| 1. |
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| 2. |
- Albert, Séréna, 1992-
(författare)
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Benthic-pelagic coupling in a changing world : Structural and functional responses of microbenthic communities to organic matter settling
- 2021
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Marine soft sediments form the second largest habitat on the planet. Organisms residing in this environment represent a vast reservoir of biodiversity, and play key roles in ecosystem processes. Most benthic organisms depend on organic matter (OM) inputs from phytoplankton in the overlying water column as food supply, but human impacts such as eutrophication and climate change are profoundly altering natural ecosystem dynamics. The consequences of changes in benthic-pelagic coupling for the biodiversity and functioning of soft-sediment communities have yet to be resolved. The aim of this thesis is to assess the role of OM settling on soft-sediments microeukaryotic (small organisms < 1 mm) and bacterial communities. The intents are two-fold, to investigate impacts on (1) community structure and diversity (chapters I, II and IV); and (2) ecosystem functioning, notably in relation to nitrogen (N) cycling (chapters I and III). Our results show that settling OM quantity and quality both had a significant impact on microeukaryotic alpha-diversity. We observed a decrease in alpha-diversity following settling of diatom-derived spring bloom OM, possibly as a result of competitive exclusion, while cyanobacteria-derived summer bloom OM did not affect alpha-diversity (chapters I and IV). We also found that high biomass of diatoms and others fast sinking phytoplankton groups in the water column led to lower microeukaryotic alpha diversity after this material settled on the seafloor (chapter IV). Presumably, following this large sedimentation event, sediment oxygen (O2) demand was strongly stimulated, excluding O2-sensitive taxa. Overall, we propose that the assembly of microeukaryotic communities was primarily mediated by OM settling quantity (chapter IV), while differences in OM quality led to significant but more subtle changes, occurring at fine taxonomic level (chapter I). The response of bacterial communities to OM settling was less pronounced, and probably restricted to the uppermost sediment layer (chapters I and IV). We did, however, observe a significant effect of OM quality on bacterial communities assembly at the sediment-water interface, with taxa favored either by diatom- or by cyanobacteria-derived OM (chapter II). This study also showed that feedback mechanisms from nutrient recycling in the sediment could play a role in this response. Finally, our results indicated a substantial influence of OM quality on N cycling at the sediment-water interface. We found that settling of fresh OM (i.e. low C:N ratio) stimulated denitrification activity (chapters I and III), while simultaneously promoting more N recycling to the water column than settling of degraded OM (i.e. high C:N ratio) did (chapter III). Altogether, our results indicate that current changes in OM settling dynamics in marine systems will likely impact microeukaryotic and, to some extent, bacterial biodiversity in soft sediments. Alterations in settling OM quality, in particular, may also affect crucial microbial processes involved in N cycling. This thesis highlights the importance of considering benthic-pelagic coupling mechanisms to better understand likely future changes in marine ecosystems.
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| 3. |
- Albert, Séréna, et al.
(författare)
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Influence of settling organic matter quantity and quality on benthic nitrogen cycling
- 2021
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Ingår i: Limnology and Oceanography. - : Wiley. - 1939-5590 .- 0024-3590. ; 66:5, s. 1882-1895
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Tidskriftsartikel (refereegranskat)abstract
- Coastal sediments are major contributors to global carbon (C) mineralization and nutrient cycling and are tightly linked to processes in the pelagic environment. In this study, we aimed to investigate the regulating potential of quantity and quality of planktonic organic matter (OM) deposition on benthic metabolism, with a particular focus on nitrogen (N) cycling processes. We simulated inputs of spring (C : N 10.9) and summer (C : N 5.6) plankton communities in high and low quantities to sediment cores, and followed oxygen consumption, nutrient fluxes as well as nitrate reduction rates, that is, denitrification and dissimilatory nitrate reduction to ammonium for 10 d. Our results demonstrate the primary importance of OM quality in determining the fate of organic N once it settles to the sediment surface. Settling of N-rich summer plankton material resulted in a ∼ twofold lower denitrification efficiency (40–56%) compared to N-poor spring plankton (88–115%). This indicates that N-rich plankton deposition favors recycling of inorganic nutrients to the water column over N-loss via denitrification. OM quantity was positively related to mineralization activity, but this neither directly affected N fluxes nor denitrification activity, highlighting the complex interplay between the OM quantity and quality in regulating N cycling. In light of these new findings, we support the use of simple qualitative indicators such as C : N ratio of OM to investigate how future changes in benthic-pelagic coupling might influence N budgets at the sediment–water interface.
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| 4. |
- Asplund, Maria. E., 1970, et al.
