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Funktion Und Evolution Pflanzlicher B-Chromosomen

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Funktion und Evolution pflanzlicher B-Chromosomen Habilitationsschrift zur Erlangung des akademischen Grades doctor agriculturarum habilitatus (Dr. agr. habil.) verteidigt am 14.12.2009 an der Naturwissenschaftlichen Fakultät III der Martin-Luther-Universität Halle-Wittenberg von Dr. agr. Andreas Houben geb. am 06.05.1963 in Magdeburg Gutachter: Prof. Dr. Klaus Pillen (Martin-Luther Universität Halle-Wittenber) Prof. Dr. Thomas Schmidt (Technische Universität Dresden) Prof. Dr. Christian Jung (Christian-Albrechts-Universität zu Kiel) Dekan der Naturwissenschaftlichen Fakultät III Prof. Dr. Peter Wycisk Inhaltsverzeichnis Seite 1. Einleitung und Fragestellung 4 2. Eigene Arbeiten (A1-A20) 7 3. Zusammenfassende Diskussion 29 3.1. Ursprung und Evolution von B-Chromosomen 29 3.2. A- und B-Chromosomen können sich in der Verteilung 33 von Eu- und Heterochromatin-typischen Histonveränderungen unterscheiden 3.3. Meiotische B-Chromosomen zeigen eine veränderte 36 Zellzyklus-abhängige Histon H3-Phosphorylierung 3.4. Segregationsverhalten und Zentromere von 37 B-Chromosomen 3.5. B-Chromosomen zeigen eine schwache 39 Transkriptionsaktivität 3.6. Brachycome dichromosomatica, eine Modellart für die 40 Analyse von B-Chromosomen 3.7. Potentielle Anwendung von B-Chromosomen in der 43 Biotechnologie 4. Zusammenfassung (in Deutsch und Englisch) 44 5. Literaturverzeichnis 52 Danksagung 58 2 Abkürzungsverzeichnis bp Basenpaar Bs B-Chromosomen 2C DNA-Gehalt des diploiden, unreplizierten Chromosomensatzes DAPI 4’,6’-Diamidino-2-Phenylindol FISH Fluoreszenz-in-situ-Hybridisierung G1 Zellzyklusphase G1 G2 Zellzyklusphase G2 ITS2 Internal-Transcribed-Spacer 2 K4 Lysin 4 K9 Lysin 9 K20 Lysin 20 K27 Lysin 27 H3 Histon H3 H4 Histon H4 mB Mikro-B-Chromosomen Mbp Megabasenpaare me methyliert me1 monomethyliert me2 dimethyliert me3 trimethyliert nB Standard-B-Chromosomen NOR Nukleolusorganisatorregion rDNA ribosomale DNA rRNA ribosomale RNA PCR Polymerase-Kettenreaktion ph Phorphorylierung RT-PCR Reverse-Transcriptase-PCR S10 Serin 10 S28 Serin 28 T3 Threonin 3, T11 Threonin 11 T32 Threonin 32 3 1. Einleitung und Fragestellung Chromosomen portionieren das Genom und ermöglichen die korrekte Weitergabe der Zellkern-lokalisierten Erbinformation. Die Genomgröße und Anzahl der Chromosomen innerhalb einer Spezies sind weitgehend konstant. Häufig ist aber die Artenevolution mit einer numerischen und strukturellen Karyotypveränderung verbunden, wodurch sich die Genomgröße und Anzahl von Chromosomen einzelner Individuen einer Art unterscheiden kann. Eine Vielzahl von pflanzlichen und tierischen Arten besitzen neben den normalen Standardchromosomen oder auch A-Chromosomen genannt, überzählige sogenannte B-Chromosomen. Diese Zusatzchromosomen unterscheiden sich von den A-Chromosomen dadurch, dass diese keine essentiellen Erbinformationen besitzen, ein nicht-mendelndes Segregationsverhalten zeigen und auch nicht mit den A-Chromosomen während der Meiose paaren (Jones and Rees, 1982; Jones, 1991a; Jones and Puertas, 1993; Beukeboom, 1994; Jones, 1993; Bougourd and Jones, 1997; Covert, 