Cable bacteria can strongly alter sediment biogeochemistry. Here, we used laboratory incubations to determine the potential impact of their activity on the cycling of iron (Fe), phosphorus (P) and sulfur (S). Microsensor depth profiles of oxygen, sulfide and pH in combination with electric potential profiling and fluorescence in situ hybridisation (FISH) analyses showed a rapid development (5 d) of cable bacteria, followed by a long period of activity (200 d). During most of the experiment, the current density correlated linearly with the oxygen demand. Sediment oxygen uptake was attributed to the activity of cable bacteria and the oxidation of reduced products from the anaerobic degradation of organic matter, such as ammonium. Pore water sulfide was low (5 uM) throughout the experiment. Sulfate reduction acted as the main source of sulfide for cable bacteria. Pore water Fe2C reached levels of up to 1.7mM during the incubations, due to the dissolution of FeS (30 %) and siderite, an Fe carbonate mineral (70 %). Following the upward diffusion of Fe2C, a surface enrichment of Fe oxides formed. Hence, besides FeS, siderite may act as a major source of Fe for Fe oxides in coastal surface sediments where cable bacteria are active. Using uXRF, we show that the enrichments in Fe oxides induced by cable bacteria are located in a thin subsurface layer of 0.3 mm. We show that similar subsurface layers enriched in Fe and P are also observed at field sites where cable bacteria were recently active and little bioturbation occurs. This suggests that such subsurface Fe oxide layers, which are not always visible to the naked eye, could potentially be a marker for recent activity of cable bacteria.

Over the last decades, hypoxia in marine coastal environments has become more and more widespread, prolonged and intense. Hypoxic events have large consequences for the functioning of benthic ecosystems. In severe cases, they may lead to complete anoxia and the presence of toxic sulfides in the sediment and bottom-water, thereby strongly affecting biological compartments of benthic marine ecosystems. Within these ecosystems, benthic foraminifera show a high diversity of ecological responses, with a wide range of adaptive life strategies. Some species are particularly resistant to hypoxia-anoxia, and consequently it is interesting to study the whole foraminiferal community as well as species-specific responses to such events. Here we investigated the temporal dynamics of living benthic foraminiferal communities (recognised by CellTracker™ Green) at two sites in the saltwater Lake Grevelingen in the Netherlands. These sites are subject to seasonal anoxia with different durations and are characterised by the presence of free sulfide (H₂S) in the uppermost part of the sediment. Our results indicate that foraminiferal communities are impacted by the presence of H₂S in their habitat, with a stronger response in the case of longer exposure times. At the deepest site (34 m), in summer 2012, 1 to 2 months of anoxia and free H₂S in the surface sediment resulted in an almost complete disappearance of the foraminiferal community. Conversely, at the shallower site (23 m), where the duration of anoxia and free H₂S was shorter (1 month or less), a dense foraminiferal community was found throughout the year except for a short period after the stressful event. Interestingly, at both sites, the foraminiferal community showed a delayed response to the onset of anoxia and free H₂S, suggesting that the combination of anoxia and free H₂S does not lead to increased mortality, but rather to strongly decreased reproduction rates. At the deepest site, where highly stressful conditions prevailed for 1 to 2 months, the recovery time of the community takes about half a year. In Lake Grevelingen, Elphidium selseyense and Elphidium magellanicum are much less affected by anoxia and free H₂S than Ammonia sp. T6. We hypothesise that this is not due to a higher tolerance for H₂S, but rather related to the seasonal availability of food sources, which could have been less suitable for Ammonia sp. T6 than for the elphidiids.

Oxygen depletion in coastal waters may lead to release of toxic sulfide from sediments. Cable bacteria can limit sulfide release by promoting iron oxide formation in sediments. Currently, it is unknown how widespread this phenomenon is. Here, we assess the abundance, activity, and biogeochemical impact of cable bacteria at 12 Baltic Sea sites. Cable bacteria were mostly absent in sediments overlain by anoxic and sulfidic bottom waters, emphasizing their dependence on oxygen or nitrate as electron acceptors. At sites that were temporarily reoxygenated, cable bacterial densities were low. At seasonally hypoxic sites, cable bacterial densities correlated linearly with the supply of sulfide. The highest densities were observed at Gulf of Finland sites with high rates of sulfate reduction. Microelectrode profiles of sulfide, oxygen, and pH indicated low or no in situ cable bacteria activity at all sites. Reactivation occurred within 5 days upon incubation of an intact sediment core from the Gulf of Finland with aerated overlying water. We found no relationship between cable bacterial densities and macrofaunal abundances, salinity, or sediment organic carbon. Our geochemical data suggest that cable bacteria promote conversion of iron monosulfides to iron oxides in the Gulf of Finland in spring, possibly explaining why bottom waters in this highly eutrophic region rarely contain sulfide in summer.

Climate variability has major implications for marine geochemical cycles and biogenic carbonate production. Therefore, past climate-driven changes in marine environments are often inferred from geochemical data of the marine carbonate archive. Proxy calibration studies are essential for the reconstruction of such past environmental changes. Here, we use the geochemical composition of living specimens of the benthic foraminifer Ammonia tepida at three sites in a seasonally hypoxic (oxygen concentration < 63 μmol/L) marine coastal system (Lake Grevelingen, the Netherlands) to explore the use of Mn/Ca as a proxy for coastal hypoxia. The study is based on samples from three stations along a depth transect, that show contrasts in the seasonal cycle of Mn ²⁺ concentrations in the pore water of the surface sediment. In general, the sediment and pore water geochemistry of the three stations in Lake Grevelingen show increased Mn ²⁺ concentrations in late winter/spring, combined with increased Mn refluxing in summer, which are due to cable bacteria activity and bottom water hypoxia/anoxia, respectively. Laser Ablation-ICP-MS (LA-ICP-MS) allowed a comparison of Mn/Ca ratios of different parts of the benthic foraminiferal test. Our results show that higher Mn/Ca ratios are registered at the deepest station, which experiences the longest and most severe seasonal periods of hypoxia/anoxia. Additionally, the signal preserved in the central part of the benthic foraminiferal tests, which is thought to reflect the entire calcification history of the analysed specimen, appears to be driven by high pore water Mn ²⁺ concentrations due to cable bacteria activity in late winter/spring. Conversely, high Mn/Ca ratios in the last chambers reflect increased Mn refluxing in the surface sediment due to summer hypoxia/anoxia. Thus, Mn/Ca ratios of A. tepida give insight into the complex spatial and temporal variability of pore water manganese.