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We also found that in elderly testes, nuclear-FOXO4 + Leydig cells expressed less 3-hydroxysteroid dehydrogenase (3-HSD), the rate-limiting enzyme in testosterone synthesis, than nuclear-FOXO4 - Leydig cells (Figure 2D)

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220 These regions include the hippocampus and entorhinal cortex, structures that play key roles in memory and language