Symplastic communication in organ formation and tissue patterning.

Communication between cells is a crucial step to coordinate organ formation and tissue patterning. In plants, the intercellular transport of metabolites and signalling molecules occur symplastically through membranous structures (named plasmodesmata) that traverse the cell wall to connect the cytoplasm and endoplasmic reticulum of neighbouring cells. This review aims to highlight the importance of symplastic communication in plant development. We revisit current literature reporting the effects of changing plasmodesmata in cell morphogenesis, organ initiation and meristem maintenance and comment on recent work involving the identification of novel plasmodesmata regulators and of mobile developmental proteins and RNA molecules. New opportunities for unravelling the dynamic regulation and function of plasmodesmata are also discussed.EPSRC (Grant ID: EP/MO27740/1)This is the author accepted manuscript. The final version is available from Elsevier via http://dx.doi.org/10.1016/j.pbi.2015.10.00

Differentiation of conductive cells: a matter of life and death

Two major conducting tissues in plants, phloem and xylem, are composed of highly specialized cell types adapted to long distance transport. Sieve elements (SEs) in the phloem display a thick cell wall, callose-rich sieve plates and low cytoplasmic density. SE differentiation is driven by selective autolysis combined with enucleation, after which the plasma membrane and some organelles are retained. By contrast, differentiation of xylem tracheary elements (TEs) involves complete clearance of the cellular components by programmed cell death followed by autolysis of the protoplast; this is accompanied by extensive deposition of lignin and cellulose in the cell wall. Emerging molecular data on TE and SE differentiation indicate a central role for NAC and MYB type transcription factors in both processes.The Y.H. laboratory is funded by the Academy of Finland Centre of Excellence programme, the Gatsby Foundation, the Biotechnology and Biological Sciences Research Council, the University of Helsinki, the European Research Council Advanced Investigator Grant Symdev (No. 323052) and Tekes (the Finnish Funding Agency for Technology and Innovation)

Strigolactone- and Karrikin-Independent SMXL Proteins Are Central Regulators of Phloem Formation

Plant stem cell niches, the meristems, require long-distance transport of energy metabolites and signaling molecules along the phloem tissue. However, currently it is unclear how specification of phloem cells is controlled. Here we show that the genes SUPPRESSOR OF MAX2 1-LIKE3 (SMXL3), SMXL4, and SMXL5 act as cell-autonomous key regulators of phloem formation in Arabidopsis thaliana. The three genes form an uncharacterized subclade of the SMXL gene family that mediates hormonal strigolactone and karrikin signaling. Strigolactones are endogenous signaling molecules regulating shoot and root branching [1] whereas exogenous karrikin molecules induce germination after wildfires [2]. Both activities depend on the F-box protein and SCF (Skp, Cullin, F-box) complex component MORE AXILLARY GROWTH2 (MAX2) [3-5]. Strigolactone and karrikin perception leads to MAX2-dependent degradation of distinct SMXL protein family members, which is key for mediating hormonal effects [6-12]. However, the nature of events immediately downstream of SMXL protein degradation and whether all SMXL proteins mediate strigolactone or karrikin signaling is unknown. In this study we demonstrate that, within the SMXL gene family, specifically SMXL3/4/5 deficiency results in strong defects in phloem formation, alteredsugar accumulation, and seedling lethality. By comparing protein stabilities, we show that SMXL3/4/5 proteins function differently to canonical strigolactone and karrikin signaling mediators, although being functionally interchangeable with those under low strigolactone/karrikin signaling conditions. Our observations reveal a fundamental mechanism of phloem formation and indicate that diversity of SMXL protein functions is essential for a steady fuelling of plant meristems.Peer reviewe

Transcriptional regulatory framework for vascular cambium development in Arabidopsis roots

