Multiple Roles of Brassinosteroid Signaling in Vascular Development.

Brassinosteroids (BRs) are plant steroid hormones that control growth and stress responses. In the context of development, BRs play diverse roles in controlling cell differentiation and tissue patterning. The vascular system, which is essential for transporting water and nutrients throughout the plant body, initially establishes a tissue pattern during primary development and then dramatically increases the number of vascular cells during secondary development.

LONESOME HIGHWAY-TARGET OF MONOPTEROS5 transcription factor complex promotes a predifferentiation state for xylem vessel differentiation in the root apical meristem by inducing the expression of VASCULAR-RELATED NAC-DOMAIN genes.

In Arabidopsis thaliana, heterodimers comprising two bHLH family proteins, LONESOME HIGHWAY (LHW) and TARGET OF MONOPTEROS5 (TMO5) or its homolog TMO5-LIKE 1 (T5L1) control vascular development in the root apical meristem (RAM). The LHW-TMO5/T5L1 complex regulates vascular cell proliferation, vascular pattern organization, and xylem vessel differentiation; however, the mechanism of preparation for xylem vessel differentiation in the RAM remains elusive. We examined the relationship between LHW-T5L1 and VASCULAR-RELATED NAC-DOMAIN (VND) genes, which are key regulators of vessel differentiation, using reverse genetics approaches.

HD-ZIP III-dependent local promotion of brassinosteroid synthesis suppresses vascular cell division in Arabidopsis root apical meristem.

Spatiotemporal control of cell division in the meristem is vital for plant growth. In the stele of the root apical meristem (RAM), procambial cells divide periclinally to increase the number of vascular cell files. Class III homeodomain leucine zipper (HD-ZIP III) proteins are key transcriptional regulators of RAM development and suppress the periclinal division of vascular cells in the stele; however, the mechanism underlying the regulation of vascular cell division by HD-ZIP III transcription factors (TFs) remains largely unknown.

Transcriptional networks regulating root vascular development.

Vascular development involves multiple processes, including the establishment of vascular stem cells (e.g. procambium/cambium cells), stem cell divisions, and cell specification.

A Positive Feedback Loop Comprising LHW-TMO5 and Local Auxin Biosynthesis Regulates Initial Vascular Development in Arabidopsis Roots.

The phytohormone auxin governs various developmental processes in plants including vascular formation. Auxin transport and biosynthesis are important factors in determining auxin distribution in tissues. Although the role of auxin transport in vein pattern formation is widely recognized, that of auxin biosynthesis in vascular development is poorly understood.

Vascular tissue development in plants.

The plant vasculature is a sophisticated system that has greatly contributed to the evolution of land plants over the past few hundred million years. The formation of the vascular system is a well-organized plant developmental process, but it is also flexible in response to environmental changes. Provascular cells arise after asymmetric cell division in early embryos and differentiate into various vascular cells, including procambial cells, which function as vascular stem cells.

LOB DOMAIN-CONTAINING PROTEIN 15 Positively Regulates Expression of VND7, a Master Regulator of Tracheary Elements.

Xylem includes xylem parenchyma cells, fibers and tracheary elements. Differentiation of tracheary elements is an irreversible process that is controlled by the master regulator VASCULAR-RELATED NAC-DOMAIN 7 (VND7). Molecular events occurring downstream of VND7 are well understood, but little is known regarding upstream regulation of VND7.

Practical Synthesis of Spermine, Thermospermine and Norspermine.

Polyamines, such as spermine (1), thermospermine (2) and norspermine (3), are widely distributed in nature, and have multiple biological activities. In addition, many of their conjugates have potential for pharmacological use. Here, we present a solid-phase synthesis using our nitrobenzenesulfonyl (Ns) strategy, which can provide 1, 2 and 3 on a gram scale.

Functional mechanism of bHLH complexes during early vascular development.

The vascular system spreads throughout the plant body. This highly organized network contains several types of cells. Vascular cell development is initiated during embryogenesis, and then vascular cells proliferate, form a vascular pattern, and commit to specific cell fates.

A Negative Feedback Loop Controlling bHLH Complexes Is Involved in Vascular Cell Division and Differentiation in the Root Apical Meristem.

Controlling cell division and differentiation in meristems is essential for proper plant growth. Two bHLH heterodimers consisting of LONESOME HIGHWAY (LHW) and TARGET OF MONOPTEROS 5 (TMO5)/TMO5-LIKE1 (T5L1) regulate periclinal cell division in vascular cells in the root apical meristem (RAM). In this study, we further investigated the functions of LHW-T5L1, finding that in addition to controlling cell division, this complex regulates xylem differentiation in the RAM via a novel negative regulatory system.

A bHLH complex activates vascular cell division via cytokinin action in root apical meristem.

