{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T02:07:18Z","timestamp":1788833238769,"version":"build-2803163510"},"reference-count":41,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2027,4,1]],"date-time":"2027-04-01T00:00:00Z","timestamp":1806537600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2027,4,1]],"date-time":"2027-04-01T00:00:00Z","timestamp":1806537600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,8,6]],"date-time":"2026-08-06T00:00:00Z","timestamp":1785974400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012226","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","award":["JUSRP202504023"],"award-info":[{"award-number":["JUSRP202504023"]}],"id":[{"id":"10.13039\/501100012226","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["22578169"],"award-info":[{"award-number":["22578169"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100013804","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100013804","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Synthetic and Systems Biotechnology"],"published-print":{"date-parts":[[2027,4]]},"DOI":"10.1016\/j.synbio.2026.08.003","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T02:38:12Z","timestamp":1787107092000},"page":"1-12","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["Combinatorial metabolic engineering of Mycobacterium neoaurum LY-2 for efficient androstenedione production from phytosterols"],"prefix":"10.1016","volume":"18","author":[{"given":"Liyang","family":"Xu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chuxian","family":"Peng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenjing","family":"Pan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingjia","family":"Tong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinsong","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhenghong","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"6","key":"10.1016\/j.synbio.2026.08.003_bib1","doi-asserted-by":"crossref","first-page":"1423","DOI":"10.1007\/s00253-012-4078-0","article-title":"Microbial steroid transformations: current state and prospects","volume":"94","author":"Donova","year":"2012","journal-title":"Appl Microbiol Biotechnol"},{"issue":"6","key":"10.1016\/j.synbio.2026.08.003_bib2","doi-asserted-by":"crossref","first-page":"688","DOI":"10.1016\/S0141-0229(03)00029-2","article-title":"Microbial conversion of steroid compounds: recent developments","volume":"32","author":"Fernandes","year":"2003","journal-title":"Enzyme Microb Technol"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1134\/S0003683807010012","article-title":"Transformation of steroids by actinobacteria: a review","volume":"43","author":"Donova","year":"2007","journal-title":"Appl Biochem Microbiol"},{"issue":"15","key":"10.1016\/j.synbio.2026.08.003_bib4","doi-asserted-by":"crossref","first-page":"6725","DOI":"10.1016\/j.biortech.2008.01.039","article-title":"Androstenedione production by biotransformation of phytosterols","volume":"99","author":"Malaviya","year":"2008","journal-title":"Bioresour Technol"},{"issue":"6","key":"10.1016\/j.synbio.2026.08.003_bib5","doi-asserted-by":"crossref","first-page":"1947","DOI":"10.1073\/pnas.0605728104","article-title":"A gene cluster encoding cholesterol catabolism in a soil actinomycete provides insight into Mycobacterium tuberculosis survival in macrophages","volume":"104","author":"Van der Geize","year":"2007","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"10.1016\/j.synbio.2026.08.003_bib6","doi-asserted-by":"crossref","first-page":"684","DOI":"10.1111\/j.1365-2958.2007.05827.x","article-title":"A highly conserved transcriptional repressor controls a large regulon involved in lipid degradation in Mycobacterium smegmatis and Mycobacterium tuberculosis","volume":"65","author":"Kendall","year":"2007","journal-title":"Mol Microbiol"},{"issue":"2","key":"10.1016\/j.synbio.2026.08.003_bib7","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1631\/jzus.B1200067","article-title":"Optimization of biotransformation from phytosterol to androstenedione by a mutant Mycobacterium neoaurum ZJUVN-08","volume":"14","author":"Zhang","year":"2013","journal-title":"J Zhejiang Univ - Sci B"},{"key":"10.1016\/j.synbio.2026.08.003_bib8","doi-asserted-by":"crossref","DOI":"10.1016\/j.ces.2020.116195","article-title":"Enhancing production of 9\u03b1-hydroxy-androst-4-ene-3, 17-dione (9-OHAD) from phytosterols by metabolic pathway engineering of mycobacteria","volume":"230","author":"Sun","year":"2021","journal-title":"Chem Eng Sci"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib9","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1186\/s13068-023-02376-2","article-title":"Unravelling and engineering an operon involved in the side-chain degradation of sterols in Mycolicibacterium neoaurum for the production of steroid synthons","volume":"16","author":"Zhao","year":"2023","journal-title":"Biotechnol Biofuels Bioprod"},{"issue":"4","key":"10.1016\/j.synbio.2026.08.003_bib10","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1007\/s43393-023-00161-8","article-title":"The function of steroid terminal monooxygenase in cholesterol or phytosterol degradation by the Mycobacterium neoaurum NRRL B-3805","volume":"3","author":"Li","year":"2023","journal-title":"Syst Microbiol Biomanuf"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib11","doi-asserted-by":"crossref","DOI":"10.1002\/biot.202300439","article-title":"Enhancing production and purity of 9-OH-AD from phytosterols by balancing metabolic flux of the side-chain degradation and 9-position hydroxylation in Mycobacterium neoaurum","volume":"19","author":"Zhu","year":"2023","journal-title":"Biotechnol J"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib12","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1007\/s00253-023-12847-z","article-title":"Rerouting phytosterol degradation pathway for directed androst-1, 4-diene-3, 17-dione microbial bioconversion","volume":"108","author":"Ke","year":"2024","journal-title":"Appl Microbiol Biotechnol"},{"key":"10.1016\/j.synbio.2026.08.003_bib13","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1016\/j.procbio.2023.06.023","article-title":"Aldolase SalA dominates C24 steroidal side-chain-cleavage in the phytosterol degradation from Mycobacterium neoaurum","volume":"131","author":"Wang","year":"2023","journal-title":"Process Biochem"},{"issue":"25","key":"10.1016\/j.synbio.2026.08.003_bib14","doi-asserted-by":"crossref","first-page":"9541","DOI":"10.1021\/acssuschemeng.5c02070","article-title":"Integrated metabolic engineering and substrate pretreatment for sustainable androstenedione production from phytosterols","volume":"13","author":"Gao","year":"2025","journal-title":"ACS Sustain Chem Eng"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib15","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1186\/s12934-022-01942-x","article-title":"Combined enhancement of the propionyl-CoA metabolic pathway for efficient androstenedione production in Mycolicibacterium neoaurum","volume":"21","author":"Su","year":"2022","journal-title":"Microb Cell Fact"},{"issue":"9","key":"10.1016\/j.synbio.2026.08.003_bib16","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1007\/s10295-012-1130-0","article-title":"Influence of hydroxypropyl-\u03b2-cyclodextrin on phytosterol biotransformation by different strains of Mycobacterium neoaurum","volume":"39","author":"Shen","year":"2012","journal-title":"J Ind Microbiol Biotechnol"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib17","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1186\/s12934-023-02052-y","article-title":"Improving the production of 9\u03b1-hydroxy-4-androstene-3, 17-dione from phytosterols by 3-ketosteroid-\u03941-dehydrogenase deletions and multiple genetic modifications in Mycobacterium fortuitum","volume":"22","author":"Liu","year":"2023","journal-title":"Microb Cell Fact"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib18","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1186\/s13068-023-02331-1","article-title":"Driving the conversion of phytosterol to 9\u03b1-hydroxy-4-androstene-3, 17-dione in Mycolicibacterium neoaurum by engineering the supply and regeneration of flavin adenine dinucleotide","volume":"16","author":"Song","year":"2023","journal-title":"Biotechnol Biofuels Bioprod"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib19","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1186\/s12934-017-0796-4","article-title":"Cofactor engineering to regulate NAD+\/NADH ratio with its application to phytosterols biotransformation","volume":"16","author":"Su","year":"2017","journal-title":"Microb Cell Fact"},{"issue":"6","key":"10.1016\/j.synbio.2026.08.003_bib20","doi-asserted-by":"crossref","first-page":"679","DOI":"10.1111\/j.1751-7915.2012.00331.x","article-title":"Catabolism