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For mitigation of climate change, all sources and sinks of greenhouse gases from the environment must be quantified and their driving factors identified. Nitrous oxide (N2O) is a strong greenhouse gas, and the contribution of aquatic systems to the global N2O budget remains poorly constrained. In this study, we measured N2O concentrations in a eutrophic coastal system, Roskilde Fjord (Denmark), and combined measurements with statistical modeling to quantify the N2O fluxes and budget in the system over a period of six months. To do so, we collected water at 15 sampling points and measured N2O concentrations along with physico-chemical water quality parameters, e.g. temperature, salinity, dissolved inorganic nitrogen and phosphorus, and silicon. We used mixed-effect regression models to predict N2O concentrations in the water from water quality parameters. We then derived N2O fluxes using well-established equations of N2O solubility and water-atmosphere exchanges. These fluxes were then put in perspective with those measured at the landscape scale by eddy-covariance at a 96 m nearby tall tower, and to those estimated from the agricultural land next to the fjord using Intergovernmental Panel on Climate Change (IPCC) guidelines. N2O concentrations in the Roskilde Fjord ranged between 2.40 and 8.05 nmol l−1. The best fitting model between water parameters and N2O concentrations in water included phosphorus and temperature. We estimated that (i) Roskilde Fjord was a sink of N2O, with a median inward flux of -0.04 nmol m−2 s−1, (ii) while the surrounding median agricultural flux was 0.13-0.18 nmol m−2 s−1, and (iii) the median landscape flux was 0.07 nmol m−2 s−1. All estimates of N2O fluxes were of the same magnitude and consistent with each other. These preliminary results need to be consolidated by further research.

Light is essential for primary production and, therefore, its attenuation controls the vertical distribution of plants and phytoplankton over the water column. The diffuse attenuation of irradiance (K) is mainly governed by the attenuation by the water itself and the concentrations of optically active substances (e.g., phytoplankton, inorganic particles and colored dissolved molecules), which makes it an important parameter for eutrophication monitoring. Over the past century, Denmark has had recurrent eutrophication events, with extreme episodic cases where anoxic conditions were observed. Since the 1980’s, eutrophication in Danish waters has been monitored with regards to the diffuse attenuation coefficient of scalar irradiance (Ko) of photosynthetically active radiation (PAR, 400–700 nm), Ko(PAR). However, radiometric measurements in Denmark are difficult in winter due to low solar zenith angle and only few light hours. On top of that, radiometric measurements in the first meters of the water column are highly affected by light refraction influenced by waves, compromising the monitoring of shallow turbid waters as in Denmark. Therefore, we developed a semi-analytical model based on data to from a spectral AC instrument (AC-S, Sea-Bird Scientific) that can estimate the underwater light field and the diffuse attenuation coefficient of downwelling irradiance, Kd(PAR). We tested two distinct approaches based on equations from the literature for estimation of Kd(PAR). The results show that modeled PAR profiles follow the overall shape of in situ radiometric profiles but with smoother profiles, especially in the surface layer (2–5 m). Along with that, the method provided robust Kd(PAR) estimates, that were strongly correlated to the reference Ko(PAR) values from in situ profiles and with low root mean square error (RMSE). Thus, AC-S data can be used to estimate the underwater light field and Ko(PAR). This will make possible to retrieve Ko(PAR) in the absence of daylight and, therefore, allow for environmental monitoring outside the daylight hours, making environmental monitoring more efficient. In addition, the method provides valuable insights into the factors controlling light attenuation.

The study is based on a very extensive data-set of physical, biological, and optical parameters from below the sea ice in the western Amundsen Basin, central Arctic Ocean, in August–September 2012 during the record low sea ice extent. The water column was strongly stratified at all stations related to salinity differences between a surface layer of reduced salinities (34). A nitrate utilization-based budget in the surface layer gave a primary production of 67.5 mg C m− 2 d− 1, which reduced to 3.9 mg C m− 2 d− 1 in August 2012. Amundsen Basin primary production rates are lower than rates determined for other Arctic Ocean deep-water basins, and also lower compared to rates on the shelf. Below ice phytoplankton was well adapted to low light conditions in the Amundsen Basin and the photosynthetic potential was high, but limited by the low nutrient fluxes induced by the strong stratification. Amundsen Basin is foreseen to be ice-free in summer in 3–4 decades, and the question whether primary production will increase when ice-free was resolved with a coupled physical-biogeochemical model. Results showed that production will increase 10 to 14 times from the present 3.9 mg C m− 2 d− 1 to 37.4 and 55.2 mg C m− 2 d− 1 for an ice-free August and July–August, respectively. The study substantiates that both present and future ice-free low production rates were related to the strong stratification, reduced nutrient fluxes, and deep lying nutrient rich waters. Low production rates and strong stratification are discussed in the view of parameters that increase this stratification as higher freshwater run off or reduce stratification as wind.

