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      Studying Secondary Growth and Bast Fiber Development: The Hemp Hypocotyl Peeks behind the Wall

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          Abstract

          Cannabis sativa L. is an annual herbaceous crop grown for the production of long extraxylary fibers, the bast fibers, rich in cellulose and used both in the textile and biocomposite sectors. Despite being herbaceous, hemp undergoes secondary growth and this is well exemplified by the hypocotyl. The hypocotyl was already shown to be a suitable model to study secondary growth in other herbaceous species, namely Arabidopsis thaliana and it shows an important practical advantage, i.e., elongation and radial thickening are temporally separated. This study focuses on the mechanisms marking the transition from primary to secondary growth in the hemp hypocotyl by analysing the suite of events accompanying vascular tissue and bast fiber development. Transcriptomics, imaging and quantification of phytohormones were carried out on four representative developmental stages (i.e., 6–9–15–20 days after sowing) to provide a comprehensive overview of the events associated with primary and secondary growth in hemp. This multidisciplinary approach provides cell wall-related snapshots of the growing hemp hypocotyl and identifies marker genes associated with the young (expansins, β-galactosidases, and transcription factors involved in light-related processes) and the older hypocotyl (secondary cell wall biosynthetic genes and transcription factors).

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          Most cited references72

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          PlantTFDB 3.0: a portal for the functional and evolutionary study of plant transcription factors

          With the aim to provide a resource for functional and evolutionary study of plant transcription factors (TFs), we updated the plant TF database PlantTFDB to version 3.0 (http://planttfdb.cbi.pku.edu.cn). After refining the TF classification pipeline, we systematically identified 129 288 TFs from 83 species, of which 67 species have genome sequences, covering main lineages of green plants. Besides the abundant annotation provided in the previous version, we generated more annotations for identified TFs, including expression, regulation, interaction, conserved elements, phenotype information, expert-curated descriptions derived from UniProt, TAIR and NCBI GeneRIF, as well as references to provide clues for functional studies of TFs. To help identify evolutionary relationship among identified TFs, we assigned 69 450 TFs into 3924 orthologous groups, and constructed 9217 phylogenetic trees for TFs within the same families or same orthologous groups, respectively. In addition, we set up a TF prediction server in this version for users to identify TFs from their own sequences.
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            Cytokinins: activity, biosynthesis, and translocation.

            Cytokinins (CKs) play a crucial role in various phases of plant growth and development, but the basic molecular mechanisms of their biosynthesis and signal transduction only recently became clear. The progress was achieved by identifying a series of key genes encoding enzymes and proteins controlling critical steps in biosynthesis, translocation, and signaling. Basic schemes for CK homeostasis and root/shoot communication at the whole-plant level can now be devised. This review summarizes recent findings on the relationship between CK structural variation and activity, distinct features in CK biosynthesis between higher plants and Agrobacterium infected plants, CK translocation at whole-plant and cellular levels, and CKs as signaling molecules for nutrient status via root-shoot communication.
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              A battery of transcription factors involved in the regulation of secondary cell wall biosynthesis in Arabidopsis.

              SECONDARY WALL-ASSOCIATED NAC DOMAIN PROTEIN1 (SND1) is a master transcriptional switch activating the developmental program of secondary wall biosynthesis. Here, we demonstrate that a battery of SND1-regulated transcription factors is required for normal secondary wall biosynthesis in Arabidopsis thaliana. The expression of 11 SND1-regulated transcription factors, namely, SND2, SND3, MYB103, MYB85, MYB52, MYB54, MYB69, MYB42, MYB43, MYB20, and KNAT7 (a Knotted1-like homeodomain protein), was developmentally associated with cells undergoing secondary wall thickening. Of these, dominant repression of SND2, SND3, MYB103, MYB85, MYB52, MYB54, and KNAT7 significantly reduced secondary wall thickening in fiber cells. Overexpression of SND2, SND3, and MYB103 increased secondary wall thickening in fibers, and overexpression of MYB85 led to ectopic deposition of lignin in epidermal and cortical cells in stems. Furthermore, SND2, SND3, MYB103, MYB85, MYB52, and MYB54 were able to induce secondary wall biosynthetic genes. Direct target analysis using the estrogen-inducible system revealed that MYB46, SND3, MYB103, and KNAT7 were direct targets of SND1 and also of its close homologs, NST1, NST2, and vessel-specific VND6 and VND7. Together, these results demonstrate that a transcriptional network consisting of SND1 and its downstream targets is involved in regulating secondary wall biosynthesis in fibers and that NST1, NST2, VND6, and VND7 are functional homologs of SND1 that regulate the same downstream targets in different cell types.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                18 November 2016
                2016
                : 7
                : 1733
                Affiliations
                [1] 1Environmental Research and Innovation Department, Luxembourg Institute of Science and Technology Esch-sur-Alzette, Luxembourg
                [2] 2Groupe de Recherche en Physiologie Végétale, Earth and Life Institute-Agronomy, Université catholique de Louvain Louvain-la-Neuve, Belgium
                [3] 3Institute of Experimental Botany, The Czech Academy of Sciences Prague, Czechia
                Author notes

                Edited by: Nicholas Provart, University of Toronto, Canada

                Reviewed by: Jedrzej Jakub Szymanski, Weizmann Institute of Science, Israel; Jake Stout, University of Manitoba, Canada

                *Correspondence: Gea Guerriero, gea.guerriero@ 123456list.lu

                This article was submitted to Plant Systems and Synthetic Biology, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2016.01733
                5114303
                27917184
                6027b815-0660-4674-be62-226a7e37053c
                Copyright © 2016 Behr, Legay, Žižková, Motyka, Dobrev, Hausman, Lutts and Guerriero.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 10 August 2016
                : 03 November 2016
                Page count
                Figures: 9, Tables: 0, Equations: 0, References: 93, Pages: 19, Words: 0
                Funding
                Funded by: Fonds National de la Recherche Luxembourg 10.13039/501100001866
                Award ID: CANCAN C13/SR/5774202
                Funded by: Grantová Agentura České Republiky 10.13039/501100001824
                Award ID: 16-14649S
                Categories
                Plant Science
                Original Research

                Plant science & Botany
                cannabis sativa,hypocotyl,bast fibers,cell wall,rna-seq,immunohistochemistry,phytohormones

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