Higher organisms possess mechanisms to maintain stem cells' proliferative and pluripotent states in stem cell niches [1]. Plants possess two types of stem cell niches in the root and shoot apical meristems, where regulatory interactions exist between stem cells and organizing cells. Recent studies provided new insights into the molecular mechanism of stem cell maintenance [2-4].

Initiation of vascular development.

The initiation of vascular development occurs during embryogenesis and the development of lateral organs, such as lateral roots and leaves. Understanding the mechanism underlying the initiation of vascular development has been an important goal of plant biologists. Auxin flow is a crucial factor involved in the initiation of vascular development.

An atypical bHLH transcription factor regulates early xylem development downstream of auxin.

The vascular system in plants, which comprises xylem, phloem and vascular stem cells, originates from provascular cells and forms a continuous network throughout the plant body. Although various aspects of vascular development have been extensively studied, the early process of vascular development remains largely unknown. LONESOME HIGHWAY (LHW), which encodes an atypical basic helix-loop-helix (bHLH) transcription factor, plays an essential role in establishing vascular cells.

A novel function of TDIF-related peptides: promotion of axillary bud formation.

Small peptides derived from the CLAVATA3/EMBRYO SURROUNDING REGION-related (CLE) gene family play a key role in various cell-cell communications in land plants. Among them, tracheary element differentiation inhibition factor (TDIF; CLE41/CLE44 peptide) and CLE42 peptide of Arabidopsis have almost identical amino acid sequences and act as inhibitors of tracheary element differentiation. In this study, we report a novel function of TDIF and CLE42.

Differentiation of Arabidopsis guard cells: analysis of the networks incorporating the basic helix-loop-helix transcription factor, FAMA.

Nearly all extant land plants possess stomata, the epidermal structures that mediate gas exchange between the plant and the environment. The developmental pathways, cell division patterns, and molecules employed in the generation of these structures are simple examples of processes used in many developmental contexts. One specific module is a set of "master regulator" basic helix-loop-helix transcription factors that regulate individual consecutive steps in stomatal development.

Arabidopsis VASCULAR-RELATED NAC-DOMAIN6 directly regulates the genes that govern programmed cell death and secondary wall formation during xylem differentiation.

Xylem consists of three types of cells: tracheary elements (TEs), parenchyma cells, and fiber cells. TE differentiation includes two essential processes, programmed cell death (PCD) and secondary cell wall formation. These two processes are tightly coupled.

Transcriptional regulation of vascular cell fates.

In vascular development, uncommitted cells differentiate into different xylem cells through vascular stem cells, such as procambial cells, during vein formation as well as embryogenesis. Cascades of transcriptional regulation of genes play crucial roles in the progress of vascular development. Auxin, cytokinin, and brassinosteroids also function in procambial cell determination, procambial maintenance, and xylem cell differentiation from procambial cells, respectively, through transcriptional regulation.

Orthologs of Arabidopsis thaliana stomatal bHLH genes and regulation of stomatal development in grasses.

Stomata are adjustable pores in the plant epidermis that regulate gas exchange between the plant and atmosphere; they are present on the aerial portions of most higher plants. Genetic pathways controlling stomatal development and distribution have been described in some detail for one dicot species, Arabidopsis, in which three paralogous bHLH transcription factors, FAMA, MUTE and SPCH, control discrete sequential stages in stomatal development. Orthologs of FAMA, MUTE and SPCH are present in other flowering plants.

Non-cell-autonomous control of vascular stem cell fate by a CLE peptide/receptor system.

Land plants evolved a long-distance transport system of water and nutrients composed of the xylem and phloem, both of which are generated from the procambium- and cambium-comprising vascular stem cells. However, little is known about the molecular mechanism of cell communication governing xylem-phloem patterning. Here, we show that a dodecapeptide (HEVHypSGHypNPISN; Hyp, 4-hydroxyproline), TDIF (tracheary element differentiation inhibitory factor), is secreted from the phloem and suppresses the differentiation of vascular stem cells into xylem cells through a leucine-rich repeat receptor-like kinase (LRR-RLK).

Regulation of the Arabidopsis root vascular initial population by LONESOME HIGHWAY.

Complex organisms consist of a multitude of cell types arranged in a precise spatial relation to each other. Arabidopsis roots generally exhibit radial tissue organization; however, within a tissue layer, cells are not identical. Specific vascular cell types are arranged in diametrically opposed longitudinal files that maximize the distance between them and create a bilaterally symmetric (diarch) root.

Transcription factor control of asymmetric cell divisions that establish the stomatal lineage.

The establishment of new cell lineages during development often requires a symmetry-breaking event. An asymmetric division in the epidermis of plants initiates a lineage that ultimately produces stomatal guard cells. Stomata are pores in the epidermis that serve as the main conduits for gas exchange between plants and the atmosphere; they are critical for photosynthesis and exert a major influence on global carbon and water cycles.