{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T06:48:28Z","timestamp":1790837308406,"version":"4.1.0"},"reference-count":43,"publisher":"Elsevier BV","issue":"10","license":[{"start":{"date-parts":[[2021,3,1]],"date-time":"2021-03-01T00:00:00Z","timestamp":1614556800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2021,3,1]],"date-time":"2021-03-01T00:00:00Z","timestamp":1614556800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2021,2,15]],"date-time":"2021-02-15T00:00:00Z","timestamp":1613347200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["1R01GM115472-01"],"award-info":[{"award-number":["1R01GM115472-01"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["1R01GM130889-01"],"award-info":[{"award-number":["1R01GM130889-01"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["5 T32 CA009009035-43"],"award-info":[{"award-number":["5 T32 CA009009035-43"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["POLQ\u2212\/\u2212"],"award-info":[{"award-number":["POLQ\u2212\/\u2212"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["POLQ\u2212\/\u2212 HCT116"],"award-info":[{"award-number":["POLQ\u2212\/\u2212 HCT116"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01CA186238"],"award-info":[{"award-number":["R01CA186238"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01CA244179"],"award-info":[{"award-number":["R01CA244179"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01GM57962"],"award-info":[{"award-number":["R01GM57962"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["TDP2\u2212\/\u2212"],"award-info":[{"award-number":["TDP2\u2212\/\u2212"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Cell Reports"],"published-print":{"date-parts":[[2021,3]]},"DOI":"10.1016\/j.celrep.2021.108820","type":"journal-article","created":{"date-parts":[[2021,3,9]],"date-time":"2021-03-09T20:58:29Z","timestamp":1615323509000},"page":"108820","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":27,"title":["Pol\u03b8 promotes the repair of 5\u2032-DNA-protein crosslinks by microhomology-mediated end-joining"],"prefix":"10.1016","volume":"34","author":[{"given":"Gurushankar","family":"Chandramouly","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuren","family":"Liao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Timur","family":"Rusanov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nikita","family":"Borisonnik","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marissa L.","family":"Calbert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tatiana","family":"Kent","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katherine","family":"Sullivan-Reed","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Umeshkumar","family":"Vekariya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ekaterina","family":"Kashkina","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tomasz","family":"Skorski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hong","family":"Yan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard T.","family":"Pomerantz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.celrep.2021.108820_bib1","doi-asserted-by":"crossref","first-page":"5743","DOI":"10.1093\/nar\/gkw326","article-title":"A role for human homologous recombination factors in suppressing microhomology-mediated end joining","volume":"44","author":"Ahrabi","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.celrep.2021.108820_bib2","doi-asserted-by":"crossref","first-page":"797","DOI":"10.1158\/0008-5472.CAN-13-1443","article-title":"Compensatory functions and interdependency of the DNA-binding domain of BRCA2 with the BRCA1-PALB2-BRCA2 complex","volume":"74","author":"Al Abo","year":"2014","journal-title":"Cancer Res."},{"key":"10.1016\/j.celrep.2021.108820_bib3","doi-asserted-by":"crossref","first-page":"e1169343","DOI":"10.1080\/23723556.2016.1169343","article-title":"BRCA1-CtIP interaction in the repair of DNA double-strand breaks","volume":"3","author":"Aparicio","year":"2016","journal-title":"Mol. Cell. Oncol."},{"key":"10.1016\/j.celrep.2021.108820_bib4","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1083\/jcb.201504005","article-title":"MRN, CtIP, and BRCA1 mediate repair of topoisomerase II-DNA adducts","volume":"212","author":"Aparicio","year":"2016","journal-title":"J.