The iron “redox battery” in sandy sediments: Its impact on organic matter remineralization and phosphorus cycling

Permeable sandy sediments cover 50-60% of the global continental shelf and are important bioreactors that regulate organic matter (OM) turnover and nutrient cycling in the coastal ocean. In sands, the dynamic porewater advection can cause rapid mass transfer and variable redox conditions, thus affecting OM remineralization pathways as well as the recycling of iron and phosphorus. In this study, North Sea sands were incubated in flow-through reactors (FTRs) to investigate biogeochemical processes under porewater advection and changing redox conditions. We found that the average rate of anaerobic OM remineralization was 12 times lower than the aerobic pathway, and Fe(III) oxyhydroxides were found as the major electron acceptors during 34 days of anoxic incubation. Abundant reduced Fe in the solid phase (expressed as Fe(II)) was measured before extensive Fe2+ release into porewater, and most of the reduced Fe (~96%) remained in the solid phase throughout the anoxic incubation. Fe(II) retained in the solid phase, either through the formation of authigenic Fe(II)-bearing minerals or adsorption, was easily re-oxidized upon exposure to O2 . Excessive P release (apart from OM remineralization) started at the beginning of the anoxic incubation and accelerated after the release of Fe2+ with a constant P/Fe2+ ratio of 0.26. After 34 days of anoxic incubation, porewater was re-oxygenated and >99% of released P was coprecipitated through Fe2+ oxidation (so-called “Fe2+ curtain”). Our results demonstrate that Fe(III)/Fe(II) in the solid phase can serve as relatively immobile and rechargeable “redox battery” under dynamic porewater advection. Due to frequent oscillation of redox conditions, the Fe “redox battery” is characteristic for permeable sediments and plays an important role in coastal OM turnover. We also suggest that P liberated before Fe2+ release can escape the “Fe2+ curtain” in porewater advection, thus potentially increasing net benthic P efflux from permeable sediments under variable redox conditions

Response of Bacterial Communities to Different Detritus Compositions in Arctic Deep-Sea Sediments

Benthic deep-sea communities are largely dependent on particle flux from surface waters. In the Arctic Ocean, environmental changes occur more rapidly than in other ocean regions, and have major effects on the export of organic matter to the deep sea. Because bacteria constitute the majority of deep-sea benthic biomass and influence global element cycles, it is important to better understand how changes in organic matter input will affect bacterial communities at the Arctic seafloor. In a multidisciplinary ex situ experiment, benthic bacterial deep-sea communities from the Long-Term Ecological Research Observatory HAUSGARTEN were supplemented with different types of habitat-related detritus (chitin, Arctic algae) and incubated for 23 days under in situ conditions. Chitin addition caused strong changes in community activity, while community structure remained similar to unfed control incubations. In contrast, the addition of phytodetritus resulted in strong changes in community composition, accompanied by increased community activity, indicating the need for adaptation in these treatments. High-throughput sequencing of the 16S rRNA gene and 16S rRNA revealed distinct taxonomic groups of potentially fast-growing, opportunistic bacteria in the different detritus treatments. Compared to the unfed control, Colwelliaceae, Psychromonadaceae, and Oceanospirillaceae increased in relative abundance in the chitin treatment, whereas Flavobacteriaceae, Marinilabiaceae, and Pseudoalteromonadaceae increased in the phytodetritus treatments. Hence, these groups may constitute indicator taxa for the different organic matter sources at this study site. In summary, differences in community structure and in the uptake and remineralization of carbon in the different treatments suggest an effect of organic matter quality on bacterial diversity as well as on carbon turnover at the seafloor, an important feedback mechanism to be considered in future climate change scenarios

Benthic element cycling on the Antarctic shelf and its potential control by sea ice cover

