{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:42:21Z","timestamp":1792788141139,"version":"4.5.0"},"reference-count":60,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,7,18]],"date-time":"2025-07-18T00:00:00Z","timestamp":1752796800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,7,18]],"date-time":"2025-07-18T00:00:00Z","timestamp":1752796800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"name":"National Research Foundation (NRF) funded by the Korean government","award":["No. RS-2024-00458958"],"award-info":[{"award-number":["No. RS-2024-00458958"]}]},{"name":"Basic Science Research Program funded by the Korean government","award":["NRF-2020R1A2C2011998"],"award-info":[{"award-number":["NRF-2020R1A2C2011998"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Biotechnol. Biofuels Bioprod."],"DOI":"10.1186\/s13068-025-02676-9","type":"journal-article","created":{"date-parts":[[2025,7,18]],"date-time":"2025-07-18T06:54:25Z","timestamp":1752821665000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":21,"title":["High-yield zeaxanthin production in Chlamydomonas reinhardtii via advanced metabolic pathway engineering"],"prefix":"10.1186","volume":"18","author":[{"given":"Junhwan","family":"Jang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas","family":"Baier","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jacob Sebastian","family":"Kneip","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olaf","family":"Kruse","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"EonSeon","family":"Jin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,7,18]]},"reference":[{"issue":"10","key":"2676_CR1","doi-asserted-by":"publisher","first-page":"1441","DOI":"10.1038\/s41592-023-02023-6","volume":"20","author":"S Dupuis","year":"2023","unstructured":"Dupuis S, Merchant SS. Chlamydomonas reinhardtii: a model for photosynthesis and so much more. Nat Methods. 2023;20(10):1441\u20132. https:\/\/doi.org\/10.1038\/s41592-023-02023-6.","journal-title":"Nat Methods"},{"issue":"4","key":"2676_CR2","doi-asserted-by":"publisher","first-page":"1500","DOI":"10.1104\/pp.010807","volume":"127","author":"CD Silflow","year":"2001","unstructured":"Silflow CD, Lefebvre PA. Assembly and motility of eukaryotic cilia and flagella. Lessons from Chlamydomonas reinhardtii. Plant Physiol. 2001;127(4):1500\u20137. https:\/\/doi.org\/10.1104\/pp.010807.","journal-title":"Plant Physiol"},{"issue":"9","key":"2676_CR3","doi-asserted-by":"publisher","first-page":"2074","DOI":"10.1021\/acssynbio.8b00251","volume":"7","author":"P Crozet","year":"2018","unstructured":"Crozet P, Navarro FJ, Willmund F, Mehrshahi P, Bakowski K, Lauersen KJ, P\u00e9rez-P\u00e9rez M-E, Auroy P, Gorchs Rovira A, Sauret-Gueto S. Birth of a photosynthetic chassis: a MoClo toolkit enabling synthetic biology in the microalga Chlamydomonas reinhardtii. ACS Synth Biol. 2018;7(9):2074\u201386. https:\/\/doi.org\/10.1021\/acssynbio.8b00251.","journal-title":"ACS Synth Biol"},{"issue":"1","key":"2676_CR4","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1083\/jcb.85.1.136","volume":"85","author":"JL Spudich","year":"1980","unstructured":"Spudich JL, Sager R. Regulation of the Chlamydomonas cell cycle by light and dark. J Cell Biol. 1980;85(1):136\u201345. https:\/\/doi.org\/10.1083\/jcb.85.1.136.","journal-title":"J Cell Biol"},{"issue":"11","key":"2676_CR5","doi-asserted-by":"publisher","first-page":"1307","DOI":"10.3390\/cells8111307","volume":"8","author":"F de Carpentier","year":"2019","unstructured":"de Carpentier F, Lemaire SD, Danon A. When unity is strength: the strategies used by Chlamydomonas to survive environmental stresses. Cells. 