1998; Camacho et al., 2000; Puertas, 2002; Jones et al., 2008). Sie treten gewöhnlich nur in einigen Individuen einer Art auf (Jones, 2004). Außerdem kann ihre Anzahl innerhalb und zwischen Individuen variieren. B- Chromosomen können extrem hohe Zahlen in natürlichen Populationen erreichen, abhängig davon, wie viele zusätzliche Chromosomen eine bestimmte Art tolerieren kann (Camacho et al., 2000). Durch diese unikalen Eigenschaften kann sich die Evolution beider Chromosomentypen unterscheiden. Da in der überwiegenden Mehrzahl die Gegenwart von B-Chromosomen, vor allem in höherer Kopienzahl, sich negativ auf die Vitalität und Fruchtbarkeit des Trägerorganismus auswirken kann, werden B-Chromosomen oftmals als parasitäre Bestandteile des Genoms eingestuft. Die Verteilung von B-Chromosomen innerhalb der Angiospermen ist nicht zufällig. Fremdbefruchtende Arten mit großen Genomen und wenigen Chromosomen besitzen bevorzugt diesen Chromosomentyp. Die Auswertung der B-Chromosomen- Verteilung zeigte auch, dass B-Chromosomen gleich oft in polyploiden und diploiden Arten auftreten (Palestis et al., 2004; Trivers et al., 2004; Levin et al., 2005). Die positive Korrelation zwischen Genomgröße und der Gegenwart von B- Chromosomen kann evtl. dadurch erklärt werden, dass großgenomige Arten 4 zusätzliche Chromosomen leichter tolerieren können (Puertas, 2002) oder, da das Genom großgenomiger Arten hauptsächlich aus nicht-kodierender DNA besteht, diese Genomkomponente die Formierung von B-Chromosomen unterstützt (Levin et al., 2005). Die Auswertung von 23.652 Angiospermen (rund 9% der geschätzten 260.000 Arten) zeigte, dass zirka 8% der Monokotyledonen und 3% der Eudicots Träger von B-Chromosomen sind. In den besonders artenreichen Poaceae und Asteraceae wurden am häufigsten Zusatzchromosomen nachgewiesen. Dagegen besitzen Liliales und Commelinales besonders viele Arten mit B-Chromosomen (Palestis et al., 2004; Levin et al., 2005). Obwohl seit der Erstbeschreibung von B-Chromosomen durch Wilson (1907) die rätselhaften Eigenschaften dieser Genomkomponente das Interesse einer Vielzahl von Forschern geweckt haben, sind noch viele Fragen zur Evolution und Funktionsweise von B-Chromosomen unbeantwortet. Insgesamt liegen dieser Habilitationsarbeit 20 Originalarbeiten zugrunde. Ziel meiner Arbeit war es: (A) den Ursprung und die Evolution von B-Chromosomen zu erkunden; (B) die Chromatinzusammensetzung von A- und B-Chromosomen zu vergleichen und chromosomale Verteilung von Eu- und Heterochromatin-typischen Histonveränderungen zu bestimmen. (C) B-Chromosomen zur Funktionsbestimmung von Zellzyklus-abhängigen Histon H3 Phosphorylierungen einzusetzen. (D) die Transkriptionsaktivität, der sogenannten „genetisch inaktiven“ B- Chromosomen erstmals molekulargenetisch zu charakterisieren. (E) Brachycome dichromosomatica als Modellart für die Analyse von B- Chromosomen zu etablieren. Zur Bearbeitung der aufgelisteten Themen wurden von mir bevorzugt B- Chromosomen des Roggens (Secale cereale L., 2n= 14 +0-8 Bs), der Australischen 5 Gänseblume (Brachycome dichromosomatica, Synonym: Brachyscome dichromosomatica, 2n= 4 +0-3 Standard Bs + 0-9 Mikro Bs), der Puschkinie (Puschkinia libanotica, 2n= ? + 0-7 Bs), und des Kleinköpfigen Pippau (Crepis capillaris, 2n= 6 +0-4 Bs) experimentell untersucht. Die B-Chromosomen des Roggens wurden untersucht, da bereits viele Vorarbeiten an diesen Chromosomen durchgeführt worden waren (Jones und Puertas, 1993) und somit die DNA-Zusammensetzung der Roggen Bs teilweise bekannt war. Aufgrund der geringen A-Chromosomenanzahl (2n = 4) und der Existenz von zwei B- Chromosomentypen war B. dichromosomatica prädestiniert, um die Evolution, DNA und Chromatinzusammensetzung von Bs zu erkunden. C. capillaris wurde ausgewählt, da diese Art B-lokalisierte rDNA Sequenzen besitzt, welche zur Analyse der Transkription und Phylogenie von B-Chromosomen geeignet sind. 6 2. Eigene Arbeiten (A1-A20) A1 Carchilan, M., Delgado, M., Ribeiro, T., Costa-Nunes, P., Caperta, A., Morais- Cecilio, L., Jones, R.N., Viegas, W., and Houben, A. (2007). Transcriptionally active heterochromatin in rye B chromosomes. Plant Cell 19, 1738-1749. A2 Cohen, S., Houben, A., and Segal, D. (2008). Extrachromosomal circular DNA derived from tandemly repeated genomic sequences in plants. Plant Journal 53, 1027-1034. A3 Gernand, D., Demidov, D., and Houben, A. (2003). The temporal and spatial pattern of histone H3 phosphorylation at serine 28 and serine 10 is similar in plants but differs between mono- and polycentric chromosomes. Cytogenet Genome Res 101, 172-176. A4 Gernand, D., Rutten, T., Varshney, A., Rubtsova, M., Prodanovic, S., Bruss, C., Kumlehn, J., Matzk, F., and Houben, A. (2005). Uniparental chromosome elimination at mitosis and interphase in wheat and pearl millet crosses involves micronucleus formation, progressive heterochromatinization, and DNA fragmentation. Plant Cell 17, 2431-2438. A5 Houben, A., and Schubert, I. (2007). Engineered plant minichromosomes: a resurrection of B chromosomes? Plant Cell 19, 2323-2327. A6 Houben, A., Demidov, D., Caperta, A.D., Karimi, R., Agueci, F., and Vlasenko, L. (2007). Phosphorylation of histone H3 in plants - A dynamic affair. Biochim Biophys Acta. A7 Houben, A., Demidov, D., Rutten, T., and Scheidtmann, K.H. (2005). Novel phosphorylation of histone H3 at threonine 11 that temporally correlates with condensation of mitotic and meiotic chromosomes in plant cells. Cytogenet Genome Res 109, 148-155. A8 Houben, A., Demidov, D., Gernand, D., Meister, A., Leach, C.R., and Schubert, I. (2003). Methylation of histone H3 in euchromatin of plant chromosomes depends on basic nuclear DNA content. Plant Journal 33, 967- 973. A9 Houben, A., Verlin, D., Leach, C.R., and Timmis, J.N. (2001). The genomic complexity of micro B chromosomes of Brachycome dichromosomatica. Chromosoma 110, 451-459. 7 A10 Houben, A., Wanner, G., Hanson, L., Verlin, D., Leach, C.R., and Timmis, J.N. (2000). Cloning and characterisation of polymorphic heterochromatic segments of Brachycome dichromosomatica. Chromosoma 109, 433-433. A11 Houben, A., Thompson, N., Ahne, R., Leach, C.R., Verlin, D., and Timmis, J.N. (1999). A monophyletic origin of the B chromosomes of Brachycome dichromosomatica (Asteraceae). Plant Syst Evol 219, 127-135. A12 Houben, A., Belyaev, N.D., Leach, C.R., and Timmis, J.N. (1997a). Differences of histone H4 acetylation and replication timing between A and B chromosomes of Brachycome dichromosomatica.
