{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T05:02:37Z","timestamp":1790830957131,"version":"4.1.0"},"reference-count":36,"publisher":"Elsevier BV","issue":"4","license":[{"start":{"date-parts":[[2026,12,1]],"date-time":"2026-12-01T00:00:00Z","timestamp":1796083200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,12,1]],"date-time":"2026-12-01T00:00:00Z","timestamp":1796083200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T00:00:00Z","timestamp":1786665600000},"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":["82071114"],"award-info":[{"award-number":["82071114"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002799","name":"Capital Medical University","doi-asserted-by":"publisher","award":["JYJF202601"],"award-info":[{"award-number":["JYJF202601"]}],"id":[{"id":"10.13039\/501100002799","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["European Journal of Cell Biology"],"published-print":{"date-parts":[[2026,12]]},"DOI":"10.1016\/j.ejcb.2026.151564","type":"journal-article","created":{"date-parts":[[2026,8,15]],"date-time":"2026-08-15T14:53:48Z","timestamp":1786805628000},"page":"151564","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["GTP-binding protein DIRAS2 inhibits ferroptosis in oral leukoplakia via regulating iron metabolism"],"prefix":"10.1016","volume":"105","author":[{"given":"Zitong","family":"Tian","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xing","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anyi","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenjing","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaofei","family":"Tang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.ejcb.2026.151564_bib1","doi-asserted-by":"crossref","DOI":"10.1016\/j.biopha.2023.116091","article-title":"LCN2: versatile players in breast cancer","volume":"171","author":"Bao","year":"2024","journal-title":"Biomed. Pharmacother."},{"key":"10.1016\/j.ejcb.2026.151564_bib2","doi-asserted-by":"crossref","DOI":"10.3389\/fcell.2025.1532097","article-title":"Expulsion of iron-rich ferritin via CD63-mediated exosome drives ferroptosis resistance in ovarian cancer cells","volume":"13","author":"Battaglia","year":"2025","journal-title":"Front. Cell Dev. Biol."},{"issue":"1","key":"10.1016\/j.ejcb.2026.151564_bib3","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1186\/s10020-025-01149-x","article-title":"Epitranscriptomic regulation of HIF-1: bidirectional regulatory pathways","volume":"31","author":"Benak","year":"2025","journal-title":"Mol. Med."},{"issue":"6","key":"10.1016\/j.ejcb.2026.151564_bib4","doi-asserted-by":"crossref","first-page":"1576","DOI":"10.18632\/oncotarget.1738","article-title":"Roles of neutrophil gelatinase-associated lipocalin (NGAL) in human cancer","volume":"5","author":"Candido","year":"2014","journal-title":"Oncotarget"},{"issue":"6","key":"10.1016\/j.ejcb.2026.151564_bib5","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1038\/nrd.2016.248","article-title":"Targeting iron metabolism in drug discovery and delivery","volume":"16","author":"Crielaard","year":"2017","journal-title":"Nat. Rev. Drug. Discov."},{"issue":"1","key":"10.1016\/j.ejcb.2026.151564_bib6","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1038\/s41368-021-00131-7","article-title":"Tumor microenvironment and immune evasion in head and neck squamous cell carcinoma","volume":"13","author":"Elmusrati","year":"2021","journal-title":"Int. J. Oral Sci."},{"issue":"5","key":"10.1016\/j.ejcb.2026.151564_bib7","doi-asserted-by":"crossref","first-page":"2088","DOI":"10.62347\/MEYW3975","article-title":"Loss of lipocalin2 confers cisplatin vulnerability through modulating NF-\u0138B mediated ferroptosis via ferroportin","volume":"14","author":"Feng","year":"2024","journal-title":"Am. J. Cancer Res."},{"issue":"7019","key":"10.1016\/j.ejcb.2026.151564_bib8","doi-asserted-by":"crossref","first-page":"917","DOI":"10.1038\/nature03104","article-title":"Lipocalin 2 mediates an innate immune response to bacterial infection by sequestrating iron","volume":"432","author":"Flo","year":"2004","journal-title":"Nature"},{"issue":"12","key":"10.1016\/j.ejcb.2026.151564_bib9","doi-asserted-by":"crossref","DOI":"10.1016\/j.bbamcr.2019.118535","article-title":"Iron homeostasis and oxidative stress: an intimate relationship","volume":"1866","author":"Galaris","year":"2019","journal-title":"Biochim. Biophys. Acta Mol. Cell Res."