{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,22]],"date-time":"2026-02-22T19:01:52Z","timestamp":1771786912803,"version":"3.50.1"},"reference-count":65,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,2,3]],"date-time":"2026-02-03T00:00:00Z","timestamp":1770076800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,2,19]],"date-time":"2026-02-19T00:00:00Z","timestamp":1771459200000},"content-version":"vor","delay-in-days":16,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-026-38338-5","type":"journal-article","created":{"date-parts":[[2026,2,3]],"date-time":"2026-02-03T22:03:09Z","timestamp":1770156189000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["DGCR8 regulates multiple processes of transcription coupled nucleotide excision repair"],"prefix":"10.1038","volume":"16","author":[{"given":"Takaaki","family":"Watanabe","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Daiyu","family":"Yoshinami","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hiroyuki","family":"Yamasaki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yukiko","family":"Tanaka","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Masato","family":"Ohtsuka","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Toshiyasu","family":"Taniguchi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2026,2,3]]},"reference":[{"key":"38338_CR1","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1016\/j.molcel.2008.01.001","volume":"29","author":"P Herrlich","year":"2008","unstructured":"Herrlich, P., Karin, M. & Weiss, C. Supreme enlightenment: damage recognition and signaling in the mammalian UV response. Mol. Cell. 29, 279\u2013290 (2008).","journal-title":"Mol. Cell."},{"key":"38338_CR2","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1146\/annurev-biochem-052621-091205","volume":"92","author":"N Nieto Moreno","year":"2023","unstructured":"Nieto Moreno, N., Olthof, A. M. & Svejstrup, J. Q. Transcription-Coupled nucleotide excision repair and the transcriptional response to UV-Induced DNA damage. Annu. Rev. Biochem. 92, 81\u2013113 (2023).","journal-title":"Annu. Rev. Biochem."},{"key":"38338_CR3","doi-asserted-by":"publisher","first-page":"958","DOI":"10.1038\/nrm2549","volume":"9","author":"PC Hanawalt","year":"2008","unstructured":"Hanawalt, P. C. & Spivak, G. Transcription-coupled DNA repair: two decades of progress and surprises. Nat. Rev. Mol. Cell. Biol. 9, 958\u2013970 (2008).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR4","doi-asserted-by":"publisher","first-page":"766","DOI":"10.1038\/s41580-019-0169-4","volume":"20","author":"H Lans","year":"2019","unstructured":"Lans, H., Hoeijmakers, J. H. J., Vermeulen, W. & Marteijn, J. A. The DNA damage response to transcription stress. Nat. Rev. Mol. Cell. Biol. 20, 766\u2013784 (2019).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR5","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1038\/nrm3822","volume":"15","author":"JA Marteijn","year":"2014","unstructured":"Marteijn, J. A., Lans, H., Vermeulen, W. & Hoeijmakers, J. H. J. Understanding nucleotide excision repair and its roles in cancer and ageing. Nat. Rev. Mol. Cell. Biol. 15, 465\u2013481 (2014).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR6","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1016\/j.celrep.2017.03.021","volume":"19","author":"PC Calses","year":"2017","unstructured":"Calses, P. C. et al. DGCR8 mediates repair of UV-Induced DNA damage independently of RNA processing. Cell. Rep. 19, 162\u2013174 (2017).","journal-title":"Cell. Rep."