and biotechnological applications of cholesterol degrading bacteria","volume":"5","author":"Garc\u00eda","year":"2012","journal-title":"Microb Biotechnol"},{"issue":"4","key":"10.1016\/j.synbio.2026.08.003_bib21","first-page":"1110","article-title":"Unraveling the structure of the mycobacterial envelope","volume":"7","author":"Daff\u00e9","year":"2019","journal-title":"Methods"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib22","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/j.chembiol.2013.11.011","article-title":"Mycolic acids: structures, biosynthesis, and beyond","volume":"21","author":"Marrakchi","year":"2014","journal-title":"Chem Biol"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib23","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1016\/j.synbio.2021.11.007","article-title":"Improving the biotransformation efficiency of soybean phytosterols in Mycolicibacterium neoaurum by the combined deletion of fbpC3 and embC in cell envelope synthesis","volume":"7","author":"Xiong","year":"2022","journal-title":"Synth Syst Biotechnol"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib24","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1186\/s12934-020-01335-y","article-title":"Enhancing the bioconversion of phytosterols to steroidal intermediates by the deficiency of kasB in the cell wall synthesis of Mycobacterium neoaurum","volume":"19","author":"Xiong","year":"2020","journal-title":"Microb Cell Fact"},{"issue":"11","key":"10.1016\/j.synbio.2026.08.003_bib25","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1007\/s11274-024-04165-x","article-title":"Improving the bioconversion of phytosterols to 9\u03b1-hydroxy-4-androstene-3, 17-dione by disruption of acyltransferase SucT and TmaT associated with the mycobacterial cell wall synthesis","volume":"40","author":"Chen","year":"2024","journal-title":"World J Microbiol Biotechnol"},{"issue":"4","key":"10.1016\/j.synbio.2026.08.003_bib26","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2- \u0394\u0394CT method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"issue":"4","key":"10.1016\/j.synbio.2026.08.003_bib27","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.ymeth.2010.01.005","article-title":"A practical approach to RT-qPCR\u2014publishing data that conform to the MIQE guidelines","volume":"50","author":"Taylor","year":"2010","journal-title":"Methods"},{"key":"10.1016\/j.synbio.2026.08.003_bib28","doi-asserted-by":"crossref","DOI":"10.1016\/j.biortech.2020.123307","article-title":"Efficient repeated batch production of androstenedione using untreated cane molasses by Mycobacterium neoaurum driven by ATP futile cycle","volume":"309","author":"Zhou","year":"2020","journal-title":"Bioresour Technol"},{"key":"10.1016\/j.synbio.2026.08.003_bib29","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/j.biortech.2019.01.144","article-title":"Efficient production of androstenedione by repeated batch fermentation in waste cooking oil media through regulating NAD+\/NADH ratio and strengthening cell vitality of Mycobacterium neoaurum","volume":"279","author":"Zhou","year":"2019","journal-title":"Bioresour Technol"},{"issue":"21","key":"10.1016\/j.synbio.2026.08.003_bib30","doi-asserted-by":"crossref","first-page":"3841","DOI":"10.3390\/molecules24213841","article-title":"Intracellular environment improvement of Mycobacterium neoaurum for enhancing androst-1, 4-diene-3, 17-dione production by manipulating NADH and reactive oxygen species levels","volume":"24","author":"Shao","year":"2019","journal-title":"Molecules"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib31","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.1186\/s40064-016-2871-4","article-title":"Efficient 9\u03b1-hydroxy-4-androstene-3, 17-dione production by engineered Bacillus subtilis co-expressing Mycobacterium neoaurum 3-ketosteroid 9\u03b1-hydroxylase and B. subtilis glucose 1-dehydrogenase with NADH regeneration","volume":"5","author":"Zhang","year":"2016","journal-title":"SpringerPlus"},{"issue":"5","key":"10.1016\/j.synbio.2026.08.003_bib32","doi-asserted-by":"crossref","first-page":"1362","DOI":"10.1099\/mic.0.034538-0","article-title":"Cholesterol utilization in mycobacteria is controlled by two TetR-type transcriptional regulators: KstR and