We measured denitrification in permeable sediments in a sealed flume tank with environmentally representative fluid flow and solute transport behavior using novel measurements. Numerical flow and reactive transport models representing the flume experiments were implemented to provide mechanistic insight into the coupled hydrodynamic and biogeochemical processes. There was broad agreement between the model results and experimental data. The model showed that the coupling between nitrification and denitrification was relatively weak in comparison to that in cohesive sediments. This was due to the direct advective transport between anoxic pore water and the overlying water column, and little interaction between the mostly oxic advective region and the underlying anoxic region. Denitrification was therefore mainly fueled by nitrate supplied from the water column. This suggests that the capacity of permeable sediments with nonmigratory ripples to remove bioavailable nitrogen from coastal ecosystems is lower than that of cohesive sediments. We conclude that while experimental measurements provide a good starting point for constraining key parameters, reactive transport models with realistic kinetic and transport parameters provide critical insight into biogeochemical processes in permeable sediment that are difficult to experimentally evaluate.
Ecological theory suggests that habitat disturbance differentially influences distributions of habitat generalist and specialist species. While well-established for macroorganisms, this theory has rarely been explored for microorganisms. Here we tested these principles in permeable (sandy) sediments, ecosystems with much spatiotemporal variation in resource availability and physicochemical conditions. Microbial community composition and function were profiled in intertidal and subtidal sediments using 16S rRNA gene amplicon sequencing and metagenomics, yielding 135 metagenome-assembled genomes. Community composition and metabolic traits modestly varied with sediment depth and sampling date. Several taxa were highly abundant and prevalent in all samples, including within the orders Woeseiales and Flavobacteriales, and classified as habitat generalists; genome reconstructions indicate these taxa are highly metabolically flexible facultative anaerobes and adapt to resource variability by using different electron donors and acceptors. In contrast, obligately anaerobic taxa such as sulfate reducers and candidate lineage MBNT15 were less abundant overall and only thrived in more stable deeper sediments. We substantiated these findings by measuring three metabolic processes in these sediments; whereas the habitat generalist-associated processes of sulfide oxidation and fermentation occurred rapidly at all depths, the specialist-associated process of sulfate reduction was restricted to deeper sediments. A manipulative experiment also confirmed habitat generalists outcompete specialist taxa during simulated habitat disturbance. Together, these findings show metabolically flexible habitat generalists become dominant in highly dynamic environments, whereas metabolically constrained specialists are restricted to narrower niches. Thus, an ecological theory describing distribution patterns for macroorganisms likely extends to microorganisms. Such findings have broad ecological and biogeochemical ramifications.
Permeable (sandy) sediments cover half of the continental margin and are major regulators of oceanic carbon cycling. The microbial communities within these highly dynamic sediments frequently shift between oxic and anoxic states, and hence are less stratified than those in cohesive (muddy) sediments. A major question is, therefore, how these communities maintain metabolism during oxic-anoxic transitions. Here, we show that molecular hydrogen (H 2 ) accumulates in silicate sand sediments due to decoupling of bacterial fermentation and respiration processes following anoxia. In situ measurements show that H 2 is 250-fold supersaturated in the water column overlying these sediments and has an isotopic composition consistent with fermentative production. Genome-resolved shotgun metagenomic profiling suggests that the sands harbour diverse and specialized microbial communities with a high abundance of [NiFe]-hydrogenase genes. Hydrogenase profiles predict that H 2 is primarily produced by facultatively fermentative bacteria, including the dominant gammaproteobacterial family Woeseiaceae, and can be consumed by aerobic respiratory bacteria. Flow-through reactor and slurry experiments consistently demonstrate that H 2 is rapidly produced by fermentation following anoxia, immediately consumed by aerobic respiration following reaeration and consumed by sulfate reduction only during prolonged anoxia. Hydrogenotrophic sulfur, nitrate and nitrite reducers were also detected, although contrary to previous hypotheses there was limited capacity for microalgal fermentation. In combination, these experiments confirm that fermentation dominates anoxic carbon mineralization in these permeable sediments and, in contrast to the case in cohesive sediments, is largely uncoupled from anaerobic respiration. Frequent changes in oxygen availability in these sediments may have selected for metabolically flexible bacteria while excluding strict anaerobes.
The balance between estuarine denitrification and dissimilatory nitrate reduction to ammonium (DNRA) is critical for determining nitrogen loads received by oceans from inland waters. We aimed to determine the factors controlling the ratio between these processes and determining whether nitrogen was generally removed or recycled in estuaries. Rates of denitrification and DNRA with depth were measured in intact sediment cores in 11 estuaries along the coast of Victoria, Australia. The estuaries studied represent a range of biogeochemical conditions, land use, and catchment size. At a pore water profile scale, the ratio of denitrification to DNRA was well predicted by a multiple regression model with nitrate concentration in the overlying water, dissolved pore water iron, and ammonium as the predictor variables. Areal denitrification rates varied from 4 to 150 μmol · m−2 · hr−1, and DNRA rates varied from 2 to 30 μmol · m−2 · hr−1, with the ratio of denitrification to DNRA spanning a range from denitrification‐dominated (denitrification/DNRA = 8.4) to DNRA‐dominated (denitrification/DNRA = 0.3). DNRA dominated at sites with high iron pools and high organic carbon to nitrate ratios. We conclude that low nitrate and high Fe2+ availability generally enhances DNRA and drives an estuary toward being N recycling, rather than N removing.
Dissolved oxygen (DO) is a key environmental variable that drives and feeds back with numerous processes. In the aquatic sediment that makes up the hyporheic zone, DO may exhibit pronounced spatial gradients and complex patterns which control the distribution of a series of redox processes. Yet, little is known regarding the dynamics of hyporheic zone DO, especially under transitional flow regimes. Considering the natural tendency of rivers to be highly responsive to external forcing, these temporal dynamics are potentially just as important and pronounced as the spatial gradients. Here we use laboratory flume experiments and multiphysics flow and reactive transport modeling to investigate surface flow controls on the depth of oxygen penetration in the bed as well as the area of oxygenated sediment. We show that the hyporheic zone DO conditions respond over time scales of hours‐to‐days when subjected to practically instantaneous surface flow perturbations. Additionally, the flume experiments demonstrate that hyporheic zone DO conditions respond faster to surface flow acceleration than to deceleration. Finally, we found that the morphology of the dissolved oxygen plume front depends on surface flow acceleration or deceleration. This study thus shows that the highly dynamic nature of typical streams and rivers drives equally dynamic redox conditions in the hyporheic zone. Because the redox conditions and their distribution within the hyporheic zone are important from biological, ecological, and contaminant perspectives, this hyporheic redox dynamism has the potential to impact system scale aquatic chemical cycles.
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