},{"key":"38338_CR7","doi-asserted-by":"publisher","first-page":"838","DOI":"10.1042\/BST20130011","volume":"41","author":"S Macias","year":"2013","unstructured":"Macias, S., Cordiner, R. A. & C\u00e1ceres, J. F. Cellular functions of the microprocessor. Biochem. Soc. Trans. 41, 838\u2013843 (2013).","journal-title":"Biochem. Soc. Trans."},{"key":"38338_CR8","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1016\/j.cell.2009.01.035","volume":"136","author":"RW Carthew","year":"2009","unstructured":"Carthew, R. W. & Sontheimer, E. J. Origins and mechanisms of MiRNAs and SiRNAs. Cell 136, 642\u2013655 (2009).","journal-title":"Cell"},{"key":"38338_CR9","doi-asserted-by":"publisher","first-page":"1100","DOI":"10.1158\/1541-7786.MCR-11-0007","volume":"9","author":"Y Wang","year":"2011","unstructured":"Wang, Y. et al. MicroRNA-138 modulates DNA damage response by repressing histone H2AX expression. Mol. Cancer Res. MCR. 9, 1100\u20131111 (2011).","journal-title":"Mol. Cancer Res. MCR"},{"key":"38338_CR10","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1038\/s41580-018-0059-1","volume":"20","author":"T Treiber","year":"2019","unstructured":"Treiber, T., Treiber, N. & Meister, G. Regulation of MicroRNA biogenesis and its crosstalk with other cellular pathways. Nat. Rev. Mol. Cell. Biol. 20, 5\u201320 (2019).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR11","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1002\/emmm.201100209","volume":"4","author":"MV Iorio","year":"2012","unstructured":"Iorio, M. V. & Croce, C. M. MicroRNA dysregulation in cancer: diagnostics, monitoring and therapeutics. A comprehensive review. EMBO Mol. Med. 4, 143\u2013159 (2012).","journal-title":"EMBO Mol. Med."},{"key":"38338_CR12","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1016\/j.tig.2023.01.009","volume":"39","author":"D Pelletier","year":"2023","unstructured":"Pelletier, D., Rivera, B., Fabian, M. R. & Foulkes, W. D. MiRNA biogenesis and inherited disorders: clinico-molecular insights. Trends Genet. TIG. 39, 401\u2013414 (2023).","journal-title":"Trends Genet. TIG"},{"key":"38338_CR13","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1038\/s41580-021-00354-w","volume":"22","author":"P Agbu","year":"2021","unstructured":"Agbu, P. & Carthew, R. W. MicroRNA-mediated regulation of glucose and lipid metabolism. Nat. Rev. Mol. Cell. Biol. 22, 425\u2013438 (2021).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR14","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1038\/nrg.2016.134","volume":"17","author":"CP Bracken","year":"2016","unstructured":"Bracken, C. P., Scott, H. S. & Goodall, G. J. A network-biology perspective of MicroRNA function and dysfunction in cancer. Nat. Rev. Genet. 17, 719\u2013732 (2016).","journal-title":"Nat. Rev. Genet."},{"key":"38338_CR15","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1038\/nrc3932","volume":"15","author":"S Lin","year":"2015","unstructured":"Lin, S. & Gregory, R. I. MicroRNA biogenesis pathways in cancer. Nat. Rev. Cancer. 15, 321\u2013333 (2015).","journal-title":"Nat. Rev. Cancer"},{"key":"38338_CR16","doi-asserted-by":"publisher","first-page":"2090","DOI":"10.1038\/emboj.2009.156","volume":"28","author":"J Pothof","year":"2009","unstructured":"Pothof, J. et al. MicroRNA-mediated gene Silencing modulates the UV-induced DNA-damage response. EMBO J. 28, 2090\u20132099 (2009).","journal-title":"EMBO J."},{"key":"38338_CR17","doi-asserted-by":"publisher","first-page":"2686","DOI":"10.1038\/jid.2010.169","volume":"130","author":"P Dziunycz","year":"2010","unstructured":"Dziunycz, P. et al. Squamous cell carcinoma of the skin shows a distinct MicroRNA profile modulated by UV radiation. J. Invest. Dermatol. 