(författare)
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Methane Emissions From Nordic Seagrass Meadow Sediments
- 2022
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Ingår i: Frontiers in Marine Science. - : Frontiers Media S.A.. - 2296-7745. ; 8
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Tidskriftsartikel (refereegranskat)abstract
- Shallow coastal soft bottoms are important carbon sinks. Submerged vegetation has been shown to sequester carbon, increase sedimentary organic carbon (C-org) and thus suppress greenhouse gas (GHG) emissions. The ongoing regression of seagrass cover in many areas of the world can therefore lead to accelerated emission of GHGs. In Nordic waters, seagrass meadows have a high capacity for carbon storage, with some areas being recognized as blue carbon hotspots. To what extent these carbon stocks lead to emission of methane (CH4) is not yet known. We investigated benthic CH4 emission (i.e., net release from the sediment) in relation to seagrass (i.e. Zostera marina) cover and sedimentary C-org content (%) during the warm summer period (when emissions are likely to be highest). Methane exchange was measured in situ with benthic chambers at nine sites distributed in three regions along a salinity gradient from similar to 6 in the Baltic Sea (Finland) to similar to 20 in Kattegat (Denmark) and similar to 26 in Skagerrak (Sweden). The net release of CH4 from seagrass sediments and adjacent unvegetated areas was generally low compared to other coastal habitats in the region (such as mussel banks and wetlands) and to other seagrass areas worldwide. The lowest net release was found in Finland. We found a positive relationship between CH4 net release and sedimentary C-org content in both seagrass meadows and unvegetated areas, whereas no clear relationship between seagrass cover and CH4 net release was observed. Overall, the data suggest that Nordic Zostera marina meadows release average levels of CH4 ranging from 0.3 to 3.0 mu g CH4 m(-2) h(-1), which is at least 12-78 times lower (CO2 equivalents) than their carbon accumulation rates previously estimated from seagrass meadows in the region, thereby not hampering their role as carbon sinks. Thus, the relatively weak CH4 emissions from Nordic Z. marina meadows will not outweigh their importance as carbon sinks under present environmental conditions.
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| 5. |
- Björk, Mats, 1960-, et al.
(författare)
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Methane emissions from macrophyte beach wrack on Baltic seashores
- 2023
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Ingår i: Ambio. - : Springer Nature. - 0044-7447 .- 1654-7209. ; 52:1, s. 171-181
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Tidskriftsartikel (refereegranskat)abstract
- Beach wrack of marine macrophytes is a natural component of many beaches. To test if such wrack emits the potent greenhouse gas methane, field measurements were made at different seasons on beach wrack depositions of different ages, exposure, and distance from the water. Methane emissions varied greatly, from 0 to 176 mg CH4-C m−2 day−1, with a clear positive correlation between emission and temperature. Dry wrack had lower emissions than wet. Using temperature data from 2016 to 2020, seasonal changes in fluxes were calculated for a natural wrack accumulation area. Such calculated average emissions were close to zero during winter, but peaked in summer, with very high emissions when daily temperatures exceeded 20 °C. We conclude that waterlogged beach wrack significantly contributes to greenhouse gas emissions and that emissions might drastically increase with increasing global temperatures. When beach wrack is collected into heaps away from the water, the emissions are however close to zero.
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| 6. |
- Bonaglia, Stefano, et al.
(författare)
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Activated carbon stimulates microbial diversity and PAH biodegradation under anaerobic conditions in oil-polluted sediments
- 2020
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Ingår i: Chemosphere. - : Elsevier BV. - 0045-6535 .- 1879-1298. ; 248
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Tidskriftsartikel (refereegranskat)abstract
- Biodegradation by microorganisms is a useful tool that helps alleviating hydrocarbon pollution in nature. Microbes are more efficient in degradation under aerobic than anaerobic conditions, but the majority of sediment by volume is generally anoxic. Incubation experiments were conducted to study the biodegradation potential of naphthalene-a common polycyclic aromatic hydrocarbon (PAH)-and the diversity of microbial communities in presence/absence of activated carbon (AC) under aerobic/anaerobic conditions. Radio-respirometry experiments with endogenous microorganisms indicated that degradation of naphthalene was strongly stimulated (96%) by the AC addition under anaerobic conditions. In aerobic conditions, however, AC had no effects on naphthalene biodegradation. Bioaugmentation tests with cultured microbial populations grown on naphthalene showed that AC further stimulated (92%) naphthalene degradation in anoxia. Analysis of the 16S rRNA gene sequences implied that sediment amendment with AC increased microbial community diversity and changed community structure. Moreover, the relative abundance of Geobacter, Thiobacillus, Sulfuricurvum, and methanogenic archaea increased sharply after amendment with AC under anaerobic conditions. These results may be explained by the fact that AC particles promoted direct interspecies electron transfer (DIET) between microorganisms involved in PAH degradation pathways. We suggest that important ecosystem functions mediated by microbes-such as hydrocarbon degradation-can be induced and that AC enrichment strategies can be exploited for facilitating bioremediation of anoxic oil-contaminated sediments and soils.
