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tetrahydrocortisone urocortisone metabolite cortisone steroid metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous hormone hormonal replacement function action endocrine endocrinology level prohormone analogue mineralocorticoid corticosteroid glucocorticosteroid steroid glucocorticoid receptor inflammation immune system autoimmune disease stress response carbohydrate protein anti-inflammatory 3d