130, 2686\u20132689 (2010).","journal-title":"J. Invest. Dermatol."},{"key":"38338_CR18","doi-asserted-by":"publisher","first-page":"2973","DOI":"10.1002\/1873-3468.13196","volume":"592","author":"SK Pong","year":"2018","unstructured":"Pong, S. K. & Gullerova, M. Noncanonical functions of MicroRNA pathway enzymes - Drosha, DGCR8, Dicer and ago proteins. FEBS Lett. 592, 2973\u20132986 (2018).","journal-title":"FEBS Lett."},{"key":"38338_CR19","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1038\/nrm3523","volume":"14","author":"D Chowdhury","year":"2013","unstructured":"Chowdhury, D., Choi, Y. E. & Brault, M. E. Charity begins at home: non-coding RNA functions in DNA repair. Nat. Rev. Mol. Cell. Biol. 14, 181\u2013189 (2013).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR20","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1038\/s41580-022-00474-x","volume":"23","author":"E Petermann","year":"2022","unstructured":"Petermann, E., Lan, L. & Zou, L. Sources, resolution and physiological relevance of R-loops and RNA\u2013DNA hybrids. Nat. Rev. Mol. Cell. Biol. 23, 521\u2013540 (2022).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR21","first-page":"504","volume":"38","author":"R Luna","year":"2024","unstructured":"Luna, R., G\u00f3mez-Gonz\u00e1lez, B. & Aguilera, A. RNA biogenesis and RNA metabolism factors as R-loop suppressors: a hidden role in genome integrity. Genes Dev. 38, 504\u2013527 (2024).","journal-title":"Genes Dev."},{"key":"38338_CR22","doi-asserted-by":"publisher","first-page":"2267","DOI":"10.1016\/j.molcel.2022.04.014","volume":"82","author":"JR Brickner","year":"2022","unstructured":"Brickner, J. R., Garzon, J. L. & Cimprich, K. A. Walking a tightrope: the complex balancing act of R-loops in genome stability. Mol. Cell. 82, 2267\u20132297 (2022).","journal-title":"Mol. Cell."},{"key":"38338_CR23","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1038\/s41556-021-00663-4","volume":"23","author":"A Marnef","year":"2021","unstructured":"Marnef, A. & Legube, G. R-loops as Janus-faced modulators of DNA repair. Nat. Cell. Biol. 23, 305\u2013313 (2021).","journal-title":"Nat. Cell. Biol."},{"key":"38338_CR24","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1016\/j.cell.2019.08.055","volume":"179","author":"T Garc\u00eda-Muse","year":"2019","unstructured":"Garc\u00eda-Muse, T., Aguilera, A. R. & Loops From physiological to pathological roles. Cell 179, 604\u2013618 (2019).","journal-title":"Cell"},{"key":"38338_CR25","doi-asserted-by":"publisher","first-page":"398","DOI":"10.1016\/j.molcel.2019.01.024","volume":"73","author":"MP Crossley","year":"2019","unstructured":"Crossley, M. P., Bocek, M. & Cimprich, K. A. R-Loops as cellular regulators and genomic threats. Mol. Cell. 73, 398\u2013411 (2019).","journal-title":"Mol. Cell."},{"key":"38338_CR26","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1038\/s41580-019-0206-3","volume":"21","author":"C Niehrs","year":"2020","unstructured":"Niehrs, C. & Luke, B. Regulatory R-loops as facilitators of gene expression and genome stability. Nat. Rev. Mol. Cell. Biol. 21, 167\u2013178 (2020).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR27","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1038\/s41580-021-00382-6","volume":"22","author":"I Malik","year":"2021","unstructured":"Malik, I., Kelley, C. P., Wang, E. T. & Todd, P. K. Molecular mechanisms underlying nucleotide repeat expansion disorders. Nat. Rev. Mol. Cell. Biol. 22, 589\u2013607 (2021).","journal-title":"Nat. Rev. Mol. Cell. Biol."