light: Insights from physiological and transcriptome analysis","volume":"599","author":"Liu","year":"2025","journal-title":"Aquaculture"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Xing, H., Zhou, N., Liu, K., Yan, X., Li, W., Sun, X., Zhang, L., Liu, F., Xu, N., and Hu, C. (2025). Nitrogen deprivation drives red motile cell formation in Haematococcus pluvialis: Physiological and transcriptomic insights. Metabolites, 15.","DOI":"10.3390\/metabo15060388"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/0960-8524(95)00002-X","article-title":"Factors responsible for astaxanthin formation in the Chlorophyte Haematococcus pluvialis","volume":"53","author":"Harker","year":"1996","journal-title":"Bioresour. Technol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1007\/s00343-024-4104-y","article-title":"Novel insights into saline stress on photosynthetic activity and astaxanthin production of Haematococcus pluvialis","volume":"43","author":"Jin","year":"2024","journal-title":"J. Oceanol. Limnol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"126214","DOI":"10.1016\/j.biortech.2021.126214","article-title":"Sodium chloride stimulates the biomass and astaxanthin production by Haematococcus pluvialis via a two-stage cultivation strategy","volume":"344","author":"Li","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"4504","DOI":"10.3390\/md12084504","article-title":"Accumulation of astaxanthin by a new Haematococcus pluvialis strain BM1 from the white sea coastal rocks (Russia)","volume":"12","author":"Chekanov","year":"2014","journal-title":"Mar. Drugs"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1134\/S0003683818060078","article-title":"Identification and morphological-physiological characterization of astaxanthin producer strains of Haematococcus pluvialis from the Black Sea region","volume":"54","author":"Chelebieva","year":"2018","journal-title":"Appl. Biochem. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"104120","DOI":"10.1016\/j.algal.2025.104120","article-title":"Exopolysaccharide from Haematococcus pluvialis enhances photosynthesis, biomass and astaxanthin accumulation","volume":"90","author":"Chen","year":"2025","journal-title":"Algal Res."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1017\/S0967026202003669","article-title":"Ultrastructural and chemical changes in the cell wall of Haematococcus pluvialis (Volvocales, Chlorophyta) during aplanospore formation","volume":"37","author":"Hagen","year":"2002","journal-title":"Eur. J. Phycol."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Li, F., Cai, M., Lin, M., Huang, X., Wang, J., Ke, H., Zheng, X., Chen, D., Wang, C., and Wu, S. (2019). Differences between motile and nonmotile cells of Haematococcus pluvialis in the production of astaxanthin at different light intensities. Mar. Drugs, 17.","DOI":"10.3390\/md17010039"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1007\/s10811-023-03132-7","article-title":"A review of the representative microalgal-derived reactive oxygen species: Possible formation, ecophysiological implications, and metabolomic-based evaluations","volume":"36","author":"Nguyen","year":"2023","journal-title":"J. Appl. Phycol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Roginskaya, M., and Razskazovskiy, Y. (2023). Oxidative DNA damage and repair: Mechanisms, mutations, and relation to diseases. Antioxidants, 12.","DOI":"10.3390\/antiox12081623"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1007\/s00425-018-3050-3","article-title":"A hypothesis about the origin of carotenoid lipid droplets in the green algae Dunaliella and Haematococcus","volume":"249","author":"Pick","year":"2018","journal-title":"Planta"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1016\/j.biortech.2016.04.112","article-title":"Simultaneous production of triacylglycerol and high-value carotenoids by the astaxanthin-producing oleaginous green microalga Chlorella zofingiensis","volume":"214","author":"Liu","year":"2016","journal-title":"Bioresour. Technol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1016\/j.tifs.2023.04.014","article-title":"Chemical inducers regulate ROS signalling to stimulate astaxanthin production in Haematococcus pluvialis under environmental stresses: A review","volume":"136","author":"Li","year":"2023","journal-title":"Trends Food Sci. Technol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"734430","DOI":"10.1016\/j.aquaculture.2019.734430","article-title":"Transcriptomic analysis unveils survival strategies of autotrophic Haematococcus pluvialis against high light stress","volume":"513","author":"Hu","year":"2019","journal-title":"Aquaculture"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"121894","DOI":"10.1016\/j.biortech.2019.121894","article-title":"Storage of starch and lipids in microalgae: Biosynthesis and manipulation by nutrients","volume":"291","author":"Ran","year":"2019","journal-title":"Bioresour. Technol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1093\/jexbot\/51.345.659","article-title":"Chlorophyll fluorescence\u2014A practical guide","volume":"51","author":"Maxwell","year":"2000","journal-title":"J. Exp. Bot."