2019;8(11):1307. https:\/\/doi.org\/10.3390\/cells8111307.","journal-title":"Cells"},{"issue":"10","key":"2676_CR6","doi-asserted-by":"publisher","first-page":"1393","DOI":"10.3390\/plants13101393","volume":"13","author":"JS Kneip","year":"2024","unstructured":"Kneip JS, Kniepkamp N, Jang J, Mortaro MG, Jin E, Kruse O, Baier T. CRISPR\/Cas9-mediated knockout of the lycopene \u03b5-cyclase for efficient astaxanthin production in the green microalga Chlamydomonas reinhardtii. Plants. 2024;13(10):1393. https:\/\/doi.org\/10.3390\/plants13101393.","journal-title":"Plants"},{"issue":"10","key":"2676_CR7","doi-asserted-by":"publisher","first-page":"2053","DOI":"10.1111\/pbi.13364","volume":"18","author":"F Perozeni","year":"2020","unstructured":"Perozeni F, Cazzaniga S, Baier T, Zanoni F, Zoccatelli G, Lauersen KJ, Wobbe L, Ballottari M. Turning a green alga red: engineering astaxanthin biosynthesis by intragenic pseudogene revival in Chlamydomonas reinhardtii. Plant Biotechnol J. 2020;18(10):2053\u201367. https:\/\/doi.org\/10.1111\/pbi.13364.","journal-title":"Plant Biotechnol J"},{"key":"2676_CR8","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12934-020-01480-4","volume":"19","author":"I Song","year":"2020","unstructured":"Song I, Kim J, Baek K, Choi Y, Shin B, Jin E. The generation of metabolic changes for the production of high-purity zeaxanthin mediated by CRISPR-Cas9 in Chlamydomonas reinhardtii. Microb Cell Fact. 2020;19:1\u20139. https:\/\/doi.org\/10.1186\/s12934-020-01480-4.","journal-title":"Microb Cell Fact"},{"issue":"1","key":"2676_CR9","doi-asserted-by":"publisher","first-page":"30620","DOI":"10.1038\/srep30620","volume":"6","author":"K Baek","year":"2016","unstructured":"Baek K, Kim DH, Jeong J, Sim SJ, Melis A, Kim J-S, Jin E, Bae S. DNA-free two-gene knockout in Chlamydomonas reinhardtii via CRISPR-Cas9 ribonucleoproteins. Sci Rep. 2016;6(1):30620. https:\/\/doi.org\/10.1038\/srep30620.","journal-title":"Sci Rep"},{"issue":"1","key":"2676_CR10","doi-asserted-by":"publisher","first-page":"6751","DOI":"10.1038\/s41467-021-27004-1","volume":"12","author":"A Ferenczi","year":"2021","unstructured":"Ferenczi A, Chew YP, Kroll E, von Koppenfels C, Hudson A, Molnar A. Mechanistic and genetic basis of single-strand templated repair at Cas12a-induced DNA breaks in Chlamydomonas reinhardtii. Nat Commun. 2021;12(1):6751. https:\/\/doi.org\/10.1038\/s41467-021-27004-1.","journal-title":"Nat Commun"},{"key":"2676_CR11","doi-asserted-by":"publisher","first-page":"128701","DOI":"10.1016\/j.biortech.2023.128701","volume":"373","author":"B-R Jeong","year":"2023","unstructured":"Jeong B-R, Jang J, Jin E. Genome engineering via gene editing technologies in microalgae. Bioresour Technol. 2023;373:128701. https:\/\/doi.org\/10.1016\/j.biortech.2023.128701.","journal-title":"Bioresour Technol"},{"issue":"1","key":"2676_CR12","doi-asserted-by":"publisher","first-page":"6269","DOI":"10.1038\/s41467-020-19983-4","volume":"11","author":"J Neupert","year":"2020","unstructured":"Neupert J, Gallaher SD, Lu Y, Strenkert D, Segal NA, Barahimipour R, Fitz-Gibbon ST, Schroda M, Merchant SS, Bock R. An epigenetic gene silencing pathway selectively acting on transgenic DNA in the green alga Chlamydomonas. Nat Commun. 2020;11(1):6269. https:\/\/doi.org\/10.1038\/s41467-020-19983-4.","journal-title":"Nat Commun"},{"issue":"2","key":"2676_CR13","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1016\/j.jbiotec.2012.10.010","volume":"167","author":"KJ Lauersen","year":"2013","unstructured":"Lauersen KJ, Berger H, Mussgnug JH, Kruse O. Efficient recombinant protein production and secretion from nuclear transgenes in Chlamydomonas reinhardtii. J Biotechnol. 