},{"key":"38338_CR28","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1016\/j.trecan.2019.08.006","volume":"5","author":"JP Wells","year":"2019","unstructured":"Wells, J. P., White, J. & Stirling, P. C. R loops and their composite cancer connections. Trends Cancer. 5, 619\u2013631 (2019).","journal-title":"Trends Cancer"},{"key":"38338_CR29","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1038\/s41588-019-0549-x","volume":"52","author":"A Abakir","year":"2020","unstructured":"Abakir, A. et al. N6-methyladenosine regulates the stability of RNA:DNA hybrids in human cells. Nat. Genet. 52, 48\u201355 (2020).","journal-title":"Nat. Genet."},{"key":"38338_CR30","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1016\/j.molcel.2020.06.017","volume":"79","author":"C Zhang","year":"2020","unstructured":"Zhang, C. et al. METTL3 and N6-Methyladenosine promote homologous Recombination-Mediated repair of DSBs by modulating DNA-RNA hybrid accumulation. Mol. Cell. 79, 425\u2013442e7 (2020).","journal-title":"Mol. Cell."},{"key":"38338_CR31","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1038\/nature14512","volume":"523","author":"M Tresini","year":"2015","unstructured":"Tresini, M. et al. The core spliceosome as target and effector of non-canonical ATM signalling. Nature 523, 53\u201358 (2015).","journal-title":"Nature"},{"key":"38338_CR32","doi-asserted-by":"publisher","first-page":"801","DOI":"10.1016\/j.molcel.2017.05.015","volume":"66","author":"AN Blackford","year":"2017","unstructured":"Blackford, A. N., Jackson, S. P., ATM, ATR & The trinity at the heart of the DNA damage response. Mol. Cell. 66, 801\u2013817 (2017).","journal-title":"Mol. Cell."},{"key":"38338_CR33","doi-asserted-by":"publisher","first-page":"8473","DOI":"10.1093\/nar\/gku565","volume":"42","author":"\u00d6Z Aydin","year":"2014","unstructured":"Aydin, \u00d6. Z. et al. Human ISWI complexes are targeted by SMARCA5 ATPase and SLIDE domains to help resolve lesion-stalled transcription. Nucleic Acids Res. 42, 8473\u20138485 (2014).","journal-title":"Nucleic Acids Res."},{"key":"38338_CR34","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1016\/j.molcel.2013.08.007","volume":"51","author":"C Dinant","year":"2013","unstructured":"Dinant, C. et al. Enhanced chromatin dynamics by FACT promotes transcriptional restart after UV-induced DNA damage. Mol. Cell. 51, 469\u2013479 (2013).","journal-title":"Mol. Cell."},{"key":"38338_CR35","doi-asserted-by":"publisher","first-page":"4011","DOI":"10.1093\/nar\/gkz055","volume":"47","author":"F Wienholz","year":"2019","unstructured":"Wienholz, F. et al. FACT subunit Spt16 controls UVSSA recruitment to lesion-stalled RNA pol II and stimulates TC-NER. Nucleic Acids Res. 47, 4011\u20134025 (2019).","journal-title":"Nucleic Acids Res."},{"key":"38338_CR36","doi-asserted-by":"publisher","first-page":"e1003407","DOI":"10.1371\/journal.pgen.1003407","volume":"9","author":"I Cho","year":"2013","unstructured":"Cho, I., Tsai, P. F., Lake, R. J., Basheer, A. & Fan, H. Y. ATP-dependent chromatin remodeling by Cockayne syndrome protein B and NAP1-like histone chaperones is required for efficient transcription-coupled DNA repair. PLoS Genet. 9, e1003407 (2013).","journal-title":"PLoS Genet."},{"key":"38338_CR37","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1093\/nar\/gkaa1162","volume":"49","author":"HJ Kang","year":"2021","unstructured":"Kang, H. J. et al. TonEBP recognizes R-loops and initiates m6A RNA methylation for R-loop resolution. Nucleic Acids Res. 49, 269\u2013284 (2021).","journal-title":"Nucleic Acids Res."