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Here, two separate processes need to be distinguished: (1) bioturbation, which is the displacement and mixing of sediment particles by benthic fauna (and flora), and (2) bioirrigation, which refers to the process of benthic organisms flushing their burrows with overlying seawater (Meysman et al. 2006). Animal burrows basically represent invaginations of the sediment surface that increase the exchange area between sediment and water (Fig. 4). Oxygenated surface water that is pumped through the burrows 24 TINA TREUDE enables oxygen penetration into sediment depths where it would be normally depleted (Revsbech et al.

Unfortunately, it is difficult to grow Pelomyxa under controlled conditions (van Bruggen et al. 1985, 1988). 2. ECTOSYMBIOTIC BACTERIA As mentioned previously, marine anaerobic ciliates are often covered by a dense layer of symbiotic bacteria. In two cases (the marine anaerobic ciliates, Metopus contortus and Caenomorpha levanderi), it was possible to demonstrate, using FISH, that these bacteria are sulfate reducers. This stands to reason as they would be able to maintain a low intracellular pH2 in the ciliates in a similar way as do methanogens (Fenchel and Ramsing 1992).

Modern oxygen minimum zones and permanent anoxic environments are introduced. In Sect. 4, biogeochemical processes during past anoxic events and during the era before the first rise of oxygen are reviewed. 2. Sediment Biogeochemistry in Modern Oxygenated Oceans In modern oxygenated oceans, a cascade of electron acceptors (oxygen, nitrate, manganese, iron, and sulfate – in this order) is utilized by prokaryotes to degrade organic matter on the ocean floor (Canfield et al. 2005; Jørgensen 2000 ) (Fig.

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