{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T21:17:55Z","timestamp":1784582275280,"version":"3.55.0"},"reference-count":50,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T00:00:00Z","timestamp":1780272000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T00:00:00Z","timestamp":1780272000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,11,27]],"date-time":"2025-11-27T00:00:00Z","timestamp":1764201600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32572515"],"award-info":[{"award-number":["32572515"]}],"id":[{"id":"10.13039\/501100001809","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":[[2026,6]]},"DOI":"10.1016\/j.synbio.2025.11.015","type":"journal-article","created":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T04:58:19Z","timestamp":1766033899000},"page":"185-196","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":2,"special_numbering":"C","title":["Formatotrophic Komagataella phaffii expressing recombinant xylanase via metabolic engineering"],"prefix":"10.1016","volume":"12","author":[{"given":"Ziwei","family":"Zhou","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bing","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenjie","family":"Cong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hualan","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu","family":"Zheng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3071-9457","authenticated-orcid":false,"given":"Jianguo","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.synbio.2025.11.015_bib1","doi-asserted-by":"crossref","DOI":"10.1093\/femsyr\/foab059","article-title":"What makes Komagataella phaffii non-conventional?","volume":"21","author":"Ata","year":"2021","journal-title":"FEMS Yeast Res"},{"key":"10.1016\/j.synbio.2025.11.015_bib2","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1038\/s41564-023-01574-w","article-title":"OPENPichia: licence-free Komagataella phaffii chassis strains and toolkit for protein expression","volume":"9","author":"Claes","year":"2024","journal-title":"Nat Microbiol"},{"key":"10.1016\/j.synbio.2025.11.015_bib3","doi-asserted-by":"crossref","first-page":"foaf020","DOI":"10.1093\/femsyr\/foaf020","article-title":"Advancing yeast metabolism for a sustainable single carbon bioeconomy","volume":"25","author":"Kuzman","year":"2025","journal-title":"FEMS Yeast Res"},{"key":"10.1016\/j.synbio.2025.11.015_bib4","doi-asserted-by":"crossref","DOI":"10.1016\/j.cej.2024.152518","article-title":"Photosynthetic cultivation of Chlamydomonas reinhardtii with formate as a novel carbon source to the protein production","volume":"493","author":"Jiang","year":"2024","journal-title":"Chem Eng J"},{"key":"10.1016\/j.synbio.2025.11.015_bib5","doi-asserted-by":"crossref","DOI":"10.1016\/j.bej.2022.108503","article-title":"Heterologous protein expression enhancement of Komagataella phaffii by ammonium formate induction based on transcriptomic analysis","volume":"185","author":"Feng","year":"2022","journal-title":"Biochem Eng J"},{"key":"10.1016\/j.synbio.2025.11.015_bib6","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.cbpa.2016.07.005","article-title":"The formate bio-economy","volume":"35","author":"Yishai","year":"2016","journal-title":"Curr Opin Chem Biol"},{"key":"10.1016\/j.synbio.2025.11.015_bib7","doi-asserted-by":"crossref","DOI":"10.1016\/j.biortech.2023.130027","article-title":"NADH-dependent formate dehydrogenase mutants for efficient carbon dioxide fixation","volume":"393","author":"Xue","year":"2024","journal-title":"Bioresour Technol"},{"key":"10.1016\/j.synbio.2025.11.015_bib8","doi-asserted-by":"crossref","first-page":"1621","DOI":"10.1007\/s12033-023-00803-1","article-title":"A systematic review of the potential of Pichia pastoris (Komagataella phaffii) as an alternative host for biologics production","volume":"66","author":"Vijayakumar","year":"2023","journal-title":"Mol Biotechnol"},{"key":"10.1016\/j.synbio.2025.11.015_bib9","doi-asserted-by":"crossref","first-page":"7801","DOI":"10.1007\/s00253-020-10793-8","article-title":"The Mut+ strain of Komagataella phaffii (Pichia pastoris) expresses PAOX1 5 and 10 times faster than Muts and Mut- strains: evidence that formaldehyde or\/and formate are true inducers of PAOX1","volume":"104","author":"Singh","year":"2020","journal-title":"Appl Microbiol Biotechnol"},{"key":"10.1016\/j.synbio.2025.11.015_bib10","doi-asserted-by":"crossref","first-page":"1515","DOI":"10.1007\/s00449-022-02760-6","article-title":"Enhancing