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Die Expression Respiratorischer Proteine Ausgewählter Die Expression respiratorischer Proteine ausgewählter Arthropodenspezies Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades des Departments Biologie der Fakultät für Mathematik, Informatik und Naturwissenschaften der Universität Hamburg vorgelegt von Beyhan Ertas aus Vechta Hamburg, August 2009 Inhaltsverzeichnis Inhaltsverzeichnis A. Einleitung ................................................................................................................. 1 1. Metalloproteine mit respiratorischer Funktion ............................................................ 1 2. Die respiratorischen Proteine der Arthropoden.......................................................... 3 2.1. Hämoglobin ..................................................................................................... 3 2.1.1. Aufbau und Struktur ................................................................................. 3 2.1.2. Charakteristika der Arthropoden-Globine ................................................. 4 2.2. Hämocyanin..................................................................................................... 6 2.2.1. Aufbau und Struktur ................................................................................. 6 2.2.2. Bedeutung und weitere Funktion(en) von Hämocyanin ............................ 9 2.2.3. Hämocyanin-Superfamilie der Arthropoden.............................................. 9 2.2.3.1. Phenoloxidasen...................................................................................11 2.2.3.2. Hexamerine.........................................................................................12 3. Hämoglobin und Hämocyanin der Crustacea............................................................14 4. Zielsetzung...............................................................................................................16 B. Material und Methoden ...........................................................................................17 1. Allgemeines..............................................................................................................17 1.1. Geräte.............................................................................................................17 1.2. Chemikalien, Kits und sonstige Materialien.....................................................18 1.3. Klone ..............................................................................................................18 1.4. Primer.............................................................................................................19 1.5. Versuchstiere..................................................................................................19 2. Verwendete Organismen und ihre Kultivierung.........................................................20 2.1. Bakterien ........................................................................................................20 2.1.1. Bakterienstämme ....................................................................................20 2.1.2. Nährmedien und Antibiotika ....................................................................20 2.1.3. Ansetzen von bakteriellen Übernachtkulturen .........................................21 2.1.4. Anlegen von bakteriellen Dauerkulturen..................................................21 2.2. Zellkultur.........................................................................................................22 2.2.1. Zelllinien..................................................................................................22 2.2.2. Nährmedien und Zusätze ........................................................................22 2.2.3. Passage von Zellkulturen ........................................................................23 2.2.4. Anlegen von Dauerkulturen.....................................................................23 2.2.5. Auftauen von Dauerkulturen....................................................................23 I Inhaltsverzeichnis 3. Molekularbiologische Methoden ...............................................................................24 3.1. Isolierung von Nukleinsäuren..........................................................................24 3.1.1. Isolierung von genomischer DNA ............................................................24 3.1.2. Isolierung von RNA .................................................................................24 3.1.2.1. Isolierung von Gesamt-RNA mit dem RNeasy Mini Kit ........................24 3.1.2.2. Isolierung von Gesamt-RNA mittels Guanidinthiocyanat-Phenol- Chloroform..........................................................................................25 3.1.2.3. Isolierung von Gesamt-RNA mit PeqGOLD TriFast TM ..........................26 3.2. Konzentrationsbestimmung von Nukleinsäuren ..............................................26 3.3. Elektrophorese von Nukleinsäuren .................................................................27 3.3.1. DNA-Gelelektrophorese ..........................................................................27 3.3.2. RNA-Gelelektrophorese ..........................................................................27 3.4. Reverse Transkription.....................................................................................28 3.5. Polymerase-Kettenreaktion.............................................................................28 3.6. RACE: Rapid amplification of cDNA-ends.......................................................29 3.7. Allgemeine Klonierungstechniken ...................................................................30 3.7.1. TA-Klonierung .........................................................................................30 3.7.2. Klonierung mit Restriktionsendonukleasen..............................................31 3.7.2.1. Spaltung von DNA mit Restriktionsendonukleasen..............................31 3.7.2.2. Ligation ...............................................................................................31 3.7.3. Transformation von E. coli .......................................................................32 3.7.4. Identifizierung positiver Transformanten via Klon-PCR-Screening ..........32 3.7.5. Isolierung von Plasmid-DNA aus Bakterien.............................................33 3.7.6. Sequenzierung........................................................................................33 4. Sequenzauswertung/Molekulare Phylogenie ............................................................34 5. Rekombinante Expression von Proteinen.................................................................35 5.1. Expression von Proteinen in Bakterien ...........................................................35 5.1.1. Durchführung von Testexpressionen.......................................................36 5.1.2. Rekombinante Expression von löslichem Hämoglobin ............................37 5.1.3. Rekombinante Expression von löslichem Hexamerin ..............................37 5.1.4. Rekombinante Expression von löslichen Typ3-Kupferproteinen..............38 5.2. Expression von Proteinen in Insektenzellen....................................................38 5.2.1. „Bac-to-Bac Baculovirus Expression System” .........................................38 5.2.1.1. Klonierung des Expressionskonstruktes..............................................39 5.2.1.2. Herstellung von rekombinanten Bacmiden in E. coli ............................39 5.2.1.3. Präparation von Bacmid-DNA .............................................................40 5.2.1.4. Herstellung des rekombinanten Primärvirus (P1).................................41 II Inhaltsverzeichnis 5.2.1.5. Amplifikation von Baculoviren..............................................................41 5.2.1.6. Rekombinante Proteinexpression in Insektenzellen.............................42 5.2.2. „Drosophila Expression System“ .............................................................43 5.2.2.1. Klonierung des Expressionskonstruktes..............................................43 5.2.2.2. Co-Transfektion von S2-Zellen............................................................43 5.2.2.3. Selektion stabil transfizierter S2-Zellen................................................44 5.2.2.4. Rekombinante Proteinexpression in S2-Zellen ....................................45 6. Allgemeine proteinbiochemische Methoden .............................................................46 6.1. Entnahme von Hämolymphe...........................................................................46 6.2. Proteinextraktion.............................................................................................47 6.2.1. Proteinextraktion aus frisch präparierten Geweben/Tieren......................47 6.2.2. Proteinextraktion aus in RNAlater stabilisierten Tieren............................47 6.2.3. Proteinextraktion aus unterschiedlichen Zellkompartimenten ..................48 6.3. Proteinbestimmung.........................................................................................48 6.3.1. Photooptische Bestimmung von Proteinkonzentrationen.........................48 6.3.2. Bradford-Assay .......................................................................................48 6.4. Polyacrylamid-Gelelektrophorese (PAGE) ......................................................49 6.4.1. SDS-PAGE .............................................................................................49 6.4.2. Native PAGE...........................................................................................50
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