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1-pyrroline-5-carboxylic cyclic imino acid rice hyperprolinemia glutamine compound pyrrole p5c metabolite stereoisomer intermediate degradation gsa amino prokaryotic eukaryote proline arginine 3d chemistry chemical biochemistry biology molecule molecular structural structure formula skeletal geometry isomer component reagent reaction production synthesis iupac metabolism biosynthesis cycle mechanism pathway catabolism heterocyclic heterocycle ring