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Marine sediments bury ∼ 160 Tg organic carbon (OC) yr−1 globally, with ∼ 90 % of the burial occurring in continental margin sediments. It is generally believed that OC is buried more efficiently in sediments underlying anoxic bottom waters. However, recent studies revealed that sediments in the central Baltic Sea exhibit very high OC mineralization rates and consequently low OC burial efficiencies (∼ 5 %–10 %), despite being overlaid by long-term anoxic bottom waters. Here, we investigate factors contributing to this unexpectedly high OC mineralization rates in the Western Gotland Basin (WGB), a sub-basin of the central Baltic Sea. We sampled five sites along a transect in the WGB, including two where organic carbon-iron (OC–Fe) associations were quantified. Sulphate reduction rate measurements indicated that OC reactivity (k) was much higher than expected for anoxic sediments. High OC loadings (i.e., OC concentrations normalized to sediment specific surface area) and low OC–Fe associations showed that physical protection of OC is limited. Overall, these results suggest that the WGB sediments receive large amounts of bulk OC relative to the supply of mineral particles, far exceeding the potential for OC physical protection. As a result, a large fraction of OC is free from associations with mineral surfaces, thus the OC reactivity is high, despite anoxic bottom waters. Overall, our results demonstrate that anoxia does not always lead to lower OC mineralization rates and increased burial efficiencies in sediments.

Location, specific topography, and hydrographic setting together with climate change and strong anthropogenic pressure are the main factors shaping the biogeochemical functioning and thus also the ecological status of the Baltic Sea. The recent decades have brought significant changes in the Baltic Sea. First, the rising nutrient loads from land in the second half of the 20th century led to eutrophication and spreading of hypoxic and anoxic areas, for which permanent stratification of the water column and limited ventilation of deep-water layers made favourable conditions. Since the 1980s the nutrient loads to the Baltic Sea have been continuously decreasing. This, however, has so far not resulted in significant improvements in oxygen availability in the deep regions, which has revealed a slow response time of the system to the reduction of the land-derived nutrient loads. Responsible for that is the low burial efficiency of phosphorus at anoxic conditions and its remobilization from sediments when conditions change from oxic to anoxic. This results in a stoichiometric excess of phosphorus available for organic-matter production, which promotes the growth of N2-fixing cyanobacteria and in turn supports eutrophication. This assessment reviews the available and published knowledge on the biogeochemical functioning of the Baltic Sea. In its content, the paper covers the aspects related to changes in carbon, nitrogen, and phosphorus (C, N, and P) external loads, their transformations in the coastal zone, changes in organic-matter production (eutrophication) and remineralization (oxygen availability), and the role of sediments in burial and turnover of C, N, and P. In addition to that, this paper focuses also on changes in the marine CO2 system, the structure and functioning of the microbial community, and the role of contaminants for biogeochemical processes. This comprehensive assessment allowed also for identifying knowledge gaps and future research needs in the field of marine biogeochemistry in the Baltic Sea.

The external phosphorus (P) loading has been halved, but the P content in the water column and the area of anoxic bottoms in Baltic proper has increased during the last 30 years. This can be explained by a temporary internal source of dissolved inorganic phosphorus (DIP) that is turned on when the water above the bottom sediment becomes anoxic. A load-response model, explaining the evolution from 1980 to 2005, suggests that the average specific DIP flux from anoxic bottoms in the Baltic proper is about 2.3 g P m⁻² year⁻¹. This is commensurable with fluxes estimated in situ from anoxic bottoms in the open Baltic proper and from hydrographic data in the deep part of Bornholm Basin. Oxygenation of anoxic bottoms, natural or manmade, may quickly turn off the internal P source from anoxic bottoms. This new P-paradigm should have far-reaching implications for abatement of eutrophication in the Baltic proper.

