Review article. Molecular biology of salt tolerance in the context of whole-plant physiology

The halobacteria are the only organisms that are tolerant of salinity at the molecular level. All other bacteria, all fungi, all plants, and all animals avoid the need for salt tolerance for most of their macromolecules by maintaining defined and conserved conditions in the cytoplasm. These conditions favour potassium over sodium, the limitation of total inorganic ion activity, and the supplementation of this where necessary with organic solutes which are metabolically neutral osmolytes that may also be osmoprotectant. The salt tolerance of an organism depends upon the range of external salinity over which it is able to sustain these conditions in the cytoplasm. There is substantial and increasing knowledge of the molecular biology and molecular genetics of the processes of ion and organic solute transport, solute synthesis, and compartmentation that underpin cell-based tolerance. Much of recent research focuses on the identification of genes and gene products that affect cell-based tolerance, commonly derived from single-cell models. There is commonly the implicit or explicit assumption that incorporation of these genes will benefit the salt tolerance of food crop species. While this essential experimental approach is giving enormous insight there should not be rash or premature expectations. The unique and overriding consideration for the salinity tolerance of terrestrial plants is the net flux of water due to transpiration and so resides at a higher level of organization. Processes that are advantageous to a single cell in an aqueous medium may be lethal to a cell in a leaf in the air. The likely impact of single structural-gene changes in ion and solute transport upon co-ordinated plant response is probably over-estimated, and recent views consider regulatory processes and multiple gene transfers. While the technical ability for plant transformation increases daily, the practicality of using transgenic plants in complex breeding programmes seems rarely to be given enough thought. If intervention at the molecular level is to lead to salt-tolerant crop plants than it will be essential to view this in the contexts of whole plants and of plant breeding. Recent indications that a relatively small number of quantitative trait loci (QTL) may govern complex physiological characters offer the most hope for the future.