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Microbiology of Phase-Separated Reactor Systems For Microbiology of phase­separated reactor systems for biomethanation at high temperatures (55 ­ 75 °C) Vorgelegt von Dipl.­Biol. Antje Rademacher Von der Fakultät III ­ Prozesswissenschaften der Technischen Universität Berlin zur Erlangung des akademischen Grades Doktor der Naturwissenschaften ­ Dr. rer. nat. ­ genehmigte Dissertation Promotionsausschuss: Vorsitzender: Prof. Dr.‐Ing. Sven‐Uwe Geißen Berichter: Prof. Dr. Ulrich Szewzyk Berichterin: Prof. Dr. Elisabeth Grohmann Berichter: Dr. Michael Klocke Tag der wissenschaftlichen Aussprache: 13.06.2013 Berlin 2013 D 83 II Die vorliegende Arbeit wurde am Leibniz-Institut für Agrartechnik Potsdam-Bornim e.V. (Abteilung Bioverfahrenstechnik) unter der Anleitung von Herrn Dr. M. Klocke in der Zeit von 2009 bis 2012 erstellt. Gefördert wurde diese Arbeit vom Bundesministerium für Bildung und Forschung (BMBF) durch den Projektträger Jülich (Förderkennzeichen 03SF0349C). III DANKSAGUNG Im Folgenden möchte ich mich herzlich bei den Menschen bedanken, die mich während meines Promotionsvorhabens begleitet und unterstützt haben. Allen voran möchte ich mich bei Herrn Dr. Michael Klocke für seine wissenschaftliche Betreuung und Unterstützung bedanken. Er war mir stets Ansprechpartner und bereicherte meine Arbeit mit Ideen, Anregungen und konstruktiver Kritik. Dank gilt auch Herrn Prof. Dr. Ulrich Szewzyk für die Begleitung meines Promotionsvorhabens, die Anregungen zu meiner Arbeit sowie für deren Begutachtung. Ebenso gilt mein Dank Frau Prof. Dr. Elisabeth Grohmann für ihre Anregungen und die konstruktive Kritik zu meiner Arbeit sowie für die Übernahme des Gutachtens. Mein herzlicher Dank gilt Dipl.-Ing. Mandy Schönberg und Dipl.-Ing. Carsten Joost für die gute Zusammenarbeit im BMBF-Projekt, für den Aufbau und Betrieb der Versuchsanlage und die Bereitstellung von Versuchsdaten für die weitere Auswertung. Des Weiteren möchte ich den Mitarbeiterinnen und Mitarbeitern der AG Analytik des ATBs für die Analyse der Reaktorproben danken, sowie Kerstin Mundt für ihre kompetente Unterstützung im Labor. Zusätzlich möchte ich den vielen weiteren Kolleginnen und Kollegen des ATBs, insbesondere der Abteilung 1, für die angenehme Arbeitsatmosphäre danken. An dieser Stelle seien Angelika H., Antje F., Beate K., Edith N., Ingo Ba., Ingo Be., Johanna K., Kay B., Kathrin H., Katrin G., Lena H., Matthias B., Sarah H., Susanne K. und Susanne T. namentlich genannt. Vielen Dank für eure Unterstützung, die fruchtvollen Gespräche und die netten Stunden. Mein weiterer Dank gilt Herrn Dr. Andreas Ulrich für seine Hilfe und Unterstützung bei der Einführung der TRFLP-Methodik am ATB. Nicht zuletzt gilt mein besonderer Dank meinen Eltern sowie meinem Freund für die andauernde Unterstützung, Geduld und den notwendigen Rückhalt während der letzten Jahre. IV EIDESSTATTLICHE ERKLÄRUNG Ich erkläre an Eides Statt, dass die vorliegende Dissertation in allen Teilen von mir selbständig angefertigt wurde und die benutzten Hilfsmittel vollständig angegeben worden sind. Veröffentlichungen von irgendwelchen Teilen der vorliegenden Dissertation sind von mir, wie umseitig dargelegt, vorgenommen