Hypoxia hampers eutrophication reduction efforts by enabling high nutrient fluxes from sediment to bottom waters. Oxygenation of hypoxic water bodies is often proposed to reduce benthic ammonium and phosphate release. This study investigates the functional response of benthic nitrate-reducing processes to a long-term engineered oxygenation effort in a density-stratified fjord with euxinic bottom waters. Oxygenation was achieved by mixing surface water with deep, euxinic water, which increased oxygen and nitrate concentrations in the deep water column. The presence of nitrate instigated benthic nitrate reduction in the newly oxidized sediments by equally stimulating denitrification and dissimilatory nitrate reduction to ammonium (DNRA). DNRA and total nitrate reduction rates, as well as the contribution of DNRA to total nitrate reduction, decreased with increasing exposure time of the sediments to oxygen. The relative importance of DNRA as a nitrate sink was correlated to nitrate concentrations, with more nitrate being reduced to ammonium at higher bottom water nitrate concentrations. Overall, engineered oxygenation decreased the net efflux of dissolved inorganic nitrogen from the sediments by stimulating net nitrate removal through denitrification.

Coastal sediments are rich in conductive particles, possibly affecting microbial processes for which acetate is a central intermediate. In the methanogenic zone, acetate is consumed by methanogens and/or syntrophic acetate-oxidizing (SAO) consortia. SAO consortia live under extreme thermodynamic pressure, and their survival depends on successful partnership. Here, we demonstrate that conductive particles enable the partnership between SAO bacteria (i.e., Geobacter spp.) and methanogens ( Methanosarcina spp.) from the coastal sediments of the Bothnian Bay of the Baltic Sea. Baltic methanogenic sediments were rich in conductive minerals, had an apparent isotopic fractionation characteristic of CO 2 -reductive methanogenesis, and were inhabited by Geobacter and Methanosarcina . As long as conductive particles were delivered, Geobacter and Methanosarcina persisted, whereas exclusion of conductive particles led to the extinction of Geobacter . Baltic Geobacter did not establish a direct electric contact with Methanosarcina , necessitating conductive particles as electrical conduits. Within SAO consortia, Geobacter was an efficient [ 13 C]acetate utilizer, accounting for 82% of the assimilation and 27% of the breakdown of acetate. Geobacter benefits from the association with the methanogen, because in the absence of an electron acceptor it can use Methanosarcina as a terminal electron sink. Consequently, inhibition of methanogenesis constrained the SAO activity of Geobacter as well. A potential benefit for Methanosarcina partnering with Geobacter is that together they competitively exclude acetoclastic methanogens like Methanothrix from an environment rich in conductive particles. Conductive particle-mediated SAO could explain the abundance of acetate oxidizers like Geobacter in the methanogenic zone of sediments where no electron acceptors other than CO 2 are available. IMPORTANCE Acetate-oxidizing bacteria are known to thrive in mutualistic consortia in which H 2 or formate is shuttled to a methane-producing Archaea partner. Here, we discovered that such bacteria could instead transfer electrons via conductive minerals. Mineral SAO (syntrophic acetate oxidation) could be a vital pathway for CO 2 -reductive methanogenesis in the environment, especially in sediments rich in conductive minerals. Mineral-facilitated SAO is therefore of potential importance for both iron and methane cycles in sediments and soils. Additionally, our observations imply that agricultural runoff or amendments with conductive chars could trigger a significant increase in methane emissions. Acetate-oxidizing bacteria are known to thrive in mutualistic consortia in which H 2 or formate is shuttled to a methane-producing Archaea partner. Here, we discovered that such bacteria could instead transfer electrons via conductive minerals. Mineral SAO (syntrophic acetate oxidation) could be a vital pathway for CO 2 -reductive methanogenesis in the environment, especially in sediments rich in conductive minerals. Mineral-facilitated SAO is therefore of potential importance for both iron and methane cycles in sediments and soils. Additionally, our observations imply that agricultural runoff or amendments with conductive chars could trigger a significant increase in methane emissions.

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 cm2) 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.

