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Analyse Des Wirtszellproteoms Und Der Expression Viraler Proteine In Analyse des Wirtszellproteoms und der Expression viraler Proteine in virusinfizierten Gewebekulturen durch quantitative Massenspektrometrie mit metabolisch eingeführten stabilen Isotopen I n a u g u r a l d i s s e r t a t i o n zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) an der Mathematisch-Naturwissenschaftlichen Fakultät der Ernst-Moritz-Arndt-Universität Greifswald vorgelegt von Martin Skiba geboren am 11.12.1980 in Bochum Greifswald, 18.11.2010 Dekan: Prof. Dr. Klaus Fesser 1. Gutachter : Prof. Dr. Dr. h.c. Thomas C. Mettenleiter 2. Gutachter: Prof. Dr. Beate Sodeik Tag der Promotion: 04.03.2011 Inhaltsverzeichnis I. Einleitung ....................................................................................................................... 1 I.1 Vorwort ...................................................................................................................... 1 I.2 Das Pseudorabiesvirus .............................................................................................. 1 I.2.1 Die Taxonomie der Herpesviren ......................................................................... 1 I.2.2 Die Pathogenese der Pseudorabiesvirus-Infektion ............................................. 2 I.2.3 Der morphologische Aufbau des Herpesvirus Partikels ...................................... 3 I.2.4 Der Genomaufbau .............................................................................................. 4 I.2.5 Der Replikationszyklus des Pseudorabiesvirus .................................................. 4 I.2.6 Die Expression herpesviraler Gene .................................................................... 6 I.2.7 Einfluss der Infektion mit Herpesviren auf die Expression und posttranslationale Modifikation zelleigener Proteine ........................................................................ 8 I.3 Das Newcastle Disease Virus ...................................................................................12 I.3.1 Taxonomische Einordnung und Pathogenese ...................................................12 I.3.2 Morphologie und Genomorganisation des NDV ................................................12 I.3.3 Die Transkription der NDV-Gene .......................................................................14 I.3.4 NDV als viraler Vektor für rekombinante Impfstoffe ...........................................15 I.4 Proteomanalyse ........................................................................................................16 I.4.1 Quantitative Proteomics ....................................................................................17 I.4.2 Proteomanalysen von Viruspartikeln .................................................................19 I.4.3 Proteomanalysen und Expressionsstudien mit PrV-infizierten Zellen ................20 II. Zielstellung ....................................................................................................................27 II.1 Erstellung einer Proteomkarte PrV-infizierter MDBK Zellen ...................................27 II.2 Proteomanalysen an Zellfraktionen nach Infektion mit PrV-Wildtyp und einer Kinase-negativen Deletionsmutante ......................................................................28 II.3 Quantitative Analyse der Proteinexpression von Fremdgenen in rekombinanten Newcastle Disease Viren ......................................................................................28 III. Zusammenfassende Darstellung und Diskussion der Ergebnisse .............................29 III.1 “Quantitative whole-cell proteome analysis of pseudorabies virus-infected cells.” .29 III.2 “Gene expression profiling of Pseudorabies virus (PrV) infected bovine cells by combination of transcript analysis and quantitative proteomic techniques.” ...........31 III.3 “Influence of Insertion Site of Avian Influenza Virus Hemagglutinin (HA) Gene Within the Newcastle Disease Virus Genome on HA.” ..........................................32 IV. Literaturverzeichnis ...................................................................................................34 V. Quantitative whole-cell proteome analysis of pseudorabies virus-infected cells. .......63 VI. Gene expression profiling of Pseudorabies virus (PrV) infected bovine cells by combination of transcript analysis and quantitative proteomic techniques. ................75 VII. Influence of Insertion Site of Avian Influenza Virus Hemagglutinin (HA) Gene Within the Newcastle Disease Virus Genome on HA Expression. ........................................83 VIII. Zusammenfassung der Dissertation ........................................................................ 