Vascular cambium, a lateral plant meristem is a central producer of woody biomass. Although a few transcription factors (TFs) have been shown to regulate cambial activity1, the phenotypes of the corresponding loss-of-function mutants are relatively modest, highlighting our limited understanding of the underlying transcriptional regulation. Here, we utilize cambium cell-specific transcript profiling followed by a combination of TF network and genetic analyses to identify 62 novel TF genotypes displaying an array of cambial phenotypes. This approach culminated in virtual loss of cambial activity when both WUSCHEL-RELATED HOMEOBOX 4 (WOX4) and KNOTTED-like from Arabidopsis thaliana 1 (KNAT1; also known as BREVIPEDICELLUS (BP) were mutated, thereby unlocking the genetic redundancy in the regulation of cambium development. We also identified TFs with dual functions in cambial cell proliferation and xylem differentiation, including WOX4, SHORT VEGETATIVE PHASE (SVP) and PETAL LOSS (PTL). Using the TF network information, we combined overexpression of the cambial activator WOX4 and removal of the putative inhibitor PTL to engineer Arabidopsis for enhanced radial growth. This line also showed ectopic cambial activity, thus further highlighting the central roles of WOX4 and PTL in cambium development.This work was supported by Finnish Centre of Excellence in Molecular Biology of Primary Producers (Academy of Finland CoE program 2014-2019) decision #271832, the Gatsby Foundation (GAT3395/PR3)), the University of Helsinki (award 799992091) and the European Research Council Advanced Investigator Grant SYMDEV (No. 323052) to Y.H.; Academy of Finland (grants #132376, #266431, #271832), University of Helsinki HiLIFE fellowship to A.P.M.; National Research Foundation of Korea (2018R1A5A1023599 and 2016R1A2B2015258) to J-Y. L

Sieve Plate Pores in the Phloem and the Unknowns of Their Formation.

Sieve pores of the sieve plates connect neighboring sieve elements to form the conducting sieve tubes of the phloem. Sieve pores are critical for phloem function. From the 1950s onwards, when electron microscopes became increasingly available, the study of their formation had been a pillar of phloem research. More recent work on sieve elements instead has largely focused on sieve tube hydraulics, phylogeny, and eco-physiology. Additionally, advanced molecular and genetic tools available for the model species Arabidopsis thaliana helped decipher several key regulatory mechanisms of early phloem development. Yet, the downstream differentiation processes which form the conductive sieve tube are still largely unknown, and our understanding of sieve pore formation has only moderately progressed. Here, we summarize our current knowledge on sieve pore formation and present relevant recent advances in related fields such as sieve element evolution, physiology, and plasmodesmata formation.Swiss National Science Foundation (FNS; P2LAP3_170862); EMBO (ALTF 1167-2017); Finnish Centre of Excellence in Molecular Biology of Primary Producers (Academy of Finland CoE program 2014-2019) decision #271832; Gatsby Foundation (GAT3395/PR3); the National Science Foundation Biotechnology and Biological Sciences Research Council grant (BB/N013158/1); University of Helsinki (award 799992091); European Research Council Advanced Investigator Grant SYMDEV (No. 323052)

Programmed cell death: new role in trimming the root tips

How is a rapid cellular turnover of the lateral root cap achieved in plants to control cap size in the growing root tips? Downstream of ANAC033/SOMBRERO, a highly organized and temporally coordinated cell death program involving BFN1 nuclease-mediated rapid corpse clearance eliminates these cells

Plant vascular development - connective tissue connecting scientists: updates and trends at the PVB 2013 conference

In July 2013, the scientific community fascinated by plant vascular biology (PVB) was cordially welcomed to Helsinki, the capital of Finland, to discuss the latest advances in the field. This meeting was the Third International Conference on PVB, following the successful meetings organized in Taiwan and the United States. Approximately 200 scientists from all around the world participated in PVB2013, which was characterized by lively scientific discussions in a relaxed atmosphere. The meeting was rich in topics and disciplines covering vascular biology from the early embryo to the mature plant, from the root tip to aerial organs, all linked by an appreciation of the crucial connective role of vascular tissues in plant growth and development. In this report, we briefly summarize some of the contributions presented during PVB2013 focusing on the development of vascular tissues (Figs 1 and 2). This special issue also collects a number of exciting papers with a central theme of plant vascular development

