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Rodent incisors grow continuously throughout life, and epithelial progenitor cells are supplied from stem cells in the cervical loop. We report that epithelial  Runx  genes are involved in the maintenance of epithelial stem cells and their subsequent continuous differentiation and therefore growth of the incisors. Core binding factor β (Cbfb) acts as a binding partner for all Runx proteins, and targeted inactivation of this molecule abrogates the activity of all Runx complexes. Mice deficient in epithelial  Cbfb  produce short incisors and display marked underdevelopment of the cervical loop and suppressed epithelial  Fgf9  expression and mesenchymal  Fgf3  and  Fgf10  expression in the cervical loop. In culture, FGF9 protein rescues these phenotypes. These findings indicate that epithelial  Runx  functions to maintain epithelial stem cells and that  Fgf9  may be a target gene of  Runx  signaling.  Cbfb  mutants also lack enamel formation and display downregulated  Shh  mRNA expression in cells differentiating into ameloblasts. Furthermore,  Fgf9  deficiency results in a proximal shift of the  Shh  expressing cell population and ectopic FGF9 protein suppresses  Shh  expression. These findings indicate that  Shh  as well as  Fgf9  expression is maintained by  Runx/Cbfb  but that  Fgf9  antagonizes  Shh  expression. The present results provide the first genetic evidence that  Runx/Cbfb  genes function in the maintenance of stem cells in developing incisors by activating  Fgf  signaling loops between the epithelium and mesenchyme. In addition,  Runx  genes also orchestrate continuous proliferation and differentiation by maintaining the expression of  Fgf9  and  Shh  mRNA. S TEM  C ELLS  2011;29:1792–1803

Core Binding Factor Beta Functions in the Maintenance of Stem Cells and Orchestrates Continuous Proliferation and Differentiation in Mouse Incisors