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Sediment-Water Coupling in Permeable Shallow Water Sediments with Special Emphasis on Carbonate Sands and the Cycling of Coral Exudates in Reef Environments

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Sediment-Water Coupling in Permeable Shallow Water Sediments with Special Emphasis on Carbonate Sands and the Cycling of Coral Exudates in Reef Environments Sediment-water coupling in permeable shallow water sediments with special emphasis on carbonate sands and the cycling of coral exudates in reef environments Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften dem Fachbereich Biologie/Chemie der Universität Bremen vorgelegt von Christian Wild Bremen August 2003 Die vorliegende Arbeit wurde in der Zeit von Oktober 2000 bis August 2003 am Max Planck Institut für marine Mikrobiologie in Bremen angefertigt. Abgabetermin: 15. August 2003. 1. Gutachter: Prof. Dr. Bo Barker Jørgensen 2. Gutachter: Prof. Dr. Wilhelm Hagen weitere Prüfer: Prof. Dr. Gunter Kirst Dr. Claudio Richter Tag des Promotionskolloquiums: 15. Oktober 2003 DANKSAGUNG Mein herzlicher Dank geht an alle, die in irgendeiner Art und Weise zur Entstehung dieser Arbeit beigetragen haben. Herrn Professor Dr. Bo Barker Jørgensen danke ich für die Begutachtung dieser Arbeit sowie hilfreiche Kommentare und Anregungen. Herrn Professor Dr. Wilhelm Hagen danke ich für die Übernahme des Zweitgutachtens, sowie Professor Dr. Gunter Kirst, Dr. Claudio Richter, Markus Billerbeck, Maren Nickel und Andrea Peiter für Ihre Bereitschaft, ebenfalls dem Prüfungskomitee anzugehören. Das größte Dankeschön geht an Professor Dr. Markus Hüttel für die Betreuung dieser Arbeit. Ich möchte ihm insbesondere danken für seine großartige Unterstützung, sein Engagement und seine zahlreichen Ideen, die entscheidend zum Gelingen dieser Arbeit beigetragen haben. Vielen Dank auch an unsere TA’s Martina Alisch und Susanne Menger, die sowohl im Feld als auch im Labor eine große Hilfe waren. Dies gilt auch für Linda Hönemann und Heiko Schütte im Labor. Stephan Kremb, Dr. Selina Ward und Dr. Ron Johnstone waren eine wertvolle Hilfe im Feld bei den Heron Island Expeditionen 2001 beziehungsweise 2002. Professor Dr. Ove Hoegh-Guldberg und das Personal der Heron Island Research Station, insbesondere Rod Forbes, haben unseren Aufenthalt möglich und sehr angenehm gemacht. Dank gebührt auch allen meinen Co-Autoren auf den verschiedenen Manuskripten dieser Doktorarbeit für die gute Zusammenarbeit und fruchtbare Diskussionen. Ich möchte mich ausserdem bei den Mitarbeitern des Max Planck Instituts bedanken, vor allem bei Perran Cook und Hans Røy für gute Anregungen und Tips, bei Bernd Stickfort für große Hilfe bei der Beschaffung exotischer Literatur, bei Jens Kallmeyer und Elke Allers für die kompetente Betreuung meines Aquariums in meiner Abwesenheit sowie insbesondere bei Maren Nickel, Perran Cook, Tomas Wilkop und Markus Billerbeck für eine schöne Zeit am MPI. Nicht zuletzt möchte ich mich bei meinen Eltern, bei meiner Schwester Friederike sowie insbesondere bei meinen Freunden Andrea, Ines, Heike, Carin, Alexandra, Ute, Stephan, Michael, Dieter, Nick, Stefan, Herbert und Tristan bedanken, die mich unterstützt haben, für mich da waren und mir gut getan haben. ZUSAMMENFASSUNG Permeable Sande bedecken große Bereiche des Meeresbodens auf dem Kontinentalschelf. Diese Gebiete sind normalerweise stark durch den Eintrag von Partikeln und organischem Material durch das angrenzende Land beeinflusst. Relativ starke Wasserströmung und Wellenbewegung können die Grenzfläche zwischen Sediment und Wasser erreichen, wo die Wasserbewegung advektiven Porenwassertransport erzeugen kann. Transport und der Austausch von Wasser in oberflächlichen Schichten von permeablen Schelfsanden sind deshalb sehr eng mit der darüber liegenden Wassersäule verknüpft. Dies könnte den Abbau von Organik in solchen permeablen Sedimenten beeinflussen. Im ersten Teil dieser Arbeit (Kapitel 2-4) haben wir die Verknüpfung zwischen Sediment und Wassersäule und seine biogeochemischen Konsequenzen in Silikatsedimenten von der deutschen Nordsee und dem nordamerikanischen Schelf untersucht. Wir fanden, dass a) die Permeabilität ein kontrollierender Faktor für den Abbau von Organik in der Oberflächenschicht von Sedimenten ist. Eine Erhöhung der Permeabilität führt auch zu einer erhöhten Mineralisierung von organischem Material (Kapitel 2). Das Ablegen von Kotpillen auf die Sedimentoberfläche von feinkörnigen Wattenmeersedimenten durch Polychaeten erhöhte die Permeabilität der Sedimentoberfläche um zwei Größenordnungen. Dies ermöglichte advektiven Transport durch die biologisch veränderte Sedimentoberfläche was zu einer erhöhten Sauerstoffverfügbarkeit und schnellerem Abbau von Organik führte. b) Porenwasserflüsse ein weit verbreiteter Prozess in permeablen Schelfsanden sind, wie wir durch eine neue in-situ Messtechnik mit Jod als Indikator feststellen konnten (Kapitel 3). Dieses Ergebnis bestätigt die