{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,21]],"date-time":"2026-07-21T10:43:33Z","timestamp":1784630613595,"version":"3.55.0"},"reference-count":40,"publisher":"Portland Press Ltd.","issue":"6","license":[{"start":{"date-parts":[[2024,5,29]],"date-time":"2024-05-29T00:00:00Z","timestamp":1716940800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"North West Cancer Research\/Mesothelioma UK","award":["NWCRMESO2023.01"],"award-info":[{"award-number":["NWCRMESO2023.01"]}]},{"DOI":"10.13039\/501100000289","name":"Cancer Research UK","doi-asserted-by":"crossref","award":["C18342\/A23390"],"award-info":[{"award-number":["C18342\/A23390"]}],"id":[{"id":"10.13039\/501100000289","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000289","name":"Cancer Research UK","doi-asserted-by":"crossref","award":["DRCRPG-Nov22\/100007"],"award-info":[{"award-number":["DRCRPG-Nov22\/100007"]}],"id":[{"id":"10.13039\/501100000289","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000357","name":"North West Cancer Research","doi-asserted-by":"crossref","award":["RDG2021.12"],"award-info":[{"award-number":["RDG2021.12"]}],"id":[{"id":"10.13039\/501100000357","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000268","name":"Biotechnology and Biological Sciences Research Council","doi-asserted-by":"crossref","award":["BB\/T008695\/1"],"award-info":[{"award-number":["BB\/T008695\/1"]}],"id":[{"id":"10.13039\/501100000268","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Medical Research Council","award":["MR\/K0019494\/1"],"award-info":[{"award-number":["MR\/K0019494\/1"]}]},{"name":"Medical Research Council","award":["MR\/R023026\/1"],"award-info":[{"award-number":["MR\/R023026\/1"]}]},{"name":"Worldwide Cancer Research","award":["24-0353"],"award-info":[{"award-number":["24-0353"]}]}],"content-domain":{"domain":["portlandpress.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2024,6,26]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Liver fibrosis is the excessive accumulation of extracellular matrix proteins that occurs in most types of chronic liver disease. At the cellular level, liver fibrosis is associated with the activation of hepatic stellate cells (HSCs) which transdifferentiate into a myofibroblast-like phenotype that is contractile, proliferative and profibrogenic. HSC transdifferentiation induces genome-wide changes in gene expression that enable the cell to adopt its profibrogenic functions. We have previously identified that the deubiquitinase ubiquitin C-terminal hydrolase 1 (UCHL1) is highly induced following HSC activation; however, the cellular targets of its deubiquitinating activity are poorly defined.<\/jats:p>\n               <jats:p>Here, we describe a role for UCHL1 in regulating the levels and activity of hypoxia-inducible factor 1 (HIF1), an oxygen-sensitive transcription factor, during HSC activation and liver fibrosis. HIF1 is elevated during HSC activation and promotes the expression of profibrotic mediator HIF target genes. Increased HIF1\u03b1 expression correlated with induction of UCHL1 mRNA and protein with HSC activation. Genetic deletion or chemical inhibition of UCHL1 impaired HIF activity through reduction of HIF1\u03b1 levels. Furthermore, our mechanistic studies have shown that UCHL1 elevates HIF activity through specific cleavage of degradative ubiquitin chains, elevates levels of pro-fibrotic gene expression and increases proliferation rates. As we also show that UCHL1 inhibition blunts fibrogenesis in a pre-clinical 3D human liver slice model of fibrosis, these results demonstrate how small molecule inhibitors of DUBs can exert therapeutic effects through modulation of HIF transcription factors in liver disease. Furthermore, inhibition of HIF activity using UCHL1 inhibitors may represent a therapeutic opportunity with other HIF-related pathologies.