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Benthic organisms in coastal sediments affect elemental cycling and control benthic-pelagic coupling through particle reworking and ventilation of their burrows. Bioirrigation and associated porewater advection create intermittently oxic regions within sediments. The spatio-temporal patterns of such biogenic redox oscillations likely respond to seasonal factors, but quantitative information on the seasonality of bioirrigator behaviors and associated redox dynamics is scarce. We examined bioirrigation by the maldanid polychaete Clymenella torquata and its impacts on sediment oxygenation patterns in permeable sediments using high-resolution planar optode oxygen imaging. In sediment mesocosms with reconstructed summer-collected sediment, the durations of pumping and resting varied inversely with temperature. The average durations of pumping and resting increased from 4 min/4 min at 21 °C, to 6 min/6 min at 12 °C, to 15 min/14 min at 5 °C. In intact cores collected in summer, irrigation patterns (3.5 min/3.5 min) were similar to those observed at 21 °C during the temperature ramp. Pumping and resting durations in intact cores collected in winter at 6 °C averaged 9 min/26 min, significantly different from patterns at comparable temperatures in the temperature ramp. Pumping patterns strongly affected the temporal patterns of redox dynamics in surrounding sediments. In addition, temperature strongly affected burrow irrigation depth (exclusively within the top ∼10 cm at 21 °C, and down to ∼20 cm at 5–6 °C with an apparent transition at ∼15 °C), indicating that the zone with dynamic redox conditions migrates vertically on a seasonal basis. The differences in pumping patterns between in- and out-of-season experiments and the effect of temperature on irrigation depth underscore the importance of conducting experiments with bioturbators in-season and at field temperatures. The observed seasonal differences in bioirrigation patterns and associated spatio-temporal redox dynamics suggest that rates and pathways of redox-sensitive diagenetic processes and benthic chemical fluxes in permeable sediments likely show considerable seasonal variation. 

Permeable sediments, which represent more than 50% of the continental shelves, have been largely neglected as a potential source of Fe in current global estimates of benthic dissolved iron Fed fluxes. There are open questions regarding the effects of a range of factors on Fed fluxes from these deposits, including seasonal dynamics and the role of bioirrigation. To address these gaps, we performed laboratory-based sediment incubation experiments with muddy sands during summer (21 °C) and winter (7 °C). We used bioirrigation mimics to inject overlying water into the permeable sediment with patterns resembling the bioirrigation activity of the prolific bioturbating polychaete,Clymenella torquata. Newly developed in-line Fe accumulators were used to estimate Fe fluxes with a recirculating set-up. We found high Fed fluxes from sandy sediments, especially in benthic chambers with simulated bioirrigation. In the winter fluxes reached 200 µmol Fed m-2 d-1 at the onset of irrigation and then decreased over the course of a 13-day experiment while in the summer fluxes from irrigated sediments reached 100 µmol Fed m-2 d-1 and remained high throughout a 7-day experiment. Despite different geochemical expressions of Fe-S cycling and resulting porewater Fed concentrations in winter and summer, large Fed fluxes were sustained during both seasons. Solid-phase and porewater concentration profiles showed that maximum concentrations of key constituents, including total solid-phase reactive Fe, and porewater Fed and ammonium, were located closer to the sediment water interface (SWI) in irrigated cores than in non-irrigated cores due to the upward advective transport of dissolved porewater constituents. This upward transport also facilitated Fed fluxes out of the sediments, especially during times of active pumping. Our study demonstrates the potential for large Fed fluxes from sandy sediments in both summer and winter, despite relatively low standing stocks of labile organic matter and porewater Fed. The primary driver of these high fluxes was advective porewater transport, in our study induced by the activity of infaunal organisms. These results suggest that permeable sediments, which dominate shelf regions, must be explicitly considered in global estimates of benthic Fed fluxes, and cannot be simply extrapolated from estimates based on muddy sediments. 

