{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T11:07:36Z","timestamp":1784286456119,"version":"3.55.0"},"reference-count":53,"publisher":"Oxford University Press (OUP)","issue":"23","license":[{"start":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T00:00:00Z","timestamp":1760140800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01HL15306801"],"award-info":[{"award-number":["R01HL15306801"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"name":"The University of Michigan Cancer Center","award":["P30CA046592"],"award-info":[{"award-number":["P30CA046592"]}]},{"name":"University of Michigan Transgenic Animal Model Core"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,11,18]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The MRE11 DNA nuclease plays central roles in the repair of DNA double-strand breaks (DSBs) as a core component of the MRE11-RAD50-NBS1 (MRN) complex. MRN localizes to chromosomal DSBs and recruits and activates the DSB repair protein kinase, ATM, which phosphorylates downstream substrates to elicit cellular DNA damage responses. Pathogenic variants in MRE11 cause the genome instability disorder ataxia-telangiectasia-like disorder (ATLD). The first ATLD patient allele identified, ATLD1, is a nonsense mutation that deletes 76 residues from the MRE11 C-terminus and markedly reduces levels of MRE11-ATLD1 and the entire MRN complex. The MRE11 C-terminus has been demonstrated to function in DNA binding, mediate protein interactions, and undergo post-translational modifications that regulate the MRE11 nuclease. We previously demonstrated that transgenic mice expressing reduced wildtype MRN levels exhibit severe phenotypes, including small body size, anemia, and DNA DSB repair defects. Thus, it is currently unknown whether low MRE11-ATLD1 levels, loss of the C-terminus, or both cause disease-associated phenotypes. In this study, we generated transgenic mouse models that express near endogenous or significantly reduced levels of MRE11-ATLD1 to determine the in vivo importance of the MRE11 C-terminus. We observe that low MRE11-ATLD1 expression leads to anemia, bone marrow failure, extramedullary hematopoiesis, and impaired lymphocyte development, similar to mice expressing low wildtype MRE11. In contrast, higher MRE11-ATLD1 expression results in a subset of moderate phenotypes, indicating that loss of the C-terminus has limited impact on MRN functions in vivo. These findings provide a foundation for predicting the clinical presentation and severity of ATLD patient phenotypes.<\/jats:p>","DOI":"10.1093\/hmg\/ddaf154","type":"journal-article","created":{"date-parts":[[2025,9,23]],"date-time":"2025-09-23T11:56:25Z","timestamp":1758628585000},"page":"1964-1978","source":"Crossref","is-referenced-by-count":1,"title":["Differential expression of a disease-associated\n                    <i>MRE11<\/i>\n                    variant reveals distinct phenotypic outcomes"],"prefix":"10.1093","volume":"34","author":[{"given":"McKenna B","family":"DeFoer","sequence":"first","affiliation":[{"name":"Department of Human Genetics, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2063, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ahmed M","family":"Mostafa","sequence":"additional","affiliation":[{"name":"Department of Pathology, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2067, Ann Arbor, MI 48109-2200,","place":["United States"]},{"name":"Department of Biochemistry, Faculty of Pharmacy, Ain Shams University , African Union Authority St., Abbassia, Cairo 11566 ,","place":["Egypt"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrea J","family":"Hartlerode","sequence":"additional","affiliation":[{"name":"Department of Human Genetics, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2063, Ann Arbor, MI 48109-2200,","place":["United States"]},{"name":"Department of Pathology, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2067, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven K","family":"Orban","sequence":"additional","affiliation":[{"name":"Department of Pathology, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2067, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Keegan","family":"McDonough","sequence":"additional","affiliation":[{"name":"Department of Human Genetics, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2063, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sophie","family":"Quirk","sequence":"additional","affiliation":[{"name":"Department