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| 7. |
- Bonaglia, Stefano
(författare)
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Benthic metabolism and sediment nitrogen cycling in Baltic sea coastal areas : the role of eutrophication, hypoxia and bioturbation
- 2012
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Licentiatavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Eutrophication is one of the greatest threats for the Baltic Sea, and one of its more critical consequences is bottom water hypoxia. Nutrient enrichment and oxygen-depletion affect both the deep central basins and a number of coastal areas, even though strategies for nutrient reduction have lately been implemented. In order to better understand why those threats are expanding and formulate more effective remediation strategies two main achievements are needed: (1) new data on benthic nutrient dynamics should be available in order to develop updated budgets for sensitive Baltic areas; (2) the main transformation processes and their regulation mechanisms (i.e. oxygen availability, presence of macrofauna, different organic loading scenarios) should be better constrained.Paper I was able to demonstrate that re-oxygenation of previously anoxic sediment has a positive effect on the ecosystem because of better retention of nutrients and efficient conversion of fixed nitrogen to nitrogen gas. Sediment colonization by the invasive genus Marenzelleria counteracts some of the positive aspects provided by benthic oxygenation (in particular, nutrient retention, N2 loss). A possible explanation for this reversal can be that Marenzelleria does stimulate anaerobic more that aerobic metabolism.Results from Paper II suggest that at the outermost stations of Himmerfjärden denitrification follows a pronounced seasonal pattern, primarily regulated by bottom water temperatures. At the innermost and impacted site oxygen level in the bottom water varies considerably during the year and causes denitrification/DNRA predominance to be the main nitrate reduction pathway. On an annual scale, the net amount of lost N2 is comparable at the four sampling sites and accounts for 96% of the total DIN discharged from the sewage treatment plant, suggesting that denitrification in the estuarine sediment acts as a major nitrogen sink for external N inputs.
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| 8. |
- Bonaglia, Stefano, et al.
(författare)
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Capping with activated carbon reduces nutrient fluxes, denitrification and meiofauna in contaminated sediments
- 2019
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Ingår i: Water Research. - : Elsevier BV. - 0043-1354 .- 1879-2448. ; 148, s. 515-525
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Tidskriftsartikel (refereegranskat)abstract
- Sediment capping with activated carbon (AC) is an effective technique used in remediation of contaminated sediments, but the ecological effects on benthic microbial activity and meiofauna communities have been largely neglected. This study presents results from a 4-week experiment investigating the influence of two powdered AC materials (bituminous coal-based and coconut shell-derived) and one control material (clay) on biogeochemical processes and meiofauna in contaminated sediments. Capping with AC induced a 62‒63% decrease in denitrification and a 66‒87 % decrease in dissimilatory nitrate reduction to ammonium (DNRA). Sediment porewater pH increased from 7.1 to 9.0 and 9.7 after addition of bituminous AC and biomass-derived AC, respectively. High pH (>8) persisted for at least two weeks in the bituminous AC and for at least 24 days in the coconut based AC, while capping with clay had no effect on pH. We observed a strong impact (nitrate fluxes being halved in presence of AC) on nitrification activity as nitrifiers are sensitive to high pH. This partly explains the significant decrease in nitrate reduction rates since denitrification was almost entirely coupled to nitrification. Total benthic metabolism estimated by sediment oxygen uptake was reduced by 30 and 43 % in presence of bituminous coal-based AC and coconut shell-derived AC, respectively. Meiofauna abundances decreased by 60‒62 % in the AC treatments. Taken together, these observations suggest that AC amendments deplete natural organic carbon, intended as food, to heterotrophic benthic communities. Phosphate efflux was 91 % lower in presence of bituminous AC compared to untreated sediment probably due to its content of aluminum (Al) oxides, which have high affinity for phosphate. This study demonstrates that capping with powdered AC produces significant effects on benthic biogeochemical fluxes, microbial processes and meiofauna abundances, which are likely due to an increase in porewater pH and to the sequestration of natural, sedimentary organic matter by AC particles.
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| 9. |
- Bonaglia, Stefano, 1983-
(författare)
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Control factors of the marine nitrogen cycle : The role of meiofauna, macrofauna, oxygen and aggregates
- 2015
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- The ocean is the most extended biome present on our planet. Recent decades have seen a dramatic increase in the number and gravity of threats impacting the ocean, including discharge of pollutants, cultural eutrophication and spread of alien species. It is essential therefore to understand how different impacts may affect the marine realm, its life forms and biogeochemical cycles. The marine nitrogen cycle is of particular importance because nitrogen is the limiting factor in the ocean and a better understanding of its reaction mechanisms and regulation is indispensable. Furthermore, new nitrogen pathways have continuously been described. The scope of this project was to better constrain cause-effect mechanisms of microbially mediated nitrogen pathways, and how these can be affected by biotic and abiotic factors.This thesis demonstrates that meiofauna, the most abundant animal group inhabiting the world’s seafloors, considerably alters nitrogen cycling by enhancing nitrogen loss from the system. In contrast, larger fauna such as the polychaete Marenzelleria spp. enhance nitrogen retention, when they invade eutrophic Baltic Sea sediments. Sediment anoxia, caused by nutrient excess, has negative consequences for ecosystem processes such as nitrogen removal because it stops nitrification, which in turn limits both denitrification and anammox. This was the case of Himmerfjärden and Byfjord, two estuarine systems affected by anthropogenic activities, such as treated sewage discharges. When Byfjord was artificially oxygenated, nitrate reduction mechanisms started just one month after pumping. However, the balance between denitrification and nitrate ammonification did not favor either nitrogen removal or its retention.Anoxia is also present in aggregates of the filamentous cyanobacteria Nodularia spumigena. This thesis shows that even in fully oxic waters, millimetric aggregates can host anaerobic nitrogen processes, with clear implications for the pelagic compartment. While the thesis contributed to our knowledge on marine nitrogen cycling, more data need to be collected and experiments performed in order to understand key processes and regulation mechanisms of element cycles in the ocean. In this way, stakeholders may follow and take decisions in order to limit the continuous flow of human metabolites and impacts on the marine environment.
