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The combination model of the above three compounds (butin:caffeic acid: luteolin = 1:4:10) was established using a combination of pharmacological and statistical techniques to promote melanogenesis in B16F10 cells in vitro, and promote melanogenesis in zebrafish in vivo through the expression of melanin-related genes TYR, MITF, KIT and DCT [118]

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Importantly, nuclear-localized FOXO4-positive Leydig cells in elderly testes showed reduced expression of 3-HSD, a rate-limiting enzyme in testosterone synthesis, implying a connection between nuclear FOXO4 and age-related decline in testosterone production [1]