Nitrogen fixation estimates for the Baltic Sea indicate high rates for the previously overlooked Bothnian Sea

Dense blooms of diazotrophic filamentous cyanobacteria are formed every summer in the Baltic Sea. We estimated their contribution to nitrogen fixation by combining two decades of cyanobacterial biovolume monitoring data with recently measured genera-specific nitrogen fixation rates. In the Bothnian Sea, estimated nitrogen fixation rates were 80 kt N year⁻¹, which has doubled during recent decades and now exceeds external loading from rivers and atmospheric deposition of 69 kt year⁻¹. The estimated contribution to the Baltic Proper was 399 kt N year⁻¹, which agrees well with previous estimates using other approaches and is greater than the external input of 374 kt N year⁻¹. Our approach can potentially be applied to continuously estimate nitrogen loads via nitrogen fixation. Those estimates are crucial for ecosystem adaptive management since internal nitrogen loading may counteract the positive effects of decreased external nutrient loading.     

Nitrogen fixation by cyanobacteria stimulates production in Baltic food webs

Filamentous, nitrogen-fixing cyanobacteria form extensive summer blooms in the Baltic Sea. Their ability to fix dissolved N₂ allows cyanobacteria to circumvent the general summer nitrogen limitation, while also generating a supply of novel bioavailable nitrogen for the food web. However, the fate of the nitrogen fixed by cyanobacteria remains unresolved, as does its importance for secondary production in the Baltic Sea. Here, we synthesize recent experimental and field studies providing strong empirical evidence that cyanobacterial nitrogen is efficiently assimilated and transferred in Baltic food webs via two major pathways: directly by grazing on fresh or decaying cyanobacteria and indirectly through the uptake by other phytoplankton and microbes of bioavailable nitrogen exuded from cyanobacterial cells. This information is an essential step toward guiding nutrient management to minimize noxious blooms without overly reducing secondary production, and ultimately most probably fish production in the Baltic Sea.     

Modeling Nutrient Transports and Exchanges of Nutrients Between Shallow Regions and the Open Baltic Sea in Present and Future Climate

We quantified horizontal transport patterns and the net exchange of nutrients between shallow regions and the open sea in the Baltic proper. A coupled biogeochemical-physical circulation model was used for transient simulations 1961-2100. The model was driven by regional downscaling of the IPCC climate change scenario A1B from two global General Circulation Models in combination with two nutrient load scenarios. Modeled nutrient transports followed mainly the large-scale internal water circulation and showed only small circulation changes in the future projections. The internal nutrient cycling and exchanges between shallow and deeper waters became intensified, and the internal removal of phosphorus became weaker in the warmer future climate. These effects counteracted the impact from nutrient load reductions according to the Baltic Sea Action Plan. The net effect of climate change and nutrient reductions was an increased net import of dissolved inorganic phosphorus to shallow areas in the Baltic proper.     

巢湖蓝藻的机械清除工艺以及藻水分离实验研究简

对巢湖蓝藻进行机械清除以及藻水分离实验研究,实验采用的浮式围栏引导-机械清除-投加剥离液辅助机械清除工艺处理量大,除藻效率高,筛网过滤-浓缩-卧螺离心机脱水成藻泥的藻水分离工艺较为理想,藻泥含水率仅为89%。2011年5—10月在巢湖运用上述方法清除湖面水华蓝藻,共处理富藻水1.6万m³,得到藻泥970 t,累计清除蓝藻106.7 t（干重）。按照所清除蓝藻的总氮、总磷的平均含量计算,相当于从湖中移除了氮6.25 t,磷2.1 t。表明在富营养湖泊中水华蓝藻大量暴发时,采用上述方法除藻,对控制蓝藻水华污染,有效降低内源氮、磷等污染物负荷具有十分重要的作用。     

巢湖蓝藻的机械清除工艺以及藻水分离实验研究

对巢湖蓝藻进行机械清除以及藻水分离实验研究,实验采用的浮式围栏引导一机械清除．投加剥离液辅助机械清除工艺处理量大,除藻效率高,筛网过滤-浓缩-卧螺离心机脱水成藻泥的藻水分离工艺较为理想,藻泥含水率仅为89％。2011年5-10月在巢湖运用上述方法清除湖面水华蓝藻,共处理富藻水1．6万m3,得到藻泥970t,累计清除蓝藻106．7t（干重）。按照所清除蓝藻的总氮、总磷的平均含量计算,相当于从湖中移除了氮6．25t,磷2．1t。表明在富营养湖泊中水华蓝藻大量暴发时,采用上述方法除藻,对控制蓝藻水华污染,有效降低内源氮、磷等污染物负荷具有十分重要的作用。     

Extremes of Temperature, Oxygen and Blooms in the Baltic Sea in a Changing Climate

In the future, the Baltic Sea ecosystem will be impacted both by climate change and by riverine and atmospheric nutrient inputs. Multi-model ensemble simulations comprising one IPCC scenario (A1B), two global climate models, two regional climate models, and three Baltic Sea ecosystem models were performed to elucidate the combined effect of climate change and changes in nutrient inputs. This study focuses on the occurrence of extreme events in the projected future climate. Results suggest that the number of days favoring cyanobacteria blooms could increase, anoxic events may become more frequent and last longer, and salinity may tend to decrease. Nutrient load reductions following the Baltic Sea Action Plan can reduce the deterioration of oxygen conditions.     