2013;167(2):101\u201310. https:\/\/doi.org\/10.1016\/j.jbiotec.2012.10.010.","journal-title":"J Biotechnol"},{"key":"2676_CR14","doi-asserted-by":"publisher","DOI":"10.1016\/j.algal.2024.103422","author":"SA Razz","year":"2024","unstructured":"Razz SA. Comprehensive overview of microalgae-derived carotenoids and their applications in diverse industries. Algal Res. 2024. https:\/\/doi.org\/10.1016\/j.algal.2024.103422.","journal-title":"Algal Res"},{"issue":"1","key":"2676_CR15","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1186\/s12934-021-01736-7","volume":"21","author":"I Song","year":"2022","unstructured":"Song I, Kim S, Kim J, Oh H, Jang J, Jeong SJ, Baek K, Shin W-S, Sim SJ, Jin E. Macular pigment-enriched oil production from genome-edited microalgae. Microb Cell Fact. 2022;21(1):27. https:\/\/doi.org\/10.1186\/s12934-021-01736-7.","journal-title":"Microb Cell Fact"},{"issue":"7","key":"2676_CR16","doi-asserted-by":"publisher","first-page":"3934","DOI":"10.1167\/iovs.10-5898","volume":"52","author":"FM Barker","year":"2011","unstructured":"Barker FM, Snodderly DM, Johnson EJ, Schalch W, Koepcke W, Gerss J, Neuringer M. Nutritional manipulation of primate retinas, V: effects of lutein, zeaxanthin, and n\u20133 fatty acids on retinal sensitivity to blue-light\u2013induced damage. Invest Ophthalmol Vis Sci. 2011;52(7):3934\u201342. https:\/\/doi.org\/10.1167\/iovs.10-5898.","journal-title":"Invest Ophthalmol Vis Sci"},{"issue":"1","key":"2676_CR17","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1146\/annurev-nutr-071715-051110","volume":"36","author":"J Mares","year":"2016","unstructured":"Mares J. Lutein and zeaxanthin isomers in eye health and disease. Annu Rev Nutr. 2016;36(1):571\u2013602. https:\/\/doi.org\/10.1146\/annurev-nutr-071715-051110.","journal-title":"Annu Rev Nutr"},{"issue":"6","key":"2676_CR18","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1016\/j.exer.2010.09.015","volume":"91","author":"LM Renzi","year":"2010","unstructured":"Renzi LM, Hammond BR. The effect of macular pigment on heterochromatic luminance contrast. Exp Eye Res. 2010;91(6):896\u2013900. https:\/\/doi.org\/10.1016\/j.exer.2010.09.015.","journal-title":"Exp Eye Res"},{"issue":"1","key":"2676_CR19","doi-asserted-by":"publisher","first-page":"476","DOI":"10.1167\/iovs.12-10411","volume":"54","author":"BR Hammond","year":"2013","unstructured":"Hammond BR, Fletcher LM, Elliott JG. Glare disability, photostress recovery, and chromatic contrast: relation to macular pigment and serum lutein and zeaxanthin. Invest Ophthalmol Vis Sci. 2013;54(1):476\u201381. https:\/\/doi.org\/10.1167\/iovs.12-10411.","journal-title":"Invest Ophthalmol Vis Sci"},{"issue":"12","key":"2676_CR20","doi-asserted-by":"publisher","first-page":"1415","DOI":"10.1001\/jamaophthalmol.2015.3590","volume":"133","author":"J Wu","year":"2015","unstructured":"Wu J, Cho E, Willett WC, Sastry SM, Schaumberg DA. Intakes of lutein, zeaxanthin, and other carotenoids and age-related macular degeneration during 2 decades of prospective follow-up. JAMA Ophthalmol. 2015;133(12):1415\u201324. https:\/\/doi.org\/10.1001\/jamaophthalmol.2015.3590.","journal-title":"JAMA Ophthalmol"},{"key":"2676_CR21","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1016\/j.preteyeres.2015.10.003","volume":"50","author":"PS Bernstein","year":"2016","unstructured":"Bernstein PS, Li B, Vachali PP, Gorusupudi A, Shyam R, Henriksen BS, Nolan JM. Lutein, zeaxanthin, and meso-zeaxanthin: the basic and clinical science underlying carotenoid-based nutritional interventions against ocular disease. Prog Retin Eye Res. 2016;50:34\u201366. https:\/\/doi.org\/10.1016\/j.preteyeres.2015.10.003.","journal-title":"Prog Retin Eye Res"},{"issue":"4","key":"2676_CR22","doi-asserted-by":"publisher","first-page":"827","DOI":"10.3390\/nu14040827","volume":"14","author":"M Mrowicka","year":"2022","unstructured":"Mrowicka M, Mrowicki J, Kucharska E, Majsterek I. Lutein and zeaxanthin and their roles in age-related macular degeneration\u2014neurodegenerative disease. Nutrients. 