The effects of previous photosynthetic activity on dark respiration rates were investigated in Heterocapsa rotundata (Dinophyceae), Rhodomonas salina (Cryptophyceae) and Thalassiosira weissflogii (Coscinodiscophyceae). Dark respiration rates were measured after 1 to 3 h of light exposure in 12 bottles where incubation irradiances ranging from 0 to 600 μmol photons m − 2 s − 1 . The results showed that the dark respiration increases with increasing light exposure (both irradiance and duration) in all exponentially growing cultures but not in two experiments with cultures in stationary phase ( T. weissflogii ). This relationship could be described by a linear model for all species, relating respiration to the previous accumulated photosynthetic product with a slope of about 0.035 h − 1 , indicating a growth-related respiration of 3.5% per hour of the newly fixed carbon. The maximum respiration rate (R max ) was estimated from both light dose and accumulated photosynthetic product using a non-linear model. Although R max and light-saturated gross photosynthesis (P g max ) varied between the three species, a similar R max : P g max ratio of ~ 0.12 was found. The maintenance respiration rates which are comparable with maintenance respiration rates from literature, varied from 0.004 to 0.02 mol O 2 g − 1 Chl h − 1 for the three species. Overall, this study suggests a diel periodicity of phytoplankton respiration driven by accumulation of newly fixed carbon in light. The consequence is a decreasing respiration rate during the night or during long dark incubations.

To analyse the potential future ecological state of estuaries located in the temperate climate (here exemplified with the Odense Fjord estuary, Denmark), we combined end-of-the-century climate change projections from four different climate models, four contrasting land use scenarios (“Agriculture for nature”, “Extensive agriculture”, “High-tech agriculture” and “Market driven agriculture”) and two different eco-hydrological models. By decomposing the variance of the model-simulated output from all scenario and model combinations, we identified the key sources of uncertainties of these future projections. There was generally a decline in the ecological state of the estuary in scenarios with a warmer climate. Strikingly, even the most nature-friendly land use scenario, where a proportion of the intensive agricultural area was converted to forest, may not be enough to counteract the negative effects of a future warmer climate on the ecological state of the estuary. The different land use scenarios were the most significant sources of uncertainty in the projections of future ecological state, followed, in order, by eco-hydrological models and climate models, albeit all three sources caused high variability in the simulated outputs. Therefore, when projecting the future state of aquatic ecosystems in a global warming context, one should at the very least consider to evaluate an ensemble of land use scenarios (nutrient loads) but ideally also include multiple eco-hydrological models and climate change projections. Our study may set precedence for future attempts to predict and quantify uncertainties of model and model input ensembles, as this will likely be key elements in future tools for decision-making processes.

Anthropogenic nutrient enrichment is changing the structure and the function of coastal ecosystems. These coastal zones are transitions between freshwater and marine systems where multiple biogeochemical processes remove, produce, and transform organic matter. The extent to which the coastal zone is merely a conduit for terrestrial (allochthonous) organic matter versus a distinct source of autochthonous organic matter fueled by eutrophication is unclear. To address this issue, we characterized the freshwater and marine dissolved organic matter (DOM) pools in a eutrophic estuary with a long water residence time (Roskilde Fjord, Denmark) over an annual cycle. We combined elemental, optical (absorbance and fluorescence), and isotopic analyses to obtain insight about the bulk properties of the DOM pool during this period. We also used sediment traps to analyze the changes related to the exchange of organic matter between the particulate organic matter and DOM fractions. The results showed that labile autochthonous DOM from in situ primary production was rapidly transformed to more recalcitrant DOM that accumulated in the estuary despite continuous exchange with the open sea. Also, parts of the particulate organic matter pool were degraded rapidly (within 24 hr) and transformed into the DOM pool. Accumulated DOM was characterized by relatively low molecular size and stable carbon isotopic value and by high protein-like fluorescence. These results indicate that autotrophic material can be a major source of specific recalcitrant DOM in eutrophic coastal waters, contributing significantly to the flux of organic carbon to the ocean.