\u00a0Cell Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib5","doi-asserted-by":"crossref","first-page":"67","DOI":"10.3390\/genes7090067","article-title":"DNA Polymerase \u03b8: A Unique Multifunctional End-Joining Machine","volume":"7","author":"Black","year":"2016","journal-title":"Genes (Basel)"},{"key":"10.1016\/j.celrep.2021.108820_bib6","doi-asserted-by":"crossref","first-page":"4423","DOI":"10.1038\/s41467-019-12272-9","article-title":"Molecular basis of microhomology-mediated end-joining by purified full-length Pol\u03b8","volume":"10","author":"Black","year":"2019","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2021.108820_bib7","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1038\/nature13771","article-title":"Sae2 promotes dsDNA endonuclease activity within Mre11-Rad50-Xrs2 to resect DNA breaks","volume":"514","author":"Cannavo","year":"2014","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2021.108820_bib8","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1038\/nature14184","article-title":"Homologous-recombination-deficient tumours are dependent on Pol\u03b8-mediated repair","volume":"518","author":"Ceccaldi","year":"2015","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2021.108820_bib9","doi-asserted-by":"crossref","first-page":"e1001005","DOI":"10.1371\/journal.pgen.1001005","article-title":"Dual roles for DNA polymerase theta in alternative end-joining repair of double-strand breaks in Drosophila","volume":"6","author":"Chan","year":"2010","journal-title":"PLoS Genet."},{"key":"10.1016\/j.celrep.2021.108820_bib10","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1016\/j.molcel.2016.10.010","article-title":"Nbs1 Converts the Human Mre11\/Rad50 Nuclease Complex into an Endo\/Exonuclease Machine Specific for Protein-DNA Adducts","volume":"64","author":"Deshpande","year":"2016","journal-title":"Mol. Cell"},{"key":"10.1016\/j.celrep.2021.108820_bib11","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1083\/jcb.200506029","article-title":"Repair of double-strand breaks by nonhomologous end joining in the absence of Mre11","volume":"171","author":"Di Virgilio","year":"2005","journal-title":"J.\u00a0Cell Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib12","doi-asserted-by":"crossref","first-page":"30842","DOI":"10.1074\/jbc.M112.393983","article-title":"Biochemical characterization of human tyrosyl-DNA phosphodiesterase 2 (TDP2\/TTRAP): a Mg(2+)\/Mn(2+)-dependent phosphodiesterase specific for the repair of topoisomerase cleavage complexes","volume":"287","author":"Gao","year":"2012","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.celrep.2021.108820_bib13","doi-asserted-by":"crossref","first-page":"e1003226","DOI":"10.1371\/journal.pgen.1003226","article-title":"TDP2-dependent non-homologous end-joining protects against topoisomerase II-induced DNA breaks and genome instability in cells and in\u00a0vivo","volume":"9","author":"G\u00f3mez-Herreros","year":"2013","journal-title":"PLoS Genet."},{"key":"10.1016\/j.celrep.2021.108820_bib14","doi-asserted-by":"crossref","first-page":"4856","DOI":"10.1038\/s41467-019-12829-8","article-title":"Genome-wide microhomologies enable precise template-free editing of biologically relevant deletion mutations","volume":"10","author":"Grajcarek","year":"2019","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2021.108820_bib15","doi-asserted-by":"crossref","first-page":"2984","DOI":"10.1158\/0008-5472.CAN-09-4040","article-title":"A small interfering RNA screen of genes involved in DNA repair identifies tumor-specific radiosensitization by POLQ knockdown","volume":"70","author":"Higgins","year":"2010","journal-title":"Cancer Res."},{"key":"10.1016\/j.celrep.2021.108820_bib16","doi-asserted-by":"crossref","first-page":"580","DOI":"10.1016\/j.molcel.2016.10.011","article-title":"Mre11 Is Essential for the Removal of Lethal Topoisomerase 2 Covalent Cleavage Complexes","volume":"64","author":"Hoa","year":"2016","journal-title":"Mol. Cell"},{"key":"10.1016\/j.celrep.2021.108820_bib17","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.dnarep.2018.10.002","article-title":"An RNAi screen in human cell lines reveals conserved DNA damage repair pathways that mitigate formaldehyde sensitivity","volume":"72","author":"Juarez","year":"2018","journal-title":"DNA Repair (Amst.)"