worden. Weiter erkläre ich, dass ich nicht schon anderweitig einmal die Promotionsabsicht angemeldet oder ein Promotionseröffnungsverfahren beantragt habe. __________________________ ___________________________ Ort, Datum Unterschrift V Teile dieser Arbeit wurden bereits wie folgt veröffentlicht: Rademacher A, Zakrzewski M, Schlüter A, Schönberg M, Szczepanowski R, Goesmann A, Pühler A & Klocke M (2012) Characterization of microbial biofilms in a thermophilic biogas system by high-throughput metagenome sequencing. FEMS Microbiology Ecology 79, 785-799, DOI: 10.1111/j.1574-6941.2011.01265.x Rademacher A, Nolte C, Schönberg M & Klocke M (2012) Temperature increases from 55 to 75 °C in a two-phase biogas reactor result in fundamental alterations within the bacterial and archaeal community structure. Applied Microbiology and Biotechnology 96, 565-576, DOI: 10.1007/s00253-012-4348-x Eikmeyer F, Rademacher A, Hanreich A, Hennig M, Jaenicke S, Maus I, Wibberg D, Zakrzewski M, Pühler A, Klocke M & Schlüter A (2013) Detailed analysis of metagenome datasets obtained from biogas-producing microbial communities residing in biogas reactors does not indicate the presence of putative pathogenic microorganisms. Biotechnology for Biofuels 6, 49, DOI:10.1186/1754-6834-6-49 Rademacher A, Hanreich A, Bergmann I & Klocke M (2012) Black-Box-Biogasreaktor - Mikrobielle Gemeinschaften zur Biogaserzeugung. Biospektrum 18, 727-729, DOI:10.1007/s12268-012-0253-1 Hanreich A, Rademacher A & Klocke M (2012) Mikrobielles Leben im Biogasreaktor – Einblicke in einen komplexen und dynamischen Mikrokosmos. In Biogas Potenziale - Erkennen, Erforschen, Erwirtschaften, p. 114-123. Potsdam, Leibniz-Institut für Agrartechnik Potsdam-Bornim e.V. BMBF-Schlussbericht (2013) Bioprozesstechnik der Hydrolyse: Verfahrenstechnische Optimierung des Bioleaching-Verfahrens unter thermophilen bis hyperthermophilen Bedingungen und Analyse der hydrolytischen Mikroflora (Förderkennzeichen 03SF0349C). eds. Schönberg M, Rademacher A, Klocke M & Linke B, Hannover, TIB/UB. VI ZUSAMMENFASSUNG In den letzten Jahren nahm die Zahl an landwirtschaftlichen Biogasanlagen in Deutschland stetig zu. Im Jahr 2011 wurden 7.215 Biogasanlagen mit einer installierten elektrischen Leistung von 2.904 MW betrieben. Thermophile Temperaturen sowie eine räumliche Trennung der Prozessphasen Hydrolyse und Acidogenese von der Methanogenese können gegenüber den häufig verwendeten, mesophilen, einphasigen Rührkesselreaktoren den Biogasprozess verbessern und stabilisieren. Für eine weitere Optimierung solcher Anlagen ist ein detailliertes Wissen über die bakterielle und archaeelle Gemeinschaft, die am Abbau der Biomasse und der nachfolgenden Methanproduktion beteiligt sind, unabdingbar. In dieser Studie wurden deshalb drei identische Leach-bed Biogasreaktorsysteme mit räumlich getrennten Prozessphasen untersucht, die jeweils mit Roggen- Ganzpflanzensilage und Stroh mit einer Verweilzeit von 21 Tagen betrieben wurden. Jedes Reaktorsystem bestand aus einem Leach-bed Reaktor (LBR), dessen Temperatur schrittweise von 55 auf 75 °C erhöht wurde, einem Prozessflüssigkeitsspeicher und einem nachgeschalteten Anaerobfilter (AF), der konstant während des gesamten Versuchs bei 55 °C betrieben