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    Lista de taxa invasores e de risco para Portugal Júlio Gaspar Reis Versão pré-publicação – maio de 2016 Imagem da capa: amêijoa-asiática Corbicula fluminea, rio da Areia, Valado dos Frades, Nazaré. Foto do autor. Como citar esta obra: Reis J (2016) Lista de taxa invasores e de risco para Portugal. Versão pré-publicação, maio de 2016. 107 pp. Júlio Gaspar Reis publica esta obra sob a licença “Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International”. http://creativecommons.org/licenses/by-nc-sa/4.0/deed.pt – 2 – ÍNDICE ÍNDICE, 3 Alternanthera caracasana (R), 16 LISTA DE ABREVIATURAS E SIGLAS, 7 Alternanthera herapungens (R), 16 INTRODUÇÃO, 8 Alternanthera nodiflora (R), 17 VÍRUS, 9 Alternanthera philoxeroides (R), 17 Ranavirus (I), 9 Amaranthus spp. (N), 17 BACTÉRIAS, 10 Amaryllis belladona (C), 17 Erwinia amylovora (I), 10 Ambrosia artemisiifolia (N), 17 [Candidatus Liberibacter africanus] (I), 10 Amorpha fruticosa (C), 17 [Candidatus Phytoplasma vitis] (I), 10 Aptenia cordifolia (C), 17 Pseudomonas syringae pv. actinidae (I), 10 Araujia sericifera (C), 18 Xanthomonas arboricola pv. pruni (R), 10 Arctotheca calendula (I), 18 Xylella fastidiosa (R), 11 Artemisia verlotiorum (N), 18 CHROMALVEOLATA, 12 Arundo donax (I), 18 Plasmopara viticola (I), 12 Aster squamatus (N), 18 PLANTAS, 13 Azolla filiculoides (I), 18 Abutilon theophrasti (N), 13 Bidens aurea (N), 18 Acacia baileyana (C), 13 Bidens frondosa (I), 19 Acacia cultriformis (E), 13 Boussingaultia cordifolia (C), 19 Acacia cyclops (I), 13 Carpobrotus edulis (I), 19 Acacia dealbata (I), 13 Cercis siliquastrum (N), 19 Acacia decurrens (E), 13 Chamaecyparis lawsoniana (C), 19 Acacia karroo (N/I), 14 Chasmanthe spp. (N), 19 Acacia longifolia (I), 14 Clethra arborea (I), 19 Acacia mearnsii (I), 14 Commelina communis (N?), 20 Acacia melanoxylon (I), 14 Conyza bilbaoana (C), 20 Acacia pycnantha (I), 14 Conyza bonariensis (I), 20 Acacia retinodes (I), 14 Conyza canadensis (I), 20 Acacia saligna (I), 14 Conyza sumatrensis (I), 20 Acacia sophorae (Labill.) R.Br.
  • Flowers of Italy's Gargano Peninsula

    Flowers of Italy's Gargano Peninsula

    Flowers of Italy's Gargano Peninsula Naturetrek Tour Report 15 - 22 April 2019 Ophrys scolopax ssp cornuta Orchis pauciflora Ophrys tenthredinifera Alpine Swift at Peschici Glanville Fritillary Report and images by Andrew Cleave Naturetrek Mingledown Barn Wolf's Lane Chawton Alton Hampshire GU34 3HJ UK T: +44 (0)1962 733051 E: [email protected] W: www.naturetrek.co.uk Tour Report Flowers of Italy's Gargano Peninsula Tour participants: Andrew Cleave & Andrew Bray (leaders) with nine Naturetrek clients. Summary We managed to fit plenty of sites and lots of plants into our week in the “Orchid Capital of Europe”, visiting a good variety of plant-rich habitats ranging from breezy coastal salt marshes and limestone sea cliffs to typical Mediterranean stony hillsides, olive groves and shady woodlands. The orchids did not fail to delight, and we found a superb selection of species during the week. The non-orchid flora was equally attractive with showy displays of Narcissi, Wild Tulips, Anemones and Irises, and plenty of the usual Mediterranean species in all the areas we visited. We were fortunate in that most of the locations we looked at were almost deserted and we were able to explore beautiful woodlands and open glades with only bird-song and cow-bells to be heard. Butterflies were spotted at most of the sites we visited, and there were plenty of other interesting insects to be found as well. Although we spent most of our time looking down at the plants, we did manage to produce a very good bird list with some exciting coastal species seen, as well as the more usual Mediterranean birds of stony hillsides and dark forests.
