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glucuronic acid glca uronic urine gum arabic gums xanthan kombucha metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor sugar proteoglycans glycoglycerolipids building block heparin chondroitin sulfate dermatan keratan hyaluronic galacturonic acids glucuronidation udpga fermentation enterohepatic circulation molecule molecular formula structural structure 3d