Carbon-to-chlorophyll a ratios (C:Chl a; weight : weight) were analyzed for 7578 coastal seawater samples collected from Danish waters from 1990 to 2014. The aim was to identify the seasonal and spatial dynamics relative to nutrient richness and to study the effect of reduced nitrogen loadings over time. C:Chl a values were lowest during winter, about 15 across all stations. During the spring, C:Chl a increased to summer values between 20 and 96, depending on the annual mean of total nitrogen concentration. An inter-annual sinusoidal model with monthly time steps described the seasonal C:Chl a pattern well. The amplitudes of the model varied inversely with the annual mean of total nitrogen. Data also showed that a reduction in nitrogen loadings to the area by ∼ 40% over the past 24 yr, resulted in a statistically significant increase in mean annual C:Chl a values of 0.8 ± 0.2 yr−1. The patterns derived from this large data set can be used to predict C:Chl a values for temperate coastal phytoplankton. Use of the empirical relationships derived from the data set improves predictions of C:Chl a values and thereby, e.g., carbon based food-web calculations and carbon-based ecosystem models, which often are validated using chlorophyll measurements.

The chromophoric dissolved organic matter (CDOM) is a significant water constituent influencing inherent and apparent optical properties of natural waters and plays a key role in ecosystem functioning. The spectral slope (S) describing the approximate exponential decline in CDOM absorption with increasing wavelength is widely used for tracing changes in the chemical composition of CDOM. The currently accepted method of characterizing CDOM absorption (i.e., fitting a simple exponential model) can lead to loss of information and large errors. We propose a better method for modeling CDOM absorption spectra based on a Gaussian decomposition approach that removes the errors associated with the choice of the spectral range used to estimate S. Using artificially generated spectra with known parameters (n = 1000), we show that our method provides robust estimates of S closely resembling the original values. On average, the error on S estimations was 0.16% for the proposed method compared to 27% and 11% for the traditional modeling approaches fitted over 300–700 nm and 240–700 nm respectively. We further demonstrate the ability of the method to decompose and model chromophores present in complex spectra from oceanic water samples from around the world. The proposed method opens avenues for long-term or cross-site comparison studies of the dynamics of the CDOM pool and constitutes a promising supplement to techniques based on CDOM fluorescence.

We used continuous measurements of dissolved oxygen (DO) in dark bottles to characterise patterns of the dark respiration rate (R dark) for three marine phytoplankton monocultures and in natural-water samples from two marine coastal systems. Furthermore, patterns of ecosystem community respiration rate were determined from open-water changes in DO in a fjord and in a lake. We considered two models of R dark to describe temporal changes in DO: Constant R dark and decreasing R dark; increasing R dark. In addition, the effect of incubation time on R dark was investigated in bottle incubations. Constant R dark was observed in short-term (12-h) bottle incubations in natural-water samples from two marine coastal systems. Declining R dark was observed in marine phytoplankton cultures and open-water measurements in a lake. Increasing R dark was observed in open-water measurements in a fjord, particularly during summer. Long-term (120-h) bottle incubations in natural-water samples showed an increase in R dark after 48 and 72 h. We show that the conventional expectation of constant rates of respiration in darkness is far from typical, because non-linear changes are common under both controlled experimental conditions, as well as for open-water measurements of ecosystem respiration.

Intensive farming has severe impacts on the chemical status of groundwater and streams and consequently on the ecological status of dependent ecosystems. Eutrophication is a widespread problem in lakes and marine waters. Common problems are hypoxia, algal blooms, fish kills, and loss of water clarity, underwater vegetation, biodiversity and recreational value. In this paper we evaluate the nitrogen (N) and phosphorus (P) concentrations of groundwater and surface water in a coastal catchment, the loadings and sources of N and P, and their effect on the ecological status of an estuary. We calculate the necessary reductions in N and P loadings to the estuary for obtaining a good ecological status, which we define based on the number of days with N and P limitation, and the corresponding stream and groundwater threshold values assuming two different management options. The calculations are performed by the combined use of empirical models and a physically based 3-D integrated hydrological model of the whole catchment. The assessment of the ecological status indicates that the N and P loads to the investigated estuary should be reduced to levels corresponding to 52 and 56% of the current loads, respectively, to restore good ecological status. Model estimates show that threshold total N (TN) concentrations should be in the range of 2.9 to 3.1 mg l−1 in inlet freshwater (streams) to Horsens estuary and 6.0 to 9.3 mg l−1 in shallow aerobic groundwater (∼ 27–41 mg l−1 of nitrate), depending on the management measures implemented in the catchment. The situation for total P (TP) is more complex, but data indicate that groundwater threshold values are not needed. The stream threshold value for TP to Horsens estuary for the selected management options is 0.084 mg l−1. Regional climate models project increasing winter precipitation and runoff in the investigated region resulting in increasing runoff and nutrient loads to the Horsens estuary and many other coastal waters if present land use and farming practices continue. Hence, lower threshold values are required in many coastal catchments in the future to ensure good status of water bodies and ecosystems.