Antarctic shelf regions are potential carbon and nutrient cycling hotspots where rapid climatic changes are projected to affect seasonal sea ice cover, water column stratification, and thus surface primary production and associated fluxes of organic carbon to the seafloor. Here, we report on surface sediment oxygen profiles and respective fluxes in combination with pore water profiles of dissolved iron (DFe) and phosphate (PO43-) from 7 stations along a 400 mile transect with variable sea ice cover and water column stratification from the East Antarctic Peninsula to the west of South Orkney Islands. Our results show that sea ice concentrations and stratification of the upper water column decreased across the transect. We defined a marginal sea ice index of 5-35% sea ice cover which was positively correlated with the benthic carbon mineralization rate. C-mineralization rates increased gradually between the heavy ice-covered station and the marginal sea ice stations from 1.1 to 7.3 mmol C m-2 d-1, respectively. The rates decreased again to 1.8 mmol C m-2 d-1 at the ice-free station, likely attributed to a deeper water column mixed layer depth, which decreases primary production and thus organic carbon export to the sediment. Iron cycling in the sediment was elevated at the marginal sea ice stations where Fe-reduction led to DFe fluxes in the pore water of up to 0.379 mmol DFe m-2 d-1, while moderate (0.068 mmol DFe m-2 d-1) and negligible fluxes were observed at ice-free and ice-covered stations, respectively. In pore waters, concentrations of DFe and PO43- were significantly correlated with almost identical flux ratios of 0.33 mol PO43- per mol DFe for most of the stations, indicating a strong control of the iron cycling on the phosphate release to the water column. The high benthic DFe and PO43- fluxes highlight the importance of sediments underlying the marginal ice zone as source for limiting nutrients to the shelf waters

The response of seagrass (Posidonia oceanica) meadow metabolism to CO2-levels and hydrodynamic exchange determined with aquatic eddy covariance

We investigated light, water velocity, and CO2 as drivers of primary production in Mediterranean seagrass (Posidonia oceanica) meadows and neighboring bare sands using the aquatic eddy covariance technique. Study locations included an open-water meadow and a nearshore meadow, the nearshore meadow being exposed to greater hydrodynamic exchange. A third meadow was located at a CO2 vent. We found that, despite the oligotrophic environment, the meadows had a remarkably high metabolic activity, up to 20 times higher than the surrounding sands. They were strongly autotrophic, with net production half of gross primary production. Thus, P. oceanica meadows are oases of productivity in an unproductive environment. Secondly, we found that turbulent oxygen fluxes above the meadow can be significantly higher in the afternoon than in the morning at the same light levels. This hysteresis can be explained by the replenishment of nighttime-depleted oxygen within the meadow during the morning. Oxygen depletion and replenishment within the meadow do not contribute to turbulent O2 flux. The hysteresis disappeared when fluxes were corrected for the O2 storage within the meadow and, consequently, accurate metabolic rate measurements require measurements of meadow oxygen content. We further argue that oxygen-depleted waters in the meadow provide a source of CO2 and inorganic nutrients for fixation, especially in the morning. Contrary to expectation, meadow metabolic activity at the CO2 vent was lower than at the other sites, with negligible net primary production

High Net Primary Production of Mediterranean seagrass (Posidonia oceanica) Meadows Determined with Aquatic Eddy Covariance

We report primary production and respiration of Posidonia oceanica meadows determined with the non-invasive aquatic eddy covariance technique. Oxygen fluxes were measured in late spring at an open-water meadow (300 m from shore), at a nearshore meadow (60 m from shore), and at an adjacent sand bed. Despite the oligotrophic environment, the meadows were highly productive and highly autotrophic. Net ecosystem production (54 to 119 mmol m-2 d-1) was about one-half of gross primary production. In adjacent sands, net primary production was a tenth- to a twentieth smaller (4.6 mmol m-2 d-1). Thus, P. oceanica meadows are an oasis of productivity in unproductive surroundings. During the night, dissolved oxygen was depleted in the open-water meadow. This caused a hysteresis where oxygen production in the late afternoon was greater than in the morning at the same irradiance. Therefore, for accurate measurements of diel primary production and respiration in this system, oxygen must be measured within the canopy. Generally, these measurements demonstrate that P. oceanica meadows fix substantially more carbon than they respire. This supports the high rate of organic carbon accumulation and export for which the ecosystem is known

Ciliary flows in corals ventilate target areas of high photosynthetic oxygen production