},{"key":"38338_CR38","doi-asserted-by":"publisher","first-page":"3123","DOI":"10.1016\/j.celrep.2017.09.010","volume":"20","author":"VA Church","year":"2017","unstructured":"Church, V. A. et al. Microprocessor recruitment to elongating RNA polymerase II is required for differential expression of MicroRNAs. Cell. Rep. 20, 3123\u20133134 (2017).","journal-title":"Cell. Rep."},{"key":"38338_CR39","doi-asserted-by":"publisher","first-page":"2104","DOI":"10.1038\/s41467-020-15903-8","volume":"11","author":"Y van der Weegen","year":"2020","unstructured":"van der Weegen, Y. et al. The cooperative action of CSB, CSA, and UVSSA target TFIIH to DNA damage-stalled RNA polymerase II. Nat. Commun. 11, 2104 (2020).","journal-title":"Nat. Commun."},{"key":"38338_CR40","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1016\/S0092-8674(03)00316-7","volume":"113","author":"R Groisman","year":"2003","unstructured":"Groisman, R. et al. The ubiquitin ligase activity in the DDB2 and CSA complexes is differentially regulated by the COP9 signalosome in response to DNA damage. Cell 113, 357\u2013367 (2003).","journal-title":"Cell"},{"key":"38338_CR41","doi-asserted-by":"publisher","first-page":"598","DOI":"10.1038\/nature17416","volume":"531","author":"S Cavadini","year":"2016","unstructured":"Cavadini, S. et al. Cullin-RING ubiquitin E3 ligase regulation by the COP9 signalosome. Nature 531, 598\u2013603 (2016).","journal-title":"Nature"},{"key":"38338_CR42","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1038\/ng.2228","volume":"44","author":"X Zhang","year":"2012","unstructured":"Zhang, X. et al. Mutations in UVSSA cause UV-sensitive syndrome and destabilize ERCC6 in transcription-coupled DNA repair. Nat. Genet. 44, 593\u2013597 (2012).","journal-title":"Nat. Genet."},{"key":"38338_CR43","doi-asserted-by":"publisher","first-page":"598","DOI":"10.1038\/ng.2230","volume":"44","author":"P Schwertman","year":"2012","unstructured":"Schwertman, P. et al. UV-sensitive syndrome protein UVSSA recruits USP7 to regulate transcription-coupled repair. Nat. Genet. 44, 598\u2013602 (2012).","journal-title":"Nat. Genet."},{"key":"38338_CR44","doi-asserted-by":"publisher","first-page":"586","DOI":"10.1038\/ng.2229","volume":"44","author":"Y Nakazawa","year":"2012","unstructured":"Nakazawa, Y. et al. Mutations in UVSSA cause UV-sensitive syndrome and impair RNA polymerase IIo processing in transcription-coupled nucleotide-excision repair. Nat. Genet. 44, 586\u2013592 (2012).","journal-title":"Nat. Genet."},{"key":"38338_CR45","doi-asserted-by":"publisher","first-page":"1994","DOI":"10.1016\/j.celrep.2014.05.013","volume":"7","author":"J Quick-Cleveland","year":"2014","unstructured":"Quick-Cleveland, J. et al. The DGCR8 RNA-binding Heme domain recognizes primary MicroRNAs by clamping the hairpin. Cell. Rep. 7, 1994\u20132005 (2014).","journal-title":"Cell. Rep."},{"key":"38338_CR46","doi-asserted-by":"publisher","first-page":"1737","DOI":"10.1038\/s41467-017-01713-y","volume":"8","author":"AC Partin","year":"2017","unstructured":"Partin, A. C. et al. Heme enables proper positioning of drosha and DGCR8 on primary MicroRNAs. Nat. Commun. 8, 1737 (2017).","journal-title":"Nat. Commun."},{"key":"38338_CR47","doi-asserted-by":"publisher","first-page":"eabd2639","DOI":"10.1126\/scisignal.abd2639","volume":"14","author":"X Jiang","year":"2021","unstructured":"Jiang, X. et al. Control of ribosomal protein synthesis by the microprocessor complex. Sci. Signal. 14, eabd2639 (2021).","journal-title":"Sci. Signal."