xylanase expression of Komagataella phaffii induced by formate through Mit1 co-expression","volume":"45","author":"Liu","year":"2022","journal-title":"Bioproc Biosyst Eng"},{"key":"10.1016\/j.synbio.2025.11.015_bib11","doi-asserted-by":"crossref","first-page":"7819","DOI":"10.1007\/s00253-022-12249-7","article-title":"Enhancing xylanase expression by Komagataella phaffii by formate as carbon source and inducer","volume":"106","author":"Liu","year":"2022","journal-title":"Appl Microbiol Biotechnol"},{"key":"10.1016\/j.synbio.2025.11.015_bib12","doi-asserted-by":"crossref","DOI":"10.1093\/synbio\/ysab020","article-title":"Enzyme engineering and in vivo testing of a formate reduction pathway","volume":"6","author":"Wang","year":"2021","journal-title":"Synth Biol"},{"key":"10.1016\/j.synbio.2025.11.015_bib13","doi-asserted-by":"crossref","first-page":"1176","DOI":"10.1016\/j.jmb.2015.02.005","article-title":"Multiple pathways for protein transport to peroxisomes","volume":"427","author":"Kim","year":"2015","journal-title":"J Mol Biol"},{"key":"10.1016\/j.synbio.2025.11.015_bib14","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1016\/j.bbrc.2018.06.008","article-title":"SKLPT imaging: efficient in\u00c2 vivo pre-evaluation of genome-editing modules using fluorescent protein with peroxisome targeting signal","volume":"503","author":"Konno","year":"2018","journal-title":"Biochem Biophys Res Commun"},{"key":"10.1016\/j.synbio.2025.11.015_bib15","doi-asserted-by":"crossref","first-page":"1161","DOI":"10.1016\/j.tibtech.2024.02.005","article-title":"Peroxisome-based metabolic engineering for biomanufacturing and agriculture","volume":"42","author":"Song","year":"2024","journal-title":"Trends Biotechnol"},{"key":"10.1016\/j.synbio.2025.11.015_bib16","doi-asserted-by":"crossref","first-page":"862","DOI":"10.1016\/j.synbio.2022.04.005","article-title":"Comparative proteomics analysis of Pichia pastoris cultivating in glucose and methanol","volume":"7","author":"Hou","year":"2022","journal-title":"Synth Sys Biotechnol"},{"key":"10.1016\/j.synbio.2025.11.015_bib17","series-title":"Molecular cloning: a laboratory manual","author":"Green","year":"2012"},{"key":"10.1016\/j.synbio.2025.11.015_bib18","article-title":"Small-scale purification of peroxisomes for analytical applications","author":"Cramer","year":"2015","journal-title":"Cold Spring Harb Protoc"},{"key":"10.1016\/j.synbio.2025.11.015_bib19","doi-asserted-by":"crossref","first-page":"3704","DOI":"10.1073\/pnas.1500545112","article-title":"Computational protein design enables a novel one-carbon assimilation pathway","volume":"112","author":"Siegel","year":"2015","journal-title":"Proc Natl Acad Sci USA"},{"key":"10.1016\/j.synbio.2025.11.015_bib20","doi-asserted-by":"crossref","first-page":"1153","DOI":"10.1111\/j.1567-1364.2007.00255.x","article-title":"Formaldehyde dehydrogenase from the recombinant yeast Hansenula polymorpha: isolation and bioanalytic application","volume":"7","author":"Demkiv","year":"2007","journal-title":"FEMS Yeast Res"},{"key":"10.1016\/j.synbio.2025.11.015_bib21","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1007\/s13765-018-0368-2","article-title":"High-level expression and characterization of Aspergillus niger ATCC 1015 xylanase B in Komagataella phaffii","volume":"61","author":"Liu","year":"2018","journal-title":"Appl Biol Chem"},{"key":"10.1016\/j.synbio.2025.11.015_bib22","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1016\/j.bej.2008.03.009","article-title":"A new fermentation strategy for S-adenosylmethionine production in recombinant Pichia pastoris","volume":"41","author":"Zhang","year":"2008","journal-title":"Biochem Eng J"},{"key":"10.1016\/j.synbio.2025.11.015_bib23","article-title":"Energy charge as an indicator of pexophagy in Pichia pastoris","volume":"8","author":"Zhang","year":"2017","journal-title":"Front Microbiol"},{"key":"10.1016\/j.synbio.2025.11.015_bib24","doi-asserted-by":"crossref","first-page":"220","DOI":"10.3390\/microorganisms10020220","article-title":"Unravelling formaldehyde metabolism in bacteria: road towards synthetic methylotrophy","volume":"10","author":"Klein","year":"2022","journal-title":"Microorganisms"},{"key":"10.1016\/j.synbio.2025.11.015_bib25","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1186\/s12915-015-0186-5","article-title":"Systems-level organization of yeast methylotrophic lifestyle","volume":"13","author":"Ru\u00dfmayer","year":"2015","journal-title":"BMC