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stachydrine betonica officinalis proline betaine l-proline teucrium polium halopithys incurva escherichia coli zwitterion food component plant metabolite human blood serum l-prolinium enantiomer d-proline methyl hygrate l-stachydrine methylproline methylbetaine metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor metabolome database hmdb 3d molecule molecular formula skeletal structural structure bioactive biochemical