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dihydroprogesterone dhp allopregnanedione endogenous progestogen neurosteroid progesterone intermediate akrs aldo-keto reductases liver mammary gland progesterone receptor agonist positive allosteric modulator gaba progestogenic rendering metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme contractor human metabolome database hmdb hormone hormonal therapy replacement function action endocrine level prohormone