Ziel dieser Dissertation ist, Wissensdefizite im Bereich der Bioirrigation von Süßwassersedimenten abzubauen. Als Untersuchungsorganismus wurde Chironomus plumosus ausgewählt weil diese weit verbreitete Larve ihre U-förmigen Röhren mit Überstandswasser durchspült und so Nahrung aus dem Wasser filtriert. Um Bioirrigation in Wohnröhren mit einem Durchmesser von ca. 1,7 mm zu untersuchen, wurden geeignete Messmethoden entwickelt, mit denen für das 4. Larvenstadium die Parameter Fließgeschwindigkeit (14,9 mm/s), Pumpzeit (33 min/h) und Pumprate (61 ml/h) gemessen wurden. Bei einer Populationsdichte von 745 Larven/m2 kann somit ein Wasservolumen äquivalent zum Volumen des Müggelsee in Berlin, innerhalb von 5 Tagen durch das Sediment gepumpt werden. Die Positronen-Emissions-Tomographie wurde für die Sedimentanalytik adaptiert und der Transport im Porenwasser analysiert. Mit den Untersuchungen wure gezeigt, dass auch in schlammigen Seesedimenten ein advektiver Transport durch Bioirrigation verursacht wird, der nicht zu vernachlässigen ist. Steigende Temperaturen resultieren aufgrund steigender Fließgeschwindigkeit in einen signifikanten Anstieg der Pumprate sowie der Eintragsrate von Überstandswasser ins Sediment. Ein abfallender Sauerstoffgehalt verlängert die Pumpzeit und führt zu einer sinkenden Fließgeschwindigkeit. Außerdem wird aus den Untersuchungen eine jahreszeitliche Variabilität der Bioirrigation sichtbar, welche unabhängig von konstanten Laborbedingungen auftritt. Mit dem Wasserstrom werden Porenwasserspezies wie SRP in den Wasserkörper abtransportiert, wohingegen SO42- und O2 aus dem Überstandswasser in das Sediment eingetragen werden. Ferner kommt es zur Oxidation von Fe2+ infolgedessen Phosphat im Sediment festgelegt wird, wie mittels P-Fraktionierung gezeigt werden konnte. Mikrobiologische Untersuchungen zeigten, dass Bioirrigation die mikrobielle Abundanz steigert, die Bakteriengemeinschaft verändert und das Potential zur enzymatischen Hydrolyse erhöht. / The aim of this thesis is to fill gaps of knowledge regarding bioirrigation in freshwater sediments. Chironomus plumosus was chosen for the investigations since the filter-feeding larva dwelling in U-shaped burrows is quite common and flushes its burrow with water from the overlying water body. To investigate bioirrigation activity in burrows of approximately 1.7 mm in diameter appropriate measurement techniques were developed. With the methods several parameters were measured for 4th stage of larvae: flow velocity (14.9 mm s-1), pumping time (33 min h-1), and pumping rate (61 ml h-1). Consequently, a water volume equivalent to the volume of Lake Müggelsee in Berlin is pumped through the sediment every 5 days by a population density of 745 larvae m-2. The nuclear medicine imaging technique Positron Emissions Tomography was adapted and used to analyze the transport in the sediment pore water. By means of the experiments, it could be shown that even in muddy lake sediments advection is a relevant transport process and should not be neglected. Rising temperatures result in increased pumping rates and increased influx rates of surface water into the sediment due to increased flow velocities in the burrows. Dropping oxygen concentrations prolong the pumping duration while the flow velocity decreases. Furthermore, experiments show a seasonal variability of bioirrigation which is independent of constant laboratory conditions. Pore water species such as SRP are transported with the water flow into the overlying water body, whereas SO42- and O2 are transported from the overlying water into the sediment. Due to the oxidation of Fe2+, phosphorus is fixed into the sediment, a result confirmed by P-fractionation. Microbiological investigations of the burrow walls demonstrated that the bioirrigation activity enhances the microbial abundance, changes the community structure, and increases the potential of enzymatic hydrolysis.