of Human Genetics, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2063, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brianna K L","family":"Ferguson","sequence":"additional","affiliation":[{"name":"Department of Human Genetics, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2063, Ann Arbor, MI 48109-2200,","place":["United States"]},{"name":"Department of Pathology, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2067, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-7792-5236","authenticated-orcid":false,"given":"David O","family":"Ferguson","sequence":"additional","affiliation":[{"name":"Department of Pathology, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2067, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7178-4258","authenticated-orcid":false,"given":"JoAnn M","family":"Sekiguchi","sequence":"additional","affiliation":[{"name":"Department of Human Genetics, University of Michigan Medical School , 109 Zina Pitcher Place, Rm 2063, Ann Arbor, MI 48109-2200,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,10,11]]},"reference":[{"key":"2026071706515531800_ref1","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1016\/S0092-8674(00)81547-0","article-title":"The DNA double-strand break repair gene hMRE11 is mutated in individuals with an ataxia-telangiectasia-like disorder","volume":"99","author":"Stewart","year":"1999","journal-title":"Cell"},{"key":"2026071706515531800_ref2","doi-asserted-by":"publisher","first-page":"796","DOI":"10.1038\/s41580-021-00394-2","article-title":"Cellular functions of the protein kinase ATM and their relevance to human disease","volume":"22","author":"Lee","year":"2021","journal-title":"Nat Rev Mol Cell Biol"},{"key":"2026071706515531800_ref3","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.cell.2008.08.017","article-title":"Mre11 dimers coordinate DNA end bridging and nuclease processing in double-strand-break repair","volume":"135","author":"Williams","year":"2008","journal-title":"Cell"},{"key":"2026071706515531800_ref4","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.1016\/S1097-2765(01)00381-1","article-title":"Human Rad50\/Mre11 is a flexible complex that can tether DNA ends","volume":"8","author":"Jager","year":"2001","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref5","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1016\/j.molcel.2022.12.003","article-title":"Cryo-EM structure of the Mre11-Rad50-Nbs1 complex reveals the molecular mechanism of scaffolding functions","volume":"83","author":"Rotheneder","year":"2023","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref6","doi-asserted-by":"publisher","first-page":"808","DOI":"10.1038\/nsmb.1639","article-title":"Multiple functions of MRN in end-joining pathways during isotype class switching","volume":"16","author":"Dinkelmann","year":"2009","journal-title":"Nat Struct Mol Biol"},{"key":"2026071706515531800_ref7","doi-asserted-by":"publisher","first-page":"819","DOI":"10.1038\/nsmb.1641","article-title":"Role of Mre11 in chromosomal nonhomologous end joining in mammalian cells","volume":"16","author":"Rass","year":"2009","journal-title":"Nat Struct Mol Biol"},{"key":"2026071706515531800_ref8","doi-asserted-by":"publisher","first-page":"814","DOI":"10.1038\/nsmb.1640","article-title":"Role of mammalian Mre11 in classical and alternative nonhomologous end joining","volume":"16","author":"Xie","year":"2009","journal-title":"Nat Struct Mol Biol"},{"key":"2026071706515531800_ref9","doi-asserted-by":"publisher","first-page":"2144","DOI":"10.1093\/nar\/gkq1175","article-title":"CtIP and MRN promote non-homologous end-joining of etoposide-induced DNA double-strand breaks in G1","volume":"39","author":"Quennet","year":"2011","journal-title":"Nucleic Acids Res"},{"key":"2026071706515531800_ref10","doi-asserted-by":"publisher","first-page":"586","DOI":"10.1038\/s41580-025-00841-4","article-title":"Mechanisms and regulation of DNA end resection in the maintenance of genome stability","volume":"26","author":"Ceccaldi","year":"2025","journal-title":"Nat Rev Mol Cell Biol"},{"key":"2026071706515531800_ref11","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1016\/j.jns.2013.11.032","article-title":"Exome sequencing reveals a novel MRE11 mutation in a patient with progressive myoclonic ataxia","volume":"337","author":"Miyamoto","year":"2014","journal-title":"J