Modeling riverine nutrient transport to the Baltic Sea: a large-scale approach

We developed for the first time a catchment model simulating simultaneously the nutrient land-sea fluxes from all 105 major watersheds within the Baltic Sea drainage area. A consistent modeling approach to all these major watersheds, i.e., a consistent handling of water fluxes (hydrological simulations) and loading functions (emission data), will facilitate a comparison of riverine nutrient transport between Baltic Sea subbasins that differ substantially. Hot spots of riverine emissions, such as from the rivers Vistula, Oder, and Daugava or from the Danish coast, can be easily demonstrated and the comparison between these hot spots, and the relatively unperturbed rivers in the northern catchments show decisionmakers where remedial actions are most effective to improve the environmental state of the Baltic Sea, and, secondly, what percentage reduction of riverine nutrient loads is possible. The relative difference between measured and simulated fluxes during the validation period was generally small. The cumulative deviation (i.e., relative bias) [Sigma(Simulated - Measured)/Sigma Measured x 100 (%)] from monitored water and nutrient fluxes amounted to +8.2% for runoff, to -2.4% for dissolved inorganic nitrogen, to +5.1% for total nitrogen, to +13% for dissolved inorganic phosphorus and to +19% for total phosphorus. Moreover, the model suggests that point sources for total phosphorus compiled by existing pollution load compilations are underestimated because of inconsistencies in calculating effluent loads from municipalities.     

Management options and effects on a marine ecosystem: assessing the future of the Baltic

We are using the coupled models in a decision support system, Nest, to evaluate the response of the marine ecosystem to changes in external loads through various management options. The models address all the seven major marine basins and the entire drainage basin of the Baltic Sea. A series of future scenarios have been developed, in close collaboration with the Helsinki Commission, to see the possible effects of improved wastewater treatment and manure handling, phosphorus-free detergents, and less intensive land use and live stocks. Improved wastewater treatment and the use of phosphorus-free detergents in the entire region would drastically decrease phosphorus loads and improve the marine environment, particularly the occurrence of cyanobacterial blooms. However, the Baltic Sea will remain eutrophic, and to reduce other effects, a substantial reduction of nitrogen emissions must be implemented. This can only be obtained in these scenarios by drastically changing land use. In a final scenario, we have turned 50% of all agricultural lands into grasslands, together with efficient wastewater treatments and a ban of phosphorus in detergents. This scenario will substantially reduce primary production and the extension of hypoxic bottoms, increase water transparency in the most eutrophied basins, and virtually eliminate extensive cyanobacterial blooms.     

The impact of benthic macrofauna for nutrient fluxes from Baltic Sea sediments

This article focuses on the ecological role of benthic macrofauna on nutrient dynamics and benthic-pelagic coupling in the Baltic Sea with relation to eutrophication. Generally, benthic macrofaunal activities have large effects on sediment biogeochemistry and often with stimulatory effects on processes that counteract eutrophication, i.e., denitrification and increased phosphorus retention of the sediment. The degree of faunal impact on such processes varies depending on faunal density and functional group composition. The effect of macrofaunal activities on sediment nutrient dynamics can also result in a higher nitrogen: phosporus ratio of the sediments efflux compared with sediments without macrofauna. Increased internal nutrient loading during eutrophication-induced anoxia is suggested to be caused both by altered sediment biogeochemical processes and through reduced or lost bioturbating macrofauna and thereby a reduced stimulatory effect from their activities on natural purification processes of the Baltic Sea ecosystem.     

A New Phosphorus Paradigm for the Baltic Proper

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.

Electronic supplementary material

The online version of this article (doi:10.1007/s13280-013-0441-3) contains supplementary material, which is available to authorized users.     

