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Chapter 2 SLOT V2 Efficiency of the Benthic Microbial Methane Filter under Varying Fluid Fluxes Dissertation zur Erlangung des Doktorgrades in den Naturwissenschaften - Doktor rerum naturalium- ‐ ‐ vorgelegt der Mathematischen-Naturwissenschaftlichen Fakultät der Christian-Albrechts-Universität zu Kiel Philip Steeb Kiel 2014 2 Hiermit erkläre ich, daß die vorliegende Doktorarbeit „Efficiency of the Benthic Microbial Methane Filter under Varying Fluid Fluxes“ nach Inhalt und Form meine eigene ist, abgesehen von der Beratung meiner Betreuerin und den bezeichneten Co-Autoren der einzelnen Kapitel. Diese Arbeit wurde noch keiner Hochschule zur Prüfung vorgelegt und alle veröffentlichen Teile sind in „Publication Outline“ aufgeführt. Die Arbeit wurde unter Einhaltung der Regeln guter wissenschaftlicher Praxis der Deutschen Forschungsgemeinschaft erstellt. ____________ _____________ Ort, Datum Philip Steeb 3 4 Abstract: Marine sediments all over the world bear methane, generated from organic matter and deposited at the seafloor. By diffusion or ascending fluids the methane is transported to the sediment surface and emitted into the oceans and further into the atmosphere, where it acts as greenhouse gas. However, most of the methane is anaerobically or aerobically oxidized by microbes in the upper sediments and in the water column. One of the most important methane oxidizing processes is the anaerobic oxidation of methane (AOM) in the microbial benthic methane filter. Consortia of anaerobic methanotroph (ANME) archeae and sulfate reducing bacteria (SRB) consume the methane with sulfate as final electron acceptor. While the methane is delivered from deeper reservoirs, sulfate originates from the seawater and has to diffuse into the sediment. AOM occurs mainly in the intersection of sulfate and methane, the sulfate methane transition zone (SMTZ). At continental margins gas hydrate bearing sediments form huge reservoirs of methane. Methane can be transported by upward migrating fluids from dewatered and dehydrated sediments. At the sediment surface, advecting fluids form so called “cold seeps”. The microbial benthic methane filter of cold seeps is well established and adapted to the flow conditions. However, increasing fluid flux can challenge the microbial benthic filter by higher methane flux and lower sulfate penetration depth, by resulted higher methane emissions. The aim of this thesis is to study efficiency and adaptability of the benthic methane filter under non-steady state conditions. For this purpose I developed a sediment- flow-through (SLOT) system to mimic the fluid regime of the benthic methane filter and to monitor long-term incubations of undisturbed sediment cores from cold seeps under varying fluid fluxes. All experiments were performed under anoxic conditions and with two parallel sediments cores under varying fluid flow conditions. The presented thesis combines field data and experimental data from sediments of different cold seep sites and their microbial benthic methane filter. First experiments with coastal gassy sediments from Eckernförde Bay constrain the possibility of SLOT to simulate fluid flow regimes of microbial benthic filters under fluid flux differences of one magnitude. Additionally, other seep related processes such as sulfide oxidation and precipitation were observed and prove the usefulness of the SLOT system for biogeochemical studies. Two neighboring mounds with different steady state conditions were studied offshore Costa Rica. These mounds differ in fluid source, fluid flux as well as methane flux. A. Both mounds exhibit a well established microbial benthic filter which is adapted to their respective flow conditions. Fluid and methane source are reflected in the isotopic signature of authigenic carbonates, which are the archive of cold seep systems. 5 Field and experimental data from the Quepos Slide offshore Costa Rica were combined to study the benthic methane filter. In the experiments the microbial benthic methane filter was highly efficient and shows a good adaption within 150-170 d response time with low methane efflux within the transition. A fluid pulse with higher methane emissions reveals how fast the methane efflux can change when fluid flux is increased. Furthermore, three different cold seep habitats from a recently discovered cold seep area at the continental margin of Chile were studied. Experiments with a bacterial mat habitat from a young seep site show a uniform fluid flow through the sediment and reveal a high adaptability of the benthic methane filter. In a clam field with high sulfide production an enormous methane efflux occured before the benthic filter was adapted to the new fluid regime. In sediments with an aerobic benthic methane filter from a tubeworm habitat, fluids were channeled by biogenic tubes. The benthic methane filter changed during the 150 d of anoxic incubation from an aerobic to an anaerobic methane sink. In the present thesis it is demonstrated for the first time how benthic methane filters adapt to increased fluid flux. The studies reveal that beside preconditions and the intensity of the variation the sediment matrix flow conditions through the sediment is prime important for the adaptability of the benthic methane filter. 