Is it good noise? The role of developmental instability in the shaping of a root system

Root architecture plays a major part in determining a root system's ability to function effectively and efficiently in its essential roles of anchorage and the capture of soil resources. The characteristics of root development that are conventionally considered to be the main determinants of root architecture are the rate, angle, and duration of root growth and the pattern of root branching. In this review, the case is made that there is an additional trait that has been largely ignored but which has a significant influence on root architecture, namely the degree to which stochasticity (or 'developmental instability') affects the developmental process. Although the intrinsic variability in the development and growth of lateral roots has been recognized for some time, in almost every study of root development this remarkable facet of root behaviour tends to be hidden beneath the veil of statistical averaging. Progress in other fields is providing intriguing insights into the phenomenon of developmental instability, how it is generated at the molecular and cellular levels and the genetic mechanisms by which it is buffered. This review will consider the existence of developmental instability in roots, its underlying causes, its effects on root architecture, and the evidence that it is under genetic control. The hypothesis will be advanced that developmental instability in roots is an adaptive trait, and its potential relevance to root function will be discussed in both an ecological and an agronomic context.