2022;14(4):827. https:\/\/doi.org\/10.3390\/nu14040827.","journal-title":"Nutrients"},{"key":"2676_CR23","doi-asserted-by":"publisher","first-page":"1460","DOI":"10.1007\/s10620-009-0824-2","volume":"54","author":"SM Chamberlain","year":"2009","unstructured":"Chamberlain SM, Hall JD, Patel J, Lee JR, Marcus DM, Sridhar S, Romero MJ, Labazi M, Caldwell RW, Bartoli M. Protective effects of the carotenoid zeaxanthin in experimental nonalcoholic steatohepatitis. Dig Dis Sci. 2009;54:1460\u20134. https:\/\/doi.org\/10.1007\/s10620-009-0824-2.","journal-title":"Dig Dis Sci"},{"issue":"2","key":"2676_CR24","doi-asserted-by":"publisher","first-page":"392","DOI":"10.1016\/s0735-1097(97)00188-5","volume":"30","author":"L Mosca","year":"1997","unstructured":"Mosca L, Rubenfire M, Mandel C, Rock C, Tarshis T, Tsai A, Pearson T. Antioxidant nutrient supplementation reduces the susceptibility of low density lipoprotein to oxidation in patients with coronary artery disease. J Am Coll Cardiol. 1997;30(2):392\u20139. https:\/\/doi.org\/10.1016\/s0735-1097(97)00188-5.","journal-title":"J Am Coll Cardiol"},{"issue":"3","key":"2676_CR25","doi-asserted-by":"publisher","first-page":"762","DOI":"10.1093\/ajcn\/85.3.762","volume":"85","author":"W Wang","year":"2007","unstructured":"Wang W, Connor SL, Johnson EJ, Klein ML, Hughes S, Connor WE. Effect of dietary lutein and zeaxanthin on plasma carotenoids and their transport in lipoproteins in age-related macular degeneration. Am J Clin Nutr. 2007;85(3):762\u20139. https:\/\/doi.org\/10.1093\/ajcn\/85.3.762.","journal-title":"Am J Clin Nutr"},{"issue":"2","key":"2676_CR26","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1046\/j.1523-1747.2003.12355.x","volume":"121","author":"S Gonz\u00e1lez","year":"2003","unstructured":"Gonz\u00e1lez S, Astner S, An W, Pathak MA, Goukassian D. Dietary lutein\/zeaxanthin decreases ultraviolet B-induced epidermal hyperproliferation and acute inflammation in hairless mice. J Invest Dermatol. 2003;121(2):399\u2013405. https:\/\/doi.org\/10.1046\/j.1523-1747.2003.12355.x.","journal-title":"J Invest Dermatol"},{"issue":"4","key":"2676_CR27","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1159\/000101807","volume":"20","author":"P Palombo","year":"2007","unstructured":"Palombo P, Fabrizi G, Ruocco V, Ruocco E, Fluhr J, Roberts R, Morganti P. Beneficial long-term effects of combined oral\/topical antioxidant treatment with the carotenoids lutein and zeaxanthin on human skin: a double-blind, placebo-controlled study. Skin Pharmacol Physiol. 2007;20(4):199\u2013210. https:\/\/doi.org\/10.1159\/000101807.","journal-title":"Skin Pharmacol Physiol"},{"key":"2676_CR28","doi-asserted-by":"publisher","DOI":"10.1016\/j.algal.2021.102266","volume":"55","author":"L Bourdon","year":"2021","unstructured":"Bourdon L, Jensen AA, Kavanagh JM, McClure DD. Microalgal production of zeaxanthin. Algal Res. 2021;55: 102266. https:\/\/doi.org\/10.1016\/j.algal.2021.102266.","journal-title":"Algal Res"},{"key":"2676_CR29","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1016\/j.fbp.2017.10.008","volume":"107","author":"J Song","year":"2018","unstructured":"Song J, Yang Q, Huang W, Xiao Y, Li D, Liu C. Optimization of trans lutein from pumpkin (Cucurbita moschata) peel by ultrasound-assisted extraction. Food Bioprod Process. 