KstR2","volume":"156","author":"Kendall","year":"2010","journal-title":"Microbiology"},{"issue":"2","key":"10.1016\/j.synbio.2026.08.003_bib33","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1016\/j.chembiol.2011.12.016","article-title":"Cholesterol catabolism by Mycobacterium tuberculosis requires transcriptional and metabolic adaptations","volume":"19","author":"Griffin","year":"2012","journal-title":"Chem Biol"},{"issue":"4","key":"10.1016\/j.synbio.2026.08.003_bib34","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1111\/j.1365-2958.2006.05297.x","article-title":"Dual role of isocitrate lyase 1 in the glyoxylate and methylcitrate cycles in Mycobacterium tuberculosis","volume":"61","author":"Gould","year":"2006","journal-title":"Mol Microbiol"},{"issue":"12","key":"10.1016\/j.synbio.2026.08.003_bib35","doi-asserted-by":"crossref","first-page":"3973","DOI":"10.1099\/mic.0.2007\/011726-0","article-title":"Role of the methylcitrate cycle in propionate metabolism and detoxification in Mycobacterium smegmatis","volume":"153","author":"Upton","year":"2007","journal-title":"Microbiology"},{"issue":"9","key":"10.1016\/j.synbio.2026.08.003_bib36","doi-asserted-by":"crossref","first-page":"1928","DOI":"10.1002\/bit.26336","article-title":"Metabolic engineering of cofactor flavin adenine dinucleotide (FAD) synthesis and regeneration in Escherichia coli for production of \u03b1\u2010keto acids","volume":"114","author":"Hou","year":"2017","journal-title":"Biotechnol Bioeng"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib37","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1038\/s41579-019-0273-7","article-title":"The mycobacterial cell envelope-a moving target","volume":"18","author":"Dulberger","year":"2020","journal-title":"Nat Rev Microbiol"},{"key":"10.1016\/j.synbio.2026.08.003_bib38","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1016\/j.jbiotec.2020.09.019","article-title":"Improving the biotransformation of phytosterols to 9\u03b1-hydroxy-4-androstene-3, 17-dione by deleting embC associated with the assembly of cell envelope in Mycobacterium neoaurum","volume":"323","author":"Xiong","year":"2020","journal-title":"J Biotechnol"},{"issue":"2","key":"10.1016\/j.synbio.2026.08.003_bib39","doi-asserted-by":"crossref","first-page":"494","DOI":"10.1007\/s12010-017-2665-3","article-title":"Efficient bioconversion of high concentration phytosterol microdispersion to 4-androstene-3, 17-dione (AD) by Mycobacterium sp. B3805","volume":"185","author":"Mancilla","year":"2018","journal-title":"Appl Biochem Biotechnol"},{"key":"10.1016\/j.synbio.2026.08.003_bib40","doi-asserted-by":"crossref","DOI":"10.1016\/j.bej.2020.107789","article-title":"A combined strategy of metabolic pathway regulation and two-step bioprocess for improved 4-androstene-3, 17-dione production with an engineered Mycobacterium neoaurum","volume":"164","author":"Chang","year":"2020","journal-title":"Biochem Eng J"},{"issue":"1","key":"10.1016\/j.synbio.2026.08.003_bib41","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1186\/s12934-021-01653-9","article-title":"Efficient conversion of phytosterols into 4-androstene-3, 17-dione and its C1, 2-dehydrogenized and 9\u03b1-hydroxylated derivatives by engineered Mycobacteria","volume":"20","author":"Li","year":"2021","journal-title":"Microb Cell Fact"}],"container-title":["Synthetic and Systems Biotechnology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405805X26001924?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405805X26001924?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T01:27:53Z","timestamp":1788830873000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2405805X26001924"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2027,4]]},"references-count":41,"alternative-id":["S2405805X26001924"],"URL":"https:\/\/doi.org\/10.1016\/j.synbio.2026.08.003","relation":{},"ISSN":["2405-805X"],"issn-type":[{"value":"2405-805X","type":"print"}],"subject":[],"published":{"date-parts":[[2027,4]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Combinatorial metabolic engineering of Mycobacterium neoaurum LY-2 for efficient androstenedione production from phytosterols","name":"articletitle","label":"Article Title"},{"value":"Synthetic and Systems Biotechnology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.synbio.2026.08.003","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co. Ltd.","name":"copyright","label":"Copyright"}]}}