wurde. Verschiedene kultivierungsunabhängige Methoden wurden genutzt um die mikrobielle Gemeinschaft im Reaktorsystem zu charakterisieren und zu quantifizieren sowie deren Dynamik zu verfolgen. Neben der Analyse von Genbibliotheken, TRFLP Fingerprintanalysen, Metagenomanalysen und qPCR Analysen wurden die Proben auch mittels Fluoreszenz in situ Hybridisierung sowie DAPI- und Propidiumiodid-Färbung untersucht. Die Ergebnisse der Studie zeigten, dass die beabsichtigte Phasentrennung zwischen der Hydrolyse und Acidogenese im LBR und der Methanogenese im AF durch die Etablierung von spezialisierten Gemeinschaften zumindest teilweise erreicht wurde. Im Vergleich zum AF wurden im LBR verstärkt hydrolytische Bakterien und somit ein erhöhtes genetisches Potential zum Abbau von Kohlenhydraten festgestellt. Während einer 21-tägigen Fermentation bei konstanter Temperatur zeigte diese bakterielle Gemeinschaft dynamische Veränderungen, die mit Konzentrationsänderungen der Fermentationsprodukte und somit dem Grad des Biomasseabaus einhergingen. Auch bei einer schrittweisen Erhöhung der Temperatur im LBR kam es zu Veränderungen in der bakteriellen Gemeinschaft. Bis zu einer Reaktortemperatur von VII 65 °C dominierten Vertreter der Clostridia. Ab 70 °C kam es zu einer verstärkten Ansiedlung von Vertretern der Klassen Bacteroidia, Thermotogae und Bacilli. Diese scheinen somit vor allem bei hyperthermophilen Temperaturen wichtig für den anaeroben Abbau von Biomasse zu sein. Entsprechende Veränderungen zeigten sich auch bei der Analyse des genetischen Potentials für den anaeroben Abbau von Biomasse. Ab einer Reaktortemperatur von 70 °C veränderte sich das genetische Potential bestimmte polysaccharidabbauende Glycosidhydrolasen zu exprimieren. Parallel dazu reduzierte sich die Abbaurate der eingebrachten Biomasse deutlich. Mittels einer Anreicherung mit Kompost konnten leichte Verbesserungen in der Reaktorleistung bei 70 °C erzielt werden, die jedoch nicht von Dauer waren. Um eine langanhaltende positive Veränderung erzielen zu können, scheint somit eine kontinuierliche Anreicherung notwendig zu sein. Die methanogene Gemeinschaft im AF zeigte nur leichte Variationen in ihrer Zusammensetzung während der Temperaturerhöhung im LBR. Sowohl Vertreter der Methanobacteriales als auch der Methanosarcinales wurden identifiziert. Es ist somit wahrscheinlich, dass verschiedene methanogene Stoffwechselwege, wie die acetoklastische und hydrogenotrophe Methanogenese, im AF parallel ablaufen. Des Weiteren wurde das Reaktorsystem hinsichtlich seines potentiellen Risikos, Wachstum und Verbreitung von Pathogenen zu gewährleisten, untersucht. Die hierfür untersuchten Metagenome zeigten jedoch, dass das Risiko für die Verbreitung von potentiellen Pathogenen durch die Ausbringung von Gärresten auf landwirtschaftlich genutzte Ackerflächen als sehr gering angesehen werden kann. Die Studie gibt Aufschluss über Struktur und Dynamik der bakteriellen und archaeellen Gemeinschaft sowie über das genetische Potential zum anaeroben Abbau von pflanzlichen Polysacchariden bei hohen Temperaturen in zweiphasigen Leach-bed Biogassystemen. Insgesamt wurde die beste Reaktorleistung bei Temperaturen
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