Dans le bassin d’Arcachon, le crustacé thalassinidé Upogebia pusilla habite préférentiellement les vasières intertidales colonisées par les zostères naines (Zostera noltei) où il trouve la stabilité sédimentaire indispensable à l’établissement durable de son terrier. Du fait de la profondeur de ce dernier (> 30 cm) et de sa grande mobilité, cette espèce ingénieure a souvent été négligée dans les études antérieures ayant pour objectif de mieux comprendre le rôle des communautés macro-benthiques dans le fonctionnement des herbiers de phanérogames et les conséquences de leur régression rapide. Ce manuscrit présente une étude intégrée du rôle d’U. pusilla dans le fonctionnement de son écosystème en s’attachant particulièrement à caractériser et quantifier les relations entre (1) ses différentes activités (fouissage, ventilation, locomotion, …), (2) les modes de bioturbation qui en résultent et (3)leurs impacts respectifs sur la dynamique biogéochimique sédimentaire. Ce travail montre qu’U. pusilla engendre un remaniement et une bioirrigation intenses de la matrice sédimentaire qui l’entoure. Sa présence stimule ainsi fortement la reminéralisation de la matière organique sédimentée et les échanges de solutés à travers l’interface eau-sédiment.Bien que l’influence d’U. pusilla sur ces processus écologiques et biogéochimiques soit très dépendante des conditions environnementales (e.g., saisonnalité, prévalence parasitaire),l’ensemble de mes résultats suggère que le déclin progressif de ses populations, conséquence directe de la dégradation de son habitat, est susceptible de fortement altérer le fonctionnement global des écosystèmes du bassin d’Arcachon. / In Arcachon bay, the endobenthic thalassinid crustacean Upogebia pusilla is tightly associated with the intertidal dwarf grass Zostera noltei providing the sediment stability required for the construction of (semi-)permanent burrows. Because of the depth and complex architecture of their burrow (> 30 cm), this high mobile engineer species have been largely ignored in previous studies aiming at better understanding the role of macrobenthic communities in the functioning of seagrass ecosystems and the consequences of their rapid decline. This manuscript presents an integrated study regarding the role played by U. pusillain the functioning of its ecosystem, with particular emphasis on the characterisation and quantification of the relationships between (1) its different activities (burrowing, ventilating,walking…), (2) both bioturbation modes and rates and (3) their respective impacts on the sedimentary biogeochemical dynamics. This work shows that mud shrimp activity leads tohigh mixing and bioirrigation of the surrounding sediment matrix, thus strongly enhancing organic matter mineralisation processes and solute exchanges across the sediment-water interface. Although the influence of U. pusilla on ecological and biogeochemical processes largely depends on environmental conditions (e.g., seasonality and parasitism), altogether my results suggest that the gradual decline of its population in Arcachon bay, as a direct consequence of benthic habitat degradation, may greatly alter the overall functioning of this vulnerable marine ecosystem.

Aquatic sediments are the recipients of a continual rain of organic debris from the water column. The decomposition reactions within the sediment and the rates of material exchange between the sediment and water column are critically moderated by the transport processes within the sediment. The sediment and solute movement induced by burrowing animals – bioturbation and bioirrigation – far exceed abiotic transport processes such as sedimentation burial and molecular diffusion. Thalassinidean shrimp are particularly abundant burrowing animals. Living in high density populations along coastlines around the world, these shrimp build complex burrow networks which they actively maintain and irrigate.¶ I used a laser scanner to map thalassinidean shrimp (Trypaea australiensis) mound formation. These experiments measured rapid two-way exchange between the sediment and depth. Subduction from the sediment surface proved to be just as important as sediment expulsion from depth, yet this is not detected by conventional direct entrapment techniques. The experiments demonstrated that a daily sampling frequency was needed to capture the extent of the two-way exchange.¶ I derived a one-dimensional non-local model accounting for the excavation, infill and collapse (EIC) of burrows. Maximum likelihood analyses were used to test the model against 210Pb and 228Th profiles taken from sediment cores in Port Phillip Bay, Melbourne. The maximum likelihood approach proved to be a useful technique for quantifying parameter confidence bounds and allowing formal comparison with a comparable biodiffusion model. The EIC model generally outperformed the biodiffusion model, and in all cases best EIC model parameter estimates required some level of burrow infill with surface material. The EIC model was expanded to two and three dimensions, which allowed the representation of lateral heterogeneity resulting from the excavation, infill and collapse of burrow structures. A synthetic dataset generated by the two-dimensional model was used to demonstrate the effects of heterogeneity and core sampling on the mixing information that can be extracted from one-dimensional sediment core data.