<\/jats:p>","DOI":"10.1042\/bsr20232147","type":"journal-article","created":{"date-parts":[[2024,5,29]],"date-time":"2024-05-29T11:53:21Z","timestamp":1716983601000},"update-policy":"https:\/\/doi.org\/10.1042\/crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["UCHL1-dependent control of hypoxia-inducible factor transcriptional activity during liver fibrosis"],"prefix":"10.1042","volume":"44","author":[{"given":"Amy","family":"Collins","sequence":"first","affiliation":[{"name":"1Newcastle Fibrosis Research Group, Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rebecca","family":"Scott","sequence":"additional","affiliation":[{"name":"1Newcastle Fibrosis Research Group, Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Caroline\u00a0L.","family":"Wilson","sequence":"additional","affiliation":[{"name":"1Newcastle Fibrosis Research Group, Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giuseppe","family":"Abbate","sequence":"additional","affiliation":[{"name":"2FibroFind Ltd, FibroFind Laboratories, Medical School, Newcastle University, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gabrielle B.","family":"Ecclestone","sequence":"additional","affiliation":[{"name":"3Department of Biochemistry, Cell and Systems Biology, Institute of Systems, Molecular and Integrative Biology University of Liverpool, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Adam\u00a0G.","family":"Albanese","sequence":"additional","affiliation":[{"name":"3Department of Biochemistry, Cell and Systems Biology, Institute of Systems, Molecular and Integrative Biology University of Liverpool, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Demi","family":"Biddles","sequence":"additional","affiliation":[{"name":"4Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne NE2 4HH, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven","family":"White","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jeremy","family":"French","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John","family":"Moir","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wasfi","family":"Alrawashdeh","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Colin","family":"Wilson","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sanjay","family":"Pandanaboyana","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John\u00a0S.","family":"Hammond","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rohan","family":"Thakkar","sequence":"additional","affiliation":[{"name":"5Department of HPB and Transplant Surgery, Freeman Hospital, Newcastle Upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fiona","family":"Oakley","sequence":"additional","affiliation":[{"name":"1Newcastle Fibrosis Research Group, Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jelena","family":"Mann","sequence":"additional","affiliation":[{"name":"1Newcastle Fibrosis Research Group, Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, U.K."},{"name":"2FibroFind Ltd, FibroFind Laboratories, Medical School, Newcastle University, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Derek A.","family":"Mann","sequence":"additional","affiliation":[{"name":"1Newcastle Fibrosis Research Group, Biosciences Institute, Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Niall S.","family":"Kenneth","sequence":"additional","affiliation":[{"name":"3Department of Biochemistry, Cell and Systems Biology, Institute of Systems, Molecular and Integrative Biology University of Liverpool, U.K."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"288","published-online":{"date-parts":[[2024,6,14]]},"reference":[{"key":"2024061410511760400_B1","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1016\/j.addr.2009.03.011","article-title":"Transcriptional regulation of hepatic stellate cells","volume":"61","author":"Mann","year":"2009","journal-title":"Adv. Drug. Deliv. Rev."},{"key":"2024061410511760400_B2","doi-asserted-by":"publisher","DOI":"10.3390\/ijms20102507","article-title":"Epigenetic mechanisms in hepatic stellate cell activation during liver fibrosis and carcinogenesis","volume":"20","author":"Barcena-Varela","year":"2019","journal-title":"Int. J. Mol. Sci."