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Although mutations in several genes involved in ribosome biogenesis have been identified in patients with TCS, the molecular mechanisms underlying their effects remain poorly understood. In this study, we analyzed the effects of TCS type 4 (TCS4)\u2010associated mutations in\n                    <jats:italic>Schizosaccharomyces pombe<\/jats:italic>\n                    by introducing R1022C or R1022S mutations into Rpa2, the second\u2010largest subunit of RNA polymerase I (Pol I). The\n                    <jats:italic>rpa2<\/jats:italic>\n                    <jats:sup>R1022C<\/jats:sup>\n                    and\n                    <jats:italic>rpa2<\/jats:italic>\n                    <jats:sup>R1022S<\/jats:sup>\n                    mutants exhibited impaired cell growth under nutrient\u2010rich conditions without affecting the Rpa2 protein levels. Furthermore, Pol I abnormally accumulated at the 5\u2032 region of rDNA in these mutants, resulting in defective 35S pre\u2010rRNA biogenesis and increased sensitivity to the Pol I inhibitor BMH\u201021. These findings highlight the essential role of the Rpa2 residues associated with TCS4 in rRNA transcription and cell growth.\n                  <\/jats:p>","DOI":"10.1002\/2211-5463.70152","type":"journal-article","created":{"date-parts":[[2026,1,16]],"date-time":"2026-01-16T11:21:26Z","timestamp":1768562486000},"page":"561-569","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Analysis of Treacher Collins syndrome 4\u2010associated mutations in\n                    <i>Schizosaccharomyces pombe<\/i>"],"prefix":"10.1002","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1134-7504","authenticated-orcid":false,"given":"Kei","family":"Kawakami","sequence":"first","affiliation":[{"name":"Department of Biochemistry Shimane University School of Medicine  Izumo 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