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The diversity and dynamics of ammonia-oxidizing bacteria (AOB) and archaea (AOA) nitrifying communities in the sediments of the eutrophic Gulf of Finland (GoF) were investigated. Using clone libraries of ammonia monooxygenase ( amoA ) gene fragments and terminal restriction fragment length polymorphism (TRFLP), we found a low richness of both AOB and AOA. The AOB amoA phylogeny matched that of AOB 16S ribosomal genes from the same samples. AOA communities were characterized by strong spatial variation while AOB communities showed notable temporal patterns. At open sea sites, where transient anoxic conditions prevail, richness of both AOA and AOB was lowest and communities were dominated by organisms with gene signatures unique to the GoF. Given the importance of nitrification as a link between the fixation of nitrogen and its removal from aquatic environments, the low diversity of ammonia-oxidizing microbes across the GoF could be of relevance for ecosystem resilience in the face of rapid global environmental changes. [ABSTRACT FROM AUTHOR]

Chemolithoautotrophic denitrification is an important mechanism of nitrogen loss in the water column of euxinic basins, but its isotope fractionation factor is not known. Sulfurimonas gotlandica GD1T, a recently isolated bacterial key player in Baltic Sea pelagic redoxcline processes, was used to determine the isotope fractionation of nitrogen and oxygen in nitrate during denitrification. Under anoxic conditions, nitrate reduction was accompanied by nitrogen and oxygen isotope fractionation of 23.8 ± 2.5° and 11.7 ± 1.1°, respectively. The isotope effect for nitrogen was in the range determined for heterotrophic denitrification, with only the absence of stirring resulting in a significant decrease of the fractionation factor. The relative increase in δ18ONO3 to δ15NNO3 did not follow the 1:1 relationship characteristic of heterotrophic, marine denitrification. Instead, δ18ONO3 increased slower than δ15NNO3, with a conserved ratio of 0.5:1. This result suggests that the periplasmic nitrate reductase (Nap) of S. gotlandica strain GD1T fractionates the N and O in nitrate differently than the membrane-bound nitrate reductase (Nar), which is generally prevalent among heterotrophic denitrifiers and is considered as the dominant driver for the observed isotope fractionation. Hence in the Baltic Sea redoxcline, other, as yet-unidentified factors likely explain the low apparent fractionation. [ABSTRACT FROM AUTHOR]

Hypoxia has occurred intermittently over the Holocene in the Baltic Sea, but the recent expansion from less than 10 000 km before 1950 to >60 000 km since 2000 is mainly caused by enhanced nutrient inputs from land and atmosphere. With worsening hypoxia, the role of sediments changes from nitrogen removal to nitrogen release as ammonium. At present, denitrification in the water column and sediments is equally important. Phosphorus is currently buried in sediments mainly in organic form, with an additional contribution of reduced Fe-phosphate minerals in the deep anoxic basins. Upon the transition to oxic conditions, a significant proportion of the organic phosphorus will be remineralized, with the phosphorus then being bound to iron oxides. This iron-oxide bound phosphorus is readily released to the water column upon the onset of hypoxia again. Important ecosystems services carried out by the benthic fauna, including biogeochemical feedback-loops and biomass production, are also lost with hypoxia. The results provide quantitative knowledge of nutrient release and recycling processes under various environmental conditions in support of decision support tools underlying the Baltic Sea Action Plan. [ABSTRACT FROM AUTHOR]

Primary production in the eutrophic Baltic Sea is limited by nitrogen availability; hence denitrification (natural transformation of nitrate to gaseous N) in the sediments is crucial in mitigating the effects of eutrophication. This study shows that dissimilatory nitrate reduction to ammonium (DNRA) process, where nitrogen is not removed but instead recycled in the system, dominates nitrate reduction in low oxygen conditions (O <110 μM), which have been persistent in the central Gulf of Finland during the past decade. The nitrogen removal rates measured in this study show that nitrogen removal has decreased in the Gulf of Finland compared to rates measured in mid-1990s and the decrease is most likely caused by the increased bottom water hypoxia. [ABSTRACT FROM AUTHOR]