Neurol Sci"},{"key":"2026071706515531800_ref12","doi-asserted-by":"publisher","DOI":"10.3390\/cells9071678","article-title":"A survey of reported disease-related mutations in the MRE11-RAD50-NBS1 complex","volume":"9","author":"Rahman","year":"2020","journal-title":"Cells"},{"key":"2026071706515531800_ref13","doi-asserted-by":"publisher","first-page":"1830","DOI":"10.1002\/ajmg.a.36546","article-title":"Hypergonadotropic hypogonadism and hypersegmented neutrophils in a patient with ataxia-telangiectasia-like disorder: potential diagnostic clues?","volume":"164A","author":"Yoshida","year":"2014","journal-title":"Am J Med Genet A"},{"key":"2026071706515531800_ref14","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1016\/j.dnarep.2010.12.002","article-title":"Two unrelated patients with MRE11A mutations and Nijmegen breakage syndrome-like severe microcephaly","volume":"10","author":"Matsumoto","year":"2011","journal-title":"DNA Repair (Amst)"},{"key":"2026071706515531800_ref15","doi-asserted-by":"publisher","first-page":"435","DOI":"10.1016\/j.jpeds.2009.02.037","article-title":"Two brothers with ataxia-telangiectasia-like disorder with lung adenocarcinoma","volume":"155","author":"Uchisaka","year":"2009","journal-title":"J Pediatr"},{"key":"2026071706515531800_ref16","doi-asserted-by":"publisher","first-page":"2155","DOI":"10.1093\/hmg\/ddh221","article-title":"MRE11 mutations and impaired ATM-dependent responses in an Italian family with ataxia-telangiectasia-like disorder","volume":"13","author":"Delia","year":"2004","journal-title":"Hum Mol Genet"},{"key":"2026071706515531800_ref17","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1016\/j.ajhg.2009.04.010","article-title":"Human RAD50 deficiency in a Nijmegen breakage syndrome-like disorder","volume":"84","author":"Waltes","year":"2009","journal-title":"Am J Hum Genet"},{"key":"2026071706515531800_ref18","doi-asserted-by":"publisher","first-page":"10121","DOI":"10.1038\/s41598-018-28370-5","article-title":"Reversible mislocalization of a disease-associated MRE11 splice variant product","volume":"8","author":"Hartlerode","year":"2018","journal-title":"Sci Rep"},{"key":"2026071706515531800_ref19","doi-asserted-by":"publisher","first-page":"1511","DOI":"10.1016\/S1097-2765(03)00455-6","article-title":"Checkpoint failure and chromosomal instability without lymphomagenesis in Mre11(ATLD1\/ATLD1) mice","volume":"12","author":"Theunissen","year":"2003","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref20","doi-asserted-by":"publisher","first-page":"9558","DOI":"10.1074\/jbc.M808906200","article-title":"Human CtIP mediates cell cycle control of DNA end resection and double strand break repair","volume":"284","author":"Huertas","year":"2009","journal-title":"J Biol Chem"},{"key":"2026071706515531800_ref21","doi-asserted-by":"publisher","first-page":"940","DOI":"10.1016\/j.molcel.2016.10.017","article-title":"Phosphorylated CtIP functions as a Co-factor of the MRE11-RAD50-NBS1 endonuclease in DNA end resection","volume":"64","author":"Anand","year":"2016","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref22","doi-asserted-by":"publisher","first-page":"2223","DOI":"10.1016\/j.molcel.2024.05.019","article-title":"Molecular insights into the activation of Mre11-Rad50 endonuclease activity by Sae2\/CtIP","volume":"84","author":"Nicolas","year":"2024","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref23","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abb2947","article-title":"Chromatin-associated MRN complex protects highly transcribing genes from genomic instability","volume":"7","author":"Salifou","year":"2021","journal-title":"Sci Adv"},{"key":"2026071706515531800_ref24","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1016\/j.cell.2023.11.022","article-title":"Metabolic regulation of homologous recombination repair by MRE11 lactylation","volume":"187","author":"Chen","year":"2024","journal-title":"Cell"},{"key":"2026071706515531800_ref25","doi-asserted-by":"publisher","first-page":"e2422720122","DOI":"10.1073\/pnas.2422720122","article-title":"ATM priming and end resection-coupled phosphorylation of MRE11 is important for fork protection and replication restart","volume":"122","author":"Zhang","year":"2025","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026071706515531800_ref26","doi-asserted-by":"publisher","first-page":"1605","DOI":"10.1093\/hmg\/ddae101","article-title":"Reduced