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orotic acid pyrimidinedione vitamin b13 carboxylic vitamin mitochondrial enzyme dihydroorotate dehydrogenase cytoplasmic pyrimidine synthesis pathway mineral carrier dietary supplements bioavailability lithium orotate supplement biological activity bioactive nutrient prescription drug food additive nutraceutica dietary aciduria acidemia urea cycle disorder metabolic ornithine transcarbamylase deficiency blood nitrogen megaloblastic anaemia metabolite metabolism biosynthesis cycle mechanism path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic endogenous 3d molecule