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Das Metallbindeprotein HIPP3 Als Regulatorische Komponente Pflanzlicher Stressantworten Und Entwicklung

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Das Metallbindeprotein HIPP3 Als Regulatorische Komponente Pflanzlicher Stressantworten Und Entwicklung Das Metallbindeprotein HIPP3 als regulatorische Komponente pflanzlicher Stressantworten und Entwicklung Dissertation zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) vorgelegt der Naturwissenschaftlichen Fakultät I – Biowissenschaften – der Martin-Luther-Universität Halle-Wittenberg von Wiebke Zschiesche geb. am 16. April 1978 in Hoyerswerda Gutachter 1: Prof. Dr. Klaus Humbeck Gutachter 2: Prof. Dr. Markus Pietzsch Gutachter 3: Prof. Dr. Kiriakos Kotzabasis Tag der öffentlichen Promotionsverteidigung: 17.12.2013 - INHALTSVERZEICHNIS - I ZUSAMMENFASSUNG .......................................................................................... 1 I SUMMARY .............................................................................................................. 3 II EINLEITUNG .......................................................................................................... 5 1. Pflanzliche Entwicklung ist abhängig von den Umweltbedingungen .......................... 5 2. Regulation der Blüten- und Samenentwicklung ............................................................ 5 3. Pflanzliche Antwort auf Pathogenbefall ......................................................................... 8 4. HIPP-Proteine ................................................................................................................ 11 4.1 HIPP-Proteine haben eine Schwermetallbindedomäne (HMA) ...................................... 13 4.2 HIPP-Proteine besitzen eine Isoprenylierungs-Sequenz ................................................ 14 III ZIELSETZUNG .................................................................................................... 16 IV ERGEBNISSE ..................................................................................................... 17 1. HIPP3, Gen und Protein ................................................................................................ 17 2. Subzelluläre Lokalisation von HIPP3 ........................................................................... 20 3. Potentielle Interaktionspartner von HIPP3 .................................................................. 23 4. Metallbindung von HIPP3 .............................................................................................. 28 5. Expressionsmuster von HIPP3 ..................................................................................... 32 5.1 Abiotischer Stress: Trockenstress ................................................................................ 33 5.2 ABA-Behandlung ........................................................................................................... 34 5.3 Biotischer Stress ........................................................................................................... 35 5.4 Informationen zur Expression von HIPP3 aus der Arabidopsis-eFP-Browser Datenbank ..................................................................................................................... 38 5.5 Co-Expressionsnetzwerk um HIPP3 .............................................................................. 40 5.6 Vorhersage von cis-Elementen im Promotor .................................................................. 42 - INHALTSVERZEICHNIS - 6. Überexpression und knockdown von HIPP3 ............................................................... 43 6.1 knockdown von HIPP3 .................................................................................................. 43 6.2 Überexpression von HIPP3 ........................................................................................... 44 6.3 Vergleich der HIPP3 Transkriptgehalte in den HIPP3-oe Linien und der hipp3-1 Mutante ......................................................................................................................... 46 7. Gene, die durch HIPP3 knockdown und Überexpression differentiell reguliert werden ............................................................................................................ 47 7.1 Überblick über differentiell regulierte Gene .................................................................... 48 7.2 Funktionelle Einordnung der differentiell regulierten Gene............................................. 49 7.3 Detaillierte Analyse der 100 am stärksten differentiell regulierten Gene ........................ 50 7.4 Detaillierte Charakterisierung der am stärksten regulierten Gene und Verifizierung der Expressionsunterschiede mittels quantitativer realtime-PCR ................................... 52 7.5 Co-Expressionsanalyse der am stärksten regulierten Gene .......................................... 