Benthic processes were measured at a coastal deposition area in the northern Baltic Sea, covering all seasons. The N2 production rates, 90–400 μmol N m−2 d−1, were highest in autumn-early winter and lowest in spring. Heterotrophic bacterial production peaked unexpectedly late in the year, indicating that in addition to the temperature, the availability of carbon compounds suitable for the heterotrophic bacteria also plays a major role in regulating the denitrification rate. Anaerobic ammonium oxidation (anammox) was measured in spring and autumn and contributed 10% and 15%, respectively, to the total N2 production. The low percentage did, however, result in a significant error in the total N2 production rate estimate, calculated using the isotope pairing technique. Anammox must be taken into account in the Gulf of Finland in future sediment nitrogen cycling research. [ABSTRACT FROM AUTHOR]

Estuaries have been suggested to have an important role in reducing the nitrogen load transported to the sea. We measured denitrification rates in six estuaries of the northern Baltic Sea. Four of them were river mouths in the Bothnian Bay (northern Gulf of Bothnia), and two were estuary bays, one in the Archipelago Sea (southern Gulf of Bothnia) and the other in the Gulf of Finland. Denitrification rates in the four river mouths varied between 330 and 905 µmol N m-2 d-1. The estuary bays at the Archipelago Sea and the Gulf of Bothnia had denitrification rates from 90 µmol N m-2 d-1 to 910 µmol N m-2 d-1 and from 230 µmol N m-2 d-1 to 320 µmol N m-2 d-1, respectively. Denitrification removed 3.6-9.0% of the total nitrogen loading in the river mouths and in the estuary bay in the Gulf of Finland, where the residence times were short. In the estuary bay with a long residence time, in the Archipelago Sea, up to 4.5% of nitrate loading and 19% of nitrogen loading were removed before entering the sea. According to our results, the sediments of the fast-flowing rivers and the estuary areas with short residence times have a limited capacity to reduce the nitrogen load to the Baltic Sea. [ABSTRACT FROM AUTHOR]

The community structure of the bacteria associated with Nodularia spumigena (Mertens) cyanobacterial aggregates in the Baltic Sea was studied with temperature gradient gel electrophoresis (TGGE), using a 16S rRNA gene fragment as a target. Various developmental stages of the aggregates and free-floating cyanobacterial filaments were sampled to reveal possible changes in associated microbial community structure during development and senescence of the aggregates. The microbial community structures of all samples differed, and the communities of young and decaying aggregates were separated by cluster analysis of the TGGE fingerprint data. Sequencing of the TGGE fragments indicated the presence of bacteria from the α-, β-, and γ-proteobacterial groups, as well as members of Cytophaga–Flexibacter–Bacteroides lineages and gram-positive Actinobacteria spp. The majority of the Nodularia-associated sequences were not closely related to previously reported 16S rDNA sequences from the Baltic Sea or any other environment. The structure of the bacterial assemblage reflects the environmental changes associated with the succession and decay of the cyanobacterial aggregates. In addition, the sequence data suggest that the N. spumigena (Mertens) blooms in the Baltic Sea may host thus far uncharacterized bacterial species. [ABSTRACT FROM AUTHOR]

Steep redoxclines form between oxic surface water and the stagnant, sulfidic hypolimnion in the eutrophied, brackish water Baltic Sea. Nitrification, denitrification, and anammox were measured at and below the redoxcline to quantify the role of water-column nitrogen processes in the overall magnitude of nitrogen removal in the Baltic Sea. Rates of nitrification were very high (up to 85 nmol N L-1 d-1) at the oxic-anoxic interface, but, surprisingly, nitrification was separated from the processes reducing nitrate to N2 (up to 810 nmol N L-1 d-1) by tens of meters in depth. N2 production was dominated by chemolithotrophic denitrification, with anammox playing only a negligible role, and limited to the water layers in which nitrite or nitrate coexisted with sulfide. The alternating oxygen concentrations in the basin induce irregular bursts of nitrogen removal. Nitrification takes place after mixing of ammonium-rich deep water with oxic water, and denitrification uses the formed nitrite and nitrate once anoxic conditions re-establish. Although removal rates can be high, conditions allowing such rates are likely short-lived. While the sedimentary denitrification rates at the shallower, oxic areas are lower, they are more constant in time, highlighting the need to avoid hypoxia, which would prevent sedimentary denitrification. [ABSTRACT FROM AUTHOR]