Modeling the Baltic Sea eutrophication in a decision support system

SANBALTS (Simple As Necessary Baltic Long-Term Large-Scale) is a model of the coupled nitrogen and phosphorus cycles. This model has been developed as an integral part of the decision support system Marine Research on Eutrophication's Nest with the overall aim to evaluate management options for reducing Baltic Sea eutrophication. Simulated nutrient and oxygen concentrations as well as transport flows and major biogeochemical fluxes can be analyzed in many different ways, including construction of detailed nutrient budgets and tracing the fate of nutrient inputs. The large amounts of data that exist for this sea makes it possible to validate model results with observations. Major biogeochemical properties of the Baltic Sea are discussed through an analyses of model sensitivity to external forcing and internal parameterizations. Model results emphasize two features that are especially important for ecosystem management: i) impacts of local measures would always be modified by the long-range transports from other regions and ii) the response to significant changes in loads would only be seen after several decades.     

Bacterial diversity and function in the Baltic Sea with an emphasis on cyanobacteria

In this article we summarize the current knowledge of Baltic Sea cyanobacteria, focusing on diversity, toxicity, and nitrogen fixation in the filamentous heterocystous taxa. We also review the recent results of our microbial diversity studies in planktonic and benthic habitats in the Baltic Sea. Based on molecular analyses, we have improved the understanding of cyanobacterial population structure by assessing genetic diversity within species that are morphologically inseparable. Moreover, we have studied microbial functions such as toxin production and nitrogen fixation in situ under different environmental conditions. Phosphorus limitation of bloom-forming, nitrogen-fixing cyanobacteria was clearly verified, emphasizing the importance of continuous efforts to reduce this element in the Baltic Sea. We have designed a rapid and reliable detection method for the toxic cyanobacterium Nodularia spumigena, which can be used to study bloom formation of this important toxin producer.     

Increasing nitrogen limitation in the Bothnian Sea,potentially caused by inflow of phosphate-rich water from the Baltic Proper

The study showed that the open water of the Bothnian Sea (BS) is likely to have shifted from altering nitrogen and phosphorous limitations of the spring bloom to more nitrogen-limited conditions during the last 20 years. This is affected by the by inflow of phosphate-rich and oxygen-depleted water from depths near the halocline in the northern Baltic Proper, where severe oxygen conditions currently cause extreme phosphate concentrations in the deep water. The change in relation between inorganic nitrogen and phosphorous in the BS occurs first in the deep water and then progresses to the surface water. The change can potentially cause increased production in the BS and more frequent cyanobacterial blooms. There does not appear to be any immediate concern in the short-term perspective for the state of the BS, but a progression of the processes may lead to a more eutrophic state of the BS.

Electronic supplementary material

The online version of this article (doi:10.1007/s13280-015-0675-3) contains supplementary material, which is available to authorized users.     

Searching efficient protection strategies for the eutrophied Gulf of Finland: the combined use of 1D and 3D modeling in assessing long-term state scenarios with high spatial resolution

An experiment combining the use of two ecosystem models was conducted to search for effective protection strategies for the Gulf of Finland (Baltic Sea). Reference and scenario simulations were first run with a one-dimensional (1D) model for seven main basins of the entire Baltic Sea until steady state was achieved. The obtained basinwise distributions of inorganic nitrogen (N) and phosphorus (P), as well as sediment labile P, were then used to initiate 5-y simulations with a three-dimensional (3D) ecosystem model. The results suggest that relatively small local load reductions (the "Finland" scenario) would improve only the state of adjacent coastal waters significantly. This would be the case, even for runs covering several decades, which clearly exceed the residence times of nutrients in the Gulf of Finland. A significant decrease from a substantial loading source to the Gulf (the "St. Petersburg" scenario) would decrease cyanobacterial biomasses in the entire Gulf of Finland and also immediately outside it. A reduction in the current Polish nutrient loads would improve the situation in the whole Baltic Proper and cause an extensive decline in cyanobacterial biomasses in the Gulf of Finland, as well. However, it would take several decades until the improvement caused by reducing loads in the "Poland" scenario is seen, while in the "St. Petersburg" scenario the corresponding time lag would only be a few years. Our results suggest that the common water protection policy in the Baltic Sea region should have the largest nutrient sources as its primary target, regardless of their location and country.     

Projected future climate change and Baltic Sea ecosystem management

Climate change is likely to have large effects on the Baltic Sea ecosystem. Simulations indicate 2–4 °C warming and 50–80 % decrease in ice cover by 2100. Precipitation may increase ~30 % in the north, causing increased land runoff of allochthonous organic matter (AOM) and organic pollutants and decreased salinity. Coupled physical–biogeochemical models indicate that, in the south, bottom-water anoxia may spread, reducing cod recruitment and increasing sediment phosphorus release, thus promoting cyanobacterial blooms. In the north, heterotrophic bacteria will be favored by AOM, while phytoplankton production may be reduced. Extra trophic levels in the food web may increase energy losses and consequently reduce fish production. Future management of the Baltic Sea must consider the effects of climate change on the ecosystem dynamics and functions, as well as the effects of anthropogenic nutrient and pollutant load. Monitoring should have a holistic approach, encompassing both autotrophic (phytoplankton) and heterotrophic (e.g., bacterial) processes.     

