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      A transcriptional view on somatic embryogenesis

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          Abstract

          Somatic embryogenesis is a form of induced plant cell totipotency where embryos develop from somatic or vegetative cells in the absence of fertilization. Somatic embryogenesis can be induced in vitro by exposing explants to stress or growth regulator treatments. Molecular genetics studies have also shown that ectopic expression of specific embryo‐ and meristem‐expressed transcription factors or loss of certain chromatin‐modifying proteins induces spontaneous somatic embryogenesis. We begin this review with a general description of the major developmental events that define plant somatic embryogenesis and then focus on the transcriptional regulation of this process in the model plant Arabidopsis thaliana (arabidopsis). We describe the different somatic embryogenesis systems developed for arabidopsis and discuss the roles of transcription factors and chromatin modifications in this process. We describe how these somatic embryogenesis factors are interconnected and how their pathways converge at the level of hormones. Furthermore, the similarities between the developmental pathways in hormone‐ and transcription‐factor‐induced tissue culture systems are reviewed in the light of our recent findings on the somatic embryo‐inducing transcription factor BABY BOOM.

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          Most cited references118

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          The Arabidopsis NFYA5 transcription factor is regulated transcriptionally and posttranscriptionally to promote drought resistance.

          Nuclear factor Y (NF-Y) is a ubiquitous transcription factor composed of three distinct subunits (NF-YA, NF-YB, and NF-YC). We found that the Arabidopsis thaliana NFYA5 transcript is strongly induced by drought stress in an abscisic acid (ABA)-dependent manner. Promoter:beta-glucuronidase analyses showed that NFYA5 was highly expressed in vascular tissues and guard cells and that part of the induction by drought was transcriptional. NFYA5 contains a target site for miR169, which targets mRNAs for cleavage or translational repression. We found that miR169 was downregulated by drought stress through an ABA-dependent pathway. Analysis of the expression of miR169 precursors showed that miR169a and miR169c were substantially downregulated by drought stress. Coexpression of miR169 and NFYA5 suggested that miR169a was more efficient than miR169c at repressing the NFYA5 mRNA level. nfya5 knockout plants and plants overexpressing miR169a showed enhanced leaf water loss and were more sensitive to drought stress than wild-type plants. By contrast, transgenic Arabidopsis plants overexpressing NFYA5 displayed reduced leaf water loss and were more resistant to drought stress than the wild type. Microarray analysis indicated that NFYA5 is crucial for the expression of a number of drought stress-responsive genes. Thus, NFYA5 is important for drought resistance, and its induction by drought stress occurs at both the transcriptional and posttranscriptional levels.
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            The PLETHORA genes mediate patterning of the Arabidopsis root stem cell niche.

            A small organizing center, the quiescent center (QC), maintains stem cells in the Arabidopsis root and defines the stem cell niche. The phytohormone auxin influences the position of this niche by an unknown mechanism. Here, we identify the PLETHORA1 (PLT1) and PLT2 genes encoding AP2 class putative transcription factors, which are essential for QC specification and stem cell activity. The PLT genes are transcribed in response to auxin accumulation and are dependent on auxin response transcription factors. Distal PLT transcript accumulation creates an overlap with the radial expression domains of SHORT-ROOT and SCARECROW, providing positional information for the stem cell niche. Furthermore, the PLT genes are activated in the basal embryo region that gives rise to hypocotyl, root, and root stem cells and, when ectopically expressed, transform apical regions to these identities. Thus, the PLT genes are key effectors for establishment of the stem cell niche during embryonic pattern formation.
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              PLETHORA proteins as dose-dependent master regulators of Arabidopsis root development.

              Factors with a graded distribution can program fields of cells in a dose-dependent manner, but no evidence has hitherto surfaced for such mechanisms in plants. In the Arabidopsis thaliana root, two PLETHORA (PLT) genes encoding AP2-domain transcription factors have been shown to maintain the activity of stem cells. Here we show that a clade of four PLT homologues is necessary for root formation. Promoter activity and protein fusions of PLT homologues display gradient distributions with maxima in the stem cell area. PLT activities are largely additive and dosage dependent. High levels of PLT activity promote stem cell identity and maintenance; lower levels promote mitotic activity of stem cell daughters; and further reduction in levels is required for cell differentiation. Our findings indicate that PLT protein dosage is translated into distinct cellular responses.
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                Author and article information

                Contributors
                kim.boutilier@wur.nl
                Journal
                Regeneration (Oxf)
                Regeneration (Oxf)
                10.1002/(ISSN)2052-4412
                REG2
                Regeneration
                John Wiley and Sons Inc. (Hoboken )
                2052-4412
                05 December 2017
                August 2017
                : 4
                : 4 ( doiID: 10.1002/reg2.2017.4.issue-4 )
                : 201-216
                Affiliations
                [ 1 ] Bioscience Wageningen University and Research Wageningen The Netherlands
                [ 2 ] Laboratory of Molecular Biology Wageningen University and Research Wageningen The Netherlands
                Author notes
                [*] [* ] Correspondence

                Kim Boutilier, Bioscience, Wageningen University and Research, PO Box 16, 6700 AA Wageningen, The Netherlands.

                Email: kim.boutilier@ 123456wur.nl

                Author information
                http://orcid.org/0000-0003-2971-2047
                http://orcid.org/0000-0001-9285-7199
                http://orcid.org/0000-0001-6110-5939
                Article
                REG291
                10.1002/reg2.91
                5743784
                29299323
                c6d3a1f9-3067-4a39-8a10-f37640e3a982
                © 2017 The Authors. Regeneration published by John Wiley & Sons Ltd.

                This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 22 July 2016
                : 15 September 2017
                : 04 October 2017
                Page count
                Figures: 3, Tables: 0, Pages: 16, Words: 14264
                Categories
                Review
                Review
                Custom metadata
                2.0
                reg291
                August 2017
                Converter:WILEY_ML3GV2_TO_NLMPMC version:5.2.8 mode:remove_FC converted:26.12.2017

                chromatin modifications,regeneration,somatic embryogenesis,totipotency,transcription factors

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