Biol"},{"key":"10.1016\/j.synbio.2025.11.015_bib26","doi-asserted-by":"crossref","first-page":"933","DOI":"10.1016\/j.cell.2020.07.010","article-title":"Converting Escherichia coli to a synthetic methylotroph growing solely on methanol","volume":"182","author":"Chen","year":"2020","journal-title":"Cell"},{"key":"10.1016\/j.synbio.2025.11.015_bib27","doi-asserted-by":"crossref","first-page":"9379","DOI":"10.1007\/s11033-022-07790-7","article-title":"Improving AOX1 promoter efficiency by overexpression of Mit1 transcription factor","volume":"49","author":"Samin","year":"2022","journal-title":"Mol Biol Rep"},{"key":"10.1016\/j.synbio.2025.11.015_bib28","doi-asserted-by":"crossref","first-page":"6245","DOI":"10.1074\/jbc.M115.692053","article-title":"Mit1 transcription factor mediates methanol signaling and regulates the alcohol oxidase 1 (AOX1) promoter in Pichia pastoris","volume":"291","author":"Xiaolong","year":"2016","journal-title":"J Biol Chem"},{"key":"10.1016\/j.synbio.2025.11.015_bib29","doi-asserted-by":"crossref","first-page":"7791","DOI":"10.1093\/nar\/gkab535","article-title":"Recombination machinery engineering facilitates metabolic engineering of the industrial yeast Pichia pastoris","volume":"49","author":"Cai","year":"2021","journal-title":"Nucleic Acids Res"},{"key":"10.1016\/j.synbio.2025.11.015_bib30","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1002\/bit.10556","article-title":"Metabolic analysis of the synthesis of high levels of intracellular human SOD in Saccharomyces cerevisiae rhSOD 2060 411 SGA122","volume":"82","author":"Gonzalez","year":"2003","journal-title":"Biotechnol Bioeng"},{"key":"10.1016\/j.synbio.2025.11.015_bib31","doi-asserted-by":"crossref","DOI":"10.1093\/femsre\/fuaf011","article-title":"Seven critical challenges in synthetic one-carbon assimilation and their potential solutions","volume":"49","author":"Puiggen\u00e9","year":"2025","journal-title":"FEMS Microbiol Rev"},{"key":"10.1016\/j.synbio.2025.11.015_bib32","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1016\/j.jbiosc.2016.03.022","article-title":"Regulation of intracellular formaldehyde toxicity during methanol metabolism of the methylotrophic yeast Pichia methanolica","volume":"122","author":"Wakayama","year":"2016","journal-title":"J Biosci Bioeng"},{"key":"10.1016\/j.synbio.2025.11.015_bib33","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1016\/j.mib.2006.01.001","article-title":"Regulatory links between carbon and nitrogen metabolism","volume":"9","author":"Commichau","year":"2006","journal-title":"Curr Opin Microbiol"},{"key":"10.1016\/j.synbio.2025.11.015_bib34","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1128\/MMBR.00038-15","article-title":"The emergence of 2-oxoglutarate as a master regulator metabolite","volume":"79","author":"Huergo","year":"2015","journal-title":"Microbiol Mol Biol Rev"},{"key":"10.1016\/j.synbio.2025.11.015_bib35","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1038\/s41929-019-0272-0","article-title":"Making quantitative sense of electromicrobial production","volume":"2","author":"Claassens","year":"2019","journal-title":"Nat Catal"},{"key":"10.1016\/j.synbio.2025.11.015_bib36","series-title":"The prokaryotes: volume 2: ecophysiology and biochemistry","first-page":"354","article-title":"Acetogenic prokaryotes","author":"Drake","year":"2006"},{"key":"10.1016\/j.synbio.2025.11.015_bib37","doi-asserted-by":"crossref","first-page":"1039","DOI":"10.1016\/j.bbabio.2012.10.013","article-title":"Design and analysis of metabolic pathways supporting formatotrophic growth for electricity-dependent cultivation of microbes","volume":"1827","author":"Bar-Even","year":"2013","journal-title":"Biochim Biophys Acta"},{"key":"10.1016\/j.synbio.2025.11.015_bib38","doi-asserted-by":"crossref","first-page":"5057","DOI":"10.1128\/JB.00228-08","article-title":"Formate as the main branch point for methylotrophic metabolism in Methylobacterium extorquens AM1","volume":"190","author":"Crowther","year":"2008","journal-title":"J Bacteriol"},{"key":"10.1016\/j.synbio.2025.11.015_bib39","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1007\/BF00406069","article-title":"Growth yields and the efficiency of oxidative phosphorylation during autotrophic growth of Paracoccus denitrificans on methanol and formate","volume":"118","author":"van Verseveld","year":"1978","journal-title":"Arch Microbiol"},{"key":"10.1016\/j.synbio.2025.11.015_bib40","first-page":"1","article-title":"Autotrophic metabolism