Tornado1 and tornado2 are required for the specification of radial and circumferential pattern in the Arabidopsis root

The cell layers of the Arabidopsis primary root are arranged in a simple radial pattern. The outermost layer is the lateral root cap and lies outside the epidermis that surrounds the ground tissue. The files of epidermal and lateral root cap cells converge on a ring of initials (lateral root cap/epidermis initial) from which the epidermal and lateral root cap tissues of the seedling are derived, once root growth is initiated after germination. Each initial gives rise to a clone of epidermal cells and a clone of lateral root cap cells. These initial divisions in the epidermal/lateral root cap initial are defective in tornado1 (trn1) and trn2 plants indicating a requirement for TRN1 and TRN2 for initial cell function. Furthermore, lateral root cap cells develop in the epidermal position in trn1 and trn2 roots indicating that TRN1 and TRN2 are required for the maintenance of the radial pattern of cell specification in the root. The death of these ectopic lateral root cap cells in the elongation zone (where lateral root cap cells normally die) results in the development of gaps in the epidermis. These observations indicate that TRN1 and TRN2 are required to maintain the distinction between the lateral root cap and epidermis and suggest that lateral root cap fate is the default state. It also suggests that TRN1 and TRN2 repress lateral root cap fate in cells in the epidermal location. Furthermore, the position-dependent pattern of root hair and non-root hair cell differentiation in the epidermis is defective in trn1 and trn2 mutants. Together these results indicate that TRN1 and TRN2 are required for the maintenance of both the radial pattern of tissue differentiation in the root and for the subsequent circumferential pattern within the epidermis.     

TRIPTYCHON,not CAPRICE,participates in feedback regulation of SCM expression in the Arabidopsis root epidermis

The Arabidopsis root epidermal cells decide their fates (root-hair cell and non-hair cell) according to their position. SCRAMBLED (SCM), an atypical leucine-rich repeat receptor-like kinase (LRR RLK) mediates the positional information to the epidermal cells enabling them to adopt the proper fate. Via feedback regulation, the SCM protein accumulates preferentially in cells adopting the root-hair cell fate. In this study, we determine that TRY, but not the related factor CPC, is responsible for this preferential SCM accumulation. We observed severe reduction of SCM::GUS expression in the try-82 mutant root, but not in the cpc-1 mutant. Furthermore, the overexpression of TRY by CaMV35S promoter caused an increase in the expression of SCM::GUS in the root epidermis. Intriguingly, the overexpression of CPC by CaMV35S promoter repressed the expression of SCM::GUS. Together, these results suggest that TRY plays a unique role in generating the appropriate spatial expression of SCM.     

A Common Position-Dependent Mechanism Controls Cell-Type Patterning and GLABRA2 Regulation in the Root and Hypocotyl Epidermis of Arabidopsis

A position-dependent pattern of epidermal cell types is produced during root development in Arabidopsis thaliana. This pattern is reflected in the expression pattern of GLABRA2 (GL2), a homeobox gene that regulates cell differentiation in the root epidermis. GL2 promoter::GUS fusions were used to show that the TTG gene, a regulator of root epidermis development, is necessary for maximal GL2 activity but is not required for the pattern of GL2 expression. Furthermore, GL2-promoter activity is influenced by expression of the myc-like maize R gene (35S::R) in Arabidopsis but is not affected by gl2 mutations. A position-dependent pattern of cell differentiation and GL2-promoter activity was also discovered in the hypocotyl epidermis that was analogous to the pattern in the root. Non-GL2-expressing cell files in the hypocotyl epidermis located outside anticlinal cortical cell walls exhibit reduced cell length and form stomata. Like the root, the hypocotyl GL2 activity was shown to be influenced by ttg and 35S::R but not by gl2. The parallel pattern of cell differentiation in the root and hypocotyl indicates that TTG and GL2 participate in a common position-dependent mechanism to control cell-type patterning throughout the apical-basal axis of the Arabidopsis seedling.     

SABRE is required for stabilization of root hair patterning in Arabidopsis thaliana

Patterned differentiation of distinct cell types is essential for the development of multicellular organisms. The root epidermis of Arabidopsis thaliana is composed of alternating files of root hair and non‐hair cells and represents a model system for studying the control of cell‐fate acquisition. Epidermal cell fate is regulated by a network of genes that translate positional information from the underlying cortical cell layer into a specific pattern of differentiated cells. While much is known about the genes of this network, new players continue to be discovered. Here we show that the SABRE (SAB) gene, known to mediate microtubule organization, anisotropic cell growth and planar polarity, has an effect on root epidermal hair cell patterning. Loss of SAB function results in ectopic root hair formation and destabilizes the expression of cell fate and differentiation markers in the root epidermis, including expression of the WEREWOLF (WER) and GLABRA2 (GL2) genes. Double mutant analysis reveal that wer and caprice (cpc) mutants, defective in core components of the epidermal patterning pathway, genetically interact with sab. This suggests that SAB may act on epidermal patterning upstream of WER and CPC. Hence, we provide evidence for a role of SAB in root epidermal patterning by affecting cell‐fate stabilization. Our work opens the door for future studies addressing SAB‐dependent functions of the cytoskeleton during root epidermal patterning.     