},{"issue":"3","key":"10.1016\/j.ejcb.2026.151564_bib10","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1016\/j.bbagen.2011.10.013","article-title":"Regulation of iron transport and the role of transferrin","volume":"1820","author":"Gkouvatsos","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"issue":"3","key":"10.1016\/j.ejcb.2026.151564_bib11","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/j.pan.2020.01.002","article-title":"Lipocalin-2 expression and function in pancreatic diseases","volume":"20","author":"Gumpper","year":"2020","journal-title":"Pancreatology"},{"issue":"12","key":"10.1016\/j.ejcb.2026.151564_bib12","doi-asserted-by":"crossref","first-page":"2591","DOI":"10.3390\/cells9122591","article-title":"Iron: an essential element of cancer metabolism","volume":"9","author":"Hsu","year":"2020","journal-title":"Cells"},{"issue":"4","key":"10.1016\/j.ejcb.2026.151564_bib13","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1007\/s10735-024-10197-2","article-title":"Induction of peroxiredoxin 1 by hypoxia promotes cellular autophagy and cell proliferation in oral leukoplakia via HIF-1\u03b1\/BNIP3 pathway","volume":"55","author":"Li","year":"2024","journal-title":"J. Mol. Histol."},{"key":"10.1016\/j.ejcb.2026.151564_bib14","doi-asserted-by":"crossref","DOI":"10.1016\/j.tice.2025.103037","article-title":"The small GTP-binding Ras-like protein DIRAS2 promotes cell proliferation in oral leukoplakia via the RAF\/MEK\/MAPK pathway","volume":"96","author":"Li","year":"2025","journal-title":"Tissue Cell"},{"issue":"7","key":"10.1016\/j.ejcb.2026.151564_bib15","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1093\/carcin\/bgw050","article-title":"Lipocalin 2 prevents oral cancer metastasis through carbonic anhydrase IX inhibition and is associated with favourable prognosis","volume":"37","author":"Lin","year":"2016","journal-title":"Carcinogenesis"},{"issue":"22","key":"10.1016\/j.ejcb.2026.151564_bib16","doi-asserted-by":"crossref","first-page":"7038","DOI":"10.1111\/febs.16059","article-title":"Signaling pathways and defense mechanisms of ferroptosis","volume":"289","author":"Liu","year":"2022","journal-title":"FEBS J."},{"issue":"8","key":"10.1016\/j.ejcb.2026.151564_bib17","doi-asserted-by":"crossref","first-page":"3027","DOI":"10.31557\/APJCP.2025.26.8.3027","article-title":"Cross-talk between hypoxia-inducible factors in driving the pathology of oral squamous cell carcinoma: an immunohistochemical analysis of the role of hypoxia-inducible factors and cancer-associated fibroblasts","volume":"26","author":"M","year":"2025","journal-title":"Asian Pac. J. Cancer Prev."},{"key":"10.1016\/j.ejcb.2026.151564_bib18","doi-asserted-by":"crossref","first-page":"7235","DOI":"10.1038\/srep07235","article-title":"Lipocalin2 as a plasma marker for tumors with hypoxic regions","volume":"4","author":"Nakamura","year":"2014","journal-title":"Sci. Rep."},{"issue":"11","key":"10.1016\/j.ejcb.2026.151564_bib19","doi-asserted-by":"crossref","first-page":"1264","DOI":"10.1038\/ng1658","article-title":"Identification of a ferrireductase required for efficient transferrin-dependent iron uptake in erythroid cells","volume":"37","author":"Ohgami","year":"2005","journal-title":"Nat. Genet."},{"issue":"9","key":"10.1016\/j.ejcb.2026.151564_bib20","doi-asserted-by":"crossref","first-page":"4131","DOI":"10.7150\/ijbs.111228","article-title":"Acute exposure to cadmium triggers NCOA4-mediated ferritinophagy and ferroptosis in never-smokers oral cancer cells","volume":"21","author":"Petriaggi","year":"2025","journal-title":"Int. J. Biol. Sci."},{"issue":"1","key":"10.1016\/j.ejcb.2026.151564_bib21","doi-asserted-by":"crossref","first-page":"449","DOI":"10.3390\/ijms24010449","article-title":"Lipid peroxidation and iron metabolism: two corner stones in the homeostasis control of ferroptosis","volume":"24","author":"Rochette","year":"2022","journal-title":"Int. J. Mol. Sci."},{"issue":"12","key":"10.1016\/j.ejcb.2026.151564_bib22","doi-asserted-by":"crossref","first-page":"2489","DOI":"10.1007\/s10147-025-02906-y","article-title":"The role of hypoxia-inducible factor-1\u03b1 on colon cancer progression and metastasis","volume":"30","author":"Saadh","year":"2025","journal-title":"Int. J. Clin. Oncol."},{"issue":"4","key":"10.1016\/j.ejcb.2026.151564_bib23","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1080\/15548627.2018.1427022","article-title":"RAS-related GTPases DIRAS1 and DIRAS2 induce autophagic cancer cell death and are required for autophagy in murine ovarian cancer cells","volume":"14","author":"Sutton","year":"2018","journal-title":"Autophagy"},{"issue":"13","key":"10.1016\/j.ejcb.2026.151564_bib24","doi-asserted-by":"crossref","first-page":"1411","DOI":"10.1177\/0022034520945242","article-title":"Association or causation? Exploring the oral microbiome and cancer links","volume":"99","author":"Teles","year":"2020","journal-title":"J. Dent. Res."},{"issue":"11","key":"10.1016\/j.ejcb.2026.151564_bib25","doi-asserted-by":"crossref","first-page":"4065","DOI":"10.1172\/JCI42004","article-title":"Lipocalin 2 is essential for chronic kidney disease progression in mice and humans","volume":"120","author":"Viau","year":"2010","journal-title":"J. Clin. Invest."