100 IX. Summary ................................................................................................................ 103 X. Anhang ................................................................................................................... 106 X.1 Abkürzungsverzeichnis ....................................................................................... 106 X.2 Publikationen ...................................................................................................... 109 X.2.1 Veröffentlichungen .......................................................................................... 109 X.2.2 Tagungsbeiträge: ............................................................................................ 109 X.3 Eidesstattliche Erklärung ..................................................................................... 110 X.4 Lebenslauf .......................................................................................................... 111 X.5 Danksagung ........................................................................................................ 112 I. Einleitung I.1 Vorwort In der vorliegenden Arbeit wurden Proteinexpressionsanalysen an Gewebekulturen durchgeführt, die mit dem Pseudorabiesvirus (PrV) oder mit rekombinanten Newcastle Disease Viren (NDV) infiziert worden waren. Die quantitative Analyse erfolgte nach zweidimensionaler Proteinelektrophorese (2DE) massenspektrometrisch unter Verwendung metabolisch eingeführter stabiler Isotope. Um in die Thematik einzuführen, sollen zunächst die verwendeten Krankheitserreger und die Grundlagen der verwendeten Protein- und Massenanalyse vorgestellt werden. I.2 Das Pseudorabiesvirus I.2.1 Die Taxonomie der Herpesviren Die Herpesviren werden in drei Familien innerhalb der Ordnung der Herpesvirales unterteilt. Zu der Familie der Herpesviridae werden die Herpesviren der Säuger, Vögel und Reptilien gezählt. Die Familie der Alloherpesviridae umfasst die Herpesviren der Fische und Amphibien, und die Familie Malacoherpesviridae enthält das Herpesvirus der Auster (Davison et al., 2009). Aufgrund ihres unterschiedlichen Wirts- und Zelltropismus werden die Vertreter der Familie der Herpesviridae weiter in die drei Unterfamilien Alpha-, Beta- und Gammaherpesvirinae klassifiziert (Roizmann et al., 1992). Die Alphaherpesviren weisen im Allgemeinen ein breites experimentelles Wirtsspektrum, einen kurzen Replikationszyklus und eine schnelle lytische Ausbreitung in der Zellkultur auf. Häufig wird eine reaktivierbare Latenz in sensorischen Ganglien des Wirts etabliert. Die Unterfamilie wird von vier Genera gebildet. Dem Genus Simplexvirus werden die humanpathogenen Herpes Simplex Viren Typ 1 und 2 (HSV-1 und -2; HHV1 und 2), sowie das Herpes B Virus (McHV1) zugeordnet. Das humanpathogene Varizella-Zoster Virus (VZV; HHV3) und die tierpathogenen Bovine Herpesviren 1 und 5 (BoHV1 und 5), Equinen Herpesviren 1 und 4 (EHV1 und 4) und das Pseudorabiesvirus (PrV; SuHV1) gehören zum Genus Varicellovirus (Roizmann et al., 1992). Die beiden weiteren Genera Mardivirus, sowie Iltovirus beinhalten u.a. die Erreger der Marek´schen Krankheit (MDV; GaHV2 und 3) bzw. der Infektiösen Laryngotracheitis (ILTV; GaHV1) der Hühner (Minson et al., 2000). Die Vertreter der Betaherpesviren zeichnen sich durch ein engeres Wirtsspektrum und einen längeren Replikationszyklus aus. Typische Merkmale sind eine langsamere Ausbreitung in der Zellkultur, häufig verbunden mit einer abnormen Vergrößerung infizierter Zellen (Zytomegalie). Latenz etabliert sich in sekretorischen Drüsen, lymphoretikulären Zellen, der Niere und in anderen Geweben. 1 Gebildet wird die Unterfamilie von den vier Genera Cytomegalovirus mit dem humanpathogenen Humanen Zytomegalievirus (HCMV; HHV5), Muromegalovirus, mit dem Murinen Zytomegalievirus (MCMV; MuHV1), Roseolovirus mit den Humanen Herpesviren Typ 6 und 7 (HHV6 und 7) und Proboscivirus mit dem Endotheliotropen Elefantenherpesvirus (EIHV1) als einzigem Vertreter. Die Gammaherpesviren besitzen ein sehr enges Wirtsspektrum und infizieren produktiv und latent B- und T-Lymphozyten. Unterteilt wird diese Unterfamilie in das Genus Lymphocryptovirus mit dem Epstein-Barr- Virus (EBV; HHV4), dem Genus Rhadinovirus mit dem Kaposi´s sarcoma associated herpesvirus (KSHV; HHV8), dem Genus Macavirus u.a. mit dem bösartigen Katarrhalfieber (CpHV2) und dem Genus Percavirus. In den Familien der Alloherpesviridae und der Malacoherpesviridae finden sich Vertreter, die zwar nur geringe Sequenzhomologien zu anderen Herpesviren aufweisen, aufgrund morphologischer und biologischer Eigenschaften aber dennoch diesen zugeordnet werden. So finden sich bei den Alloherpesviridae Herpesviren der Fische und der Amphibien mit den bekanntesten Vertretern Channel Catfish Virus (IcHV1), Koi-Herpesvirus (CyHV3) und den Froschherpesviren RaHV1 und 2.
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