Impact of Climate Change on Ecological Quality Indicators and Biogeochemical Fluxes in the Baltic Sea: A Multi-Model Ensemble Study

Multi-model ensemble simulations using three coupled physical-biogeochemical models were performed to calculate the combined impact of projected future climate change and plausible nutrient load changes on biogeochemical cycles in the Baltic Sea. Climate projections for 1961-2099 were combined with four nutrient load scenarios ranging from a pessimistic business-as-usual to a more optimistic case following the Helsinki Commission's (HELCOM) Baltic Sea Action Plan (BSAP). The model results suggest that in a future climate, water quality, characterized by ecological quality indicators like winter nutrient, summer bottom oxygen, and annual mean phytoplankton concentrations as well as annual mean Secchi depth (water transparency), will be deteriorated compared to present conditions. In case of nutrient load reductions required by the BSAP, water quality is only slightly improved. Based on the analysis of biogeochemical fluxes, we find that in warmer and more anoxic waters, internal feedbacks could be reinforced. Increased phosphorus fluxes out of the sediments, reduced denitrification efficiency and increased nitrogen fixation may partly counteract nutrient load abatement strategies.     

Hypoxia in the Baltic Sea: Biogeochemical Cycles, Benthic Fauna, and Management

Hypoxia has occurred intermittently over the Holocene in the Baltic Sea, but the recent expansion from less than 10 000 km² before 1950 to >60 000 km² 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.     

Nutrient mitigation capacity in Mississippi Delta, USA drainage ditches

Eutrophication and hypoxia within aquatic systems are a serious international concern. Various management practices have been proposed to help alleviate nutrient loads transported to the Gulf of Mexico and other high-profile aquatic systems. The current study examined the nutrient mitigation capacity of a vegetated (V) and non-vegetated (NV) agricultural drainage ditch of similar size and landform in the Mississippi Delta. While no statistically significant differences in ammonium, nitrate, or dissolved inorganic phosphorus mitigation between the two ditches existed, there were significant differences in total inorganic phosphorus percent load reductions (V: 36% ± 4; NV: 71% ± 4). However, both agricultural drainage ditches were able to mitigate nutrients, thus reducing the load reaching downstream aquatic receiving systems. Further studies examining ecosystem dynamics within drainage ditches such as sediment and plant nutrient partitioning, as well as microbial processes involved, are needed to provide a better understanding of natural nutrient variability, seasonality and flux.     

MEAD: an interdisciplinary study of the marine effects of atmospheric deposition in the Kattegat

This paper summarises the results of the EU funded MEAD project, an interdisciplinary study of the effects of atmospheric nitrogen deposition on the Kattegat Sea between Denmark and Sweden. The study considers emissions of reactive nitrogen gases, their transport, transformations, deposition and effects on algal growth together with management options to reduce these effects. We conclude that atmospheric deposition is an important source of fixed nitrogen to the region particularly in summer, when nitrogen is the limiting nutrient for phytoplankton growth, and contributes to the overall eutrophication pressures in this region. However, we also conclude that it is unlikely that atmospheric deposition can, on its own, induce algal blooms in this region. A reduction of atmospheric nitrogen loads to this region will require strategies to reduce emissions of ammonia from local agriculture and Europe wide reductions in nitrous oxide emissions.     

Future Nutrient Load Scenarios for the Baltic Sea Due to Climate and Lifestyle Changes

Dynamic model simulations of the future climate and projections of future lifestyles within the Baltic Sea Drainage Basin (BSDB) were considered in this study to estimate potential trends in future nutrient loads to the Baltic Sea. Total nitrogen and total phosphorus loads were estimated using a simple proxy based only on human population (to account for nutrient sources) and stream discharges (to account for nutrient transport). This population-discharge proxy provided a good estimate for nutrient loads across the seven sub-basins of the BSDB considered. All climate scenarios considered here produced increased nutrient loads to the Baltic Sea over the next 100 years. There was variation between the climate scenarios such that sub-basin and regional differences were seen in future nutrient runoff depending on the climate model and scenario considered. Regardless, the results of this study indicate that changes in lifestyle brought about through shifts in consumption and population potentially overshadow the climate effects on future nutrient runoff for the entire BSDB. Regionally, however, lifestyle changes appear relatively more important in the southern regions of the BSDB while climatic changes appear more important in the northern regions with regards to future increases in nutrient loads. From a whole-ecosystem management perspective of the BSDB, this implies that implementation of improved and targeted management practices can still bring about improved conditions in the Baltic Sea in the face of a warmer and wetter future climate.