Fine submicron organic particles can represent an important fraction of the dissolved organic matter (DOM) pool in aquatic ecosystems and their optical properties differ from those normally considered dissolved (

In optically complex inland waters, the underwater attenuation of photosynthetically active radiation (PAR) is controlled by a variable combination of absorption and scattering components of the lake or river water. Here we applied a photon budget approach to identify the main optical components affecting PAR attenuation in Lake St. Charles, a drinking water reservoir for Quebec City, Canada. This analysis showed the dominant role of colored dissolved organic matter (CDOM) absorption (average of 44% of total absorption during the sampling period), but with large changes over depth in the absolute and relative contribution of the individual absorption components (water, nonalgal particulates, phytoplankton and CDOM) to PAR attenuation. This pronounced vertical variation occurred because of the large spectral changes in the light field with depth, and it strongly affected the average in situ diffuse absorption coefficients in the water column. For example, the diffuse absorption coefficient for pure-water in the ambient light field was 10-fold higher than the value previously measured in the blue open ocean and erroneously applied to lakes and coastal waters. Photon absorption budget calculations for a range of limnological conditions confirmed that phytoplankton had little direct influence on underwater light, even at chlorophyll a values above those observed during harmful algal blooms in the lake. These results imply that traditional measures of water quality such as Secchi depth and radiometric transparency do not provide a meaningful estimate of the biological state of the water column in CDOM-colored lakes and reservoirs.

Monitoring data (1999 to 2012) and data from a 2 wk field study at a seasonally strat- ified station in the Aarhus Bight near the Danish coast were used to demonstrate that the vertical distribution of photosynthesis influences both water column oxygen conditions and the fate of the organic material produced. The primary production (PP) occurring below the surface layer, i.e. in the pycnocline-bottom layer (PBL), is shown to contribute significantly to total PP. Oxygen concen- trations in the PBL are shown to correlate significantly with the deep primary production (DPP) as well as with salinity, phosphate loading, wind and transparency in the surface layer. The phyto- plankton communities detected in the surface layer and PBL during the field study were very dif- ferent. Large cells, especially Ceratium spp., dominated in the PBL, while small diatoms, mainly Proboscis alata, dominated in surface waters. On the basis of chlorophyll-normalised photo - synthetic parameters and variable fluorescence, it is shown that the 2 populations were physio - logically distinct. The population in the PBL was photosynthetically active and adapted/ acclimated to lower light than the population in the surface layer. Sinking rates (based on sedi- ment trap collections) of carbon and nitrogen were highest in the PBL. Lyngsgaard et al. (2014; Limnol Oceanogr 59:1679�1690) have demonstrated that the vertical distribution of PP in this region is influenced by anthropogenic nutrient loading. Thus, the present study indicates that eutrophication effects may include changes in the structure of planktonic food webs and element cycling in the water column, both brought about through an altered vertical distribution of PP.

