Modelling of root growth and bending in two dimensions.

A special co-ordinate system is developed for modelling the gravitropic bending of plant roots. It is based on the Local Theory of Curves in differential geometry and describes, in one dimension, growth events that may actually occur in two, or even three, dimensions. With knowledge of the spatial distributions of relative elemental growth rates (RELELs) for the upper and lower flanks of a gravistimulated root, and also their temporal dependencies, it is possible to compute the development of curvature along the root and hence describe the time-course of gravitropic bending. In addition, the RELEL distributions give information about the velocity field and the basipetal displacement of points along the root's surface. According to the Fundamental Theorem of Local Curve Theory, the x and y co-ordinates of the root in its bending plane are then determined from the associated values of local curvature and local velocity. With the aid of this model, possible mathematical growth functions that correspond to biological mechanisms involved in differential growth can be tested. Hence, the model can help not only to distinguish the role of various physiological or biophysical parameters in the bending process, but also to validate hypotheses that make assumptions concerning their relative importance. However, since the model is constructed at the level of the organ and treats the root as a fluid continuum, none of the parameters relate to cellular behaviour; the parameters must instead necessarily apply to properties that impinge on the behaviour of the external boundary of the root.