2018;107:104\u201312. https:\/\/doi.org\/10.1016\/j.fbp.2017.10.008.","journal-title":"Food Bioprod Process"},{"key":"2676_CR30","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1016\/j.fitote.2015.12.009","volume":"109","author":"R Manikandan","year":"2016","unstructured":"Manikandan R, Thiagarajan R, Goutham G, Arumugam M, Beulaja M, Rastrelli L, Skalicka-Wo\u017aniak K, Habtemariam S, Orhan IE, Nabavi SF. Zeaxanthin and ocular health, from bench to bedside. Fitoterapia. 2016;109:58\u201366. https:\/\/doi.org\/10.1016\/j.fitote.2015.12.009.","journal-title":"Fitoterapia"},{"issue":"1","key":"2676_CR31","doi-asserted-by":"publisher","first-page":"62","DOI":"10.1016\/j.gendis.2021.02.009","volume":"9","author":"Y Deng","year":"2022","unstructured":"Deng Y, Qiao L, Du M, Qu C, Wan L, Li J, Huang L. Age-related macular degeneration: epidemiology, genetics, pathophysiology, diagnosis, and targeted therapy. Genes Dis. 2022;9(1):62\u201379. https:\/\/doi.org\/10.1016\/j.gendis.2021.02.009.","journal-title":"Genes Dis"},{"issue":"1","key":"2676_CR32","doi-asserted-by":"publisher","first-page":"2222545","DOI":"10.1080\/07853890.2023.2222545","volume":"55","author":"I Korva-Gurung","year":"2023","unstructured":"Korva-Gurung I, Kubin A-M, Ohtonen P, Hautala N. Incidence and prevalence of neovascular age-related macular degeneration: 15-year epidemiological study in a population-based cohort in Finland. Ann Med. 2023;55(1):2222545. https:\/\/doi.org\/10.1080\/07853890.2023.2222545.","journal-title":"Ann Med"},{"key":"2676_CR33","unstructured":"Synthetic zeaxanthin market size and projections [Internet]. 2024 [cited 9 Jun 2025]. https:\/\/www.marketresearchintellect.com\/product\/global-synthetic-zeaxanthin-market-size-and-forecast\/."},{"issue":"4","key":"2676_CR34","doi-asserted-by":"publisher","first-page":"891","DOI":"10.1021\/acs.jafc.7b05400","volume":"66","author":"W Huang","year":"2018","unstructured":"Huang W, Lin Y, He M, Gong Y, Huang J. Induced high-yield production of zeaxanthin, lutein, and \u03b2-carotene by a mutant of Chlorella zofingiensis. J Agric Food Chem. 2018;66(4):891\u20137. https:\/\/doi.org\/10.1021\/acs.jafc.7b05400.","journal-title":"J Agric Food Chem"},{"issue":"3","key":"2676_CR35","doi-asserted-by":"publisher","first-page":"820","DOI":"10.1021\/acssynbio.2c00616","volume":"12","author":"S Amendola","year":"2023","unstructured":"Amendola S, Kneip JS, Meyer F, Perozeni F, Cazzaniga S, Lauersen KJ, Ballottari M, Baier T. Metabolic engineering for efficient ketocarotenoid accumulation in the green microalga Chlamydomonas reinhardtii. ACS Synth Biol. 2023;12(3):820\u201331. https:\/\/doi.org\/10.1021\/acssynbio.2c00616.","journal-title":"ACS Synth Biol"},{"issue":"6","key":"2676_CR36","doi-asserted-by":"publisher","first-page":"189","DOI":"10.3390\/md15060189","volume":"15","author":"M Kim","year":"2017","unstructured":"Kim M, Ahn J, Jeon H, Jin E. Development of a Dunaliella tertiolecta strain with increased zeaxanthin content using random mutagenesis. Mar Drugs. 2017;15(6):189. https:\/\/doi.org\/10.3390\/md15060189.","journal-title":"Mar Drugs"},{"issue":"2","key":"2676_CR37","doi-asserted-by":"publisher","first-page":"e04361","DOI":"10.1128\/spectrum.04361-22","volume":"11","author":"H-H Chen","year":"2023","unstructured":"Chen H-H, Liang M-H, Ye Z-W, Zhu Y-H, Jiang J-G. Engineering the \u03b2-carotene metabolic pathway of microalgae Dunaliella to confirm its carotenoid synthesis pattern in comparison to bacteria and plants. Microbiol Spectr. 2023;11(2):e04361-e14322. https:\/\/doi.org\/10.1128\/spectrum.04361-22.","journal-title":"Microbiol Spectr"},{"issue":"11","key":"2676_CR38","doi-asserted-by":"publisher","DOI":"10.1002\/pld3.185","volume":"3","author":"S Bethmann","year":"2019","unstructured":"Bethmann S, Melzer M, Schwarz N, Jahns P. The zeaxanthin epoxidase is degraded along with the D1 protein during photoinhibition of photosystem II. Plant Direct. 