¶ Burrow irrigation brings oxygenated water into burrow depths, and can affect the nitrogen cycle by increasing the rates of coupled nitrification and denitrification reactions. I modelled the nitrogen chemistry in the annulus of sediment surrounding an irrigated burrow using a radially-symmetrical diffusion model. The model was applied to three published case studies involving thalassinidean shrimp experiments and to field data from Port Phillip Bay. The results highlighted divergences between current theoretical understanding and laboratory and field measurements. The model further demonstrated potential limitations of measurements of burrow characteristics and animal behaviour in narrow laboratory tanks. Activities of burrowing animals had been hypothesised to contribute to high denitrification rates within Port Phillip Bay. Modelling work in this thesis suggests that the model burrow density required to explain these high denitrification rates is not consistent with the sampled density of thalassinidean shrimp in the Bay, although dense burrows of other animals are likely to be important. Limitations of one-dimensional representations of nitrogen diagenesis were explored via comparisons between one-dimensional models and the full cylinder model.

Persistent organic pollutants (POPs) are hydrophobic substances that readily sorb to organic matter in particles and colloids instead of being freely dissolved in the water phase. This sorption affects the bio­availability and environmental transport of the POPs. The major part of this thesis concerns the role of sediments as secondary sources of POPs. As the primary emissions decrease, contaminated sediments where POPs have accumulated can become the main source of contamination. If the contaminated sediment by time becomes covered with cleaner layers, the POPs are buried and no longer in contact with the aquatic environment. Experiments in this thesis showed, however, that new invading species can alter the sediment-water dynamics as a result of their bioturbation, i.e. mixing of sediment particles and pore-water. Marenzelleria spp., invading species in the Baltic Sea that burrow deeper than native species, were found to increase the remobilization of buried contaminants. The sediment-to-water flux was inversely related to the burial depth (2-10 cm) of the POP congeners (polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers) and also inversely related to the hydrophobicity of the congener. The flux was therefore most pronounced for less hydrophobic contaminants, which was linked to the bioirrigating behaviour of these species. Marenzelleria spp. also accumulated the buried POPs and increased concentrations in surface sedi­ment. Contaminants previously considered buried at a ’safe’ depth can thus be remobilized as a result of the invasion of Marenzelleria spp. in the Baltic Sea. One method to decrease the remobilization of contaminants from sediments is ’capping’, i.e. a layer of clean material is placed as a cap on the sediment. By amending the cap with active materials, which sequester the POPs and decrease their availability, thinner layers can be used (’active capping’ or ’thin-layer capping’). Results from an experiment with thin-layer capping using different active materials (activated carbon (AC) and kraft lignin) showed that both the sediment-to-water flux and the bioaccumulation by benthic species of polychlorinated dibenzo-p-dioxins and dibenzofurans (PCDD/Fs), hexachlorobenzene (HCB) and octachlorostyrene (OCS) decreased with increased thick­ness of the cap layer (0.5-5 cm). Amendments with active materials further increased the cap efficiency. AC was more efficient than kraft lignin, and a 3 cm cap with 3.3% AC reduced the flux and bioaccumulation with ~90%. The reduction of the sediment-to-water flux was inversely related to the hydrophobicity of the POP, and reductions in the flux had similar magnitudes as reductions in the concentration in deep-burrowing polychaetes, demonstrating the importance of bioturbation for sediment-to-water transport. In a one-year study on the levels of PCDD/Fs, PCBs, and HCB in a coastal area of the Baltic Sea, the correlations between the POP levels and the levels of particles and organic carbon in the water were found to differ for POPs of different structure and hydrophobicity. The levels of PCDD/Fs decreased to one third in May, which could be related to the increased sedimentation, i.e. water-to-sediment transport, during spring bloom.