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Papers In Press, published online ahead of print January 23, 2006
J. Biol. Chem, 10.1074/jbc.M510682200
Submitted on September 30, 2005
Revised on December 12, 2005
Accepted on January 23, 2006

Activation of C/EBPalpha expression by C/EBPbeta during adipogenesis requires a PPARgamma-associated repression of HDAC1 at the C/EBPalpha gene promoter

Ying Zuo, Li Qiang, and Stephen R. Farmer

Biochemistry Dept., Boston University School of Medicine, Boston, MA 02118

Corresponding Author: farmer{at}biochem.bumc.bu.edu

Studies have shown that C/EBPbeta can stimulate adipogenesis in non-committed fibroblasts by activating expression of PPARgamma . Other investigations have established a role for C/EBPalpha as well as PPARgamma in orchestrating the complex program of adipogenic gene expression during terminal preadipocyte differentiation. Consequently, it is important to identify factors regulating transcription of the C/EBPalpha gene. In this report, we demonstrate that inhibition of PPARgamma activity by exposure of 3T3-L1 preadipocytes to a potent and selective PPARgamma antagonist inhibits adipogenesis, but also blocks the activation of C/EBPalpha expression at the onset of differentiation. Ectopic expression of C/EBPbeta in Swiss 3T3 mouse fibroblasts (Swiss-LAP cells) induces PPARgamma expression without any significant enhancement of C/EBPalpha expression. Treatment of Swiss-LAP cells with a PPARgamma agonist induces adipogenesis, which includes activation of C/EBPalpha expression. To further establish a role for PPARgamma in regulating C/EBPalpha expression, we expressed C/EBPbeta in PPARgamma -deficient mouse embryo fibroblasts (MEFs). The data show that C/EBPbeta is capable of inducing PPARgamma in PPARgamma +/- MEFs, which leads to activation of adipogenesis including C/EBPalpha expression following exposure to a PPARgamma ligand. In contrast, C/EBPbeta is not able to induce C/EBPalpha expression or adipogenesis in PPARgamma -/- MEFs. ChIP analysis reveals that C/EBPbeta is bound to the minimal promoter of the C/EBPalpha gene in association with HDAC1 in unstimulated Swiss-LAP cells. Exposure of the cells to a PPARgamma ligand dislodges HDAC1 from the C/EBPalpha gene promoter, which involves degradation of HDAC1 in the 26S proteasome. These data suggest that C/EBPbeta activates a single unified pathway of adipogenesis involving its stimulation of PPARgamma expression, which then activates C/EBPalpha expression by dislodging HDAC1 from the promoter for degradation in the proteasome.


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