Groundwater-borne contaminants such as nutrients, dissolved organic carbon (DOC), coloured dissolved organic matter (CDOM) and pesticides can have an impact the biological quality of lakes. The sources of pollutants can, however, be difficult to identify due to high heterogeneity in groundwater flow patterns. This study presents a novel approach for fast hydrological surveys of small groundwater-fed lakes using multiple groundwater-borne tracers. Water samples were collected from the lake and temporary groundwater wells, installed every 50 m within a distance of 5–45 m to the shore, were analysed for tracer concentrations of CDOM, DOC, total dissolved nitrogen (TDN, groundwater only), total nitrogen (TN, lake only), total dissolved phosphorus (TDP, groundwater only), total phosphorus (TP, lake only), δ18O ∕ δ16O isotope ratios and fluorescent dissolved organic matter (FDOM) components derived from parallel factor analysis (PARAFAC). The isolation of groundwater recharge areas was based on δ18O measurements and areas with a high groundwater recharge rate were identified using a microbially influenced FDOM component. Groundwater discharge sites and the fractions of water delivered from the individual sites were isolated with the Community Assembly via Trait Selection model (CATS). The CATS model utilized tracer measurements of TDP, TDN, DOC and CDOM from the groundwater samples and related these to the tracer measurements of TN, TP, DOC and CDOM in the lake. A direct comparison between the lake and the inflowing groundwater was possible as degradation rates of the tracers in the lake were taken into account and related to a range of water retention times (WRTs) of the lake (0.25–3.5 years in 0.25-year increments). These estimations showed that WRTs above 2 years required a higher tracer concentration of inflowing water than found in any of the groundwater wells around the lake. From the estimations of inflowing tracer concentration, the CATS model isolated groundwater discharge sites located mainly in the eastern part of the lake with a single site in the southern part. Observations from the eastern part of the lake revealed an impermeable clay layer that promotes discharge during heavy precipitation events, which would otherwise be difficult to identify using traditional hydrological methods. In comparison to the lake concentrations, high tracer concentrations in the southern part showed that only a smaller fraction of water could originate from this area, thereby confirming the model results. A Euclidean cluster analysis of δ18O isotopes identified recharge sites corresponding to areas adjacent to drainage channels, and a cluster analysis of the microbially influenced FDOM component C4 further identified five sites that showed a tendency towards high groundwater recharge rate. In conclusion, it was found that this methodology can be applied to smaller lakes within a short time frame, providing useful information regarding the WRT of the lake and more importantly the groundwater recharge and discharge sites around the lake. Thus, it is a tool for specific management of the catchment.

The seasonal variation in phytoplankton activity is determined by analysing 1385 primary production (PP) profiles, chlorophyll a (Chl) concentration profiles and phytoplankton carbon biomass concentrations (C) from the period 1998-2012. The data was collected at six different stations in the Baltic Sea transition zone (BSTZ) which is a location with strong seasonal production patterns with light as the key parameter controlling this productivity. We show that the use of Chl as a proxy for phytoplankton activity strongly overestimates the contribution from the spring production to annual pelagic carbon flow. Spring (February and March) Chl comprised 16-30% of the total annual Chl produced, whereas spring C was much lower (8-23%) compared to the annual C. Spring PP accounted for 10-18% of the total annual PP, while the July-August production contributed 26-33%, i.e. within the time frame when zooplankton biomass and grazing pressure are highest. That is, Chl failed in this study to reflect the importance of the high summer PP. A better proxy for biomass may be C, which correlated well with the seasonal pattern of PP (Pearson correlation, p

Based on an extensive literature survey containing more than 12,000 paired measurements of dissolved organic carbon (DOC) concentrations and absorption of chromophoric dissolved organic matter (CDOM) distributed over four continents and seven oceans, we described the global distribution and transformation of dissolved organic matter (DOM) along the aquatic continuum across rivers and lakes to oceans. A strong log-linear relationship (R2 = 0.92) between DOC concentration and CDOM absorption at 350 nm was observed at a global scale, but was found to be ecosystem-dependent at local and regional scales. Our results reveal that as DOM is transported towards the oceans, the robustness of the observed relation decreases rapidly (R2 from 0.94 to 0.44) indicating a gradual decoupling between DOC and CDOM. This likely reflects the decreased connectivity between the landscape and DOM along the aquatic continuum. To support this hypothesis, we used the DOC-specific UV absorbance (SUVA) to characterize the reactivity of the DOM pool which decreased from 4.9 to 1.7 m2 × gC −1 along the aquatic continuum. Across the continuum, a piecewise linear regression showed that the observed decrease of SUVA occurred more rapidly in freshwater ecosystems compared to marine water ecosystems, suggesting that the different degradation processes act preferentially on CDOM rather than carbon content. The observed change in the DOM characteristics along the aquatic continuum also suggests that the terrestrial DOM pool is gradually becoming less reactive, which has profound consequences on cycling of organic carbon in aquatic ecosystems.

The study is based on CTD and water sampling in the Eurasian Basin of the Arctic Ocean in August–September 2012. The research icebreaker Oden and a helicopter were platforms for the sampling. We here report data for inorganic and organic bound nutrient (NO3 −, NO2 −, NH4 +, PO4 3−, Si(OH)4, DON, DOP) concentrations and chlorophyll a between 10 m and 200 m at 29 stations and down to about 4500 m at 13 stations. The data set also includes below sea ice samples (1-2 m) sampling at 31 stations. A total number of 15 different water masses were identified, and their N:P relationship showed that 13 of these clearly originated from the Atlantic Ocean.

