991 64-4238 (V.R.F.)Mannitol, a major photosynthetic product and transport carbohydrate in many plants, accounts for approximately 50% of the carbon fixed by celery (Apium graveolens 1.) leaves. Previous subfractionation studies of celery leaves indicated that the enzymes for mannitol synthesis were located in the cytosol, but these data are inconsistent with that published for the sites of sugar alcohol synthesis in other families and taxa, including apple (Malus) and a brown alga (fucus). Using antibodies to a key synthetic enzyme, NADPH-dependent mannose-6-phosphate reductase (MCPR), and immunocytochemical techniques, we have resolved both the intercellular and intracellular sites of mannitol synthesis. In leaves, MCPR was found only in cells containing ribulose-l,5-bisphosphate carboxylase/oxygenase. MCPR was almost exclusively cytosolic in these cells, with the nucleus being the only organelle to show labeling. The key step in transport carbohydrate biosynthesis that is catalyzed by MCPR displays no apparent preferential association with vascular tissues or with the bundle sheath. These results show that MCPR and, thus, mannitol synthesis are closely associated with the distribution of photosynthetic carbon metabolism in celery leaves. The principal role of MCPR is, therefore, in the assimilation of carbon being exported from the chloroplast, and i t seems unlikely that this enzyme plays even an indirect role in phloem loading of mannitol.The acyclic sugar alcohols (polyols) are second only to the sugars as principal products of carbon assimilation and account for an estimated 30% of global primary production (Bieleski, 1982). In a11 polyol-producing higher plants studied to date, Suc is also a major photoassimilate. In some species, sucrosyl-oligosaccharides (eg. raffinose) are found in the phloem sap in addition to the polyol and SUC. The nature of primary production and transport in higher plant polyol producers raises fundamental questions as to why such an apparently complex system has evolved when Suc alone performs adequately in many other species. Questions are also raised as to how carbon partitioning into polyols and other primary products is regulated at a biochemical, cellular, and tissue level. Celery (Apium graveolens L.) is a fine model in which to study such questions, because mature leaves