Control of Asymmetric Cell Divisions during Root Ground Tissue Maturation

Controlling the production of diverse cell/tissue types is essential for the development of multicellular organisms such as animals and plants. The Arabidopsis thaliana root, which contains distinct cells/tissues along longitudinal and radial axes, has served as an elegant model to investigate how genetic programs and environmental signals interact to produce different cell/tissue types. In the root, a series of asymmetric cell divisions (ACDs) give rise to three ground tissue layers at maturity (endodermis, middle cortex, and cortex). Because the middle cortex is formed by a periclinal (parallel to the axis) ACD of the endodermis around 7 to 14 days post-germination, middle cortex formation is used as a parameter to assess maturation of the root ground tissue. Molecular, genetic, and physiological studies have revealed that the control of the timing and extent of middle cortex formation during root maturation relies on the interaction of plant hormones and transcription factors. In particular, abscisic acid and gibberellin act synergistically to regulate the timing and extent of middle cortex formation, unlike their typical antagonism. The SHORT-ROOT, SCARECROW, SCARECROW-LIKE 3, and DELLA transcription factors, all of which belong to the plant-specific GRAS family, play key roles in the regulation of middle cortex formation. Recently, two additional transcription factors, SEUSS and GA- AND ABA-RESPONSIVE ZINC FINGER, have also been characterized during ground tissue maturation. In this review, we provide a detailed account of the regulatory networks that control the timing and extent of middle cortex formation during post-embryonic root development.