Positional Signaling and Expression of ENHANCER OF TRY AND CPC1 Are Tuned to Increase Root Hair Density in Response to Phosphate Deficiency in Arabidopsis thaliana

Phosphate (Pi) deficiency induces a multitude of responses aimed at improving the acquisition of Pi, including an increased density of root hairs. To understand the mechanisms involved in Pi deficiency-induced alterations of the root hair phenotype in Arabidopsis (Arabidopsis thaliana), we analyzed the patterning and length of root epidermal cells under control and Pi-deficient conditions in wild-type plants and in four mutants defective in the expression of master regulators of cell fate, CAPRICE (CPC), ENHANCER OF TRY AND CPC 1 (ETC1), WEREWOLF (WER) and SCRAMBLED (SCM). From this analysis we deduced that the longitudinal cell length of root epidermal cells is dependent on the correct perception of a positional signal (‘cortical bias’) in both control and Pi-deficient plants; mutants defective in the receptor of the signal, SCM, produced short cells characteristic of root hair-forming cells (trichoblasts). Simulating the effect of cortical bias on the time-evolving probability of cell fate supports a scenario in which a compromised positional signal delays the time point at which non-hair cells opt out the default trichoblast pathway, resulting in short, trichoblast-like non-hair cells. Collectively, our data show that Pi-deficient plants increase root hair density by the formation of shorter cells, resulting in a higher frequency of hairs per unit root length, and additional trichoblast cell fate assignment via increased expression of ETC1.     

Distinct and overlapping roles of single-repeat MYB genes in root epidermal patterning

Cell specification in the root epidermis of Arabidopsis generates a position-dependent pattern of root-hair cells and non-hair cells. Here we conduct a comprehensive analysis of the five members of a single-repeat R3 MYB gene family, including CAPRICE (CPC), TRIPTYCHON (TRY), ENHANCER of TRY and CPC 1, 2, and 3 (ETC1, ETC2, and ETC3), and study their role and functional relationship in root epidermal cell specification. Based on genetic and expression analyses, CPC, TRY and ETC1, but not ETC2 or ETC3, promote the hair cell fate by inhibiting non-hair specification. Further, we find that single-repeat MYB activity is required for epidermal patterning throughout root development, beginning during embryogenesis. We also identify a novel regulatory interaction whereby GLABRA2 (GL2) promotes TRY (but not CPC or ETC1) expression in the root epidermis, which generates a second lateral inhibition feedback loop. Gene fusion experiments combining CPC regulatory elements with protein-coding regions of each single-repeat MYB gene suggest that all five proteins are functionally similar, although TRY and ETC2 exhibit distinctions from CPC/ETC1/ETC3. These results provide new insight into the function of these single-repeat MYBs and suggest that divergence of their regulatory sequences is largely responsible for their distinct roles in epidermal cell patterning.     

Induction of epidermal cell fate in Arabidopsis shoots

Land plants have evolved a cuticle-bearing epidermis to protect themselves from environmental stress and pathogen attack. Despite its important role, little is known about the molecular mechanisms regulating shoot epidermal cell identity. In a recent study, we found that the Arabidopsis thaliana ATML1 gene is possibly a master regulator of shoot epidermal cell fate. We revealed that ATML1 has the ability to confer shoot epidermis-related traits to non-epidermal cells of the seedlings. These data are consistent with the previous loss-of-function mutant analyses, which implied a positive role of ATML1 in epidermal cell differentiation. Importantly, ectopic epidermal cells induced in ATML1-overexpressing lines provide a novel tool to assess the intrinsic properties of epidermal cells and to study epistatic interactions among genes involved in epidermal/mesophyll differentiation. Using this system, we obtained data revealing that ATML1 negatively influenced mesophyll cell fate. In addition, we provided a working model of how division planes in epidermal cells are determined.     

植物根毛生长发育及分子调控机理

植物根毛是植物吸收营养的主要器官, 了解根毛的发生、发育及遗传规律, 能对植物的养分吸收研究提供有利依据。文章旨在介绍植物根毛形态发生特性、发育生长过程及分子调控机理的研究进展, 利用比较基因组学方法研究农作物根毛形态和功能, 及有目的性的对根生长发育进行调控提供参考。研究发现植物根毛发育有反馈侧向抑制(lateral inhibition with feedback)和位置决定模式(position-dependent pattern of cell differentiation)两种方式。拟南芥根表皮细胞是以位置方式决定毛或非毛细胞发育类型, 已成为研究植物细胞命运和分化的模型。目前, 已经鉴定出控制根毛发育的基因, 包括一些转录因子如MYB家族蛋白TRIPTYCHON(TRY)、CAPRICE(CPC)和basic Helix-Loop-Helix (bHLH)蛋白GLABRA3、ENHANCER OF GLABRA3(EGL3)及WD-repeat蛋白等基因。最后针对根毛研究前景提出展望。     

GEM,a Novel Factor in the Coordination of Cell Division to Cell Fate Decisions in the Arabidopsis Epidermis

Cell division and cell fate decisions are highly regulated processes that need to be coordinated both spatially and temporally for correct plant growth and development. Gaining a deeper molecular and cellular understanding of these links is especially relevant for plant biology since, unlike in animals, formation of new organs is a process that takes place after embryogenesis and continues throughout the entire plant lifespan. The recent identification of a novel factor, GEM, has provided a molecular framework that coordinates cell division to cell fate in the Arabidopsis epidermis. GEM is an inhibitor of cell division through interacting with CDT1, a DNA replication protein. It also inhibits the expression of the homeobox GLABRA2 (GL2) gene that determines the hair/non-hair fate and the pavement/trichome fate in the root and leaf epidermis, respectively. GEM seems to be crucial in controlling the balance of activating/repressing histone modifications at its target promoters.Key Words: cell division, cell cycle, cell fate, GEM, GLABRA2, CDT1, DNA replication, chromatin, histone methylation, gene expression, root hair, Arabidopsis, plant