Phosphorus fertilisation (eutrophication) is expanding oxygen depletion in coastal systems worldwide. Under low-oxygen bottom water conditions, phosphorus release from the sediment is elevated, which further stimulates primary production. It is commonly assumed that re-oxygenation could break this “vicious cycle” by increasing the sedimentary phosphorus retention. Recently, a deep-water inflow into the Baltic Sea created a natural in situ experiment that allowed us to investigate if temporary re-oxygenation stimulates sedimentary retention of dissolved inorganic phosphorus (DIP). Surprisingly, during this 3-year study, we observed a transient but considerable increase, rather than a decrease, in the sediment efflux of DIP and other dissolved biogenic compounds. This suggested that the oxygenated inflow elevated the organic matter degradation in the sediment, likely due to an increase in organic matter supply to the deeper basins, potentially combined with a transient stimulation of the mineralisation efficiency. As a result, the net sedimentary DIP release per m2 was 56 %–112 % higher over the years following the re-oxygenation than before. In contrast to previous assumptions, our results show that inflows of oxygenated water to anoxic bottom waters can increase the sedimentary phosphorus release.

BackgroundDespite a clear link between aging and cancer, there has been inconclusive evidence on how biological age (BA) may be associated with cancer incidence.MethodsWe studied 308,156 UK Biobank participants with no history of cancer at enrolment. Using 18 age-associated clinical biomarkers, we computed three BA measures (Klemera-Doubal method [KDM], PhenoAge, homeostatic dysregulation [HD]) and assessed their associations with incidence of any cancer and five common cancers (breast, prostate, lung, colorectal, and melanoma) using Cox proportional-hazards models.ResultsA total of 35,426 incident cancers were documented during a median follow-up of 10.9 years. Adjusting for common cancer risk factors, 1-standard deviation (SD) increment in the age-adjusted KDM (hazard ratio = 1.04, 95% confidence interval = 1.03–1.05), age-adjusted PhenoAge (1.09, 1.07–1.10), and HD (1.02, 1.01–1.03) was significantly associated with a higher risk of any cancer. All BA measures were also associated with increased risks of lung and colorectal cancers, but only PhenoAge was associated with breast cancer risk. Furthermore, we observed an inverse association between BA measures and prostate cancer, although it was attenuated after removing glycated hemoglobin and serum glucose from the BA algorithms.ConclusionsAdvanced BA quantified by clinical biomarkers is associated with increased risks of any cancer, lung cancer, and colorectal cancer.

Sedimentary recycling of phosphorus is a key aspect of eutrophication. Here, we present data on benthic fluxes of dissolved inorganic phosphorus (DIP) from the Baltic Sea, an area with a long eutrophication history. The presented dataset contains 498 individual fluxes measured in situ with three types of benthic chamber landers at 59 stations over 20 years, and data cover most of the Baltic Sea subbasins (Hylén et al., 2025, https://doi.org/10.5281/zenodo.14812160). The dataset further contains information about bottom-water dissolved oxygen (O2) concentrations, sedimentary organic carbon (OC) content and sediment type. The DIP fluxes differ considerably between basins depending on OC loading and the level of O2 depletion and generally increase from the coast to the central basins. Several stations have been visited on multiple occasions, also at times with different O2 concentrations, which enables investigation of the immediate effects of shifting bottom-water O2 concentrations on the benthic DIP release. The Baltic Sea-wide benthic DIP release is estimated to be 389–484 kton yr−1 based on a data integration based on sediment type and O2 conditions during three years with varying extents of hypoxia and anoxia (2004, 2013 and 2018). The dataset reveals a lack of flux measurements in winter months, coastal areas, and sandy and coarse sediments; these should be targeted in future studies. Overall, intercomparisons between samplings and landers as well as rigorous data evaluation show that the data are of high quality. As such, this data set will, alone and together with other environmental data, be important for marine management and studies on mechanisms in benthic phosphorus cycling.

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 (N2) production by denitrification and anammox (range: 53–360 µmol N m−2 day−1) were comparable to those from Arctic and subarctic sediments worldwide (range: 34–344 µmol N m−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.