Many plants accumulate organic osmolytes in response to the imposition of environmental stresses that cause cellular dehydration. Although an adaptive role for these compounds in mediating osmotic adjustment and protecting subcellular structure has become a central dogma in stress physiology, the evidence in favour of this hypothesis is largely correlative. Transgenic plants engineered to accumulate proline, mannitol, fructans, trehalose, glycine betaine or ononitol exhibit marginal improvements in salt and/or drought tolerance. While these studies do not dismiss causative relationships between osmolyte levels and stress tolerance, the absolute osmolyte concentrations in these plants are unlikely to mediate osmotic adjustment. Metabolic benefits of osmolyte accumulation may augment the classically accepted roles of these compounds. In re-assessing the functional significance of compatible solute accumulation, it is suggested that proline and glycine betaine synthesis may buffer cellular redox potential. Disturbances in hexose sensing in transgenic plants engineered to produce trehalose, fructans or mannitol may be an important contributory factor to the stress-tolerant phenotypes observed. Associated effects on photoassimilate allocation between root and shoot tissues may also be involved. Whether or not osmolyte transport between subcellular compartments or different organs represents a bottleneck that limits stress tolerance at the whole-plant level is presently unclear. None the less, if osmolyte metabolism impinges on hexose or redox signalling, then it may be important in long-range signal transmission throughout the plant.Key-words: betaine; cold stress; drought; fructans; mannitol; osmolytes; proline; salinity; sugar signalling; trehalose.
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INTRODUCTIONEnvironmental stress is the major factor limiting plant productivity . Abiotic stresses which cause depletion of cellular water (drought, high soil salinity and temperature extremes) are responsible for the greatest agricultural losses. Upon exposure to these prevalent stresses, many plants accumulate organic osmolytes, most commonly polyhydroxylic compounds (saccharides and polyhydric alcohols) and zwitterionic alkylamines (amino acids and quaternary ammonium compounds).Several recent reviews discuss osmolyte accumulation in plants (Ingram & Bartels 1996;Serrano 1996). It is generally accepted that the increase in cellular osmolarity which results from the accumulation of non-toxic (thus 'compatible') osmotically active solutes is accompanied by the influx of water into, or at least a reduced efflux from, cells, thus providing the turgor necessary for cell expansion. None the less, a conclusive demonstration that osmotic adjustment contributes to fitness in stressful environments has yet to be achieved (Munns 1993). Since all subcellular structures must exist in an aqueous environment, tolerance to dehydration also depends on the ability of cells to maintain membrane integrity and prevent protein denaturation. Hypotheses that attribut...