The redoxclines that form between the oxic and anoxic water layers in the central Baltic Sea are sites of intensive nitrogen cycling. To gain better understanding of nitrification, we measured the biogeochemical properties along with potential nitrification rates and analyzed the assemblages of ammonia-oxidizing bacteria and archaea using functional gene microarrays. To estimate nitrification in the entire water column, we constructed a regression model for the nitrification rates and applied it to the conditions prevailing in the area in 2008–2012. The highest ammonia oxidation rates were found in a thin layer at the top of the redoxcline and the rates quickly decreased below detection limit when oxygen was exhausted. This is probably because extensive suboxic layers, which are known to harbor pelagic nitrification, are formed only for short periods after inflows in the Baltic Sea. The nitrification rates were some of the highest measured in the water columns, but the thickness of the layer where conditions were favorable for nitrification, was very small and it remained fairly stable between years. However, the depth of the nitrification layer varied substantially between years, particularly in the eastern Gotland Basin (EGB) due to turbulence in the water column. The ammonia oxidizer communities clustered differently between the eastern and western Gotland Basin (WGB) and the composition of ammonia-oxidizing assemblages correlated with the environmental variables. The ammonia oxidizer community composition was more even in the EGB, which may be related to physical instability of the redoxcline that does not allow predominance of a single archetype, whereas in the WGB, where the position of the redoxcline is more constant, the ammonia-oxidizing community was less even. Overall the ammonia-oxidizing communities in the Baltic Sea redoxclines were very evenly distributed compared to other marine environments where microarrays have been applied previously. [ABSTRACT FROM AUTHOR]

Hypoxia, a growing worldwide problem, has been intermittently present in the modern Baltic Sea since its formation ca. 8000 cal. yr BP. However, both the spatial extent and intensity of hypoxia have increased with anthropogenic eutrophication due to nutrient inputs. Physical processes, which control stratification and the renewal of oxygen in bottom waters, are important constraints on the formation and maintenance of hypoxia. Climate controlled inflows of saline water from the North Sea through the Danish Straits is a critical controlling factor governing the spatial extent and duration of hypoxia. Hypoxia regulates the biogeochemical cycles of both phosphorus (P) and nitrogen (N) in the water column and sediments. Significant amounts of P are currently released from sediments, an order of magnitude larger than anthropogenic inputs. The Baltic Sea is unique for coastal marine ecosystems experiencing N losses in hypoxic waters below the halocline. Although benthic communities in the Baltic Sea are naturally constrained by salinity gradients, hypoxia has resulted in habitat loss over vast areas and the elimination of benthic fauna, and has severely disrupted benthic food webs. Nutrient load reductions are needed to reduce the extent, severity, and effects of hypoxia. [ABSTRACT FROM AUTHOR]

Abstract: The ecosystem of the Gulf of Finland is currently dominated by internal phosphorus loading from sediments. The internal load is highly redox sensitive, and its successful modelling on basin-wide scale requires a simplified description of the sediment process. We present here an approach in which redox-sensitive sediment processes are directly linked to the decomposition of carbon instead of the oxygen concentration in near-bottom water. Mineralisation of organic carbon is known to be the major factor controlling sediment nutrient cycling, including denitrification and Fe(III) oxide reduction, giving rise to high phosphorus fluxes from anoxic sediments. Our sediment process description requires only four main parameters, which are here identified by using in situ CO2, dissolved inorganic nitrogen (DIN) and dissolved inorganic phosphorus (DIP) flux measurements carried out by Göteborg University landers. The model was tested with the aid of time series of denitrification and DIP flux rates measured in the western Gulf of Finland. Modelled near-bottom and surface nutrient concentrations were compared with monitoring data from both the eastern and western Gulf of Finland. The model simulations showed that the average net ecosystem production entering the sediment surface from the euphotic layer was 49g C m−2 a−1. This organic load induced an average denitrification rate of 2.5 g N m−2 a−1 and DIP flux of 0.67 g P m−2 a−1, corresponding to 20,200 t P a−1 for the whole Gulf of Finland. The model was able to describe the seasonality of denitrification and sediment DIP flux with high precision. Further, the modelled near-bottom and surface nutrient concentrations were compatible with the available data. The results indicate that, on the scales important for coastal and open sea conditions, our simple sediment process description works well. The new tool will help us to use 3D models to study the effects of external load on the production and decomposition of organic matter, and on subsequent benthic nutrient fluxes. [Copyright &y& Elsevier]