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We found that <jats:italic>Atp6i<\/jats:italic> deficient mice (<jats:italic>Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup>) arrested tooth root formation, indicated by truncated Hertwig\u2019s epithelial root sheath (HERS) progression. Furthermore, Atp6i deficiency significantly reduced the proliferation and differentiation of radicular odontogenic cells responsible for root formation<jats:italic>. Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> mice had largely decreased expression of odontoblast differentiation marker gene expression profiles (<jats:italic>Col1a1, Nfic, Dspp, and Osx<\/jats:italic>) in the alveolar bone. <jats:italic>Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> mice sample RNA-seq analysis results showed decreased expression levels of odontoblast markers. Additionally, there was a significant reduction in Smad2\/3 activation, inhibiting transforming growth factor-\u03b2 (TGF-\u03b2) signaling in <jats:italic>Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> odontoblasts. Through treating pulp precursor cells with <jats:italic>Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> or wild-type OC bone resorption-conditioned medium, we found the latter medium to promote odontoblast differentiation, as shown by increased odontoblast differentiation marker genes expression (<jats:italic>Nfic, Dspp, Osx<\/jats:italic>, and <jats:italic>Runx2)<\/jats:italic>. This increased expression was significantly blocked by anti-TGF-\u03b21 antibody neutralization, whereas odontoblast differentiation and Smad2\/3 activation were significantly attenuated by <jats:italic>Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> OC conditioned medium. Importantly, ectopic TGF-\u03b21 partially rescued root development and root dentin deposition of <jats:italic>Atp6i<\/jats:italic><jats:sup><jats:italic>\u2212\/\u2212<\/jats:italic><\/jats:sup> mice tooth germs were transplanted under mouse kidney capsules. Collectively, our novel data shows that the prevention of TGF-\u03b21 release from the alveolar bone matrix due to OC dysfunction may lead to osteopetrosis-associated root formation via impaired radicular odontoblast differentiation. As such, this study uncovers TGF-\u03b21 \/Smad2\/3 as a key signaling pathway regulating odontoblast differentiation and tooth root formation and may contribute to future therapeutic approaches to tooth root regeneration.<\/jats:p>","DOI":"10.1038\/s41368-023-00235-2","type":"journal-article","created":{"date-parts":[[2023,8,21]],"date-time":"2023-08-21T03:08:56Z","timestamp":1692587336000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":13,"title":["Atp6i deficient mouse model uncovers transforming growth factor-\u03b21 \/Smad2\/3 as a key signaling pathway regulating odontoblast differentiation and tooth root formation"],"prefix":"10.1038","volume":"15","author":[{"given":"Jue","family":"Wang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abigail","family":"McVicar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yilin","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hong-Wen","family":"Deng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhihe","family":"Zhao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yi-Ping","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2023,8,21]]},"reference":[{"key":"235_CR1","doi-asserted-by":"crossref","first-page":"2940","DOI":"10.2741\/1750","volume":"10","author":"KU Ogbureke","year":"2005","unstructured":"Ogbureke, K. 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