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Senescent Leydig Cell Elimination & Testosterone Restoration in Aged Mice (Aging / PMC, 2020) FOXO4-DRI Disrupts FOXO4-p53 in Senescent Leydig Cells and Improves Testicular Microenvironment A study published in Aging (and indexed on PMC) demonstrated that FOXO4 is specifically expressed in human Leydig cells, with nuclear translocation of FOXO4 in elderly men correlating with decreased testosterone synthesis