},{"issue":"3","key":"10.1016\/j.ejcb.2026.151564_bib26","doi-asserted-by":"crossref","first-page":"1111","DOI":"10.1007\/s10072-024-07832-x","article-title":"Hypoxia-inducible factor-1 as targets for neuroprotection: from ferroptosis to Parkinson's disease","volume":"46","author":"Wang","year":"2025","journal-title":"Neurol. Sci."},{"issue":"1","key":"10.1016\/j.ejcb.2026.151564_bib27","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1038\/s41420-023-01753-y","article-title":"Ferritinophagy: research advance and clinical significance in cancers","volume":"9","author":"Wang","year":"2023","journal-title":"Cell Death Discov."},{"issue":"11","key":"10.1016\/j.ejcb.2026.151564_bib28","doi-asserted-by":"crossref","first-page":"865","DOI":"10.1038\/s41419-024-07260-x","article-title":"Lipocalin-2 promotes CKD vascular calcification by aggravating VSMCs ferroptosis through NCOA4\/FTH1-mediated ferritinophagy","volume":"15","author":"Wang","year":"2024","journal-title":"Cell Death Dis."},{"issue":"8","key":"10.1016\/j.ejcb.2026.151564_bib29","doi-asserted-by":"crossref","first-page":"1862","DOI":"10.1111\/odi.13704","article-title":"Oral potentially malignant disorders: a consensus report from an international seminar on nomenclature and classification, convened by the WHO Collaborating Centre for Oral Cancer","volume":"27","author":"Warnakulasuriya","year":"2021","journal-title":"Oral Dis."},{"issue":"7","key":"10.1016\/j.ejcb.2026.151564_bib30","doi-asserted-by":"crossref","first-page":"2062","DOI":"10.1080\/15548627.2023.2170960","article-title":"ATM orchestrates ferritinophagy and ferroptosis by phosphorylating NCOA4","volume":"19","author":"Wu","year":"2023","journal-title":"Autophagy"},{"key":"10.1016\/j.ejcb.2026.151564_bib31","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1146\/annurev-nutr-071816-064559","article-title":"Lipocalin 2: an emerging player in iron homeostasis and inflammation","volume":"37","author":"Xiao","year":"2017","journal-title":"Annu. Rev. Nutr."},{"key":"10.1016\/j.ejcb.2026.151564_bib32","doi-asserted-by":"crossref","DOI":"10.1016\/j.pharmthera.2023.108385","article-title":"Lipocalin family proteins and their diverse roles in cardiovascular disease","volume":"244","author":"Yang","year":"2023","journal-title":"Pharmacol. Ther."},{"issue":"1","key":"10.1016\/j.ejcb.2026.151564_bib33","doi-asserted-by":"crossref","first-page":"7333","DOI":"10.1038\/s41467-021-27452-9","article-title":"A targetable LIFR-NF-\u03baB-LCN2 axis controls liver tumorigenesis and vulnerability to ferroptosis","volume":"12","author":"Yao","year":"2021","journal-title":"Nat. Commun."},{"issue":"1","key":"10.1016\/j.ejcb.2026.151564_bib34","doi-asserted-by":"crossref","first-page":"26877","DOI":"10.1038\/s41598-024-78552-7","article-title":"FGF6 inhibits oral squamous cell carcinoma progression by regulating PI3K\/AKT and MAPK pathways","volume":"14","author":"Zhang","year":"2024","journal-title":"Sci. Rep."},{"issue":"9","key":"10.1016\/j.ejcb.2026.151564_bib35","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1038\/s41419-024-07013-w","article-title":"RAB17 promotes endometrial cancer progression by inhibiting TFRC-dependent ferroptosis","volume":"15","author":"Zhou","year":"2024","journal-title":"Cell Death Dis."},{"issue":"21","key":"10.1016\/j.ejcb.2026.151564_bib36","doi-asserted-by":"crossref","first-page":"5159","DOI":"10.3390\/cancers15215159","article-title":"Lipocalin-2: a nurturer of tumor progression and a novel candidate for targeted cancer therapy","volume":"15","author":"\u017divalj","year":"2023","journal-title":"Cancers"}],"container-title":["European Journal of Cell Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S017193352600035X?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S017193352600035X?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T04:35:56Z","timestamp":1790829356000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S017193352600035X"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,12]]},"references-count":36,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2026,12]]}},"alternative-id":["S017193352600035X"],"URL":"https:\/\/doi.org\/10.1016\/j.ejcb.2026.151564","relation":{},"ISSN":["0171-9335"],"issn-type":[{"value":"0171-9335","type":"print"}],"subject":[],"published":{"date-parts":[[2026,12]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"GTP-binding protein DIRAS2 inhibits ferroptosis in oral leukoplakia via regulating iron metabolism","name":"articletitle","label":"Article Title"},{"value":"European Journal of Cell Biology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.ejcb.2026.151564","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier GmbH.","name":"copyright","label":"Copyright"}],"article-number":"151564"}}