57 8. Die Überexpression von HIPP3 bewirkt eine verzögerte Entwicklung der Infloreszenz ............................................................................................................. 63 V DISKUSSION ....................................................................................................... 65 1. HIPP3 ist ein isoprenyliertes Protein, das Zink bindet und im Kern lokalisiert ist .................................................................................................................. 65 2. HIPP3 interagiert mit regulatorischen Proteinen der Blüten- und Samenentwicklung ....................................................................................................... 67 3. HIPP3 wird bei Pathogenbefall induziert und bei abiotischem Stress reprimiert ...................................................................................................................... 69 4. HIPP3 als möglicher Regulator in Signalwegen der Blüten- und Samenentwicklung, und der Pathogenabwehr ............................................................ 70 VI MATERIAL UND METHODEN ........................................................................... 76 1. Material ........................................................................................................................... 76 1.1 Pflanzenmaterial ............................................................................................................ 76 - ii - - INHALTSVERZEICHNIS - 1.2 Bakterien und Hefe Stämme .......................................................................................... 76 1.3 Chemikalien, Enzyme, „Kits“ .......................................................................................... 78 1.4 Oligonukleotide .............................................................................................................. 78 2. Methoden ....................................................................................................................... 82 2.1 Generelle molekularbiologische Methoden und Datenbanken ....................................... 82 2.2 Pflanzenanzucht und Stressbehandlungen .................................................................... 82 2.3 Lokalisationsstudien im heterologen Zwiebelepidermis-System .................................... 84 2.4 Hefe-Zwei-Hybrid Experimente ...................................................................................... 86 2.5 Identifizierung des von HIPP3 gebundenen Metalls ....................................................... 87 2.6 Expressionsstudien ....................................................................................................... 90 VII LITERATURVERZEICHNIS ............................................................................... 93 VIII ABKÜRZUNGSVERZEICHNIS ....................................................................... 108 IX ABBILDUNGSVERZEICHNIS .......................................................................... 111 X TABELLENVERZEICHNIS ................................................................................ 113 APPENDIX ............................................................................................................. 114 - iii - - ZUSAMMENFASSUNG - I ZUSAMMENFASSUNG HIPP Proteine (Heavy Metal associated Isoprenylated Plant Proteins) sind durch ein C- terminales Isoprenylierungsmotiv und durch mindestens eine Schwermetall bindende HMA- Domäne (PFAM accession number: PF00403.6) gekennzeichnet. In Arabidopsis thaliana konnten auf der Basis von Sequenzanalysen 46 HIPP-Proteine identifiziert werden. Über die Funktionen der Proteine dieser für Pflanzen spezifischen Proteinfamilie ist bis jetzt wenig bekannt. Für einige HIPP-Proteine konnte eine Funktion in der Schwermetallhomöostase gezeigt werden. Andere Publikationen deuten aber auch auf eine regulatorische Funktion in Signalwegen der biotischen und abiotischen Stressantwort hin. Im Fokus der vorliegenden Arbeit steht ein bisher noch nicht näher untersuchtes HIPP Protein, HIPP3 (At5g60800), das sich neben dem Vorhandensein von zwei HMA-Domänen und dem Isoprenylierungsmotiv zusätzlich durch vier Kernlokalisierungssequenzen (NLS) auszeichnet. Die Kernlokalisation von HIPP3 konnte über transiente Expression eines chimären GFP::HIPP3 Proteins in Epidermiszellen der Zwiebel gezeigt werden. Zusätzliche Signale im Cytoplasma deuten auf eine duale subzelluläre Lokalisation hin. In einem ersten Ansatz wurden im Hefe-Zwei-Hybrid-System die Arogenat-Dehydratase (At1g11790), die Serin/Threonin Proteinkinase WNK8 (At5g41990) und der Transkriptionsrepressor ASIL1 (At1g54060) als potentielle Interaktionspartner von HIPP3 identifiziert. Um zu klären, ob und welche Schwermetalle von HIPP3 gebunden werden, wurde HIPP3 mit N-terminalem hexa- His-Tag in E. coli exprimiert und die gebundenen Metalle nach chromatographischer Aufreinigung
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