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      Pollen differentiation as well as pollen tube guidance and discharge are independent of the presence of gametes

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          ABSTRACT

          After meiosis, an unequal cell division generates the male gamete lineage in flowering plants. The generative cell will undergo a second division, giving rise to the two gametes, i.e. the sperm cells. The other cell will develop into the vegetative cell that plays a crucial role in pollen tube formation and sperm delivery. Recently, the vegetative cell has been suggested to be important for programming of the chromatin state in sperm cells and/or the resulting fertilization products. Blocking the initial unequal division genetically, we first highlight that the default differentiation state after male meiosis is a vegetative fate, which is consistent with earlier work. We find that uni-nucleated mutant microspores differentiated as wild-type vegetative cells, including chromatin remodeling and the transcriptional activation of transposable elements. Moreover, live-cell imaging revealed that this vegetative cell is sufficient for the correct guidance of the pollen tube to the female gametes. Hence, we conclude that vegetative cell differentiation and function does not depend on the formation or presence of the actual gametes but rather on external signals or a cell-autonomous pace keeper.

          Abstract

          Summary: Cell biological analyses in Arabidopsis show that the vegetative cell differentiates without the presence of the actual gametes, and is solely sufficient for pollen tube germination, guidance, ovule penetration and pollen tube discharge.

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          Molecular mechanisms underlying RB protein function.

          Inactivation of the RB protein is one of the most fundamental events in cancer. Coming to a molecular understanding of its function in normal cells and how it impedes cancer development has been challenging. Historically, the ability of RB to regulate the cell cycle placed it in a central role in proliferative control, and research focused on RB regulation of the E2F family of transcription factors. Remarkably, several recent studies have found additional tumour-suppressor functions of RB, including alternative roles in the cell cycle, maintenance of genome stability and apoptosis. These advances and new structural studies are combining to define the multifunctionality of RB.
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            Active DNA demethylation in plant companion cells reinforces transposon methylation in gametes.

            The Arabidopsis thaliana central cell, the companion cell of the egg, undergoes DNA demethylation before fertilization, but the targeting preferences, mechanism, and biological significance of this process remain unclear. Here, we show that active DNA demethylation mediated by the DEMETER DNA glycosylase accounts for all of the demethylation in the central cell and preferentially targets small, AT-rich, and nucleosome-depleted euchromatic transposable elements. The vegetative cell, the companion cell of sperm, also undergoes DEMETER-dependent demethylation of similar sequences, and lack of DEMETER in vegetative cells causes reduced small RNA-directed DNA methylation of transposons in sperm. Our results demonstrate that demethylation in companion cells reinforces transposon methylation in plant gametes and likely contributes to stable silencing of transposable elements across generations.
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              Control of male gametophyte development.

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                Author and article information

                Journal
                Development
                Development
                DEV
                develop
                Development (Cambridge, England)
                The Company of Biologists Ltd
                0950-1991
                1477-9129
                1 January 2018
                1 January 2018
                : 145
                : 1
                : dev152645
                Affiliations
                [1 ]Department of Molecular Mechanisms of Phenotypic Plasticity, Institut de Biologie Moléculaire des Plantes du CNRS, IBMP-CNRS - UPR2357, Université de Strasbourg , 12 Rue du Général Zimmer, 67084 Strasbourg Cedex, France
                [2 ]Section for Genetics and Evolutionary Biology, Department of Biosciences, University of Oslo , 0316 Oslo, Norway
                [3 ]University of Hamburg, Biozentrum Klein Flottbek , Department of Developmental Biology, 22609 Hamburg, Germany
                [4 ]Institute of Transformative Bio-Molecules (WPI-ITbM), Nagoya University , Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8601, Japan
                [5 ]JST ERATO Higashiyama Live-Holonics Project, Nagoya University , Furo-cho, Chikusa-ku, Nagoya, Aichi 464-8602, Japan
                Author notes
                [*]

                These authors contributed equally to this work

                []Author for correspondence ( arp.schnittger@ 123456uni-hamburg.de )
                Author information
                http://orcid.org/0000-0002-1247-4612
                http://orcid.org/0000-0002-4156-2056
                http://orcid.org/0000-0001-7067-0091
                Article
                DEV152645
                10.1242/dev.152645
                5825867
                29217755
                82733f98-8dc5-4a49-8b24-83ea77e2aec6
                © 2018. Published by The Company of Biologists Ltd

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( http://creativecommons.org/licenses/by/3.0), which permits unrestricted use, distribution and reproduction in any medium provided that the original work is properly attributed.

                History
                : 28 March 2017
                : 27 November 2017
                Funding
                Funded by: Studienstiftung des Deutschen Volkes, http://dx.doi.org/10.13039/501100004350;
                Funded by: Freie und Hansestadt Hamburg;
                Funded by: Japan Society for the Promotion of Science, http://dx.doi.org/10.13039/501100001691;
                Award ID: 13J09285
                Funded by: Ministry of Education, Culture, Sports, Science and Technology, http://dx.doi.org/10.13039/501100001700;
                Award ID: 16H06465
                Award ID: 16H06464
                Award ID: 16K21727
                Funded by: Japan Advanced Plant Science Network;
                Funded by: European Research Council, http://dx.doi.org/10.13039/100010663;
                Funded by: Norges forskningsråd, http://dx.doi.org/10.13039/501100005416;
                Award ID: 214052/F20
                Categories
                201
                Research Report

                Developmental biology
                arabidopsis,gametophyte,fertilization,chromatin,transposons
                Developmental biology
                arabidopsis, gametophyte, fertilization, chromatin, transposons

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