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p18-INK4c acts as a negative regulator of the cell cycle at the early G1 phase ( Cdkn2c / Tp53 double knockout mice displayed higher percentage of PCNA (a marker of proliferation expressed in S phase) -positive tubules compared with that in wild type and Cdkn2c -null mice, which indicated Cdkn2c / Tp53 double knockout mice existed active proliferation and continuation of the mitosis phase in spermatogonia, and thus a delay in meiosis entry ( Cdkn2c null mice showed a significantly increased sperm counts as compared to Tp53 null or wild type, while Cdkn2c / Tp53 double knockout mice had much lower sperm counts, which might attributed to the increased apoptosis induced by DNA damage secondary to prolongation of meiosis entry ( 2.2 The role of p53 in spermatocytes A1-type spermatogonia are transformed into preleptotene spermatocytes after six divisions and subsequently enter meiosis ( As the guardian of the genome, p53 appears to be duty-bound to respond to DNA damage during spermatogenesis

4 The broader harms related to alcohol use also show concerning trends

Statements Author contributions LB, AA and NH: contributed to structuring the review, discussing the literature and the knowledge gaps