Plant vascular biology 2013: vascular trafficking

About 200 researchers from around the world attended the Third International Conference on Plant Vascular Biology (PVB 2013) held in July 2013 at the Rantapuisto Conference Center, in Helsinki, Finland (http://www.pvb2013.org). The plant vascular system, which connects every organ in the mature plant, continues to attract the interest of researchers representing a wide range of disciplines, including development, physiology, systems biology, and computational biology. At the meeting, participants discussed the latest research advances in vascular development, long- and short-distance vascular transport and long-distance signalling in plant defence, in addition to providing a context for how these studies intersect with each other. The meeting provided an opportunity for researchers working across a broad range of fields to share ideas and to discuss future directions in the expanding field of vascular biology. In this report, the latest advances in understanding the mechanism of vascular trafficking presented at the meeting have been summarized

Molecular control of cell specification and cell differentiation during procambial development

Land plants develop vascular tissues that enable the long-distance transport of water and nutrients in xylem and phloem, provide mechanical support for their vertical growth, and produce cells in radial growth. Vascular tissues are produced in many parts of the plant and during different developmental stages. Early vascular development is focused in procambial meristems, and in some species it continues during the secondary phase of plant development in cambial meristems. In this review, we highlight recent progress in understanding procambial development. This involves the analysis of stem cell–like properties of procambial tissues, specification of xylem and phloem, and differentiation of the conductive tissues. Several major plant hormones, small-RNA species, and transcriptional networks play a role in vascular development. We describe current approaches to integrating these networks as well as their potential role in explaining the diversity and evolution of plant vascular systems

Structural Imaging of Native Cryo-Preserved Secondary Cell Walls Reveals the Presence of Macrofibrils and Their Formation Requires Normal Cellulose, Lignin and Xylan Biosynthesis.

The woody secondary cell walls of plants are the largest repository of renewable carbon biopolymers on the planet. These walls are made principally from cellulose and hemicelluloses and are impregnated with lignin. Despite their importance as the main load bearing structure for plant growth, as well as their industrial importance as both a material and energy source, the precise arrangement of these constituents within the cell wall is not yet fully understood. We have adapted low temperature scanning electron microscopy (cryo-SEM) for imaging the nanoscale architecture of angiosperm and gymnosperm cell walls in their native hydrated state. Our work confirms that cell wall macrofibrils, cylindrical structures with a diameter exceeding 10 nm, are a common feature of the native hardwood and softwood samples. We have observed these same structures in Arabidopsis thaliana secondary cell walls, enabling macrofibrils to be compared between mutant lines that are perturbed in cellulose, hemicellulose, and lignin formation. Our analysis indicates that the macrofibrils in Arabidopsis cell walls are dependent upon the proper biosynthesis, or composed, of cellulose, xylan, and lignin. This study establishes that cryo-SEM is a useful additional approach for investigating the native nanoscale architecture and composition of hardwood and softwood secondary cell walls and demonstrates the applicability of Arabidopsis genetic resources to relate fibril structure with wall composition and biosynthesis.This work was supported by the Leverhulme Trust Centre for Natural Material Innovation. Analysis of Arabidopsis mutant plants was supported as part of The Center for LignoCellulose Structure and Formation, an Energy Frontier Research Center funded by the U.S. Department of Energy (DOE), Office of Science, Basic Energy Sciences (BES), under Award # DE-SC0001090. JJL was in receipt of a studentship from the Biotechnology and Biological Sciences Research Council (BBSRC) of the UK as part of the Cambridge BBSRC-DTP Programme (Reference BB/J014540/1). JJL is currently supported by a grant from the National Science Centre, Poland as part of the SONATINA 3 programme (project number 2019/32/C/NZ3/00392). MB is employed in YH’s team through the European Research Council Advanced Investigator Grant SYMDEV (No. 323052). YH laboratory is funded by the Finnish Centre of Excellence in Molecular Biology of Primary Producers (Academy of Finland CoE program 2014-2019) (decision #271832); the Gatsby Foundation (GAT3395/PR3); the National Science Foundation Biotechnology and Biological Sciences Research Council grant (BB/N013158/1); University of Helsinki (award 799992091), and the European Research Council Advanced Investigator Grant SYMDEV (No. 323052). OMT was a recipient of an iCASE studentship from the BBSRC (Reference BB/M015432/1). The cryo-SEM facility at the Sainsbury Laboratory is supported by the Gatsby Charitable Foundatio