Asymmetically labeled sucrose, 14C(fructosyl)sucrose, was used to determine whether sucrose undergoes extracellar hydrolysis duing phloem trandocation in the sugar beet, Beta vulgaris. In addition, the metabolsm of variom sugars accmulated and trasocated was determined in varios gions of the plant. These proceses were studied in detached regions as well as in the intact, trnlocating plant in the source leaf, along the trslocation path, and in a rapidly growing sink leaf nd storage beet. Tbe data show that, unlike sucrose accumulation into the sink tisse of sugarcane, ucrose is neither hydrolzyed prior to phloem loading or during tansit, nor is it extrcellularty hydrolyzed during accumulation into sink leaves or the storage beet.Although it is well established that photosynthetically derived assimilates are transported between source and sink regions along a gradient of hydrostatic pressure, relatively little is known about the cellular and metabolic correlates of the translocation process. The translocation system is generally divided into three structurally and physiologically distinct regions: the source, path, and sink regions. Recent structural and physiological studies (5,10,17,18) have indicated that the path region is relatively passive in the translocation process, in that the driving force for transport resides in metabolism-dependent processes operating in the source and sink regions.From a mechanistic viewpoint, the major emphasis in the phloem transport literature has been toward elucidating the role of the path at the expense of source and sink metabolism, particularly at the cellular level. Recent studies by Geiger and co-workers (12, 13) have investigated events operating in the source leaf of Beta vulgaris during the loading of sucrose into the phloem prior to translocation. What has emerged from these studies and from the more recent studies by Giaquinta (14,15 (19).Hydrolysis of sucrose to hexoses by a free space invertase prior to accumulation quite convincingly occurs in sugarcane tissue. The paucity of information on this subject in other species makes it difficult to determine whether these events constitute a general mechanism of sink metabolism of translocate or whether they are characteristic for the sugarcane system.The literature on the role of extracellular invertase in phloem translocation in both source and sink regions is controversial. Part of the discrepancy and variability probably arises from the observation that invertase activity can be induced in some tissues by physical dissection or prolonged washing (8). Another possibility is that extracellular invertase as related to translocation is not universally present in various species and that sucrose need not be extracellularly hydrolyzed prior to membrane transport into the sink tissue.The form in which sucrose is transported across cell membranes during phloem translocation is important in elucidating the nature and selectivity of membrane carriers involved in phloem loading and unloading of sugars in the source and s...