levels of MRE11 cause disease phenotypes distinct from ataxia telangiectasia-like disorder","volume":"33","author":"Hartlerode","year":"2024","journal-title":"Hum Mol Genet"},{"key":"2026071706515531800_ref27","doi-asserted-by":"publisher","first-page":"701","DOI":"10.1001\/jama.2024.25349","article-title":"Essential Thrombocythemia: a review","volume":"333","author":"Tefferi","year":"2025","journal-title":"JAMA"},{"key":"2026071706515531800_ref28","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1186\/1750-1172-7-13","article-title":"Nijmegen breakage syndrome (NBS)","volume":"7","author":"Chrzanowska","year":"2012","journal-title":"Orphanet J Rare Dis"},{"key":"2026071706515531800_ref29","doi-asserted-by":"publisher","first-page":"101225","DOI":"10.1016\/j.blre.2024.101225","article-title":"Longitudinal clinical manifestations of Fanconi anemia: a systematized review","volume":"68","author":"Hoover","year":"2024","journal-title":"Blood Rev"},{"key":"2026071706515531800_ref30","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1038\/nri3570","article-title":"Orchestrating B cell lymphopoiesis through interplay of IL-7 receptor and pre-B cell receptor signalling","volume":"14","author":"Clark","year":"2014","journal-title":"Nat Rev Immunol"},{"key":"2026071706515531800_ref31","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.exphem.2021.10.002","article-title":"Hematopoietic stem and progenitor cells outside the bone marrow: where, when, and why","volume":"104","author":"Mende","year":"2021","journal-title":"Exp Hematol"},{"key":"2026071706515531800_ref32","doi-asserted-by":"publisher","DOI":"10.3390\/jcm10245831","article-title":"Extramedullary Hematopoiesis of the liver and spleen","volume":"10","author":"Cenariu","year":"2021","journal-title":"J Clin Med"},{"key":"2026071706515531800_ref33","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1038\/nsmb.2212","article-title":"Mre11 regulates CtIP-dependent double-strand break repair by interaction with CDK2","volume":"19","author":"Buis","year":"2012","journal-title":"Nat Struct Mol Biol"},{"key":"2026071706515531800_ref34","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1038\/nrm3546","article-title":"The ATM protein kinase: regulating the cellular response to genotoxic stress, and more","volume":"14","author":"Shiloh","year":"2013","journal-title":"Nat Rev Mol Cell Biol"},{"key":"2026071706515531800_ref35","doi-asserted-by":"publisher","first-page":"736","DOI":"10.1038\/nsmb.3072","article-title":"Recruitment and activation of the ATM kinase in the absence of DNA-damage sensors","volume":"22","author":"Hartlerode","year":"2015","journal-title":"Nat Struct Mol Biol"},{"key":"2026071706515531800_ref36","doi-asserted-by":"publisher","first-page":"3497","DOI":"10.1242\/jcs.111.23.3497","article-title":"Histone H3 phosphorylation is required for the initiation, but not maintenance, of mammalian chromosome condensation","volume":"111","author":"Van Hooser","year":"1998","journal-title":"J Cell Sci"},{"key":"2026071706515531800_ref37","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/j.cell.2008.08.015","article-title":"Mre11 nuclease activity has essential roles in DNA repair and genomic stability distinct from ATM activation","volume":"135","author":"Buis","year":"2008","journal-title":"Cell"},{"key":"2026071706515531800_ref38","doi-asserted-by":"publisher","first-page":"108559","DOI":"10.1016\/j.celrep.2020.108559","article-title":"A disease-causing single amino acid deletion in the coiled-coil domain of RAD50 impairs MRE11 complex functions in yeast and humans","volume":"33","author":"Chansel-Da Cruz","year":"2020","journal-title":"Cell Rep"},{"key":"2026071706515531800_ref39","doi-asserted-by":"publisher","first-page":"2136","DOI":"10.1007\/s10875-023-01591-8","article-title":"Bone marrow failure and immunodeficiency associated with human RAD50 variants","volume":"43","author":"Takagi","year":"2023","journal-title":"J Clin Immunol"},{"key":"2026071706515531800_ref40","doi-asserted-by":"publisher","first-page":"1013","DOI":"10.1016\/j.molcel.2020.11.040","article-title":"A surge of DNA damage links transcriptional reprogramming and hematopoietic deficit in Fanconi Anemia","volume":"80","author":"Shen","year":"2020","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref41","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1038\/s12276-024-01192-4","article-title":"Extramedullary