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| 10. |
- Bonaglia, Stefano, et al.
(författare)
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Denitrification and DNRA at the Baltic Sea oxic-anoxic interface : Substrate spectrum and kinetics
- 2016
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Ingår i: Limnology and Oceanography. - : Wiley. - 0024-3590 .- 1939-5590. ; 61:5, s. 1900-1915
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Tidskriftsartikel (refereegranskat)abstract
- The dependence of denitrification and dissimilatory nitrate reduction to ammonium (DNRA) on different electron donors was tested in the nitrate-containing layer immediately below the oxic-anoxic interface (OAI) at three stations in the central anoxic basins of the Baltic Sea. Additionally, pathways and rates of fixed nitrogen transformation were investigated with N-15 incubation techniques without addition of donors. Denitrification and anammox were always detected, but denitrification rates were higher than anammox rates. DNRA occurred at two sites and rates were two orders of magnitude lower than denitrification rates. Separate additions of dissolved organic carbon and sulfide stimulated rates without time lag indicating that both organotrophic and lithotrophic bacterial populations were simultaneously active and that they could carry out denitrification or DNRA. Manganese addition stimulated denitrification and DNRA at one station, but it is not clear whether this was due to a direct or indirect effect. Ammonium oxidation to nitrite was detected on one occasion. During denitrification, the production of nitrous oxide (N2O) was as important as dinitrogen (N-2) production. A high ratio of N2O to N-2 production at one site may be due to copper limitation, which inhibits the last denitrification step. These data demonstrate the coexistence of a range of oxidative and reductive nitrogen cycling processes at the Baltic OAI and suggest that the dominant electron donor supporting denitrification and DNRA is organic matter. Organotrophic denitrification is more important for nitrogen budgets than previously thought, but the large temporal variability in rates calls for long-term seasonal studies.
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| 11. |
- Bonaglia, Stefano, et al.
(författare)
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EFFECT OF MEIOFAUNA ON BENTHIC ELEMENT CYCLING IN A BALTIC SEA COASTAL AREA
- 2013
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Konferensbidrag (refereegranskat)abstract
- We have studied the role of meiofaunal communities for nutrient cycling and organic matter mineralization in coastal sediments of the Baltic Sea. Although meiofauna is orders of magnitude more abundant than macrofauna and has commonly a much more diverse community structure, its importance for sediment biogeochemical pathways is poorly understood because of objective experimental difficulties when manipulating meiofauna communities due to small body sizes (0.04 to 1 mm) and inherent fragility. We used a density extraction method to separate intact and living metazoans from sediment and tested the effect of low meiofauna and high meiofauna abundances in the presence and absence of macrofauna for exchange rates of nutrients, O2, DIC, N2, and CH4. High abundances of meiofauna stimulated O2 uptake and the net N2 efflux by 16% and 34%, respectively, but did not change oxygen penetration depths significantly. By contrast, macrofauna increased oxygen penetration depths by 21% and stimulated methane emissions by a factor of 8. These results demonstrate the importance of meiofauna in the regulation of aerobic and anaerobic microbial processes and benthic fluxes in marine sediments.
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| 12. |
- Bonaglia, Stefano, et al.
(författare)
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Effect of reoxygenation and Marenzelleria spp. bioturbation on Baltic Sea sediment metabolism
- 2013
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Ingår i: Marine Ecology Progress Series. - : Inter Research. - 0171-8630 .- 1616-1599. ; 482, s. 43-55
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Tidskriftsartikel (refereegranskat)abstract
- Nutrient reduction and the improvement of bottom water oxygen concentrations are thought to be key factors in the recovery of eutrophic aquatic ecosystems. The effects of reoxygenation and bioturbation of natural hypoxic sediments in the Baltic Sea were studied using a mesocosm experiment. Anoxic sediment box cores were collected from 100 m depth in Kanholmsfjärden (Stockholm Archipelago) and maintained in flow-through mesocosms with 3 treatments: (1) hypoxic: supplied with hypoxic water; (2) normoxic: supplied with oxic water; and (3) Marenzelleria: supplied with oxic water and the polychaete Marenzelleria spp. (2000 ind. m–2). After a 7 wk long conditioning period, net fluxes of dissolved O2, CH4, Fe2+, Mn2+, NH4+, NO2-, NO3-, PO43- and H4SiO4, and rates of nitrate ammonification (DNRA), denitrification and anammox were determined. Phosphate was taken up by the sediment in all treatments, and the uptake was highest in the normoxic treatment with Marenzelleria. Normoxic conditions stimulated the denitrification rate by a factor of 5. Denitrification efficiency was highest under normoxia (50%), intermediate in bioturbated sediments (16%), and very low in hypoxic sediments (4%). The shift from hypoxic to normoxic conditions resulted in a significantly higher retention of NH4+, H4SiO4 and Mn2+ in the sediment, but the bioturbation by Marenzelleria reversed this effect. Results from our study suggest that bioturbation by Marenzelleria stimulates the exchange of solutes between sediment and bottom water through irrigation and enhances bacterial sulfate reduction in the burrow walls. The latter may have a toxic effect on nitrifying bacteria, which, in turn, suppresses denitrification rates.
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| 13. |
- Bonaglia, Stefano, 1983, et al.
