Isolation,assay, biosynthesis,metabolism, uptake and translocation,and function of proline in plant cells and tissues

The amino acid L-proline has been the subject of intensive research during the past ten to fifteen years. This stems from the observations that it incorporates into peptide linkage thereby serving as a precursor to peptidyl-bound L-hydroxyproline, a constituent of “extensin,” and that it accumulates when some plants are exposed to diverse biological and environmental stresses. The contents of selected papers which have been published during the last quarter of a century regarding the isolation, assay, biosynthesis, metabolism, transport and function of L-proline within various plant tissues and their cells are both interpreted and summarized in this review. Occasionally, relevant information from animal and bacterial systems concerning these topics is included. Hydroxyproline-containing proteins are not considered. L-proline was reported to be a constituent of leaves as early as the 1950’s. Since then, it and its analogues have been extracted from the organs of a variety of plants. The analogues include: methyl-hydroxylproline; 4-methylene-DL-proline; L-azetidine-2-carboxylic acid; 2,3,cis-3,4-trans-dihydroxy-L-proline; L-pipecolic acid and 4-trans-hydroxypro-line. L-proline can be both detected and quantified by colorimetric, combined fluorometric-amino acid analyzer and gas Chromatographic procedures. L-proline may be synthesized from L-glutamic acid via the following biosynthetic pathway: L-glutamic acid \(\underrightarrow {\gamma - glutamic acid kinase}\) γ-glutamyl phosphate \(\underrightarrow {\gamma - glutamyl phosphate reductase}\) γ-glutamyl semialdehyde \(\underrightarrow {spontaneous cyclization}\) Δ′-pyrroline-5-Carboxylate (P5C) \(\underrightarrow {P5C reductase}\) L-proline. Proline can also originate from L-arginine and L-ornithine. Biosynthesis from the latter compound proceeds either through the γ-glutamyl semialdehyde and pyrroline-5-carboxylate pathway or alternatively a α-keto-δ-aminovaleric and pyrroline-2-carboxylate pathway. The metabolism of L-proline most likely involves the reverse of the biosynthetic pathway with an initial prolyl dehydrogenaseor prolyl oxidasemediated conversion of L-proline to Δ′-pyrroline-5-carboxylate. The metabolism of L-proline has been demonstrated to occur in excised tissues and cell free extracts, cell suspension cultures and reproductive structures. Little is known about the mechanism by which L-proline is taken up by cultured plant cells and excised tissues. Once within the plant Lproline can be translocated through the phloem at velocities similar to those for carbon dioxide assimilates. In addition to serving as a substrate for peptidyl-bound hydroxyproline, L-proline may function as an adaptation to diverse biological and environmental stresses, a cryoprotectant, a nitrogen pool, a precursor for chlorophyll synthesis upon relief of stress, a regulator together with L-histidine of fertility and sterility and/or a substrate for respiration.