hematopoiesis in cancer","volume":"56","author":"Barisas","year":"2024","journal-title":"Exp Mol Med"},{"key":"2026071706515531800_ref42","doi-asserted-by":"publisher","first-page":"1811","DOI":"10.1007\/s00439-022-02462-9","article-title":"Fanconi anemia: current insights regarding epidemiology, cancer, and DNA repair","volume":"141","author":"Peake","year":"2022","journal-title":"Hum Genet"},{"key":"2026071706515531800_ref43","doi-asserted-by":"publisher","first-page":"1962","DOI":"10.1126\/science.290.5498.1962","article-title":"Response to RAG-mediated VDJ cleavage by NBS1 and gamma-H2AX","volume":"290","author":"Chen","year":"2000","journal-title":"Science"},{"key":"2026071706515531800_ref44","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1101\/gad.956902","article-title":"Sensing of intermediates in V(D)J recombination by ATM","volume":"16","author":"Perkins","year":"2002","journal-title":"Genes Dev"},{"key":"2026071706515531800_ref45","doi-asserted-by":"publisher","first-page":"669","DOI":"10.1084\/jem.20081326","article-title":"MRN complex function in the repair of chromosomal rag-mediated DNA double-strand breaks","volume":"206","author":"Helmink","year":"2009","journal-title":"J Exp Med"},{"key":"2026071706515531800_ref46","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/j.molcel.2009.03.009","article-title":"Roles for NBS1 in alternative nonhomologous end-joining of V(D)J recombination intermediates","volume":"34","author":"Deriano","year":"2009","journal-title":"Mol Cell"},{"key":"2026071706515531800_ref47","doi-asserted-by":"publisher","first-page":"5572","DOI":"10.1128\/MCB.00917-10","article-title":"Dual functions of Nbs1 in the repair of DNA breaks and proliferation ensure proper V(D)J recombination and T-cell development","volume":"30","author":"Saidi","year":"2010","journal-title":"Mol Cell Biol"},{"key":"2026071706515531800_ref48","doi-asserted-by":"publisher","first-page":"6412","DOI":"10.1093\/emboj\/17.21.6412","article-title":"Distinct roles of two separable in vitro activities of yeast Mre11 in mitotic and meiotic recombination","volume":"17","author":"Furuse","year":"1998","journal-title":"EMBO J"},{"key":"2026071706515531800_ref49","doi-asserted-by":"publisher","first-page":"5759","DOI":"10.1038\/s41467-023-41544-8","article-title":"Genome-wide analysis of DNA-PK-bound MRN cleavage products supports a sequential model of DSB repair pathway choice","volume":"14","author":"Deshpande","year":"2023","journal-title":"Nat Commun"},{"key":"2026071706515531800_ref50","doi-asserted-by":"publisher","first-page":"5146","DOI":"10.1093\/hmg\/ddt368","article-title":"Disease-associated MRE11 mutants impact ATM\/ATR DNA damage signaling by distinct mechanisms","volume":"22","author":"Regal","year":"2013","journal-title":"Hum Mol Genet"},{"key":"2026071706515531800_ref51","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1007\/978-3-642-20792-1_6","volume-title":"Advanced Protocols for Animal Transgenesis: An ISTT Manual","author":"Becker","year":"2011"},{"key":"2026071706515531800_ref52","volume-title":"Manipulating the Mouse Embryo; A laboratory Manual","author":"Hogan","year":"1994"},{"key":"2026071706515531800_ref53","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1016\/S0092-8674(02)00747-X","article-title":"Convergence of the fanconi anemia and ataxia telangiectasia signaling pathways","volume":"109","author":"Taniguchi","year":"2002","journal-title":"Cell"}],"container-title":["Human Molecular Genetics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/hmg\/article-pdf\/34\/23\/1964\/64639847\/ddaf154.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/hmg\/article-pdf\/34\/23\/1964\/64639847\/ddaf154.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,17]],"date-time":"2026-07-17T10:52:02Z","timestamp":1784285522000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/hmg\/article\/34\/23\/1964\/8283586"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,11]]},"references-count":53,"journal-issue":{"issue":"23","published-print":{"date-parts":[[2025,11,18]]}},"URL":"https:\/\/doi.org\/10.1093\/hmg\/ddaf154","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.07.15.664809","asserted-by":"object"}]},"ISSN":["0964-6906","1460-2083"],"issn-type":[{"value":"0964-6906","type":"print"},{"value":"1460-2083","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2025,12,1]]},"published":{"date-parts":[[2025,10,11]]}}}