},{"key":"2024061410511760400_B3","doi-asserted-by":"publisher","first-page":"397","DOI":"10.1038\/nrgastro.2017.38","article-title":"Mechanisms of hepatic stellate cell activation","volume":"14","author":"Tsuchida","year":"2017","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"2024061410511760400_B4","doi-asserted-by":"publisher","first-page":"1421","DOI":"10.1016\/j.jhep.2015.07.034","article-title":"Ubiquitin C-terminal hydrolase 1: A novel functional marker for liver myofibroblasts and a therapeutic target in chronic liver disease","volume":"63","author":"Wilson","year":"2015","journal-title":"J. Hepatol."},{"key":"2024061410511760400_B5","doi-asserted-by":"publisher","first-page":"174045","DOI":"10.1016\/j.ejphar.2021.174045","article-title":"UCHL1 inhibition attenuates cardiac fibrosis via modulation of nuclear factor-kappaB signaling in fibroblasts","volume":"900","author":"Gong","year":"2021","journal-title":"Eur. J. Pharmacol."},{"key":"2024061410511760400_B6","doi-asserted-by":"publisher","first-page":"12020","DOI":"10.1021\/jacs.0c04527","article-title":"Discovery of a potent and selective covalent inhibitor and activity-based probe for the deubiquitylating enzyme UCHL1, with antifibrotic activity","volume":"142","author":"Panyain","year":"2020","journal-title":"J. Am. Chem. Soc."},{"key":"2024061410511760400_B7","first-page":"12500","article-title":"UCH-L1 involved in regulating the degradation of EGFR and promoting malignant properties in drug-resistant breast cancer","volume":"8","author":"Jin","year":"2015","journal-title":"Int. J. Clin. Exp. Pathol."},{"key":"2024061410511760400_B8","doi-asserted-by":"publisher","first-page":"1460","DOI":"10.1158\/1078-0432.CCR-19-1373","article-title":"Deubiquitinase activity profiling identifies UCHL1 as a candidate oncoprotein that promotes TGFbeta-induced breast cancer metastasis","volume":"26","author":"Liu","year":"2020","journal-title":"Clin. Cancer Res."},{"key":"2024061410511760400_B9","doi-asserted-by":"publisher","first-page":"6153","DOI":"10.1038\/ncomms7153","article-title":"UCHL1 provides diagnostic and antimetastatic strategies due to its deubiquitinating effect on HIF-1alpha","volume":"6","author":"Goto","year":"2015","journal-title":"Nat. Commun."},{"key":"2024061410511760400_B10","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1146\/annurev.biochem.74.082803.133142","article-title":"Proline hydroxylation and gene expression","volume":"74","author":"Kaelin","year":"2005","journal-title":"Annu. Rev. Biochem."},{"key":"2024061410511760400_B11","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1016\/j.molcel.2008.04.009","article-title":"Oxygen sensing by metazoans: the central role of the HIF hydroxylase pathway","volume":"30","author":"Kaelin","year":"2008","journal-title":"Mol. Cell."},{"key":"2024061410511760400_B12","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.biochi.2014.10.013","article-title":"Hypoxia-inducible factor-1alpha in hepatic fibrosis: a promising therapeutic target","volume":"108","author":"Zhan","year":"2015","journal-title":"Biochimie"},{"key":"2024061410511760400_B13","doi-asserted-by":"publisher","first-page":"G582","DOI":"10.1152\/ajpgi.90368.2008","article-title":"Reduced liver fibrosis in hypoxia-inducible factor-1alpha-deficient mice","volume":"296","author":"Moon","year":"2009","journal-title":"Am. J. Physiol. Gastrointest. Liver Physiol."},{"key":"2024061410511760400_B14","doi-asserted-by":"publisher","first-page":"41038","DOI":"10.1038\/srep41038","article-title":"Reduction of hepatic fibrosis by overexpression of von Hippel-Lindau protein in experimental models of chronic liver disease","volume":"7","author":"Wang","year":"2017","journal-title":"Sci. Rep."},{"key":"2024061410511760400_B15","doi-asserted-by":"publisher","first-page":"2826","DOI":"10.1016\/j.ajpath.2018.08.003","article-title":"Loss of prolyl-hydroxylase 1 protects against biliary fibrosis via attenuated activation of hepatic stellate cells","volume":"188","author":"Strowitzki","year":"2018","journal-title":"Am. J. Pathol."},{"key":"2024061410511760400_B16","doi-asserted-by":"publisher","first-page":"622","DOI":"10.1002\/hep.25497","article-title":"Hypoxia and hypoxia inducible factors: diverse roles in liver diseases","volume":"55","author":"Nath","year":"2012","journal-title":"Hepatology"},{"key":"2024061410511760400_B17","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1016\/j.jcmgh.2015.09.005","article-title":"Role of hypoxia-inducible factors in the development of liver fibrosis","volume":"1","author":"Roth","year":"2015","journal-title":"Cell Mol. Gastroenterol. Hepatol."