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Molekulare Untersuchungen Zur Evolution Der Aeolidida (Mollusca

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Molekulare Untersuchungen Zur Evolution Der Aeolidida (Mollusca Molekulare Untersuchungen zur Evolution der Aeolidida (Mollusca, Gastropoda, Nudibranchia, Cladobranchia) und zur Evolution einer sekundären Symbiose mit Symbiodinium (Dinoflagellata) in den Aeolidida Dissertation zur Erlangung des Doktorgrades der Fakultät der Mathematik und Naturwissenschaften der Bergischen Universität Wuppertal angefertigt am Lehrstuhl für Zoologie und Biologiedidaktik vorgelegt von Dipl.-Biol. Sabrina Bleidißel Wuppertal, im Dezember 2010 Die Dissertation kann wie folgt zitiert werden: urn:nbn:de:hbz:468-20110509-151022-7 [http://nbn-resolving.de/urn/resolver.pl?urn=urn%3Anbn%3Ade%3Ahbz%3A468-20110509-151022-7] Erstgutachterin & Betreuerin: Professorin Dr. A. Preisfeld Zweitgutachterin: Professorin Dr. H. Wägele Inhaltsverzeichnis Zusammenfassung .............................................................................................................. 1 1. Einleitung ......................................................................................................................... 3 1.1 „Schmetterlinge“ der Meere .......................................................................................... 3 1.1.1 Die Systematische Stellung und die Biologie der Aeolidida ................................... 7 1.1.2 Bisheriger Kenntnisstand zur Evolution innerhalb der Aeolidida ............................ 9 1.2 Die Systematische Stellung und die Biologie der Cnidaria als Futter-organismen der Aeolidida ..................................................................................................................... 13 1.3 Die Systematische Stellung und Biologie der Endosymbionten: Symbiodinium ......... 15 1.4 Molekulare Analyse .................................................................................................... 17 1.4.1 Molekulare Marker für phylogenetische Analysen in der molekularen Systematik 17 1.4.2 Molekulare Marker zur Identifizierung von Symbiodinium .................................... 19 1.5 Ziele der Arbeit ........................................................................................................... 20 2. Material und Methoden ................................................................................................. 21 2.1 Herkunft des Tiermaterials ......................................................................................... 21 2.2 Molekulare Methoden ................................................................................................. 28 2.2.1 Isolierung der DNA ............................................................................................... 28 2.2.2 Elektrophoretische Auftrennung von DNA ........................................................... 28 2.2.3 PCR Bedingungen ............................................................................................... 29 2.2.4 Aufreinigung der PCR Produkte ........................................................................... 33 2.2.5 Klonierung der PCR Produkte .............................................................................. 33 2.2.6 "Colony"-PCR zur Charakterisierung der Inserts ................................................. 35 2.2.7 Aufreinigung und dauerhafte Lagerung der Klone ............................................... 35 2.2.8 Restriktionsanalyse .............................................................................................. 35 2.2.9 Sequenzierung der Plasmide und Erstellung der Konsensussequenzen ............. 36 2.2.10 Überprüfung der DNA-Sequenzen ..................................................................... 36 2.2.11 Zuordnung der Symbiodinium-Sequenzen ......................................................... 36 2.2.12 Die quantitative Real-Time-PCR ........................................................................ 37 2.3 Phylogenetische Analyse der molekularen Daten ...................................................... 39 2.3.1 Alinierung ............................................................................................................. 40 2.3.2 Konkatenierte Datensätze .................................................................................... 41 2.3.3 Alinierung der plastidären LSU rDNA ................................................................... 42 2.3.4 Statistische Verfahren .......................................................................................... 42 I 2.3.4.1 Basenzusammensetzung .............................................................................. 42 2.3.4.2 Sequenzlängen der SSU rDNA ...................................................................... 43 2.3.4.3 Substitutionssättigung .................................................................................... 43 2.3.4.4 Sequenzdivergenz ......................................................................................... 43 2.3.4.5 Bestimmung der relativen Substitutionsraten................................................. 44 2.3.4.6 Bootstrap Test ............................................................................................... 44 2.3.4.7 Topologische Tests ........................................................................................ 44 2.3.5 Methoden zur Stammbaumrekonstruktion ........................................................... 44 2.3.5.1 Wahl der Außengruppe für die Phylogenese der Aeolidida ........................... 44 2.3.5.2 Distanzverfahren ............................................................................................ 45 2.3.5.3 Maximum-Parsimony ..................................................................................... 45 2.3.5.4 Maximum-Likelihood ...................................................................................... 45 2.3.5.5 Bayesianische Analyse .................................................................................. 46 2.3.5.6 Sekundärstrukturanalysen der SSU rRNA ..................................................... 46 2.4 Etablierung verschiedener Symbiosesysteme im Aquarium ....................................... 47 2.5 Identifizierung der Futterorganismen .......................................................................... 48 2.6 Fluoreszenzmessung als Methode zur Detektion von Photosyntheseaktivität ........... 49 3. Ergebnisse ..................................................................................................................... 53 3.1 Rekonstruktion der Phylogenese der Aeolidida .......................................................... 53 3.1.1 Alinierungen der Sequenzen ................................................................................ 53 3.1.2 Stammbaumrekonstruktion der Aeolidida ............................................................ 56 3.1.3 Sequenzanalysen ................................................................................................ 63 3.1.4 Zusammenfassung der Ergebnisse zur Rekonstruktion der Phylogenese der Aeolidida .................................................................................................................. 79 3.2 Rekonstruktion der Phylogenese des Genus Phyllodesmium .................................... 80 3.2.1 Alinierung der Sequenzen .................................................................................... 80 3.2.2 Stammbaumrekonstruktion des Genus Phyllodesmium ....................................... 80 3.2.3 Nahrungsanalysen von Phyllodesmium ............................................................... 82 3.2.4 Zusammenhang zwischen Nahrungsaufnahme und Form der Cerata ................. 85 3.2.5 Zusammenfassung der Ergebnisse zur Rekonstruktion der Phylogenese des Genus Phyllodesmium ............................................................................................. 87 3.3 Symbiosesysteme ...................................................................................................... 88 3.3.1 Phyllodesmium briareum ...................................................................................... 88 3.3.1.1 Etablierung des Systems ............................................................................... 88 II 3.3.1.2 Photosynthetische Aktivität der Symbionten in Phyllodesmium briareum ...... 90 3.3.2 Aeolidiella stephanieae ........................................................................................ 93 3.3.2.1 Etablierung des Systems ............................................................................... 93 3.3.2.2 Photosynthetische Aktivität der Symbionten in Aeolidiella stephanieae ........ 93 3.3.3 Vergleichende Beobachtungen zur Nahrungsaufnahme zwischen Aeolidida und nicht-Aeolidida: Gleiche Nahrung – anderes Verhalten bei der Nahrungsaufnahme - keine Symbiose ........................................................................................................ 96 3.3.4 Zusammenfassung der untersuchten Symbiose-Systeme ................................... 98 3.4 Identifizierung der Symbionten ................................................................................... 99 3.5 Relative Quantifizierung der Symbionten ................................................................. 101 3.5.1 Zusammensetzung der Symbionten-Clades in Phyllodesmium briareum und verschiedener Futterkorallen .................................................................................. 103 3.5.2 Zusammenfassung der Quantifizierung der Symbionten ................................... 110 4. Diskussion ..................................................................................................................
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