(författare)
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High methane emissions from an anoxic fjord driven by mixing and oxygenation : High methane emissions from fjords
- 2022
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Ingår i: Limnology and Oceanography Letters. - : Wiley. - 2378-2242. ; 7:5, s. 392-400
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Tidskriftsartikel (refereegranskat)abstract
- Oceanic methane (CH4) budgets lack data from high-latitude fjords that often behave as intermittently anoxic ecosystems with potentially high methane emissions. We conducted 15 expeditions and 49 in situ lander deployments in an anoxic Scandinavian fjord between 2009 and 2021. Benthic fluxes were highest at the deepest anoxic site (average 516μmol CH4 m−2 d−1), supporting bottom water methane exceeding 5000nM. Natural and engineered mixing events displaced methane-rich bottom waters, enhancing upper water concentrations and driving high sea–air flux reaching 641μmol CH4 m−2 d−1. Mixing also reduced pelagic methane oxidation from 70% to 20% of all methane sources into the fjord. Upscaling of literature fluxes combined with our results suggests that fjords globally emit 1.0±0.8 Tg CH4 yr−1. Despite their small global area, fjords are hotspots of methane release. We suggest that ongoing deoxygenation and global change will enhance methane emissions from fjords.
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| 14. |
- Bonaglia, Stefano, et al.
(författare)
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Meiofauna improve oxygenation and accelerate sulfide removal in the seasonally hypoxic seabed
- 2020
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Ingår i: Marine Environmental Research. - : Elsevier BV. - 0141-1136 .- 1879-0291. ; 159
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Tidskriftsartikel (refereegranskat)abstract
- Oxygen depleted areas are widespread in the marine realm. Unlike macrofauna, meiofauna are abundant in hypoxic sediments. We studied to what extent meiofauna affect oxygen availability, sulfide removal and microbial communities. Meiofauna were extracted alive and added to intact sediments simulating abundance gradients previously reported in the area. A total of 324 porewater microprofiles were recorded over a 3-week incubation period and microbial community structure and cable bacteria densities were determined at the end of the experiment. At high abundances meiofauna activity deepened oxygen penetration by 85%, 59%, and 62% after 5, 14, and 22 days, respectively, compared to control sediment with scarce meiofauna. After 6 days, meiofauna increased the volume of oxidized, sulfide-free sediment by 68% and reduced sulfide fluxes from 8.8 to 0.4 mmol m(-2) d(-1). After 15 days, the difference with the control attenuated due to the presence of a cable bacteria population, which facilitated sulfides oxidation in all treatments. 16S rRNA gene analysis revealed that meiofauna affected microbial community structure (beta diversity). Thus, meiofauna bioturbation plays an important role in deepening oxygen penetration, counteracting euxinia and in structuring microbial diversity of hypoxic sediments. Co-existence with cable bacteria demonstrates neutralism interaction between these two ecosystem engineers.
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| 15. |
- Bonaglia, Stefano, et al.
(författare)
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Meiofauna increases bacterial denitrification in marine sediments
- 2014
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Ingår i: Nature Communications. - : Springer Science and Business Media LLC. - 2041-1723. ; 5, s. 5133-
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Tidskriftsartikel (refereegranskat)abstract
- Denitrification is a critical process that can alleviate the effects of excessive nitrogen availability in aquatic ecosystems subject to eutrophication. An important part of denitrification occurs in benthic systems where bioturbation by meiofauna (invertebrates <1mm) and its effect on element cycling are still not well understood. Here we study the quantitative impact of meiofauna populations of different abundance and diversity, in the presence and absence of macrofauna, on nitrate reduction, carbon mineralization and methane fluxes. In sediments with abundant and diverse meiofauna, denitrification is double that in sediments with low meiofauna, suggesting that meiofauna bioturbation has a stimulating effect on nitrifying and denitrifying bacteria. However, high meiofauna densities in the presence of bivalves do not stimulate denitrification, while dissimilatory nitrate reduction to ammonium rate and methane efflux are significantly enhanced. We demonstrate that the ecological interactions between meio-, macrofauna and bacteria are important in regulating nitrogen cycling in soft-sediment ecosystems.
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| 16. |
- Bonaglia, Stefano, 1983, et al.
(författare)
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Meiofauna Shaping Biogeochemical Processes
- 2023
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Ingår i: New Horizons in Meiobenthos Research. - Cham : Springer. - 9783031216220
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Bokkapitel (refereegranskat)abstract
- Biogeochemical processes at the sediment–water interface are essential for the functioning of marine ecosystems. It is a central question in benthic ecology how these processes are controlled and mediated by biotic factors. Particularly, the role of meiobenthos, the most abundant and diverse faunal component in these systems, is little understood and requires more attention. In this chapter, we discuss the impact of meiofauna bioturbation in marine sediments on significant mechanisms and processes in (a) carbon degradation and oxygen penetration, (b) sulfide dynamics, and (c) nitrogen cycling. Particularly in the growing hypoxic areas of the seafloor where meiofauna is often the only animal group present, the role and bioturbative activities of this central component of the benthos need further scrutiny regarding the decrease of oxygen and increase of toxic hydrogen sulfide. These knowledge gaps in the interaction between meiofauna and marine biogeochemistry are the background for our concluding outlines: We present current research frontiers in order to assess the role of meiofauna as regulators of geochemical processes and microbial activities. These goals require combination of quantitative and qualitative meiobenthos investigations with state-of-the-art experimental work.