},{"key":"38338_CR48","doi-asserted-by":"publisher","first-page":"721","DOI":"10.1016\/j.molcel.2015.01.004","volume":"57","author":"Q Yang","year":"2015","unstructured":"Yang, Q. et al. Stress induces p38 MAPK-mediated phosphorylation and Inhibition of Drosha-dependent cell survival. Mol. Cell. 57, 721\u2013734 (2015).","journal-title":"Mol. Cell."},{"key":"38338_CR49","doi-asserted-by":"publisher","first-page":"777","DOI":"10.1016\/j.molcel.2014.10.020","volume":"56","author":"J Sollier","year":"2014","unstructured":"Sollier, J. et al. Transcription-coupled nucleotide excision repair factors promote R-loop-induced genome instability. Mol. Cell. 56, 777\u2013785 (2014).","journal-title":"Mol. Cell."},{"key":"38338_CR50","doi-asserted-by":"publisher","first-page":"1347","DOI":"10.1128\/jb.178.5.1347-1350.1996","volume":"178","author":"DR Koehler","year":"1996","unstructured":"Koehler, D. R., Courcelle, J. & Hanawalt, P. C. Kinetics of pyrimidine(6\u2009\u2013\u20094)pyrimidone photoproduct repair in Escherichia coli. J. Bacteriol. 178, 1347\u20131350 (1996).","journal-title":"J. Bacteriol."},{"key":"38338_CR51","doi-asserted-by":"publisher","first-page":"6015","DOI":"10.1093\/nar\/gkz359","volume":"47","author":"D Paul","year":"2019","unstructured":"Paul, D. et al. Structure and mechanism of pyrimidine-pyrimidone (6\u2009\u2013\u20094) photoproduct recognition by the Rad4\/XPC nucleotide excision repair complex. Nucleic Acids Res. 47, 6015\u20136028 (2019).","journal-title":"Nucleic Acids Res."},{"key":"38338_CR52","doi-asserted-by":"crossref","unstructured":"Adebali, O., Sancar, A. & Selby, C. P. Dynamics of transcription-coupled repair of cyclobutane pyrimidine dimers and (6\u2009\u2013\u20094) photoproducts in Escherichia coli. Proc. Natl. Acad. Sci. U. S. A. 121, e2416877121 (2024).","DOI":"10.1073\/pnas.2416877121"},{"key":"38338_CR53","doi-asserted-by":"publisher","first-page":"12806","DOI":"10.1073\/pnas.1917196117","volume":"117","author":"KF Hung","year":"2020","unstructured":"Hung, K. F., Sidorova, J. M., Nghiem, P. & Kawasumi, M. The 6\u2009\u2013\u20094 photoproduct is the trigger of UV-induced replication blockage and ATR activation. Proc. Natl. Acad. Sci. U S A. 117, 12806\u201312816 (2020).","journal-title":"Proc. Natl. Acad. Sci. U S A"},{"key":"38338_CR54","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1038\/nature08199","volume":"460","author":"HI Suzuki","year":"2009","unstructured":"Suzuki, H. I. et al. Modulation of MicroRNA processing by p53. Nature 460, 529\u2013533 (2009).","journal-title":"Nature"},{"key":"38338_CR55","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1038\/ng1303","volume":"36","author":"MC Moreira","year":"2004","unstructured":"Moreira, M. C. et al. Senataxin, the ortholog of a yeast RNA helicase, is mutant in ataxia-ocular apraxia 2. Nat. Genet. 36, 225\u2013227 (2004).","journal-title":"Nat. Genet."},{"key":"38338_CR56","doi-asserted-by":"publisher","first-page":"1128","DOI":"10.1086\/421054","volume":"74","author":"YZ Chen","year":"2004","unstructured":"Chen, Y. Z. et al. DNA\/RNA helicase gene mutations in a form of juvenile amyotrophic lateral sclerosis (ALS4). Am. J. Hum. Genet. 74, 1128\u20131135 (2004).","journal-title":"Am. J. Hum. Genet."},{"key":"38338_CR57","doi-asserted-by":"publisher","first-page":"113779","DOI":"10.1016\/j.celrep.2024.113779","volume":"43","author":"J Zhang","year":"2024","unstructured":"Zhang, J. et al. The ARID1A-METTL3-m6A axis ensures effective RNase H1-mediated resolution of R-loops and genome stability. Cell. Rep. 43, 113779 (2024).","journal-title":"Cell. Rep."