2019;3(11): e00185. https:\/\/doi.org\/10.1002\/pld3.185.","journal-title":"Plant Direct"},{"issue":"3","key":"2676_CR39","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1002\/bit.26499","volume":"115","author":"K Baek","year":"2018","unstructured":"Baek K, Yu J, Jeong J, Sim SJ, Bae S, Jin E. Photoautotrophic production of macular pigment in a Chlamydomonas reinhardtii strain generated by using DNA-free CRISPR-Cas9 RNP-mediated mutagenesis. Biotechnol Bioeng. 2018;115(3):719\u201328. https:\/\/doi.org\/10.1002\/bit.26499.","journal-title":"Biotechnol Bioeng"},{"issue":"1","key":"2676_CR40","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1186\/s13068-024-02483-8","volume":"17","author":"M Kim","year":"2024","unstructured":"Kim M, Cazzaniga S, Jang J, Pivato M, Kim G, Ballottari M, Jin E. Photoautotrophic cultivation of a Chlamydomonas reinhardtii mutant with zeaxanthin as the sole xanthophyll. Biotechnol Biofuels Bioprod. 2024;17(1):41. https:\/\/doi.org\/10.1186\/s13068-024-02483-8.","journal-title":"Biotechnol Biofuels Bioprod"},{"issue":"38","key":"2676_CR41","doi-asserted-by":"publisher","first-page":"11965","DOI":"10.1021\/jacs.8b04844","volume":"140","author":"S Park","year":"2018","unstructured":"Park S, Fischer AL, Steen CJ, Iwai M, Morris JM, Walla PJ, Niyogi KK, Fleming GR. Chlorophyll-carotenoid excitation energy transfer in high-light-exposed thylakoid membranes investigated by snapshot transient absorption spectroscopy. J Am Chem Soc. 2018;140(38):11965\u201373. https:\/\/doi.org\/10.1021\/jacs.8b04844.","journal-title":"J Am Chem Soc"},{"issue":"4","key":"2676_CR42","doi-asserted-by":"publisher","first-page":"760","DOI":"10.1111\/j.1399-3054.1997.tb00003.x","volume":"100","author":"H Paulsen","year":"1997","unstructured":"Paulsen H. Pigment ligation to proteins of the photosynthetic apparatus in higher plants. Physiol Plant. 1997;100(4):760\u20138. https:\/\/doi.org\/10.1111\/j.1399-3054.1997.tb00003.x.","journal-title":"Physiol Plant"},{"key":"2676_CR43","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1016\/j.jtice.2017.04.020","volume":"79","author":"C-Y Chen","year":"2017","unstructured":"Chen C-Y, Ho S-H, Liu C-C, Chang J-S. Enhancing lutein production with Chlorella sorokiniana Mb-1 by optimizing acetate and nitrate concentrations under mixotrophic growth. J Taiwan Inst Chem Eng. 2017;79:88\u201396. https:\/\/doi.org\/10.1016\/j.jtice.2017.04.020.","journal-title":"J Taiwan Inst Chem Eng"},{"key":"2676_CR44","doi-asserted-by":"publisher","first-page":"1545","DOI":"10.1007\/s10811-018-1666-0","volume":"31","author":"Z Zhang","year":"2019","unstructured":"Zhang Z, Tan Y, Wang W, Bai W, Fan J, Huang J, Wan M, Li Y. Efficient heterotrophic cultivation of Chlamydomonas reinhardtii. J Appl Phycol. 2019;31:1545\u201354. https:\/\/doi.org\/10.1007\/s10811-018-1666-0.","journal-title":"J Appl Phycol"},{"key":"2676_CR45","doi-asserted-by":"publisher","DOI":"10.1016\/j.chemosphere.2021.131475","volume":"285","author":"J Liu","year":"2021","unstructured":"Liu J, Yin J, Ge Y, Han H, Liu M, Gao F. Improved lipid productivity of Scenedesmus obliquus with high nutrient removal efficiency by mixotrophic cultivation in actual municipal wastewater. Chemosphere. 2021;285: 131475. https:\/\/doi.org\/10.1016\/j.chemosphere.2021.131475.","journal-title":"Chemosphere"},{"issue":"4","key":"2676_CR46","doi-asserted-by":"publisher","first-page":"992","DOI":"10.1111\/j.1529-8817.2012.01189.x","volume":"48","author":"H Qiao","year":"2012","unstructured":"Qiao H, Wang G, Liu K, Gu W. Short-term effects of acetate and microaerobic conditions on photosynthesis and respiration in chlorella sorokiniana gxnn 01 (chlorophyta) 1. J Phycol. 