Long-term paternal exercise could reduce the expression of H19 and alter several metabolic genes including GLUT4, FOXO1 and pyruvate dehydrogenase kinase 4, in offspring skeletal muscle in C57BL/6J male mice, and offspring mice whose fathers were exposed to a long-term exercise regimen exhibited lower energy expenditure and an increased risk of obesity and insulin resistance on a HFD [164,169]. Here, FOXO1 is linked to obesity disorder.