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Osmotisch Regulierte Biosynthese Der Osmotisch regulierte Biosynthese der kompatiblen Solute Ectoin und Hydroxyectoin in Salibacillus salexigens: Biochemische Charakterisierung der Ectoin-Hydroxylase EctD und Identifizierung ihres Strukturgenes Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) dem Fachbereich Biologie der Philipps-Universität Marburg vorgelegt von Jan Bursy aus Löbau Marburg/Lahn, 2005 Die Untersuchungen zu der vorliegenden Arbeit wurden in der Zeit von Oktober 2001 bis Oktober 2004 im Laboratorium für Mikrobiologie der Philipps-Universität Marburg unter der Leitung von Prof. Dr. Erhard Bremer durchgeführt. Vom Fachbereich Biologie der Philipps-Universität Marburg als Dissertation angenommen am: 26.08.2005 Erstgutachter: Prof. Dr. Erhard Bremer Zweitgutachter: Prof. Dr. Wolfgang Buckel Tag der mündlichen Prüfung: 14.09.2005 Im Zusammenhang mit der vorliegenden Arbeit wurden folgende Manuskripte zur Veröffentlichung vorbereitet: Bursy, J., A. J. Pierik, N. Pica und E. Bremer (2005). Osmotically induced synthesis of the compatible solute hydroxyectoine is mediated by an evolutionarily conserved ectoine hydroxylase. Manuskript. Bursy, J., H. Hartmann, A. U. Kuhlmann, A. J. Pierik und E. Bremer (2005). Ectoine hydroxylase from Streptomyces coelicolor A3(2): characterization of an enzyme responsible for the biosynthesis of the compatible solute hydroxyectoine. Manuskript. Kuhlmann, A. U., J. Bursy, S. Gimpel, T. Hoffmann und E. Bremer (2005). Synthesis of the compatible solute ectoine in Virgibacillus pantothenticus is triggered by two environmental stimuli: high salt and low temperature. Manuskript. Inhaltsverzeichnis 1 I Inhaltsverzeichnis I Inhaltsverzeichnis..................................................................................................................1 Tabellenverzeichnis..............................................................................................................4 Abbildungsverzeichnis.........................................................................................................4 Abkürzungen ........................................................................................................................6 II Zusammenfassung.................................................................................................................7 III Einleitung ..............................................................................................................................9 1 Mikrobielle Anpassung an variierende Umweltbedingungen.....................................9 1.1 Die Bedeutung von Wasser, Osmose und Turgor.....................................................11 1.2 Anpassung von Mikroorganismen an hyperosmotische Wachstumsbedingungen ...12 1.3 Eigenschaften, Struktur und Wirkungsweise kompatibler Solute.............................14 1.4 Die initiale Stressantwort: Kalium-Aufnahme unter hyperosmolaren Bedingungen 19 1.5 Die zweite Anpassungsphase: Aufnahme und Synthese kompatibler Solute ...........20 1.6 Ectoin und Hydroxyectoin........................................................................................29 2 Efflux osmotisch aktiver Substanzen nach einem hypoosmotischen Schock: Schutz vor extremem Turgor ................................................................................................39 3 Perzeption osmotischer Veränderungen und regulatorische Mechanismen..............40 4 Verwendung kompatibler Solute als Kohlenstoff- und Stickstoffquelle ..................42 5 Salibacillus salexigens ..............................................................................................43 5.1 Moderat halophile und halotolerante Mikroorganismen...........................................43 IV Ziel der Arbeit .....................................................................................................................45 V Material und Methoden .......................................................................................................46 1 Chemikalien und Reagenzien....................................................................................46 1.1 Gase...........................................................................................................................46 1.2 Referenz-Substanzen für die HPLC-Analyse............................................................46 2 Mikroorganismen, Plasmide und Oligonukleotide....................................................47 3 Kulturmedien, Zusätze und Wachstumsbedingungen...............................................50 3.1 Medien.......................................................................................................................50 3.2 Sterilisation................................................................................................................52 3.3 Wachstumsbedingungen............................................................................................52 4 Molekularbiologische und genetische Methoden......................................................53 4.1 Agarose-Gelelektrophorese.......................................................................................53 4.2 Präparation von Nukleinsäuren.................................................................................53 4.3 DNA-DNA-Hybridisierung (Southern-Blot)............................................................55 4.4 RNA-RNA-Hybridisierung (Northern-Blot).............................................................56 4.5 Identifikation von mRNA-Startpunkten mittels Primer Extensions-Analyse...........57 5 Konstruktion von Plasmiden, Genbanken und Bakterienstämmen...........................57 5.1 Restriktion von DNA................................................................................................57 5.2 Ligationen..................................................................................................................58 5.3 Transformation von E. coli........................................................................................58 5.4 Polymerasekettenreaktion (PCR)..............................................................................58 5.5 Klonierung mit dem TOPO TA®- Cloning-Kit.........................................................58 5.6 DNA-Sequenzierung .................................................................................................59 5.7 Herstellung einer S. salexigens Genbank und Klonierung von ectABC....................59 5.8 Klonierung von ectD .................................................................................................60 5.9 Konstruktion eines Expressionssystemes für die Ectoin-Hydroxylase EctD............60 6 Biochemische Methoden...........................................................................................61 Inhaltsverzeichnis 2 6.1 Proteine......................................................................................................................61 6.2 HPLC Analyse...........................................................................................................69 7 Computergestützte Analyse von DNA- und Proteinsequenzen ................................71 7.1 Zugriffsnummern der Nukleotidsequenzen...............................................................71 VI Ergebnisse ...........................................................................................................................72 1 De novo Biosynthese von Ectoin und Hydroxyectoin in Salibacillus salexigens....72 1.1 Die Ectoin-Biosynthese in Abhängigkeit von der Salinität des Mediums................72 1.2 Hydroxyectoin-Biosynthese in Abhängigkeit von der Wachstumsphase .................73 2 Isolierung und Charakterisierung der Ectoin-Biosynthese-Gene ectABC aus Salibacillus salexigens ..............................................................................................74 2.1 Klonierung der Ectoin-Biosynthese-Gene ectABC ...................................................74 2.2 Sequenzanalyse der ectABC-Gene und der abgeleiteten EctABC Proteine aus S. salexigens ..................................................................................................................76 2.3 Transkriptionelle Analyse der Ectoin-Biosynthesegene ectABC..............................79 2.4 Der Transkriptionsstart der ectABC-mRNA und Analyse des Promotors ................80 3 Das Ectoin-Hydroxylase-Gen ectD aus Salibacillus salexigens...............................81 3.1 Isolierung und Charakterisierung des Ectoin-Hydroxylase-Genes ectD aus S. salexigens ..................................................................................................................82 4 Biochemische und physiologische Charakterisierung der Ectoin-Hydroxylase EctD aus S. salexigens........................................................................................................94 4.1 Entwicklung eines Aktivitäts-Testes zum Nachweis des EctD-Enzymes.................94 4.2 Reinigung der Ectoin Hydroxylase EctD aus S. salexigens......................................95 4.3 Bestimmung der aminoterminalen Aminosäuresequenz der Ectoin-Hydroxylase ...97 4.4 Bestimmung der molekularen Masse der Ectoin Hydroxylase EctD........................97 4.5 Eindimensionale 1H-NMR-Spektroskopie zur Produktbestätigung..........................98 4.6 Kinetische Parameter der Ectoin-Hydroxylase EctD aus S. salexigens....................99 4.7 Synthese der Ectoin-Hydroxylase EctD in S. salexigens ........................................110 4.8 Heterologe Produktion der S. salexigens Ectoin-Hydroxylase
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