2012;48(4):992\u20131001. https:\/\/doi.org\/10.1111\/j.1529-8817.2012.01189.x.","journal-title":"J Phycol"},{"key":"2676_CR47","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1016\/j.algal.2018.05.006","volume":"33","author":"FJ Fields","year":"2018","unstructured":"Fields FJ, Ostrand JT, Mayfield SP. Fed-batch mixotrophic cultivation of Chlamydomonas reinhardtii for high-density cultures. Algal Res. 2018;33:109\u201317. https:\/\/doi.org\/10.1016\/j.algal.2018.05.006.","journal-title":"Algal Res"},{"issue":"4","key":"2676_CR48","doi-asserted-by":"publisher","first-page":"1439","DOI":"10.1104\/pp.122.4.1439","volume":"122","author":"PB Heifetz","year":"2000","unstructured":"Heifetz PB, Forster B, Osmond CB, Giles LJ, Boynton JE. Effects of acetate on facultative autotrophy in Chlamydomonas reinhardtii assessed by photosynthetic measurements and stable isotope analyses. Plant Physiol. 2000;122(4):1439\u201346. https:\/\/doi.org\/10.1104\/pp.122.4.1439.","journal-title":"Plant Physiol"},{"issue":"2","key":"2676_CR49","doi-asserted-by":"publisher","first-page":"793","DOI":"10.1021\/jf902628j","volume":"58","author":"KH Cha","year":"2010","unstructured":"Cha KH, Lee HJ, Koo SY, Song D-G, Lee D-U, Pan C-H. Optimization of pressurized liquid extraction of carotenoids and chlorophylls from Chlorella vulgaris. J Agric Food Chem. 2010;58(2):793\u20137. https:\/\/doi.org\/10.1021\/jf902628j.","journal-title":"J Agric Food Chem"},{"key":"2676_CR50","doi-asserted-by":"publisher","first-page":"1269","DOI":"10.1007\/s00232-014-9688-2","volume":"247","author":"E Luengo","year":"2014","unstructured":"Luengo E, Cond\u00f3n-Abanto S, \u00c1lvarez I, Raso J. Effect of pulsed electric field treatments on permeabilization and extraction of pigments from Chlorella vulgaris. J Membr Biol. 2014;247:1269\u201377. https:\/\/doi.org\/10.1007\/s00232-014-9688-2.","journal-title":"J Membr Biol"},{"issue":"6","key":"2676_CR51","doi-asserted-by":"publisher","first-page":"1140","DOI":"10.1111\/j.1365-313X.2008.03746.x","volume":"57","author":"J Neupert","year":"2009","unstructured":"Neupert J, Karcher D, Bock R. Generation of Chlamydomonas strains that efficiently express nuclear transgenes. Plant J. 2009;57(6):1140\u201350. https:\/\/doi.org\/10.1111\/j.1365-313X.2008.03746.x.","journal-title":"Plant J"},{"issue":"6","key":"2676_CR52","doi-asserted-by":"publisher","first-page":"1665","DOI":"10.1073\/pnas.54.6.1665","volume":"54","author":"DS Gorman","year":"1965","unstructured":"Gorman DS, Levine R. Cytochrome f and plastocyanin: their sequence in the photosynthetic electron transport chain of Chlamydomonas reinhardi. PNAS. 1965;54(6):1665\u20139. https:\/\/doi.org\/10.1073\/pnas.54.6.1665.","journal-title":"PNAS"},{"key":"2676_CR53","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.00306","volume":"11","author":"J Kim","year":"2020","unstructured":"Kim J, Lee S, Baek K, Jin E. Site-specific gene knock-out and on-site heterologous gene overexpression in Chlamydomonas reinhardtii via a CRISPR-Cas9-mediated knock-in method. Front Plant Sci. 2020;11: 509810. https:\/\/doi.org\/10.3389\/fpls.2020.00306.","journal-title":"Front Plant Sci"},{"issue":"5","key":"2676_CR54","doi-asserted-by":"publisher","first-page":"e0232594","DOI":"10.1371\/journal.pone.0232594","volume":"15","author":"T Picariello","year":"2020","unstructured":"Picariello T, Hou Y, Kubo T, McNeill NA, Yanagisawa H-A, Oda T, Witman GB. TIM, a targeted insertional mutagenesis method utilizing CRISPR\/Cas9 in Chlamydomonas reinhardtii. PLoS ONE. 2020;15(5):e0232594. https:\/\/doi.org\/10.1371\/journal.pone.0232594.","journal-title":"PLoS ONE"},{"issue":"5","key":"2676_CR55","doi-asserted-by":"publisher","first-page":"837","DOI":"10.3390\/cells11050837","volume":"11","author":"RA Freudenberg","year":"2022","unstructured":"Freudenberg RA, Wittemeier L, Einhaus A, Baier T, Kruse O. The spermidine synthase gene SPD1: A novel auxotrophic marker for Chlamydomonas reinhardtii designed by enhanced CRISPR\/Cas9 gene editing. Cells. 