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| 17. |
- Bonaglia, Stefano, et al.
(författare)
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Methane fluxes from coastal sediments are enhanced by macrofauna
- 2017
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Ingår i: Scientific Reports. - : Springer Science and Business Media LLC. - 2045-2322. ; 7:1
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Tidskriftsartikel (refereegranskat)abstract
- Methane and nitrous oxide are potent greenhouse gases (GHGs) that contribute to climate change. Coastal sediments are important GHG producers, but the contribution of macrofauna (benthic invertebrates larger than 1 mm) inhabiting them is currently unknown. Through a combination of trace gas, isotope, and molecular analyses, we studied the direct and indirect contribution of two macrofaunal groups, polychaetes and bivalves, to methane and nitrous oxide fluxes from coastal sediments. Our results indicate that macrofauna increases benthic methane efflux by a factor of up to eight, potentially accounting for an estimated 9.5% of total emissions from the Baltic Sea. Polychaetes indirectly enhance methane efflux through bioturbation, while bivalves have a direct effect on methane release. Bivalves host archaeal methanogenic symbionts carrying out preferentially hydrogenotrophic methanogenesis, as suggested by analysis of methane isotopes. Low temperatures (8 °C) also stimulate production of nitrous oxide, which is consumed by benthic denitrifying bacteria before it reaches the water column. We show that macrofauna contributes to GHG production and that the extent is dependent on lineage. Thus, macrofauna may play an important, but overlooked role in regulating GHG production and exchange in coastal sediment ecosystems.
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| 18. |
- Bonaglia, Stefano, et al.
(författare)
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Seasonal benthic nutrient cycling in a Baltic sea estuary
- 2012
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Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract
- Decades of urban, industrial, and agricultural discharge of nitrogen and phosphorus to the Baltic Sea have contributed to the spreading of water column hypoxia and annual widespread cyanobacteria blooms. Central to mitigating Baltic Sea eutrophication is to resolve how much reduction strategies of external N and P loading are offset by internal loading of the Baltic through nutrient recycling from the sediment. We investigated the seasonal variation of benthic nitrogen and phosphorus cycling in an estuary of the Baltic impacted by decades of sewage discharge. Sediment nutrient fluxes, denitrification, Anammox, DNRA, potential nitrification, and total and diffusive oxygen uptake (TOU/DOU) were quantified with 15N-tracer methods and microsensor profiling. Data indicate benthic net efflux of ammonium and phosphorus during the summer months, decreasing N2 loss with increasing organic matter content, and benthic N/P regeneration with a ratio of 3 to 7 compared to the sewage discharge N/P of ≈ 25, and a significant contribution (6 to 25%) of Anammox to N2 loss. On average benthic denitrification and Anammox may reduce the N load to the estuary by up to 54%.
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| 19. |
- Bonaglia, Stefano, 1983-, et al.
(författare)
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Seasonal oxygen, nitrogen and phosphorus benthic cycling along an impacted Baltic Sea estuary: regulation and spatial patterns
- 2014
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Ingår i: Biogeochemistry. - : Springer Science and Business Media LLC. - 0168-2563 .- 1573-515X. ; 119:1-3, s. 139-160
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Tidskriftsartikel (refereegranskat)abstract
- The regulatory roles of temperature, eutrophication and oxygen availability on benthic nitrogen (N) cycling and the stoichiometry of regenerated nitrogen and phosphorus (P) were explored along a Baltic Sea estuary affected by treated sewage discharge. Rates of sediment denitrification, anammox, dissimilatory nitrate reduction to ammonium (DNRA), nutrient exchange, oxygen (O2) uptake and penetration were measured seasonally. Sediments not affected by the nutrient plume released by the sewage treatment plant (STP) showed a strong seasonality in rates of O2 uptake and coupled nitrification-denitrification, with anammox never accounting for more than 20% of the total dinitrogen (N2) production. N cycling in sediments close to the STP was highly dependent on oxygen availability, which masked temperature-related effects. These sediments switched from low N loss and high ammonium (NH4+) efflux under hypoxic conditions in the fall, to a major N loss system in the winter when the sediment surface was oxidized. In the fall DNRA outcompeted denitrification as the main nitrate (NO3-) reduction pathway, resulting in N recycling and potential spreading of eutrophication. A comparison with historical records of nutrient discharge and denitrification indicated that the total N loss in the estuary has been tightly coupled to the total amount of nutrient discharge from the STP. Changes in dissolved inorganic nitrogen (DIN) released from the STP agreed well with variations in sedimentary N2 removal. This indicates that denitrification and anammox efficiently counterbalance N loading in the estuary across the range of historical and present-day anthropogenic nutrient discharge. Overall low N/P ratios of the regenerated nutrient fluxes impose strong N limitation for the pelagic system and generate a high potential for nuisance cyanobacterial blooms.
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| 20. |
- Bonaglia, Stefano, et al.
