Regulation of growth anisotropy in well-watered and water-stressed maize roots .1. Spatial distribution of longitudinal, radial, and tangential expansion rates

As a system to study the regulation of growth anisotropy, we studied thinning of the primary root of maize (Zea mays L.) occurring developmentally or induced by water stress. Seedlings were transplanted into vermiculite at a water potential of approximately -0.03 MPa (well-watered) or -1.6 MPa (water-stressed). The diameter of roots in both treatments decreased with time after transplanting; the water-stressed roots became substantially thinner than well-watered roots at steady state, showing that root thinning is a genuine response to water stress. To analyze the thinning responses we quantified cell numbers and the spatial profiles of longitudinal, radial, and tangential expansion rates separately for the cortex and stele. The results showed that there was no zone of isotropic expansion and the degree of anisotropy varied greatly with position and treatment. Thinning over time in well-watered roots was caused by rates of radial and tangential expansion being too low to maintain the shape of the root. In response to low water potential, cell number in both tissues was unchanged radially but increased tangentially, which shows that thinning was caused wholly by reduced cell expansion. Water stress substantially decreased rates of tangential and radial expansion in both the stele and cortex, but only in the apical 5 mm of the root; basal to this, rates were similar in well-watered and water-stressed roots. By contrast, as in previous studies, longitudinal expansion was identical between the treatments in the apical 3 mm but in water-stressed roots was inhibited at more basal locations. The results show that expansion in longitudinal and radial directions can be regulated independently.