Surfing along the root ground tissue gene network

Organization of tissues in Arabidopsis thaliana root is made of, from outside in, epidermis, cortex, middle cortex, endodermis, pericycle and vascular tissues. Cortex, middle cortex and endodermis form the ground tissue (GT) system. Functional and molecular characterization of GT patterning mutants' properties has greatly increased our understanding of fundamental processes of plant root development. These studies have demonstrated GT is an elegant model that can be used to study how different cell types and cell fates are specified. This review analyzes GT mutants to provide a detailed account of the molecular network that regulates GT formation in A. thaliana. The most recent results indicate an unexpectedly complex network of transcription factors, epigenetic and hormonal controls that play crucial roles in GT development. Major differences exist between GT formation in dicots and monocots, particularly in the model plant rice, opening the way for evo-devo of GT formation in angiosperm. In rice, adaptation to submergence relies on a multilayered cortex. Moreover, variation in the number of cortex cell layers is also observed between the five root types. A mechanism of control for cortical cell number should then exist in rice and it remains to be determined if any of the Arabidopsis thaliana identified GT network members are also involved in this process in rice. Alternatively, a totally different network may have been invented. However, first available results suggest functional conservation in rice of at least two transcription factors, SHORT ROOT (SHR) and SCARECROW (SCR), involved in ground tissue formation in Arabidopsis.