Intertidal permeable sediments are crucial sites of organic matter remineralization. These sediments likely have a large capacity to produce reactive oxygen species (ROS) because of shifting oxic-anoxic interfaces and intense iron-sulfur cycling. Here, we show that high concentrations of the ROS hydrogen peroxide are present in intertidal sediments using microsensors, and chemiluminescent analysis on extracted porewater. We furthermore investigate the effect of ROS on potential rates of microbial degradation processes in intertidal surface sediments after transient oxygenation, using slurries that transitioned from oxic to anoxic conditions. Enzymatic removal of ROS strongly increases rates of aerobic respiration, sulfate reduction and hydrogen accumulation. We conclude that ROS are formed in sediments, and subsequently moderate microbial mineralization process rates. Although sulfate reduction is completely inhibited in the oxic period, it resumes immediately upon anoxia. This study demonstrates the strong effects of ROS and transient oxygenation on the biogeochemistry of intertidal sediments.

Photogranules are spherical aggregates formed of complex phototrophic ecosystems with potential for “aeration-free” wastewater treatment. Photogranules from a sequencing batch reactor were investigated by fluorescence microscopy, 16S/18S rRNA gene amplicon sequencing, microsensors, and stable- and radioisotope incubations to determine the granules’ composition, nutrient distribution, and light, carbon, and nitrogen budgets. The photogranules were biologically and chemically stratified, with filamentous cyanobacteria arranged in discrete layers and forming a scaffold to which other organisms were attached. Oxygen, nitrate, and light gradients were also detectable. Photosynthetic activity and nitrification were both predominantly restricted to the outer 500 µm, but while photosynthesis was relatively insensitive to the oxygen and nutrient (ammonium, phosphate, acetate) concentrations tested, nitrification was highly sensitive. Oxygen was cycled internally, with oxygen produced through photosynthesis rapidly consumed by aerobic respiration and nitrification. Oxygen production and consumption were well balanced. Similarly, nitrogen was cycled through paired nitrification and denitrification, and carbon was exchanged through photosynthesis and respiration. Our findings highlight that photogranules are complete, complex ecosystems with multiple linked nutrient cycles and will aid engineering decisions in photogranular wastewater treatment.

<p>The dominant pathway for of dissimilatory nitrogen metabolism, dissimilatory nitrate reduction to ammonium (DNRA) versus denitrification, remains challenging to predict in costal sediments. In an intertidal sand flat in the Wadden Sea, the capacity for DRNA seems to relate directly to the capacity for sulfate reduction. Part of this capacity appears to be due to microorganisms capable of perform both DNRA and sulfate reduction switching metabolisms according to their environment. It has previously been hypothesized that the capacity for DNRA may help sulfate reducers cope with oxygen stress, as well as eliminate toxic nitrite from their surroundings. We investigated the effect of transient oxygen exposure on denitrification, DNRA, and sulfate reduction, as well as the impact of nitrate exposure on sulfate reducers subjected to oxygen exposure. We found that in incubations containing nitrate, denitrification was not affected by transient oxygen exposure while DNRA and sulfate reduction were both depressed. Since DNRA and sulfate reduction were simultaneously depressed, this change is not due to sulfate reducers switching their metabolism. In contrast, in the absence of nitrate, transient oxygen exposure did not affect the rate of sulfate reduction upon anoxia. As expected, the rate of sulfate reduction in the absence of nitrate was higher than in the presence of nitrate. Together, this suggests that an intermediate of nitrate reduction other than nitrite, which was higher in the anoxic group, affects the activity of sulfate reducers.</p>

The oscillating redox conditions that characterize coastal sandy sediments foster microbial communities capable of respiring oxygen and nitrate simultaneously, thereby increasing the potential for organic matter remineralization, nitrogen (N)-loss and emissions of the greenhouse gas nitrous oxide. It is unknown to what extent these conditions also lead to overlaps between dissimilatory nitrate and sulfate respiration. Here, we show that sulfate and nitrate respiration co-occur in the surface sediments of an intertidal sand flat. Furthermore, we found strong correlations between dissimilatory nitrite reduction to ammonium (DNRA) and sulfate reduction rates. Until now, the nitrogen and sulfur cycles were assumed to be mainly linked in marine sediments by the activity of nitrate-reducing sulfide oxidisers. However, transcriptomic analyses revealed that the functional marker gene for DNRA (nrfA) was more associated with microorganisms known to reduce sulfate rather than oxidise sulfide. Our results suggest that when nitrate is supplied to the sediment community upon tidal inundation, part of the sulfate reducing community may switch respiratory strategy to DNRA. Therefore increases in sulfate reduction rate in-situ may result in enhanced DNRA and reduced denitrification rates. Intriguingly, the shift from denitrification to DNRA did not influence the amount of N2O produced by the denitrifying community. Our results imply that microorganisms classically considered as sulfate reducers control the potential for DNRA within coastal sediments when redox conditions oscillate and therefore retain ammonium that would otherwise be removed by denitrification, exacerbating eutrophication.

<p>We investigated the influence of reactive oxygen species (ROS) on microbial mineralization in intertidal permeable sediments. These sediments are crucial for coastal carbon cycling. Permeable intertidal sediments are further prone to variable surface oxygenation and active iron-sulfur cycling, and are therefore likely sites of intense ROS formation. We incubated sediment slurries from an intertidal sandflat in the German Wadden Sea over a transition to anoxic conditions, and found that removal of ROS by enzymes increased rates of aerobic and anaerobic respiration, including sulfate reduction. We additionally found high concentrations of the ROS hydrogen peroxide in sediment porewaters. Sulfate reduction was absent during the oxic period, but directly resumed upon anoxia.</p><p>This study shows the regulating effect of ROS on microbial mineralization and the impact of ROS and transient oxygenation on marine sediment biogeochemistry.</p>