of formate by Thiobacillus Strain A2","volume":"114","author":"Kelly","year":"1979","journal-title":"Microbiology"},{"key":"10.1016\/j.synbio.2025.11.015_bib41","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.ymben.2024.10.007","article-title":"Evolution-assisted engineering of E. coli enables growth on formic acid at ambient CO2 via the Serine Threonine Cycle","volume":"88","author":"Wenk","year":"2025","journal-title":"Metab Eng"},{"key":"10.1016\/j.synbio.2025.11.015_bib42","doi-asserted-by":"crossref","first-page":"5908","DOI":"10.1038\/s41467-025-61001-y","article-title":"Fast growth and high-titer bioproduction from renewable formate via metal-dependent formate dehydrogenase in Escherichia coli","volume":"16","author":"Cowan","year":"2025","journal-title":"Nat Commun"},{"key":"10.1016\/j.synbio.2025.11.015_bib43","doi-asserted-by":"crossref","first-page":"454","DOI":"10.3390\/microorganisms8030454","article-title":"Formate utilization by the Crenarchaeon Desulfurococcus amylolyticus","volume":"8","author":"Ergal","year":"2020","journal-title":"Microorganisms"},{"key":"10.1016\/j.synbio.2025.11.015_bib44","doi-asserted-by":"crossref","first-page":"7754","DOI":"10.1038\/s41467-023-43610-7","article-title":"The oxygen-tolerant reductive glycine pathway assimilates methanol, formate and CO2 in the yeast Komagataella phaffii","volume":"14","author":"Mitic","year":"2023","journal-title":"Nat Commun"},{"key":"10.1016\/j.synbio.2025.11.015_bib45","doi-asserted-by":"crossref","first-page":"646","DOI":"10.1038\/s41564-025-01941-9","article-title":"One-carbon fixation via the synthetic reductive glycine pathway exceeds yield of the Calvin cycle","volume":"10","author":"Dronsella","year":"2025","journal-title":"Nat Microbiol"},{"key":"10.1016\/j.synbio.2025.11.015_bib46","doi-asserted-by":"crossref","first-page":"7758","DOI":"10.1038\/s41467-023-43631-2","article-title":"Discovery and remodeling of Vibrio natriegens as a microbial platform for efficient formic acid biorefinery","volume":"14","author":"Tian","year":"2023","journal-title":"Nat Commun"},{"key":"10.1016\/j.synbio.2025.11.015_bib47","doi-asserted-by":"crossref","first-page":"650","DOI":"10.1016\/j.synbio.2025.03.001","article-title":"System-level characterization of engineered and evolved formatotrophic E. coli strains","volume":"10","author":"Yilmaz","year":"2025","journal-title":"Synth Sys Biotechnol"},{"key":"10.1016\/j.synbio.2025.11.015_bib48","doi-asserted-by":"crossref","first-page":"1177","DOI":"10.1016\/j.biotechadv.2015.05.008","article-title":"Cultivation strategies to enhance productivity of Pichia pastoris: a review","volume":"33","author":"Looser","year":"2015","journal-title":"Biotechnol Adv"},{"key":"10.1016\/j.synbio.2025.11.015_bib49","doi-asserted-by":"crossref","first-page":"1466","DOI":"10.3390\/microorganisms10071466","article-title":"Role of dissimilative pathway of Komagataella phaffii (Pichia pastoris): Formaldehyde toxicity and energy metabolism","volume":"10","author":"Berrios","year":"2022","journal-title":"Microorganisms"},{"key":"10.1016\/j.synbio.2025.11.015_bib50","article-title":"Beyond alcohol oxidase: the methylotrophic yeast Komagataella phaffii utilizes methanol also with its native alcohol dehydrogenase Adh2","volume":"21","author":"Domen","year":"2021","journal-title":"FEMS Yeast Res"}],"container-title":["Synthetic and Systems Biotechnology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405805X25002029?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2405805X25002029?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T23:58:02Z","timestamp":1779235082000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2405805X25002029"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6]]},"references-count":50,"alternative-id":["S2405805X25002029"],"URL":"https:\/\/doi.org\/10.1016\/j.synbio.2025.11.015","relation":{},"ISSN":["2405-805X"],"issn-type":[{"value":"2405-805X","type":"print"}],"subject":[],"published":{"date-parts":[[2026,6]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Formatotrophic Komagataella phaffii expressing recombinant xylanase via metabolic engineering","name":"articletitle","label":"Article Title"},{"value":"Synthetic and Systems Biotechnology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.synbio.2025.11.015","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2025 The Authors. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co. Ltd.","name":"copyright","label":"Copyright"}]}}