},{"key":"38338_CR58","doi-asserted-by":"publisher","first-page":"103697","DOI":"10.1016\/j.dnarep.2024.103697","volume":"140","author":"SY Choi","year":"2024","unstructured":"Choi, S. Y. The roles of TonEBP in the DNA damage response: from DNA damage bypass to R-loop resolution. DNA Repair. 140, 103697 (2024).","journal-title":"DNA Repair."},{"key":"38338_CR59","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1038\/s41573-022-00558-5","volume":"22","author":"AABA da Costa","year":"2023","unstructured":"da Costa, A. A. B. A., Chowdhury, D., Shapiro, G. I., D\u2019Andrea, A. D. & Konstantinopoulos, P. A. Targeting replication stress in cancer therapy. Nat. Rev. Drug Discov. 22, 38\u201358 (2023).","journal-title":"Nat. Rev. Drug Discov"},{"key":"38338_CR60","doi-asserted-by":"publisher","first-page":"4039","DOI":"10.1038\/ncomms5039","volume":"5","author":"GT Torrezan","year":"2014","unstructured":"Torrezan, G. T. et al. Recurrent somatic mutation in DROSHA induces MicroRNA profile changes in Wilms tumour. Nat. Commun. 5, 4039 (2014).","journal-title":"Nat. Commun."},{"key":"38338_CR61","doi-asserted-by":"publisher","first-page":"1479","DOI":"10.1172\/JCI130206","volume":"130","author":"B Rivera","year":"2020","unstructured":"Rivera, B. et al. DGCR8 microprocessor defect characterizes Familial multinodular goiter with schwannomatosis. J. Clin. Invest. 130, 1479\u20131490 (2020).","journal-title":"J. Clin. Invest."},{"key":"38338_CR62","doi-asserted-by":"publisher","first-page":"2452","DOI":"10.1038\/s41596-019-0187-x","volume":"14","author":"CB Gurumurthy","year":"2019","unstructured":"Gurumurthy, C. B. et al. Creation of CRISPR-based germline-genome-engineered mice without ex vivo handling of zygotes by i-GONAD. Nat. Protoc. 14, 2452\u20132482 (2019).","journal-title":"Nat. Protoc."},{"key":"38338_CR63","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1186\/s13059-018-1400-x","volume":"19","author":"M Ohtsuka","year":"2018","unstructured":"Ohtsuka, M. et al. i-GONAD: a robust method for in situ germline genome engineering using CRISPR nucleases. Genome Biol. 19, 25 (2018).","journal-title":"Genome Biol."},{"key":"38338_CR64","doi-asserted-by":"publisher","first-page":"2223","DOI":"10.1016\/j.celrep.2017.10.103","volume":"21","author":"T Watanabe","year":"2017","unstructured":"Watanabe, T. et al. Impediment of replication forks by long Non-coding RNA provokes chromosomal rearrangements by Error-Prone restart. Cell. Rep. 21, 2223\u20132235 (2017).","journal-title":"Cell. Rep."},{"key":"38338_CR65","doi-asserted-by":"crossref","unstructured":"Watanabe, T. & Taniguchi, T. Semi-automated image acquisition and analyses for broad users utilizing macro keyboards. Microsc (Oxf). 74, 437\u2013442 (2025).","DOI":"10.1093\/jmicro\/dfaf018"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-38338-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-38338-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-38338-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,22]],"date-time":"2026-02-22T18:03:22Z","timestamp":1771783402000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-38338-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,3]]},"references-count":65,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["38338"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-38338-5","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,3]]},"assertion":[{"value":"29 September 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 January 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"7112"}}