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1146\/annurev.arplant.59.032607.092759","article-title":"Chlorophyll fluorescence: A probe of photosynthesis in vivo","volume":"59","author":"Baker","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/s11120-016-0246-x","article-title":"Modulation of photosynthetic activity and photoprotection in Haematococcus pluvialis cells during their conversion into haematocysts and back","volume":"128","author":"Chekanov","year":"2016","journal-title":"Photosynth. Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1515\/znc-1998-1-217","article-title":"Does astaxanthin protect Haematococcus against light damage?","volume":"53","author":"Fan","year":"1998","journal-title":"Z. Naturforsch. C J. Biosci."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Shuang, S.-P., Zhang, J.-Y., Cun, Z., Wu, H.-M., Hong, J., and Chen, J.-W. (2022). A comparison of photoprotective mechanism in different light-demanding plants under dynamic light conditions. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.819843"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"106213","DOI":"10.1016\/j.envexpbot.2025.106213","article-title":"Acclimation to high light intensity increases leaf photosynthetic capacity and NPQ kinetics, but barely affects photosynthetic induction rate","volume":"237","author":"Shao","year":"2025","journal-title":"Environ. Exp. Bot."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1967","DOI":"10.1111\/nph.70121","article-title":"Non-photochemical quenching (NPQ) in photoprotection: Insights into NPQ levels required to avoid photoinactivation and photoinhibition","volume":"246","author":"Zuo","year":"2025","journal-title":"New Phytol."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Zhao, X., Chen, T., Feng, B., Zhang, C., Peng, S., Zhang, X., Fu, G., and Tao, L. (2017). Non-photochemical quenching plays a key role in light acclimation of rice plants differing in leaf color. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01968"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Yoshida, K., Seger, A., Corkill, M., Heil, P., Karsh, K., McMinn, A., and Suzuki, K. (2021). Low Fe availability for photosynthesis of sea-ice algae: Ex situ incubation of the ice diatom Fragilariopsis cylindrus in low-Fe sea ice using an ice tank. Front. Mar. Sci., 8.","DOI":"10.3389\/fmars.2021.632087"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"101144","DOI":"10.1016\/j.afres.2025.101144","article-title":"Optimized two-stage process of Haematococcus sp. for enhanced astaxanthin and essential fatty acids accumulation","volume":"5","author":"Samperi","year":"2025","journal-title":"Appl. Food Res."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"125736","DOI":"10.1016\/j.biortech.2021.125736","article-title":"Using green alga Haematococcus pluvialis for astaxanthin and lipid co-production: Advances and outlook","volume":"340","author":"Ren","year":"2021","journal-title":"Bioresour. Technol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1007\/s13668-021-00363-3","article-title":"Dietary polyunsaturated fatty acids (PUFAs): Uses and potential health benefits","volume":"10","author":"Kapoor","year":"2021","journal-title":"Curr. Nutr. Rep."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1495","DOI":"10.1007\/s00253-012-3940-4","article-title":"Patterns of carbohydrate and fatty acid changes under nitrogen starvation in the microalgae Haematococcus pluvialis and Nannochloropsis sp.","volume":"94","author":"Recht","year":"2012","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"30387","DOI":"10.1074\/jbc.M114.555144","article-title":"Metabolite profiling and integrative modeling reveal metabolic constraints for carbon partitioning under nitrogen starvation in the green algae Haematococcus pluvialis","volume":"289","author":"Recht","year":"2014","journal-title":"J. Biol. Chem."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1093\/aob\/mcm050","article-title":"Changes in growth and activity of enzymes involved in nitrate reduction and ammonium assimilation in tomato seedlings in response to NaCl stress","volume":"99","author":"Debouba","year":"2007","journal-title":"Ann. Bot."