(författare)
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Sulfide oxidation in deep Baltic Sea sediments upon oxygenation and colonization by macrofauna
- 2019
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Ingår i: Marine Biology. - : Springer Science and Business Media LLC. - 0025-3162 .- 1432-1793. ; 166
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Tidskriftsartikel (refereegranskat)abstract
- Coastal and shelf sediments affected by transient or long-term bottom water anoxia and sulfidic conditions undergo drastic changes in macrofauna communities and abundances. This study investigates how early colonization by two macrofaunal functional traits (epifauna vs. infauna) affects oxygen, sulfide, and pH dynamics in anoxic sediment upon recent bottom water oxygenation. Large mesocosms (area 900 cm(2)) with 150-m-deep Baltic Sea soft sediments were exposed to three treatments: (1) no animals; (2) addition of 170 polychaetes (Marenzelleria arctia); (3) addition of 181 amphipods (Monoporeia affinis). Porewater chemistry was investigated repeatedly by microsensor profiling over a period of 65 days. Colonization by macrofauna did not significantly deepen penetration of oxygen compared to the animal-free sediment. Bioturbation by M. affinis increased the volume of the oxidized, sulfide-free sediment by 66% compared to the animal-free control already after 13 days of incubation. By the end of the experiment M. affinis and M. arctia increased the oxidized sediment volume by 87 and 35%, respectively. Higher efficiency of epifaunal amphipods in removing hydrogen sulfide than deep-burrowing polychaetes is likely due to more substantial re-oxidation of manganese and/or nitrogen compounds associated with amphipod mixing activity. Our results thus indicate that early colonization of different functional groups might have important implications for the later colonization by benthic macrofauna, meiofauna and microbial communities that benefit from oxidized and sulfide-free sediments.
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| 21. |
- Bonaglia, Stefano, et al.
(författare)
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The fate of fixed nitrogen in marine sediments with low organic loading : an in situ study
- 2017
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Ingår i: Biogeosciences. - : Copernicus GmbH. - 1726-4170 .- 1726-4189. ; 14:2, s. 285-300
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Tidskriftsartikel (refereegranskat)abstract
- Over the last decades, the impact of human activities on the global nitrogen (N) cycle has drastically increased. Consequently, benthic N cycling has mainly been studied in anthropogenically impacted estuaries and coasts, while in oligotrophic systems its understanding is still scarce. Here we report on benthic solute fluxes and on rates of denitrification, anammox, and dissimilatory nitrate reduction to ammonium (DNRA) studied by in situ incubations with benthic chamber landers during two cruises to the Gulf of Bothnia (GOB), a cold, oligotrophic basin located in the northern part of the Baltic Sea. Rates of N burial were also inferred to investigate the fate of fixed N in these sediments. Most of the total dissolved fixed nitrogen (TDN) diffusing to the water column was composed of organic N. Average rates of dinitrogen (N-2) production by denitrification and anammox (range: 53-360 mu mol Nm(-2) day(-1)) were comparable to those from Arctic and subarctic sediments worldwide (range: 34-344 mu mol Nm(-2) day(-1)). Anammox accounted for 18-26% of the total N2 production. Absence of free hydrogen sulfide and low concentrations of dissolved iron in sediment pore water suggested that denitrification and DNRA were driven by organic matter oxidation rather than chemolithotrophy. DNRA was as important as denitrification at a shallow, coastal station situated in the northern Bothnian Bay. At this pristine and fully oxygenated site, ammonium regeneration through DNRA contributed more than one-third to the TDN efflux and accounted, on average, for 45% of total nitrate reduction. At the offshore stations, the proportion of DNRA in relation to denitrification was lower (0-16% of total nitrate reduction). Median value and range of benthic DNRA rates from the GOB were comparable to those from the southern and central eutrophic Baltic Sea and other temperate estuaries and coasts in Europe. Therefore, our results contrast with the view that DNRA is negligible in cold and well-oxygenated sediments with low organic carbon loading. However, the mechanisms behind the variability in DNRA rates between our sites were not resolved. The GOB sediments were a major source (237 kt yr(-1), which corresponds to 184% of the external N load) of fixed N to the water column through recycling mechanisms. To our knowledge, our study is the first to document the simultaneous contribution of denitrification, DNRA, anammox, and TDN recycling combined with in situ measurements.
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| 22. |
- Bradshaw, Clare, et al.
(författare)
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Physical Disturbance by Bottom Trawling Suspends Particulate Matter and Alters Biogeochemical Processes on and Near the Seafloor
- 2021
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Ingår i: Frontiers in Marine Science. - : Frontiers Media SA. - 2296-7745. ; 8
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Tidskriftsartikel (refereegranskat)abstract
- 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.
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| 23. |
- Broman, Elias, et al.