Cable bacteria that are capable of transporting electrons on centimeter scales have been found in a variety of sediment types, where their activity can strongly influence diagenetic reactions and elemental cycling. In this study, the patterns of spatial and temporal colonization of surficial sediment by cable bacteria were revealed in two-dimensions by planar pH and H₂S optical sensors for the first time. The characteristic sediment surface pH maximum zones begin to develop from isolated micro-regions and spread horizontally within 5 days, with lateral spreading rates from 0.3 to ~ 1.2 cm day⁻¹. Electrogenic anodic zones in the anoxic sediments are characterized by low pH, and the coupled pH minima also expand with time. H₂S heterogeneities in accordance with electrogenic colonization are also observed. Cable bacteria cell abundance in oxic surface sediment (0–0.25 cm) kept almost constant during the colonization period; however, subsurface cell abundance apparently increased as electrogenic activity expanded across the entire surface. Changes in cell abundance are consistent with filament coiling and growth in the anodic zone (i.e., cathodic snorkels). The spreading mechanism for the sediment pH–H₂S fingerprints and the cable bacteria abundance dynamics suggest that once favorable microenvironments are established, filamentous cable bacteria aggregate or locally activate electrogenic metabolism. Different development dynamics in otherwise similar sediment suggests that the accessibility of reductant (e.g., dissolved phase sulfide) is critical in controlling the growth of cable bacteria.

 

Benthic animals profoundly influence the cycling and storage of carbon and other elements in marine systems, particularly in coastal sediments. Recent climate change has altered the distribution and abundance of many seafloor taxa and modified the vertical exchange of materials between ocean and sediment layers. Here, we examine how climate change could alter animal-mediated biogeochemical cycling in ocean sediments. The fossil record shows repeated major responses from the benthos during mass extinctions and global carbon perturbations, including reduced diversity, dominance of simple trace fossils, decreased burrow size and bioturbation intensity, and nonrandom extinction of trophic groups. The broad dispersal capacity of many extant benthic species facilitates poleward shifts corresponding to their environmental niche as overlying water warms. Evidence suggests that locally persistent populations will likely respond to environmental shifts through either failure to respond or genetic adaptation rather than via phenotypic plasticity. Regional and global ocean models insufficiently integrate changes in benthic biological activity and their feedbacks on sedimentary biogeochemical processes. The emergence of bioturbation, ventilation, and seafloor-habitat maps and progress in our mechanistic understanding of organism–sediment interactions enable incorporation of potential effects of climate change on benthic macrofaunal mediation of elemental cycles into regional and global ocean biogeochemical models. 

Electrogenic cable bacteria can couple spatially separated redox reaction zones in marine sediments using multicellular filaments as electron conductors. Reported as generally absent from disturbed sediments, we have found subsurface cable aggregations associated with tubes of the parchment worm Chaetopterus variopedatus in otherwise intensely bioturbated deposits. Cable bacteria tap into tubes, which act as oxygenated conduits, creating a three-dimensional conducting network extending decimeters into sulfidic deposits. By elevating pH, promoting Mn, Fe-oxide precipitation in tube linings, and depleting S around tubes, they enhance tube preservation and favorable biogeochemical conditions within the tube. The presence of disseminated filaments a few cells in length away from oxygenated interfaces and the reported ability of cable bacteria to use a range of redox reaction couples suggest that these microbes are ubiquitous facultative opportunists and that long filaments are an end-member morphological adaptation to relatively stable redox domains. 

Benthic iron (Fe) fluxes from continental shelf sediments are an important source of Fe to the global ocean, yet the magnitude of these fluxes is not well constrained. Processing of Fe in sediments is of particular importance in the Arctic Ocean, which has a large shelf area and Fe limitation of primary productivity. In the Arctic fjords of Svalbard, glacial weathering delivers high volumes of Fe‐rich sediment to the fjord benthos. Benthic redox cycling of Fe proceeds through multiple pathways of reduction (i.e., dissimilatory iron reduction and reduction by hydrogen sulfide) and re‐oxidation. There are few estimates of the magnitude and controlling factors of the benthic Fe flux in Arctic fjords. We collected cores from two Svalbard fjords (Kongsfjorden and Lilliehöökfjorden), measured dissolved Fe²⁺concentrations using a two‐dimensional sensor, and analyzed iron, manganese, carbon, and sulfur species to study benthic Fe fluxes. Benthic fluxes of Fe²⁺vary throughout the fjords, with a “sweet spot” mid‐fjord controlled by the availability of organic carbon linked to sedimentation rates. The flux is also impacted by fjord circulation and sea ice cover, which influence overall mineralization rates in the sediment. Due to ongoing Arctic warming, we predict an increase in the benthic Fe²⁺flux with reduced sea ice cover in some fjords and a decrease in the Fe²⁺flux with the retreat of tidewater glaciers in other regions. Decreasing benthic Fe²⁺fluxes in fjords may exacerbate Fe limitation of primary productivity in the Arctic Ocean.