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/j.enzmictec.2009.07.006","article-title":"Proteomics analysis of proteins in green alga Haematococcus lacustris (Chlorophyceae) expressed under combined stress of nitrogen starvation and high irradiance","volume":"45","author":"Tran","year":"2009","journal-title":"Enzym. Microb. Technol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"109487","DOI":"10.1016\/j.enzmictec.2019.109487","article-title":"Transcriptome analysis of Haematococcus pluvialis of multiple defensive systems against nitrogen starvation","volume":"134","author":"Zhao","year":"2020","journal-title":"Enzym. Microb. Technol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"e00128-20","DOI":"10.1128\/MRA.00128-20","article-title":"Draft genome sequence of the astaxanthin-producing microalga Haematococcus lacustris strain NIES-144","volume":"9","author":"Morimoto","year":"2020","journal-title":"Microbiol. Resour. Announc."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/S0981-9428(00)00167-4","article-title":"Nucleotide sugars and glycosyltransferases for synthesis of cell wall matrix polysaccharides","volume":"38","author":"Gibeaut","year":"2000","journal-title":"Plant Physiol. Biochem."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/S1360-1385(96)80019-7","article-title":"The role of xanthophyll cycle carotenoids in the protection of photosynthesis","volume":"1","author":"Adams","year":"1996","journal-title":"Trends Plant Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1105\/tpc.10.7.1121","article-title":"Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion","volume":"10","author":"Niyogi","year":"1998","journal-title":"Plant Cell"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.bbabio.2011.04.012","article-title":"The role of the xanthophyll cycle and of lutein in photoprotection of photosystem II","volume":"1817","author":"Jahns","year":"2012","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"466","DOI":"10.1016\/j.bbabio.2010.01.001","article-title":"Identification of a slowly inducible zeaxanthin-dependent component of non-photochemical quenching of chlorophyll fluorescence generated under steady-state conditions in Arabidopsis","volume":"1797","author":"Nilkens","year":"2010","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1429","DOI":"10.1016\/j.bbagen.2019.05.002","article-title":"Non-photochemical quenching in the cells of the carotenogenic chlorophyte Haematococcus lacustris under favorable conditions and under stress","volume":"1863","author":"Chekanov","year":"2019","journal-title":"Biochim. Biophys. Acta Gen. Subj."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1104\/pp.19.00955","article-title":"Nucleotide metabolism in plants","volume":"182","author":"Witte","year":"2020","journal-title":"Plant Physiol."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Hazeem, L.J., Kuku, G., Dewailly, E., Slomianny, C., Barras, A., Hamdi, A., Boukherroub, R., Culha, M., and Bououdina, M. (2019). Toxicity effect of silver nanoparticles on photosynthetic pigment content, growth, ROS production and ultrastructural changes of microalgae Chlorella vulgaris. Nanomaterials, 9.","DOI":"10.3390\/nano9070914"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"124418","DOI":"10.1016\/j.biortech.2020.124418","article-title":"Gamma-aminobutyric acid facilitates the simultaneous production of biomass, astaxanthin and lipids in Haematococcus pluvialis under salinity and high-light stress conditions","volume":"320","author":"Li","year":"2021","journal-title":"Bioresour. Technol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/0304-3800(88)90057-9","article-title":"A model for the relationship between light intensity and the rate of photosynthesis in phytoplankton","volume":"42","author":"Eilers","year":"1988","journal-title":"Ecol. Modell."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.algal.2016.09.019","article-title":"Metabolomic profiling of the astaxanthin accumulation process induced by high light in Haematococcus pluvialis","volume":"20","author":"Lv","year":"2016","journal-title":"Algal Res."}],"container-title":["Marine Drugs"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1660-3397\/24\/8\/285\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T04:10:58Z","timestamp":1787631058000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1660-3397\/24\/8\/285"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,19]]},"references-count":81,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["md24080285"],"URL":"https:\/\/doi.org\/10.3390\/md24080285","relation":{},"ISSN":["1660-3397"],"issn-type":[{"value":"1660-3397","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,19]]}}}