(författare)
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Active DNRA and denitrification in oxic hypereutrophic waters
- 2021
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Ingår i: Water Research. - : Elsevier BV. - 0043-1354 .- 1879-2448. ; 194
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Tidskriftsartikel (refereegranskat)abstract
- Since the start of synthetic fertilizer production more than a hundred years ago, the coastal ocean has been exposed to increasing nutrient loading, which has led to eutrophication and extensive algal blooms. Such hypereutrophic waters might harbor anaerobic nitrogen (N) cycling processes due to low-oxygen mi- croniches associated with abundant organic particles, but studies on nitrate reduction in coastal pelagic environments are scarce. Here, we report on 15 N isotope-labeling experiments, metagenome, and RT-qPCR data from a large hypereutrophic lagoon indicating that dissimilatory nitrate reduction to ammonium (DNRA) and denitrification were active processes, even though the bulk water was fully oxygenated ( > 224 μM O 2 ). DNRA in the bottom water corresponded to 83% of whole-ecosystem DNRA (water + sedi- ment), while denitrification was predominant in the sediment. Microbial taxa important for DNRA accord- ing to the metagenomic data were dominated by Bacteroidetes (genus Parabacteroides ) and Proteobac- teria (genus Wolinella ), while denitrification was mainly associated with proteobacterial genera Pseu- domonas, Achromobacter , and Brucella . The metagenomic and microscopy data suggest that these anaero- bic processes were likely occurring in low-oxygen microniches related to extensive growth of filamentous cyanobacteria, including diazotrophic Dolichospermum and non-diazotrophic Planktothrix . By summing the total nitrate fluxes through DNRA and denitrification, it results that DNRA retains approximately one fifth (19%) of the fixed N that goes through the nitrate pool. This is noteworthy as DNRA represents thus a very important recycling mechanism for fixed N, which sustains algal proliferation and leads to further enhancement of eutrophication in these endangered ecosystems.
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| 24. |
- Broman, Elias, 1985-, et al.
(författare)
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Biotic interactions between benthic infauna and aerobic methanotrophs mediate methane fluxes from coastal sediments
- 2024
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Ingår i: The ISME journal. - 1751-7370 .- 1751-7362. ; 18:1
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Tidskriftsartikel (refereegranskat)abstract
- Coastal ecosystems dominate oceanic methane (CH4) emissions. However, there is limited knowledge about how biotic interactions between infauna and aerobic methanotrophs (i.e. CH4 oxidizing bacteria) drive the spatial-temporal dynamics of these emissions. Here, we investigated the role of meio- and macrofauna in mediating CH4 sediment-water fluxes and aerobic methanotrophic activity that can oxidize significant portions of CH4. We show that macrofauna increases CH4 fluxes by enhancing vertical solute transport through bioturbation, but this effect is somewhat offset by high meiofauna abundance. The increase in CH4 flux reduces CH4 pore-water availability, resulting in lower abundance and activity of aerobic methanotrophs, an effect that counterbalances the potential stimulation of these bacteria by higher oxygen flux to the sediment via bioturbation. These findings indicate that a larger than previously thought portion of CH4 emissions from coastal ecosystems is due to faunal activity and multiple complex interactions with methanotrophs.
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| 25. |
- Broman, Elias, 1985-, et al.
(författare)
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Cyanophage Diversity and Community Structure in Dead Zone Sediments
- 2021
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Ingår i: mSphere. - : American Society for Microbiology. - 2379-5042. ; 6:2
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Tidskriftsartikel (refereegranskat)abstract
- Up to 20% of prokaryotic organisms in the oceans are estimated to die every day due to viral infection and lysis. Viruses can therefore alter microbial diversity, community structure, and biogeochemical processes driven by these organisms. Cyanophages are viruses that infect and lyse cyanobacterial cells, adding bioavailable carbon and nutrients into the environment. Cyanobacteria are photosynthesizing bacteria, with some species capable of N-2 fixation, which are known to form large blooms as well as resistant resting cells known as akinetes. Here, we investigated cyanophage diversity and community structure plus cyanobacteria in dead zone sediments. We sampled surface sediments and sequenced DNA and RNA, along an oxygen gradient-representing oxic, hypoxic, and anoxic conditions-in one of the world's largest dead zones located in the Baltic Sea. Cyanophages were detected at all stations and, based on partial genome contigs, had a higher alpha diversity and different beta diversity in the hypoxic-anoxic sediments, suggesting that cyanobacteria in dead zone sediments and/or environmental conditions select for specific cyanophages. Some of these cyanophages can infect cyanobacteria with potential consequences for gene expression related to their photosystem and phosphate regulation. Top cyanobacterial genera detected in the anoxic sediment included Dolichospermum/Anabaena, Synechococcus, and Cyanobium. RNA transcripts classified to cyanobacteria were associated with numerous pathways, including anaerobic carbon metabolism and N-2 fixation. Cyanobacterial blooms are known to fuel oxygen-depleted ecosystems with phosphorus (so-called internal loading), and our cyanophage data indicate the potential for viral lysis of cyanobacteria which might explain the high nutrient turnover in these environments. IMPORTANCE Cyanophages are viruses that target cyanobacteria and directly control their abundance via viral lysis. Cyanobacteria are known to cause large blooms in water bodies, substantially contributing to oxygen depletion in bottom waters resulting in areas called dead zones. Our knowledge of cyanophages in dead zones is very scarce, and so far, no studies have assembled partial cyanophage genomes and investigated their associated cyanobacteria in these dark and anoxic sediments. Here, we present the first study using DNA and RNA sequencing to investigate in situ diversity of cyanophages and cyanobacteria in dead zones. Our study shows that dead zone sediments contain different cyanophages compared to oxic sediments and suggest that these viruses are able to affect cyanobacterial photosystem and phosphate regulation. Furthermore, cyanophage-controlled lysis of cyanobacteria might also increase the turnover of carbon, phosphorus, and nitrogen in these oxygen-free environments at the bottom of the sea.
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