Hypothese, dass ausgedehnte sandige Schelfgebiete als riesiges Filtersystem fungieren können. Versuche mit Durchfluss-Säulen demonstrierten, dass die Sauerstoffaufnahme des inkubierten Sediments positiv mit der Durchflussgeschwindigkeit des Porenwassers korreliert war. c) gelöstes organisches Material (DOM) in diesen Sanden sehr effektiv abgebaut werden kann (Kapitel 4). Obwohl die Bakterienzahlen pro Sedimentvolumen aufgrund der kleinen Sandkörneroberfläche relativ gering waren, scheinen die wenigen Bakterien, die auf den Sandkörnern leben, den mechanischen Sandfilter in einen biokatalytischen Filter zu verwandeln. Die biokatalytische Effizienz von Karbonatsanden, die den Meeresboden der meisten Korallenriffe bedecken, übertrifft die von Silikatsanden mit einer ähnlichen Korngrößenverteilung. Dies liegt an einer höheren Permeabilität, einer höheren spezifischen Oberfläche und höheren Bakteriendichten in Karbonatsanden. Der Kernteil dieser Arbeit (Kapitel 5-9) beschäftigt sich deshalb mit Transportmechanismen und Reaktionen in Korallensanden und den Prozessen, die die Wassersäule und diese Karbonatsande miteinander verknüpfen. Feld und Laborarbeiten wurden vor allem auf Heron Island in der australischen Great Barrier Reef Region durchgeführt. In-situ und Laborexperimente mit benthischen Kammern zeigten einen sehr aktiven sedimentären Matabolismus in Karbonatsanden des -2 -1 Riffs. Die Sauerstoffverbrauchsraten gemessen in-situ waren 65 ± 15 mmol O2 m d und stiegen mit erhöhter Permeabilität und advektivem Porenwasseraustausch an (Kapitel 5). Diese hohen Oxidationsraten in permeablen Karbonatsanden deuten darauf hin, dass diese Sedimente einen wichtigen Beitrag für den Abbau und die Regeneration von organischem Material in Rifflebensräumen leisten. Wir fanden ausserdem, dass transparente exopolymere Partikel (TEP) und klebrige marine snow- Aggregate in Korallenriffen häufig und weit verbreitet sind. Diese Aggregate können das Leben und die Stoffwechselprozesse auf und im Meeresboden beeinflussen, weil sie die Sedimentationsraten erhöhen und vielfach absedimentieren (Kapitel 6). Schleime, die ständig von Riffkorallen in die Wassersäule entlassen werden, in besonders grossen Mengen bei Umweltstress, verändern sich zum Teil zu diesen marine snow- Aggregaten und spielen eine wichtige Rolle für den Energietransport von Korallen zum benthischen Lebensraum. In-situ Inkubationen von Korallenschleim in benthischen Kammern zeigten, dass dieses organische Substrat von den Karbonatsanden mit einer Umsatzrate von mindestens 7% des zugegebenen Kohlenstoffs pro Stunde abgebaut wird (Kapitel 7). Durch die Klebrigkeit des Korallenschleims werden zusätzlich Partikel aus der Wassersäule eingefangen und so die Sinkgeschwindigkeit der entstehenden Aggregate erhöht. Dies führt zu einer Sedimentation dieses partikulären Materials in der Rifflagune. Der Transport von Energie und Nährstoffen durch Korallenschleim kann 2-26%, 1-22% und 2-5% des C, N und P Bedarfs der Lagunensedimente abdecken, oder sogar mehr unter bestimmten Bedingungen. Dieser Mechanismus reduziert auch den Verlust von Organik für das Riffsystem (Kapitel 8). Hinzu kommt, dass das jährliche massenweise Entlassen von Geschlechtsprodukten durch Riffkorallen Konsequenzen für den Metabolismus des Sediments hat. Die Sauerstoffverbrauchsraten des Sediments, in-situ gemessen vor, während und nach einem solchen Ereignis, waren für 9 Tage nach dem Ereignis signifikant erhöht verglichen mit den Werten, die vorher gemessen wurden. Dabei wurden Sauerstoffverbrauchsraten von bis zu -2 -1 230 mmol O2 m d registriert (Kapitel 9). Das massenhafte Entlassen von organischem Material während dieses Ereignisses hat somit einen starken jahreszeitlichen Einfluss auf die sedimentären Prozesse, was darauf hindeutet, dass Teile der in die Wassersäule entlassenen Korallengeschlechtsprodukte schnell, entweder direkt oder indirekt (z.B. über Nahrungskettenprozesse) die Sedimentoberfläche erreicht haben und letztendlich auch in den Karbonatsanden abgebaut wurden. Organisches Material, welches von Riffkorallen in die Wassersäule entlassen wird, hat demzufolge eine wichtige Funktion für die Ernährung der mikrobiellen Gemeinschaften und Fauna des Sediments. Dies liegt an einer sehr engen Verknüpfung zwischen Wassersäule und Sediment in Riff- Ökosystemen. Zusammenfassend kann gesagt werden, dass advektiver Transport, der den pelagischen und benthischen Lebensraum verbindet, eine größere Wichtigkeit für Stoffkreisläufe in flachen Schelfgebieten zu haben scheint als bisher gedacht. Der Lebensraum Korallenriff stellt eine Extremsituation dar, weil durch Wind oder Wellen erzeugte Druckgradienten in der Lage sind die Filtration von Material durch Karbonatsande stark zu erhöhen. Dies macht diese aktiven Sande zu riesigen katalytischen Bioreaktoren, was zur Erklärung der hohen Produktivität von Riff-Ökosystemen beitragen könnte. THESIS ABSTRACT Permeable sands are covering large parts of the
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