  • Latin for Gardeners: Over 3,000 Plant Names Explained and Explored

    Latin for Gardeners: Over 3,000 Plant Names Explained and Explored

    L ATIN for GARDENERS ACANTHUS bear’s breeches Lorraine Harrison is the author of several books, including Inspiring Sussex Gardeners, The Shaker Book of the Garden, How to Read Gardens, and A Potted History of Vegetables: A Kitchen Cornucopia. The University of Chicago Press, Chicago 60637 © 2012 Quid Publishing Conceived, designed and produced by Quid Publishing Level 4, Sheridan House 114 Western Road Hove BN3 1DD England Designed by Lindsey Johns All rights reserved. Published 2012. Printed in China 22 21 20 19 18 17 16 15 14 13 1 2 3 4 5 ISBN-13: 978-0-226-00919-3 (cloth) ISBN-13: 978-0-226-00922-3 (e-book) Library of Congress Cataloging-in-Publication Data Harrison, Lorraine. Latin for gardeners : over 3,000 plant names explained and explored / Lorraine Harrison. pages ; cm ISBN 978-0-226-00919-3 (cloth : alkaline paper) — ISBN (invalid) 978-0-226-00922-3 (e-book) 1. Latin language—Etymology—Names—Dictionaries. 2. Latin language—Technical Latin—Dictionaries. 3. Plants—Nomenclature—Dictionaries—Latin. 4. Plants—History. I. Title. PA2387.H37 2012 580.1’4—dc23 2012020837 ∞ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). L ATIN for GARDENERS Over 3,000 Plant Names Explained and Explored LORRAINE HARRISON The University of Chicago Press Contents Preface 6 How to Use This Book 8 A Short History of Botanical Latin 9 Jasminum, Botanical Latin for Beginners 10 jasmine (p. 116) An Introduction to the A–Z Listings 13 THE A-Z LISTINGS OF LatIN PlaNT NAMES A from a- to azureus 14 B from babylonicus to byzantinus 37 C from cacaliifolius to cytisoides 45 D from dactyliferus to dyerianum 69 E from e- to eyriesii 79 F from fabaceus to futilis 85 G from gaditanus to gymnocarpus 94 H from haastii to hystrix 102 I from ibericus to ixocarpus 109 J from jacobaeus to juvenilis 115 K from kamtschaticus to kurdicus 117 L from labiatus to lysimachioides 118 Tropaeolum majus, M from macedonicus to myrtifolius 129 nasturtium (p.
  • Bulbous Plants in the Iberian Peninsula. the Collection of the Royal Botanical Garden Madrid

    Bulbous Plants in the Iberian Peninsula. the Collection of the Royal Botanical Garden Madrid

    Horticulture International Journal Research Article Open Access Bulbous plants in the Iberian Peninsula. The collection of the Royal Botanical Garden Madrid Abstract Volume 4 Issue 1 - 2020 168 species of bulbous from Liliaceae, Amaryllidaceae and Iridaceae families are found Silvia Villegas, Carlos Aedo, Ramón Morales in the Iberian Peninsula and the Balearic Islands, 16 of them are introduced, numerous Real Jardín Botánico, CSIC. Plaza de Murillo 2. 28014 Madrid, endemic and subendemic (also from the other side of the Pyrenees in France). The living Spain bulb collection at the Royal Botanic Garden Madrid conserves and manages most of the species growing in the wild in the Iberian Peninsula. Correspondence: Ramón Morales, Real Jardín Botánico, CSIC. Plaza de Murillo 2. 28014 Madrid, Spain, Tel 914203017, Fax Keywords: bulbous, living collection, iberian peninsula, liliaceae, amaryllidaceae, 914200157, Email iridaceae Received: December 20, 2019 | Published: January 30, 2020 Introduction Iridaceae family have recently established themselves in the wild, like Ixia paniculata, Watsonia meriana, Freesia refracta, Freesia alba The geophites in the sense of Raunkiaer are herbaceous perennial and Crocosmia × crocosmiflora. Sisyrinchium, with 2 species living plants with its perennating buds below the ground. They can be in the Iberian Peninsula, has been introduced from North America rhizomes, tubers, bulbs and corms, these last solid bulbs almost and occurs in a few localities in Spain. All the bulbous plants species 1 without cataphylls. These types of plants are very important in the of the three families are included in Table 2. In the case of endemic Mediterranean region, because their biological cycle is well adapted (e) and subendemic (s), subspecies are also included.