},{"key":"10.1016\/j.celrep.2021.108820_bib18","doi-asserted-by":"crossref","first-page":"3615","DOI":"10.1038\/s41467-020-17455-3","article-title":"BRCA1-associated structural variations are a consequence of polymerase theta-mediated end-joining","volume":"11","author":"Kamp","year":"2020","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2021.108820_bib19","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1038\/nsmb.2961","article-title":"Mechanism of microhomology-mediated end-joining promoted by human DNA polymerase \u03b8","volume":"22","author":"Kent","year":"2015","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib20","doi-asserted-by":"crossref","first-page":"e13740","DOI":"10.7554\/eLife.13740","article-title":"Polymerase \u03b8 is a robust terminal transferase that oscillates between three different mechanisms during end-joining","volume":"5","author":"Kent","year":"2016","journal-title":"eLife"},{"key":"10.1016\/j.celrep.2021.108820_bib21","doi-asserted-by":"crossref","first-page":"2585","DOI":"10.1128\/MCB.19.4.2585","article-title":"Ku-dependent nonhomologous DNA end joining in Xenopus egg extracts","volume":"19","author":"Labhart","year":"1999","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib22","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1016\/j.molcel.2018.11.024","article-title":"Replication-Coupled DNA-Protein Crosslink Repair by SPRTN and the Proteasome in Xenopus Egg Extracts","volume":"73","author":"Larsen","year":"2019","journal-title":"Mol. Cell"},{"key":"10.1016\/j.celrep.2021.108820_bib23","first-page":"e134","article-title":"Enriching CRISPR-Cas9 targeted cells by co-targeting the HPRT gene","volume":"43","author":"Liao","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.celrep.2021.108820_bib24","doi-asserted-by":"crossref","first-page":"5689","DOI":"10.1093\/nar\/gkw274","article-title":"The structure of ends determines the pathway choice and Mre11 nuclease dependency of DNA double-strand break repair","volume":"44","author":"Liao","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.celrep.2021.108820_bib25","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1016\/j.sbi.2018.03.019","article-title":"Terminal deoxynucleotidyltransferase: the story of an untemplated DNA polymerase capable of DNA bridging and templated synthesis across strands","volume":"53","author":"Loc\u2019h","year":"2018","journal-title":"Curr. Opin. Struct. Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib26","doi-asserted-by":"crossref","first-page":"1256","DOI":"10.1073\/pnas.1521597113","article-title":"Ligase I and ligase III mediate the DNA double-strand break ligation in alternative end-joining","volume":"113","author":"Lu","year":"2016","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.celrep.2021.108820_bib27","doi-asserted-by":"crossref","first-page":"1022","DOI":"10.1016\/j.molcel.2014.04.011","article-title":"Catalytic and noncatalytic roles of the CtIP endonuclease in double-strand break end resection","volume":"54","author":"Makharashvili","year":"2014","journal-title":"Mol. Cell"},{"key":"10.1016\/j.celrep.2021.108820_bib28","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1038\/nature14157","article-title":"Mammalian polymerase \u03b8 promotes alternative NHEJ and suppresses recombination","volume":"518","author":"Mateos-Gomez","year":"2015","journal-title":"Nature"},{"key":"10.1016\/j.celrep.2021.108820_bib29","doi-asserted-by":"crossref","first-page":"1116","DOI":"10.1038\/nsmb.3494","article-title":"The helicase domain of Pol\u03b8 counteracts RPA to promote alt-NHEJ","volume":"24","author":"Mateos-Gomez","year":"2017","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib30","doi-asserted-by":"crossref","first-page":"e1000828","DOI":"10.1371\/journal.pgen.1000828","article-title":"Collaborative action of Brca1 and CtIP in elimination of covalent modifications from double-strand breaks to facilitate subsequent break repair","volume":"6","author":"Nakamura","year":"2010","journal-title":"PLoS Genet."},{"key":"10.1016\/j.celrep.2021.108820_bib31","doi-asserted-by":"crossref","first-page":"27065","DOI":"10.1074\/jbc.M109.019174","article-title":"Homologous recombination but not nucleotide excision repair plays a pivotal role in tolerance of DNA-protein cross-links in mammalian cells","volume":"284","author":"Nakano","year":"2009","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.celrep.2021.108820_bib32","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1016\/j.dnarep.2014.03.020","article-title":"Tyrosyl-DNA-phosphodiesterases (TDP1 and TDP2)","volume":"19","author":"Pommier","year":"2014","journal-title":"DNA Repair (Amst.)"},{"key":"10.1016\/j.celrep.2021.108820_bib33","doi-asserted-by":"crossref","first-page":"3634","DOI":"10.15252\/embj.201796948","article-title":"Mutational signatures of non-homologous and polymerase theta-mediated end-joining in embryonic stem cells","volume":"36","author":"Schimmel","year":"2017","journal-title":"EMBO J."