Most tropical corals live in symbiosis with Symbiodiniaceae algae whose photosynthetic production of oxygen (O2) may lead to excess O2 in the diffusive boundary layer (DBL) above the coral surface. When flow is low, cilia-induced mixing of the coral DBL is vital to remove excess O2 and prevent oxidative stress that may lead to coral bleaching and mortality. Here, we combined particle image velocimetry using O2-sensitive nanoparticles (sensPIV) with chlorophyll (Chla)-sensitive hyperspectral imaging to visualize the microscale distribution and dynamics of ciliary flows and O2 in the coral DBL in relation to the distribution of Symbiodiniaceae Chla in the tissue of the reef building coral, Porites lutea. Curiously, we found an inverse relation between O2 in the DBL and Chla in the underlying tissue, with patches of high O2 in the DBL above low Chla in the underlying tissue surrounding the polyp mouth areas and pockets of low O2 concentrations in the DBL above high Chla in the coenosarc tissue connecting neighboring polyps. The spatial segregation of Chla and O2 is related to ciliary-induced flows, causing a lateral redistribution of O2 in the DBL. In a 2D transport-reaction model of the coral DBL, we show that the enhanced O2 transport allocates parts of the O2 surplus to areas containing less chla, which minimizes oxidative stress. Cilary flows thus confer a spatially complex mass transfer in the coral DBL, which may play an important role in mitigating oxidative stress and bleaching in corals

Ciliary vortex flows and oxygen dynamics in the coral boundary layer

The exchange of metabolites between environment and coral tissue depends on the flux across the diffusive boundary layer (DBL) surrounding the tissue. Cilia covering the coral tissue have been shown to create vortices that enhance mixing in the DBL in stagnant water. To study the role of cilia under simulated ambient currents, we designed a new light-sheet microscopy based flow chamber setup. Microparticle velocimetry was combined with high-resolution oxygen profiling in the coral Porites lutea under varying current and light conditions with natural and arrested cilia beating. Cilia-generated vortices in the lower DBL mitigated extreme oxygen concentrations close to the tissue surface. Under light and arrested cilia, oxygen surplus at the tissue surface increased to 350 µM above ambient, in contrast to 25 µM under ciliary beating. Oxygen shortage in darkness decreased from 120 µM (cilia arrested) to 86 µM (cilia active) below ambient. Ciliary redistribution of oxygen had no effect on the photosynthetic efficiency of the photosymbionts and overall oxygen flux across the DBL indicating that oxygen production and consumption was not affected. We found that corals actively change their environment and suggest that ciliary flows serve predominantly as a homeostatic control mechanism which may play a crucial role in coral stress response and resilience

Extensive nitrogen loss from permeable sediments off North-West Africa

The upwelling area off North-West Africa is characterized by high export production, high nitrate and low oxygen concentration in bottom waters. The underlying sediment consists of sands that cover most of the continental shelf. Due to their permeability sands allow for fast advective pore water transport and can exhibit high rates of nitrogen (N) loss via denitrification as reported for anthropogenically eutrophied regions. However, N loss from sands underlying naturally eutrophied waters is not well studied, and in particular, N loss from the North-West African shelf is poorly constrained. During two research cruises in April/May 2010/2011, sediment was sampled along the North-West African shelf and volumetric denitrification rates were measured in sediment layers down to 8 cm depth using slurry incubations with 15N-labeled nitrate. Areal N loss was calculated by integrating volumetric rates down to the nitrate penetration depth derived from pore water profiles. Areal N loss was neither correlated with water depth nor with bottom water concentrations of nitrate and oxygen but was strongly dependent on sediment grain size and permeability. The derived empirical relation between benthic N loss and grains size suggests that pore water advection is an important regulating parameter for benthic denitrification in sands and further allowed extrapolating rates to an area of 53,000 km2 using detailed sediment maps. Denitrification from this region amounts to 995 kt yr-1 (average 3.6 mmol m-2 d-1) which is 4 times higher than previous estimates based on diffusive pore water transport. Sandy sediments cover 50-60% of the continental shelf and thus may contribute significantly to the global benthic N loss

Aquatic Respiration Rate Measurements at Low Oxygen Concentrations

Despite its huge ecological importance, microbial oxygen respiration in pelagic waters is little studied, primarily due to methodological difficulties. Respiration measurements are challenging because of the required high resolution of oxygen concentration measurements. Recent improvements in oxygen sensing techniques bear great potential to overcome these limitations. Here we compare 3 different methods to measure oxygen consumption rates at low oxygen concentrations, utilizing amperometric Clark type sensors (STOX), optical sensors (optodes), and mass spectrometry in combination with 18-18O2 labeling. Oxygen concentrations and consumption rates agreed well between the different methods when applied in the same experimental setting. Oxygen consumption rates between 30 and 400 nmol L−1 h−1 were measured with high precision and relative standard errors of less than 3%. Rate detection limits in the range of 1 nmol L−1 h−1 were suitable for rate determinations in open ocean water and were lowest at the lowest applied O2 concentration