Asymmetry and cell polarity in root development.

Within living systems, striking juxtapositions in symmetry and asymmetry can be observed and the superficial appearance of symmetric organization often gives way to cellular asymmetries at higher resolution. It is frequently asymmetry and polarity that fascinate and challenge developmental biologists. In multicellular eukaryotes, cell polarity and asymmetry are essential for diverse cellular, tissue, and organismal level function and physiology and are particularly crucial for developmental processes. In plants, where cells are surrounded by rigid cell walls, asymmetric cell divisions are the foundation of pattern formation and differential cell fate specification. Thus, cellular asymmetry is a key feature of plant biology and in the plant root the consequences of these asymmetries are elegantly displayed. Yet despite the frequency of asymmetric (formative) cell divisions, cell/tissue polarity and the proposed roles for directional signaling in these processes, polarly localized proteins, beyond those involved in auxin or nutrient transport, are exceedingly rare. Indeed, although half of the asymmetric cell divisions in root patterning are oriented parallel to the axis of growth, laterally localized proteins directly involved in patterning are largely missing in action. Here, various asymmetric cell divisions and cellular and structural polarities observed in roots are highlighted and discussed in the context of the proposed roles for positional and/or directional signaling in these processes. The importance of directional signaling and the weight given to polarity in the root-shoot axis is contrasted with how little we currently understand about laterally oriented asymmetry and polarity in the root.