Induced expression of C EBP and C EBP , two factors that contribute to PPAR induction. Because arsenic efficaciously inhibits adipogenesis induced by this PPAR agonist, it can be concluded that upstream signaling events are not required for the inhibitory effects of arsenic on differentiation. In cells induced to undergo adipogenesis, treatment with arsenic tips the balance away from differentiation and toward proliferation. At the molecular level, adipogenesis is inhibited by phosphorylation of PPAR by MAP kinase Hu et al., 1996 ; , and arsenic potentially increases MAPK activity secondary to down regulated MKP-1 expression Trouba et al., 2000a ; . Therefore, it is conceivable that arsenic blocks adipogenesis simply by maintaining cells in a state of proliferative activity. This leads to a major question of whether arsenic inhibits adipogenesis indirectly by its ability to maintain cells in a mitogenically competent state, or by a more direct mechanism involving adipogenic processes. We found that UO126mediated inhibition of mitogenic signaling does not abrogate the differentiation inhibition effect of arsenic, thereby supporting the hypothesis that arsenic interferes with the molecular events leading to differentiation. For full phenotypic differentiation of adipocytes, expression of both PPAR and C EBP is required El-Jack et al., 1999 ; . In addition to playing a pivotal role in establishment of insulinsensitive glucose transport, C EBP is necessary for maintenance of post-mitotic growth arrest in adipocytes Tao and Umek 2000; Timchenko et al., 1996 ; . Downregulation of C EBP , accomplished by expression of antisense RNA, enables quiescent cells to reenter the cell cycle Tao and Umek 2000 ; , thereby illustrating the importance of C EBP in maintaining proliferative inactivity. Because C EBP positively regulates protein levels of p21 Waf1 Cip1, Timchenko et al., 1996 ; , the abnormally low level of p21 Waf1 Cip1 seen in arsenic-treated cells Fig 5 ; might be a consequence of arsenic-induced down regulation of C EBP . Using animals genetically engineered to lack C EBP , a correlation has been made between loss of C EBP , decreased p21 Waf1 Cip1 levels, and unregulated cell proliferation in vivo Timchenko et al., 1997 ; . There is increasing evidence that C EBPs, including C EBP , are important in regulating epidermal differentiation Maytin and Habener 1998; Maytin et al., 1999; Oh and Smart 1998; Swart et al., 1997; Zhu et al., 1999 ; . C EBP and C EBP are greatly reduced in squamous cell carcinomas when compared to control cells Oh and Smart 1998 ; , establishing a link between changes in C EBP expression and malignant transformation. In light of the fact that arsenic causes primarily skin cancer, arsenic-induced alterations in C EBP could contribute to inhibition of differentiation. Like other members of the family of cyclin-dependent kinase inhibitors, p21 Cip1 Waf1 plays an important role in regulating cell cycle progression by inhibiting the cyclin-dependent kinases CDKs ; . A putative target gene of C EBP , p21 Cip1 Waf1 is upregulated following induction of C EBP Heath et al., 2000; Timchenko et al., 1996 ; . Furthermore, the proliferative.

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