Hormones and the growth of plants in response to gravity

Plants are remarkable amongst living things in that all their parts (leaves, stems, roots and flowers) can orientate their position in response to gravity. Roots normally grow downwards towards the stimulus, shoots upwards away from the stimulus, while leaves and special kinds of stems like underground rhizomes or aerial runners have adaptations to grow horizontally, approximately at right angles to the gravitational field. This directional positioning of the organ in relation to gravity is achieved by a regulation of the growth of enlarging cells below the meristems or within the organ. Mature tissues in which further cell growth is precluded are therefore unable to reorientate if their position in the field is changed. Under conditions of zero gravity, roots and shoots continue to grow in opposite directions and the orientation of laterals with respect to the apex is essentially normal. This demonstrates the inherent polarity of plant cells and the internal correlative growth regulation that each organ exerts upon its neighbours. The perception of gravity, involving statoliths, membranes and "wound" ethylene, is discussed, together with the mechanisms by which the subsequent growth responses can be mediated by changes in endogenous hormones. Evidence for how such hormonal changes can lead to modifications of the rate, extent and reorientation of cell growth is reviewed for several geotropically responding systems.