2022;11(5):837. https:\/\/doi.org\/10.3390\/cells11050837.","journal-title":"Cells"},{"key":"2676_CR56","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1007\/s00709-009-0036-9","volume":"235","author":"M Cao","year":"2009","unstructured":"Cao M, Fu Y, Guo Y, Pan J. Chlamydomonas (chlorophyceae) colony PCR. Protoplasma. 2009;235:107\u201310. https:\/\/doi.org\/10.1007\/s00709-009-0036-9.","journal-title":"Protoplasma"},{"issue":"4","key":"2676_CR57","doi-asserted-by":"publisher","first-page":"545","DOI":"10.1111\/tpj.12165","volume":"74","author":"BA Rasala","year":"2013","unstructured":"Rasala BA, Barrera DJ, Ng J, Plucinak TM, Rosenberg JN, Weeks DP, Oyler GA, Peterson TC, Haerizadeh F, Mayfield SP. Expanding the spectral palette of fluorescent proteins for the green microalga Chlamydomonas reinhardtii. Plant J. 2013;74(4):545\u201356. https:\/\/doi.org\/10.1111\/tpj.12165.","journal-title":"Plant J"},{"key":"2676_CR58","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1016\/j.ymben.2016.07.013","volume":"38","author":"KJ Lauersen","year":"2016","unstructured":"Lauersen KJ, Baier T, Wichmann J, W\u00f6rdenweber R, Mussgnug JH, H\u00fcbner W, Huser T, Kruse O. Efficient phototrophic production of a high-value sesquiterpenoid from the eukaryotic microalga Chlamydomonas reinhardtii. Metab Eng. 2016;38:331\u201343. https:\/\/doi.org\/10.1016\/j.ymben.2016.07.013.","journal-title":"Metab Eng"},{"key":"2676_CR59","doi-asserted-by":"publisher","unstructured":"Wellburn A, Lichtenthaler H, editors. Formulae and program to determine total carotenoids and chlorophylls a and b of leaf extracts in different solvents. In: SpringerPlus Photosynthesis, Brussels, Belgium, August 1\u20136, Springer; 1983, vol 2. https:\/\/doi.org\/10.1007\/978-94-017-6368-4_3.","DOI":"10.1007\/978-94-017-6368-4_3"},{"key":"2676_CR60","doi-asserted-by":"publisher","first-page":"474","DOI":"10.1186\/2193-1801-3-474","volume":"3","author":"HI Assaad","year":"2014","unstructured":"Assaad HI, Zhou L, Carroll RJ, Wu G. Rapid publication-ready MS-Word tables for one-way ANOVA. Springerplus. 2014;3:474. https:\/\/doi.org\/10.1186\/2193-1801-3-474.","journal-title":"Springerplus"}],"container-title":["Biotechnology for Biofuels and Bioproducts"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13068-025-02676-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13068-025-02676-9\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13068-025-02676-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,7,18]],"date-time":"2025-07-18T19:02:35Z","timestamp":1752865355000},"score":1,"resource":{"primary":{"URL":"https:\/\/biotechnologyforbiofuels.biomedcentral.com\/articles\/10.1186\/s13068-025-02676-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,18]]},"references-count":60,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["2676"],"URL":"https:\/\/doi.org\/10.1186\/s13068-025-02676-9","relation":{},"ISSN":["2731-3654"],"issn-type":[{"value":"2731-3654","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,18]]},"assertion":[{"value":"25 March 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 June 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 July 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that they have no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"77"}}