THE CONTROL OF CELL EXPANSION IN ROOTS

The expansion of roots is considered at the level of the single cell. The water relations of cell expansion are discussed. Water entry, solute import and cell wall properties are considered as possible regulatory points. It is argued that root cell expansion can be understood in terms of cell turgor pressure and the physical properties of the cell wall, provided solute supply is not limiting. Various measurements of cell wall properties in roots are presented and the assumptions underlying their measurements are presented. It is concluded that cell wall properties must be measured over short time periods to prevent alterations in wall properties during the experiment. The radial location of the load-bearing layers is discussed and it is concluded that, unlike aerial tissue, growth is limited by the properties of the inner layer of the root cortex. Evidence is presented to show that cell wall properties can change both during development and following turgor perturbation. In general, however, turgor itself is tightly regulated, particularly towards the root tip. A number of environmental situations are presented in which root growth is altered. The mechanism of the alteration is discussed at the single cell level. These 'stresses' include osmotic stress, low temperature and soil compaction. In many cases the alteration of root growth is consistent with changes in the cell wall properties of the growing cells. Severe stress, resulting in near cessation of root cell extension, can result in a change (usually an increase) in turgor pressure. The change in turgor pressure of the cells in the growing zone is smaller than that which would be expected from a continuation of an unstressed solute import rate. This exemplifies both the change in cell wall properties and the tight turgor homeostasis of root tips. The biochemical processes which underlie the modulation of cell wall properties are presented as they are currently understood in roots. Measurements of the chemical composition of the wall have not revealed any useful differences which can explain the developmental or stress-induced changes in cell wall properties. Recent work on cell wall enzymes and proteins may provide information about control of cross-linkages within the wall. In the last section the relative importance of apoplastic and symplastic solute transport to the expanding cells is considered. At present the consensus appears to favour the symplastic route, but the apoplastic pathway may also operate, possibly as a scavenging mechanism for leaked ions. The regulation of turgor pressure by linking solute import with wall loosening is discussed.