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Proline Synthesis In plants subjected to osmotic stress, proline is mainly synthesized from glutamic acid by D1-pyrroline-5-carboxylate (P5C) by two consecutive reductions catalyzed by P5C synthase (P5CS), which is a rate-limiting enzyme of the biosynthetic pathway in complex plants and P5C reductase (P5CR) (Verbruggen and Hermans, 2008)