},{"key":"2024061410511760400_B18","doi-asserted-by":"publisher","first-page":"286","DOI":"10.3389\/fonc.2017.00286","article-title":"Non-canonical mechanisms regulating hypoxia-inducible factor 1 alpha in cancer","volume":"7","author":"Iommarini","year":"2017","journal-title":"Front Oncol."},{"key":"2024061410511760400_B19","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1016\/j.biocel.2004.08.012","article-title":"Hypoxia-inducible factor 1: regulation by hypoxic and non-hypoxic activators","volume":"37","author":"Dery","year":"2005","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"2024061410511760400_B20","doi-asserted-by":"publisher","first-page":"9336","DOI":"10.1093\/nar\/gkx549","article-title":"XIAP upregulates expression of HIF target genes by targeting HIF1alpha for Lys63-linked polyubiquitination","volume":"45","author":"Park","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"2024061410511760400_B21","doi-asserted-by":"publisher","first-page":"3878","DOI":"10.1093\/nar\/gky127","article-title":"PERK\/eIF2alpha signaling inhibits HIF-induced gene expression during the unfolded protein response via YB1-dependent regulation of HIF1alpha translation","volume":"46","author":"Ivanova","year":"2018","journal-title":"Nucleic Acids Res."},{"key":"2024061410511760400_B22","doi-asserted-by":"publisher","first-page":"e45","DOI":"10.1093\/nar\/29.9.e45","article-title":"A new mathematical model for relative quantification in real-time RT-PCR","volume":"29","author":"Pfaffl","year":"2001","journal-title":"Nucleic Acids Res."},{"key":"2024061410511760400_B23","doi-asserted-by":"publisher","first-page":"1377","DOI":"10.1002\/hep.30651","article-title":"A bioreactor technology for modeling fibrosis in human and rodent precision-cut liver slices","volume":"70","author":"Paish","year":"2019","journal-title":"Hepatology"},{"key":"2024061410511760400_B24","doi-asserted-by":"publisher","first-page":"3489","DOI":"10.1073\/pnas.1222732110","article-title":"Recessive loss of function of the neuronal ubiquitin hydrolase UCHL1 leads to early-onset progressive neurodegeneration","volume":"110","author":"Bilguvar","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"2024061410511760400_B25","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1111\/cas.14236","article-title":"Ubiquitin carboxyl-terminal hydrolase L1 promotes hypoxia-inducible factor 1-dependent tumor cell malignancy in spheroid models","volume":"111","author":"Li","year":"2020","journal-title":"Cancer Sci."},{"key":"2024061410511760400_B26","doi-asserted-by":"publisher","first-page":"6879","DOI":"10.1038\/s41598-017-06605-1","article-title":"UCHL1-HIF-1 axis-mediated antioxidant property of cancer cells as a therapeutic target for radiosensitization","volume":"7","author":"Nakashima","year":"2017","journal-title":"Sci. Rep."},{"key":"2024061410511760400_B27","doi-asserted-by":"publisher","first-page":"129","DOI":"10.3389\/fnagi.2014.00129","article-title":"Loss of UCHL1 promotes age-related degenerative changes in the enteric nervous system","volume":"6","author":"Coulombe","year":"2014","journal-title":"Front Aging Neurosci."},{"key":"2024061410511760400_B28","doi-asserted-by":"publisher","first-page":"837","DOI":"10.1016\/j.chembiol.2003.08.010","article-title":"Discovery of inhibitors that elucidate the role of UCH-L1 activity in the H1299 lung cancer cell line","volume":"10","author":"Liu","year":"2003","journal-title":"Chem. Biol."},{"key":"2024061410511760400_B29","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1042\/BJ20081055","article-title":"Regulation of gene expression by hypoxia","volume":"414","author":"Kenneth","year":"2008","journal-title":"Biochem. J."},{"key":"2024061410511760400_B30","doi-asserted-by":"publisher","first-page":"108303","DOI":"10.1016\/j.pharmthera.2022.108303","article-title":"The intricate interplay between HIFs, ROS, and the ubiquitin system in the tumor hypoxic microenvironment","volume":"240","author":"Wang","year":"2022","journal-title":"Pharmacol Ther."