},{"key":"10.1016\/j.celrep.2021.108820_bib34","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1016\/j.tibs.2015.08.006","article-title":"Microhomology-Mediated End Joining: A Back-up Survival Mechanism or Dedicated Pathway?","volume":"40","author":"Sfeir","year":"2015","journal-title":"Trends Biochem. Sci."},{"key":"10.1016\/j.celrep.2021.108820_bib35","doi-asserted-by":"crossref","first-page":"1372","DOI":"10.1038\/nsmb.2423","article-title":"Structural basis for recognition of 5\u2032-phosphotyrosine adducts by Tdp2","volume":"19","author":"Shi","year":"2012","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib36","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1038\/nrm.2017.56","article-title":"Mechanisms of DNA-protein crosslink repair","volume":"18","author":"Stingele","year":"2017","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.celrep.2021.108820_bib37","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1146\/annurev-biochem-062917-012415","article-title":"The MRE11-RAD50-NBS1 Complex Conducts the Orchestration of Damage Signaling and Outcomes to Stress in DNA Replication and Repair","volume":"87","author":"Syed","year":"2018","journal-title":"Annu. Rev. Biochem."},{"key":"10.1016\/j.celrep.2021.108820_bib38","doi-asserted-by":"crossref","first-page":"7078","DOI":"10.1158\/0008-5472.CAN-07-0601","article-title":"BRCA1- and BRCA2-deficient cells are sensitive to etoposide-induced DNA double-strand breaks via topoisomerase II","volume":"67","author":"Treszezamsky","year":"2007","journal-title":"Cancer Res."},{"key":"10.1016\/j.celrep.2021.108820_bib39","doi-asserted-by":"crossref","first-page":"7720","DOI":"10.1073\/pnas.1213431110","article-title":"Microhomology-mediated End Joining and Homologous Recombination share the initial end resection step to repair DNA double-strand breaks in mammalian cells","volume":"110","author":"Truong","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.celrep.2021.108820_bib40","doi-asserted-by":"crossref","first-page":"7394","DOI":"10.1038\/ncomms8394","article-title":"Polymerase \u0398 is a key driver of genome evolution and of CRISPR\/Cas9-mediated mutagenesis","volume":"6","author":"van Schendel","year":"2015","journal-title":"Nat. Commun."},{"key":"10.1016\/j.celrep.2021.108820_bib41","doi-asserted-by":"crossref","first-page":"3909","DOI":"10.1074\/jbc.RA118.005188","article-title":"DNA polymerase \u03b8 (POLQ) is important for repair of DNA double-strand breaks caused by fork collapse","volume":"294","author":"Wang","year":"2019","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.celrep.2021.108820_bib42","doi-asserted-by":"crossref","first-page":"e1004654","DOI":"10.1371\/journal.pgen.1004654","article-title":"Mechanism of suppression of chromosomal instability by DNA polymerase POLQ","volume":"10","author":"Yousefzadeh","year":"2014","journal-title":"PLoS Genet."},{"key":"10.1016\/j.celrep.2021.108820_bib43","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1038\/nsmb.1940","article-title":"An essential role for CtIP in chromosomal translocation formation through an alternative end-joining pathway","volume":"18","author":"Zhang","year":"2011","journal-title":"Nat. Struct. Mol. Biol."}],"container-title":["Cell Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2211124721001340?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S2211124721001340?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,10,31]],"date-time":"2025-10-31T11:14:20Z","timestamp":1761909260000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S2211124721001340"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,3]]},"references-count":43,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2021,3]]}},"alternative-id":["S2211124721001340"],"URL":"https:\/\/doi.org\/10.1016\/j.celrep.2021.108820","relation":{},"ISSN":["2211-1247"],"issn-type":[{"value":"2211-1247","type":"print"}],"subject":[],"published":{"date-parts":[[2021,3]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Pol\u03b8 promotes the repair of 5\u2032-DNA-protein crosslinks by microhomology-mediated end-joining","name":"articletitle","label":"Article Title"},{"value":"Cell Reports","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.celrep.2021.108820","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2021 The Authors.","name":"copyright","label":"Copyright"}],"article-number":"108820"}}