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Einfluss Von Wolbachia-Endosymbionten Auf Populationsstruktur, Prägame Isolation Und Speziationsmechanismen Bei Altica Lythri

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Einfluss Von Wolbachia-Endosymbionten Auf Populationsstruktur, Prägame Isolation Und Speziationsmechanismen Bei Altica Lythri Einfluss von Wolbachia-Endosymbionten auf Populationsstruktur, prägame Isolation und Speziationsmechanismen bei Altica lythri AUBÉ, 1843 (Coleoptera, Chrysomelidae) Dissertation Zur Erlangung der Würde des Doktors der Naturwissenschaften des Fachbereichs Biologie, der Fakultät für Mathematik, Informatik und Naturwissenschaften, der Universität Hamburg vorgelegt von Regina Jäckel aus Hamburg Hamburg, im April 2011 ________________________________________________ 2 Inhaltsverzeichnis ZUSAMMENFASSUNG .......................................................................................... 6 ZIELSETZUNG DER ARBEIT................................................................................. 8 1. EINLEITUNG.................................................................................................... 9 1.1. Allgemeine Anmerkungen zur Gattung Altica .................................................................................9 1.2. Besondere Merkmale der Gattung Altica .......................................................................................10 1.3. Altica lythri: Eine der häufigsten Arten und ihre Besonderheiten................................................11 1.4. Wolbachia-Infektionen bei Altica.....................................................................................................15 1.5. Cytoplasmatische Inkompatibilität: Speziation durch Wolbachia? .............................................20 1.6. Wolbachia-Infektionen und mitochondrielle DNA.........................................................................23 1.7. Untersuchungen zur Gattung Altica und zum Einfluss von Wolbachia .......................................24 1.7.1. Einfluss von Wolbachia-Infektionen auf phylogenetische Muster: Bisherige Ergebnisse .....24 1.7.2. Phylogenie, Populationen, Einfluss von Wolbachia: Weiterführende Untersuchungen ........27 1.7.2.1. Analyse der Polyphylie: Mitochondrielle und nukleäre Daten.............................................27 1.7.2.2. Phylogenetische Analyse des Taxons Wolbachia....................................................................29 1.7.2.3. Geographische Analyse der Haplotypen-Verteilung von Altica lythri.................................30 1.7.2.4. Test auf cytoplasmatische Inkompatibilität bei Altica lythri................................................31 1.7.2.5. Karyologische Untersuchungen zu Polyploidie......................................................................31 2. MATERIAL UND METHODEN....................................................................... 33 2.1. Altica und Wolbachia: Phylogenetische Untersuchungen..............................................................33 2.1.1. Vorbereitung der Proben ............................................................................................................33 2.1.2. DNA-Extraktion und Amplifizierung ........................................................................................36 2.1.3. Sequenzierung ..............................................................................................................................37 2.1.4. Phylogenetische Analyse..............................................................................................................38 2.2. Geographische Analyse der Haplotypen-Verteilung bei Altica lythri...........................................40 2.2.1. Vorbereitung der Proben ............................................................................................................40 2.2.2. DNA-Extraktion und Amplifizierung ........................................................................................41 2.2.3. Restriktionsanalyse (RFLP) und Auswertung der Daten.........................................................42 2.3. Test auf cytoplasmatische Inkompatibilität ...................................................................................43 2.3.1. Aufzucht der Tiere für die Verpaarungsexperimente ..............................................................43 2.3.2. Bestimmung der mtDNA-Haplotypen und der Wolbachia-Infektion......................................45 2.3.2.1. Vorbereitung der Proben und DNA-Extraktion....................................................................45 2.3.2.2. Amplifizierung der mtDNA (COI) und der bakteriellen Gene ftsZ und wsp......................45 2.3.2.3. Restriktionsanalyse (RFLP) für Altica-COI und Wolbachia-wsp..........................................46 2.3.3. Durchführung der Verpaarungsexperimente ............................................................................47 2.4. Karyologische Untersuchung der Art Altica lythri........................................................................47 3. ERGEBNISSE................................................................................................ 50 3.1. Phylogenetische Untersuchungen: Die drei Genome von Altica lythri .........................................50 3.1.1. Die Phylogenie der Gattung Altica auf der Grundlage mitochondrieller Daten ....................50 3 3.1.1.1. Analyse der Sequenzen.............................................................................................................50 3.1.1.2. Phylogenetische Beziehungen .................................................................................................51 3.1.2. Überprüfung der mitochondriellen Phylogenie mittels nukleärer Genabschnitte .................56 3.1.2.1. EF1α-Intron .............................................................................................................................56 3.1.2.2. ITS2...........................................................................................................................................56 A Analyse der Sequenzen.................................................................................................................57 B Vergleich der nukleären mit der mitochondriellen Phylogenie ................................................59 C Probleme bei der Amplifikation von ITS2..................................................................................61 3.1.3. Wolbachia-Infektionen bei Altica................................................................................................61 3.1.3.1. Analyse des Wolbachia-Infektionsstatus anhand von ftsZ....................................................61 A Infektionsfrequenzen ....................................................................................................................63 B Analyse der ftsZ-Sequenzen .........................................................................................................64 C Bestimmung der Infektionstypen ................................................................................................64 3.1.3.2. Vergleich der Phylogenien von Altica und Wolbachia ..........................................................69 A Assoziation zwischen Wolbachia-Stamm und mtDNA bei Altica lythri ....................................69 B Haplotyp, Wolbachia-Infektion und das Geschlechterverhältnis bei Altica.............................72 C Analyse auf Tendenzen zu Koevolution zwischen Altica und Wolbachia.................................73 3.2. Geographische Analyse der Haplotypen-Verteilung bei Altica lythri...........................................75 3.2.1. Daten zur Häufigkeit von Männchen und Weibchen................................................................75 3.2.2. Geographische Verteilung der Haplotypen ...............................................................................77 3.2.3. Der Zusammenhang zwischen Haplotyp und Geschlecht ........................................................83 3.2.4. Haplotyp-Netzwerk und Nachweis weiterer mtDNA-Varianten .............................................87 3.2.5. Auflösung von "HT2" auf der Grundlage der Restriktionspolymorphismen........................89 3.3. Test auf cytoplasmatische Inkompatibilität ...................................................................................91 3.3.1. Analyse von mtDNA und Wolbachia-Infektion der Muttertiere..............................................91 3.3.1.1. Bestimmung der mtDNA-Haplotypen ...................................................................................92 3.3.1.2. Bestimmung des Wolbachia-Infektionstyps...........................................................................94 3.3.2. Anzahl der Nachkommen .............................................................................................................95 3.3.3. Mitochondrieller Haplotyp und Anteil männlicher Nachkommen ...........................................97 3.3.4. Ergebnisse der systematischen Verpaarungen ...........................................................................98 3.3.4.1. Verpaarungstypen: Test auf cytoplasmatische Inkompatibilität ........................................98 3.3.4.2. Infektionstypen: Produktivität und Fekundität der Weibchen........................................100 3.3.4.3. Prüfung auf Parthenogenese................................................................................................103 3.3.4.4. Zeitliche Komponente ..........................................................................................................103 3.4. Karyologische Untersuchung der Art Altica lythri...........................................................................104 4 DISKUSSION ................................................................................................. 107 4.1. Phylogenetische Untersuchungen: Analyse
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