Effects of abiotic stresses on sorbitol biosynthesis and metabolism in tomato (Solanum lycopersicum).

Polyols such as sorbitol and ribitol are a class of compatible solutes in plants that may play roles in tolerance to abiotic stresses. This study investigated the effects of water stress on sorbitol biosynthesis and metabolism and sorbitol and ribitol accumulation in tomato (Solanum lycopersicum L.). Water stress induced by withholding water and by using polyethylene glycol as a root incubation solution to mimic water stress, and NaCl stress were applied to wild-type (WT) and three genetically-modified lines of tomato (cv. Ailsa Craig), a control vector line TR22, and 2 sorbitol dehydrogenase (sdh) antisense lines TR45 and TR49. Sorbitol and ribitol content, as well as the enzymatic activities, protein accumulation, and gene expression patterns of the key sorbitol cycle enzymes aldose-6-phosphate reductase (A6PR), aldose reductase (AR), and sorbitol dehydrogenase (SDH), were measured in mature leaves. In response to the stresses, both sorbitol and ribitol accumulated in leaf tissue, most significantly in the sdh antisense lines. A6PR, characterised for the first time in this work, and AR both exhibited increased enzymatic activity correlated with sorbitol accumulation during the stress treatments, with SDH also increasing in WT and TR22 to metabolise sorbitol, reducing the content to control levels within 3 days after re-watering. In the sdh antisense lines, the lack of significant SDH activity resulted in the increased sorbitol and ribitol content above WT levels. The results highlighted a role for both A6PR and AR in biosynthesis of sorbitol in tomato where the high activity of both enzymes was associated with sorbitol accumulation. Although both A6PR and AR are aldo-keto reductases and use NADPH as a co-factor, the AR-specific inhibitor sorbinil inhibited AR only indicating that they are different enzymes. The determination that sorbitol, and perhaps ribitol as well, plays a role in abiotic responses in tomato provides a cornerstone for future studies examining how they impact tomato tolerance to abiotic stresses, and if their alteration could improve stress tolerance.

The role of SORBITOL DEHYDROGENASE in Arabidopsis thaliana.

SORBITOL DEHYDROGENASE (SDH, EC 1.1.1.14) catalyses the interconversion of polyols and ketoses (e.g. sorbitol ⟷ fructose). Using two independent Arabidopsis thaliana (L.) Heynh. sdh knockout mutants, we show that SDH (At5g51970) plays a primary role in sorbitol metabolism as well as an unexpected role in ribitol metabolism. Sorbitol content increased in both wild-type (WT) and mutant plant leaves during drought stress, but mutants showed a dramatically different phenotype, dying even if rewatered. The lack of functional SDH in mutant plants was accompanied by accumulation of foliar sorbitol and at least 10-fold more ribitol, neither of which decreased in mutant plants after rewatering. In addition, mutant plants were uniquely sensitive to ribitol in a concentration-dependent manner, which either prevented them from completing seed germination or inhibited seedling development, effects not observed with other polyols or with ribitol-treated WT plants. Ribitol catabolism may occur solely through SDH in A. thaliana, though at only 30% the rate of that for sorbitol. The results indicate a role for SDH in metabolism of sorbitol to fructose and in ribitol conversion to ribulose in A. thaliana during recovery from drought stress.

Tissue-specific expression of SORBITOL DEHYDROGENASE in apple fruit during early development.

Sorbitol, the primary photosynthate and translocated carbohydrate in apple (Malusxdomestica Borkh.), is converted to fructose by sorbitol dehydrogenase (SDH; EC 1.1.1.14) which is active in apple fruit throughout development. In the apple genome, nine SDH genes have been isolated and their sequences characterized, but their individual expression patterns during apple fruit set and development have not been determined. The objective of this work was to ascertain if SDH genes are differentially expressed and how their patterns of expression may relate to SDH activity in apple seed and cortex during early fruit development. Seed SDH activity was found to be much higher than cortex SDH activity per mg and g fresh weight (FW), and seed SDH activity contributed significantly to whole fruit SDH activity during weeks 2-5 after bloom. Five of the nine SDH genes present in the apple genome were expressed in apple fruit. Two SDH genes, SDH1 and SDH3, were expressed in both seed and cortex tissues. SDH2 expression was limited to cortex, while SDH6 and SDH9 were expressed in seed tissues only. SDH isomeric proteins of different pI values were detected in apple fruit. SDH isomers with pI values of 4.2, 4.8, 5.5, and 6.3 were found in seeds, and SDH isomers with pI values of 5.5, 6.3, 7.3, and 8.3 were found in cortex. The present work is the first to show that SDH is highly active in apple seed and that SDH genes are differentially expressed in seed and cortex during early development.