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Premature cellular senescence can be triggered by overexpression of several oncogenes, such as HRasG12V, as well as loss of tumor suppressors, such as PTEN, NF1, VHL, and ARF. Overexpression of Ras or loss of PTEN leads to ARF/p53 activation and cellular senescence. However, inactivation of Skp2 in combination with PTEN or ARF inactivation leads to an induction of p21, p27, and ATF4 in an ARF/p53-independent manner.