6 Zusammenfassung Marine Sedimente enthalten überall auf der Welt Methan, welches aus organischem Material gebildet wurde. Dieses Methan kann durch Diffusion oder mit aufsteigenden Fluiden in die Ozeane und von dort aus weiter in die Atmosphäre gelangen, wo es als Treibhausgas wirkt. Allerdings wird das meiste Methan im Sediment als auch in der Wassersäule aerob oder anaerob oxidiert. Eine der wichtigsten Reaktionen zur Methan oxidation ist die anaerobe Methanoxidation (AOM) innerhalb des benthischen Methanfilters. Konsortien aus anaeroben methanotrophen Archaen (ANME) und sulfatreduzierende Bakterien oxidieren das Methan mit Sulfat als abschließendem Elektronenakzeptor. Während das Methan aus tieferen Speichern aufsteigt, erfolgt der Sulfatnachschub per Diffusion aus dem Meereswasser. AOM findet größtenteils in der Übergangszone von Sulfat und Methan statt, der Sulfat- Methan-Übergangszone (SMTZ). An den Kontinentalhängen bilden Sedimente, welche Gashydrate enthalten riesige Methanspeicher. Methan kann durch aufsteigende Fluide aus entwässerten und dehydrierten Sedimenten aufwärts transportiert werden. An der Sedimentoberfläche formen diese ausströmenden Fluide so genannte „Cold Seeps“. Der benthische Methanfilter ist in diesen Cold Seeps gut ausgeprägt und an die Fließbedingungen angepasst. Erhöhte Fließgeschwindigkeiten, welche einen höheren Methanfluss sowie eine geringere Eindringtiefe des Sulfats zur Folge haben, können den benthischen Methanfilter stark strapazieren. Das Ziel dieser Doktorarbeit ist die Effizienz und die Anpassungsfähigkeit des benthischen Methanfilters unter nicht stationären Bedingungen zu untersuchen. Aus diesem Grund habe ich ein Durchflusssystem (SLOT System) entwickelt um die Fließbedingungen des benthischen Methanfilters zu imitieren und Langzeituntersuchengen von ungestörten Sedimenten mit verschiedenen Volumenströmen aufzuzeichnen. Alle Versuche wurden unter anoxischen Bedingungen und mit zwei parallelen Sedimentkernen, welche unterschiedliche Fließbedingungen hatten, ausgeführt. Die vorliegenden Doktorarbeit vereint Feld- und Versuchsmesswerte von Cold Seep Sedimenten unterschiedlicher Standorte und ihren benthischen Methanfiltern. Erste Versuche mit gasigen Küstensedimenten aus der Eckernförder Bucht belegen die Eignung des SLOT Systems Flussbedingungen des mikrobisch benthischen Methanfillters bei Volumenflußunterschieden von einer Größenordung zu simulieren. Zusätzlich konnten andere mit Cold Seeps verbundene Prozesse wie Sulfidoxidation und -präzipitation beobachtet werden. Dies zeigt die Nutzen des SLOT-Systems für biogeochemische Untersuchungen. Zwei benachbarte „Mounds“ mit unterschiedlichen stationären Zuständen wurden vor Costa Rica untersucht. Diese Mounds unterschieden sich in Fluidquelle, Fluidströmung und Methanfluss. Beide enthielten einen gut ausgebildeten benthischen Methanfilter, der sich an die jeweiligenFlussbedingungen angepasst 7 hatte. Fluid- und Methanquellen spiegelten sich in der Isotopensignatur der autochthonen Karbonate wider, welche das Archiv der Cold Seep Geschichte darstellen. Feld- und Versuchsergebnisse vom Quepos Slide vor Costa Rica wurden kombiniertum den benthischen Methanfilter zu untersuchen. Der mikrobielle benthische Methanfilter war sehr effizient und konnte sich mit einer Reaktionszeit von 150-170 Tagen an die geänderten Fließbedingungen anpassen. Während dieser Anpassungszeit kam es zu geringen Methanausfluss. Durch einen Fluidimpuls welcher einen hohen Methanausfluss zur Folge hatte, wurde verdeutlicht wie schnell sich der Methanausfluss ändern kann wenn das Flussvolumen ansteigt Des Weiteren wurden drei Habitate eines kürzlich am chilenischen Kontinentalhang entdeckten Cold Seep-Gebiets untersucht. Versuche mit Sedimenten aus einem Bakterienmattenhabitat bei einer jungen Seepstelle zeigen einen gleichmäßigen Fluidfluss durch das Sediment und verdeutlichen somit eine hohe Anpassungsfähigkeit des benthischen Methanfilters. In Sedimenten aus einem Muschelfeld mit hoher Sulfidproduktion gab es einen enormen Methanausfluß bevor sich der benthische Methanfilter an die neuen Flussbedingungen angepaßt hatte. In Sedimenten mit einem aeroben Methanfilter von einen Röhrenwurmfeld wurden die Fluide durch die biogenen Röhren kanalisiert. Der benthische Methanfilter wandelte sich
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