SHORT-ROOT regulates vascular patterning,but not apical meristematic activity in the Arabidopsis root through cytokinin homeostasis

SHORT-ROOT (SHR) is a key regulator of radial patterning and stem-cell renewal in the Arabidopsis root. Although SHR is expressed in the stele, its function in the vascular tissue was not recognized until recently. In shr, the protoxylem is missing due to the loss of expression of microRNA165A (miR165A) and microRNA166B (miR165B). shr is also defective in lateral root formation, but the mechanism remains unclear. To dissect the SHR developmental pathway, we recently have identified its direct targets at the genome scale by chromatin immunoprecipitation followed by microarray analysis (ChIP-chip). In further studies, we have shown that SHR regulates cytokinin homeostasis through cytokinin oxidase 3 and that this role of SHR is critical to vascular patterning in the root. In this communication we report that SHR also regulates miR165A and miR166B indirectly through its effect on cytokinin homeostasis. Although cytokinin is inhibitory to root growth, the root-apical-meristem defect in shr was not alleviated by reduction of endogenous cytokinin. These results together suggest that SHR regulates vascular patterning, but not root apical meristematic activity, through cytokinin homeostasis.     

拟南芥根的辐射形态相关基因SHORT-ROOT 研究进展

从模式植物拟南芥中克隆得到的SHORT-ROOT基因(SHR)被证明参与根部形态建成途径。目前已知SHR是与根辐射形态直接相关的重要调控因子, 同时也参与维持根尖分生组织的活性。SHR既作为转录因子启动下游基因的表达, 又作为短程信号调节根的发育。本文综述了SHR相关研究进展, 并展望其研究前景。     

拟南芥根的辐射形态相关基因SHORT-ROOT研究进展

从模式植物拟南芥中克隆得到的SHORT-ROOT基因（SHR）被证明参与根部形态建成途径。目前已知SHR是与根辐射形态直接相关的重要调控因子,同时也参与维持根尖分生组织的活性。SHR既作为转录因子启动下游基因的表达,又作为短程信号调节根的发育。本文综述了SHR相关研究进展,并展望其研究前景。     

CDC48B facilitates the intercellular trafficking of SHORT-ROOT during radial patterning in roots

CELL DIVISION CONTROL PROTEIN48 (CDC48) is essential for membrane fusion, protein degradation, and other cellular processes. Here, we revealed the crucial role of CDC48B in regulating periclinal cell division in roots by analyzing the recessive gen1 mutant. We identified the GEN1 gene through map-based cloning and verified that GEN1 encodes CDC48B. gen1 showed severely inhibited root growth, increased periclinal cell division in the endodermis, defective middle cortex (MC) formation, and altered ground tissue patterning in roots. Consistent with these phenotypes, CYCLIND 6;1(CYCD6;1), a periclinal cell division marker, was upregulated in gen1 compared to Col-0. The ratio of SHR_pro:SHR-GFP fluorescence in pre-dividing nuclei vs. the adjacent stele decreased by 33% in gen1, indicating that the trafficking of SHORT-ROOT (SHR) decreased in gen1 when endodermal cells started to divide. These findings suggest that the loss of function of CDC48B inhibits the intercellular trafficking of SHR from the stele to the endodermis, thereby decreasing SHR accumulation in the endodermis. These findings shed light on the crucial role of CDC48B in regulating periclinal cell division in roots.     

The root endodermis: A hub of developmental signals and nutrient flow

The root endodermis is the cylindrical boundary that separates the inner vascular tissue from the outer cortex and functions as an apoplasmic barrier for selective nutrient uptake. Recent developmental and cell biological studies have started to reveal the mechanisms by which this single cell layer serves as a key regulatory module of root growth, tissue patterning and nutrient flow, which in concert support the plant’s ability to survive in a terrestrial habitat. This review provides an overview of the key factors that contribute to the functioning of the root endodermis and discusses how this single cell layer dictates root growth and tissue patterning.