Density and size-dependent bioturbation effects of the infaunal polychaete Nephtys incisa on sediment biogeochemistry and solute exchange

The impact of bioturbation on the geochemistry of aquatic sediments is known to depend on the benthic infauna species that are present. However, burrowing and activity patterns of each species may also change during the different stages of a life cycle. In this study, we examined the effects of four size classes of the polychaete Nephtys incisa on burrow networks and sediment biogeochemistry. In our experimental aquaria, the total biovolume (~ biomass) of Nephtys was kept constant, but different age classes were introduced, so the size and abundance varied between treatments. Despite differences in the geometry of burrow networks (due to varying density and size of burrows as revealed by X-radiography), the transport of nonreactive solutes (Br–) showed little difference between treatments. In contrast, the depth distribution of reactive solutes (Fe2+, Mn2+, TPO3– 4, TCO2, O2, pH) depended on oxidized sediment volumes and on spatial micro-heterogeneity related to burrowing patterns. Net fluxes of O2, TCO2, and NO– 3 fluxes were strongly affected by age-dependent burrowing patterns. Carbonate dissolution and remineralization rates (reflected by TCO 2fluxes) were enhanced as the size of individuals increased. NO– 3fluxes showed progressive change from dominance of nitrification (release) to denitrification (uptake) as burrow densities decreased with larger individuals. We conclude that different age-size classes of a single species at identical biovolume affect biogeo- chemical cycling differently, due to changes in burrow sizes and burrow densities. Because of redox reaction coupling associated with burrow geometries (Fe2+, Mn2+ oxidation patterns), similar magnitudes of nonlocal transport may be a misleading indicator of biogenic impacts. Our observations demonstrate that biogeochemical impacts must be evaluated in the context of size (age-) specific traits and population densities rather than biomass or biovolume alone.

Physical Disturbance by Bottom Trawling Suspends Particulate Matter and Alters Biogeochemical Processes on and Near the Seafloor

Bottom trawling is known to affect benthic faunal communities but its effects on sediment suspension and seabed biogeochemistry are less well described. In addition, few studies have been carried out in the Baltic Sea, despite decades of trawling in this unique brackish environment and the frequent occurrence of trawling in areas where hypoxia and low and variable salinity already act as ecosystem stressors. We measured the physical and biogeochemical impacts of an otter trawl on a muddy Baltic seabed. Multibeam bathymetry revealed a 36 m-wide trawl track, comprising parallel furrows and sediment piles caused by the trawl doors and shallower grooves from the groundgear, that displaced 1,000 m3 (500 t) sediment and suspended 9.5 t sediment per km of track. The trawl doors had less effect than the rest of the gear in terms of total sediment mass but per m2 the doors had 5× the displacement and 2× the suspension effect, due to their greater penetration and hydrodynamic drag. The suspended sediment spread >1 km away over the following 3–4 days, creating a 5–10 m thick layer of turbid bottom water. Turbidity reached 4.3 NTU (7 mgDW L–1), 550 m from the track, 20 h post-trawling. Particulate Al, Ti, Fe, P, and Mn were correlated with the spatio-temporal pattern of suspension. There was a pulse of dissolved N, P, and Mn to a height of 10 m above the seabed within a few hundred meters of the track, 2 h post-trawling. Dissolved methane concentrations were elevated in the water for at least 20 h. Sediment biogeochemistry in the door track was still perturbed after 48 h, with a decreased oxygen penetration depth and nutrient and oxygen fluxes across the sediment-water interface. These results clearly show the physical effects of bottom trawling, both on seabed topography (on the scale of km and years) and on sediment and particle suspension (on the scale of km and days-weeks). Alterations to biogeochemical processes suggest that, where bottom trawling is frequent, sediment biogeochemistry may not have time to recover between disturbance events and elevated turbidity may persist, even outside the trawled area.

Impact of bottom trawling on sediment biogeochemistry: a modelling approach

Bottom trawling in shelf seas can occur more than 10 times per year for a given location. This affects the benthic metabolism, through a mortality of the macrofauna, resuspension of organic matter from the sediment, and alterations of the physical sediment structure. However, the trawling impacts on organic carbon mineralization and associated processes are not well known. Using a modelling approach, the effects of increasing trawling frequencies on early diagenesis were studied in five different sedimentary environments, simulating the effects of a deeper-penetrating gear (e.g. a tickler chain beam trawl) versus a shallower, more variable penetrating gear (e.g. an electric pulse trawl). Trawling events strongly increased oxygen and nitrate concentrations in surface sediment layers and led to significantly lower amounts of ammonium (43 %–99 % reduction) and organic carbon in the top 10 cm of the sediment (62 %–96 % reduction). As a result, total mineralization rates in the sediment were decreased by up to 28 %. The effect on different mineralization processes differed both between sediment types and between trawling frequencies. The shallow-penetrating gear had a slightly smaller effect on benthic denitrification than the deeper-penetrating gear, but there were no statistically different results between gear types for all other parameters. Denitrification was reduced by 69 % in a fine sandy sediment, whereas nitrogen removal nearly doubled in a highly eutrophic mud. This suggests that even relatively low penetration depths from bottom fishing gears generate significant biogeochemical alterations. Physical organic carbon removal through trawl-induced resuspension of sediments, exacerbated by a removal of bioturbating macrofauna, was identified as the main cause of the changes in the mineralization process.

Biogeochemical impact of cable bacteria on coastal Black Sea sediment

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 µM) throughout the experiment. Sulfate reduction acted as the main source of sulfide for cable bacteria. Pore water Fe2+ reached levels of up to 1.7 mM during the incubations, due to the dissolution of FeS (30 %) and siderite, an Fe carbonate mineral (70 %). Following the upward diffusion of Fe2+, 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 µXRF, 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.

Polychaete Invasion May Lead to Biogeochemical Change in Host Marine Environment

Marine invasive species may modify their host environment by altering ecosystem biogeochemistry. We hypothesized that the invasive polychaete Marenzelleria viridis in Baltic Sea areas increases sulfate reduction (SR) in sediment micro-zones surrounding its burrow. Consequently, higher free porewater sulfide (H2S) is expected in sediments dominated by M. viridis than in corresponding sediments inhabited by the native polychaete Hediste diversicolor. In a thin-aquaria experiment, we found high SR rates (220 to 539 nmol cm−3 d−1) around the burrow walls of M. viridis as well as in surface and subsurface sediments with overall rates 2-fold higher than in defaunated control sediment. Similarly, an in situ survey revealed subsurface porewater H2S peaks moving upward towards the sediment surface in M. viridis inhabited areas. Accordingly, 50–85% higher porewater H2S was found almost year-round in these areas compared with H. diversicolor inhabited areas, suggesting that the invasion of M. viridis probably led to a substantial change in sediment biogeochemistry. In conclusion, M. viridis stimulates SR in sediment micro-zones and increases H2S in coastal sediments. Such change to more reducing conditions after the invasion may have critical environmental implications on, e.g., the distribution of H2S intolerant flora and fauna species.