},{"key":"2024061410511760400_B31","doi-asserted-by":"publisher","first-page":"6735","DOI":"10.1021\/bi960099f","article-title":"Substrate binding and catalysis by ubiquitin C-terminal hydrolases: identification of two active site residues","volume":"35","author":"Larsen","year":"1996","journal-title":"Biochemistry"},{"key":"2024061410511760400_B32","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1007\/s00109-016-1408-1","article-title":"Hypoxia-inducible factors as molecular targets for liver diseases","volume":"94","author":"Ju","year":"2016","journal-title":"J. Mol. Med. (Berl)"},{"key":"2024061410511760400_B33","doi-asserted-by":"publisher","DOI":"10.3390\/cells8060598","article-title":"Involvement of E3 ligases and deubiquitinases in the control of HIF-alpha subunit abundance","volume":"8","author":"Kubaichuk","year":"2019","journal-title":"Cells"},{"key":"2024061410511760400_B34","doi-asserted-by":"publisher","first-page":"2453","DOI":"10.1042\/BCJ20160082","article-title":"Ubiquitin C-terminal hydrolase L1 (UCH-L1): structure, distribution and roles in brain function and dysfunction","volume":"473","author":"Bishop","year":"2016","journal-title":"Biochem. J."},{"key":"2024061410511760400_B35","doi-asserted-by":"publisher","first-page":"123706","DOI":"10.1155\/2012\/123706","article-title":"Ubiquitin C-terminal hydrolase l1 in tumorigenesis","volume":"2012","author":"Hurst-Kennedy","year":"2012","journal-title":"Biochem. Res. Int."},{"key":"2024061410511760400_B36","doi-asserted-by":"publisher","first-page":"4448","DOI":"10.1038\/s41598-017-04259-7","article-title":"Activation of hepatic stellate cells by the ubiquitin C-terminal hydrolase 1 protein secreted from hepatitis C virus-infected hepatocytes","volume":"7","author":"Cheng","year":"2017","journal-title":"Sci. Rep."},{"key":"2024061410511760400_B37","doi-asserted-by":"publisher","first-page":"10604","DOI":"10.1038\/s41598-020-67746-4","article-title":"Ubiquitin C-terminal hydrolase L1 (UCHL1) regulates post-myocardial infarction cardiac fibrosis through glucose-regulated protein of 78 kDa (GRP78)","volume":"10","author":"Lei","year":"2020","journal-title":"Sci. Rep."},{"key":"2024061410511760400_B38","doi-asserted-by":"publisher","first-page":"1161","DOI":"10.1016\/j.stem.2022.07.006","article-title":"Wound healing, fibroblast heterogeneity, and fibrosis","volume":"29","author":"Talbott","year":"2022","journal-title":"Cell Stem Cell."},{"key":"2024061410511760400_B39","doi-asserted-by":"publisher","first-page":"6619","DOI":"10.1038\/s41467-022-34395-2","article-title":"Pan-cancer single-cell analysis reveals the heterogeneity and plasticity of cancer-associated fibroblasts in the tumor microenvironment","volume":"13","author":"Luo","year":"2022","journal-title":"Nat. Commun."},{"key":"2024061410511760400_B40","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1016\/j.matbio.2015.03.005","article-title":"The role of TIMPs in regulation of extracellular matrix proteolysis","volume":"44-46","author":"Arpino","year":"2015","journal-title":"Matrix Biol."}],"container-title":["Bioscience Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/portlandpress.com\/bioscirep\/article-pdf\/doi\/10.1042\/BSR20232147\/958093\/bsr-2023-2147.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/portlandpress.com\/bioscirep\/article-pdf\/doi\/10.1042\/BSR20232147\/958093\/bsr-2023-2147.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,6,14]],"date-time":"2024-06-14T10:51:49Z","timestamp":1718362309000},"score":1,"resource":{"primary":{"URL":"https:\/\/portlandpress.com\/bioscirep\/article\/44\/6\/BSR20232147\/234510\/UCHL1-dependent-control-of-hypoxia-inducible"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,6]]},"references-count":40,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2024,6,26]]}},"URL":"https:\/\/doi.org\/10.1042\/bsr20232147","relation":{},"ISSN":["0144-8463","1573-4935"],"issn-type":[{"value":"0144-8463","type":"print"},{"value":"1573-4935","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2024,6]]},"published":{"date-parts":[[2024,6]]},"article-number":"BSR20232147"}}