Production and bioavailability of dissolved organic matter (DOM) were followed during a year in the nutrient-rich estuary, Roskilde Fjord (RF), and the more oligotrophic strait, Great Belt (GB), in Denmark. Bioavailability of dissolved organic carbon (DOC), nitrogen (DON), and phosphorous (DOP) was determined during incubations over six months. Overall, RF had three to five times larger pools of total nitrogen (TN) and total phosphorous (TP) and five to eight times higher concentrations of inorganic nutrients compared to GB. However, the allocation of carbon, nitrogen, and phosphorous into different pools were remarkably similar between the two systems. DON and DOP contributed with about equal relative fractions in the two systems: 72 +/- A 13% of total nitrogen and 21 +/- A 12% of total phosphorous. The average bioavailability of DOM was 25 +/- A 15, 17 +/- A 5.5, and 49 +/- A 29% for carbon, nitrogen, and phosphorous, respectively. The observed release of DIN from degradation of DON amounted to between 0.1 (RF winter) and 14 times (GB summer) the loadings from land and contributed with half of the total input of bioavailable nitrogen during summer. Hence, this study shows that nitrogen in DOM is important for the nitrogen cycling, especially during summer. The sum of inorganic nutrients, particulate organic matter, and bioavailable DOM (the dynamic pools of nutrients) accounted for 42 and 92% of nitrogen, and phosphorous, respectively, and was remarkably similar between the two systems compared to the difference in nutrient richness. It is hypothesized that the pelagic metabolism of nutrients in marine systems dictates a rather uniform distribution of the different fractions of nitrogen and phosphorous containing compounds regardless of eutrophication level.

Anthropogenic nitrogen (N)-loading has decreased significantly in the Baltic Sea Transition Zone over the past two decades. We show that the vertical distribution of primary production (PP) changed as a function of landbased N-loading using 1385 water column photosynthesis estimates, in which photosynthetic parameters were determined both in the surface water layer and in the pycnocline-bottom layer (PBL) at six stations near the Danish coast between 1998 and 2012. Total annual PP and surface layer PP (SPP) correlate positively with landbased N-loading from Denmark (p , 0.003). The percentage of annual PP occurring in the PBL (denoted as deep primary production, DPP) varied annually between 6% and 30% (mean 5 17%). The absolute magnitude of the DPP, as well as its relative proportion of total water column PP, correlates negatively with N-loading (p , 0.009 and p , 0.0003, respectively). Thus, SPP decreases in response to decreased N-loading, while DPP increases. Land-based N-loadings also correlate positively with the light attenuation coefficient (R2 5 0.39, p , 0.05), which may in part explain the response in DPP to changes in N-loading. DPP occurs in active phytoplankton communities acclimated and/or adapted to low light and producing oxygen in the PBL water. Primary production (PP) is an important factor in structuring marine ecosystems, and changes in PP in response to increased nutrient loading have been identified as being responsible for symptoms of eutrophication in coastal marine systems (Nixon 1995; Cloern 2001; Smith 2003). After recognizing the relationship between anthropogenic nutrient loading and eutrophication, many countries have initiated programs to reduce nutrient enrichment (Boesch 2002) with the expectation that marine PP will respond to a reduction in land-based nutrient loading. In some areas, enough data have now been collected to allow researchers to examine the extent to which this expectation has been realized. The Baltic Sea Transition Zone (BSTZ), which comprises the Kattegat and the Belt Seas and thus forms the connection between the Baltic Sea and the Skagerrak, is such an area. It is a shallow, stratified, and temperate marine system with dynamic hydrography. Surface salinity varies in the region from 10 to 14 in the southern part of the region and from 20 to 25 in the northern Kattegat

Nitrogen (N) fixation is fueling planktonic production in a multitude of aquatic environments. In meso- and poly-haline estuaries, however, the contribution of N by pelagic N₂ fixation is believed to be insignificant due to the high input of N from land and the presumed absence of active N₂-fixing organisms. Here we report N₂ fixation rates, nifH gene composition and nifH gene transcript abundance for key diazotrophic groups over 1 year in two contrasting, temperate, estuarine systems: Roskilde Fjord (RF) and the Great Belt (GB) strait. Annual pelagic N₂ fixation rates averaged 17 and 61 mmol N m(-2) per year at the two sites, respectively. In RF, N₂ fixation was mainly accompanied by transcripts related to heterotrophic (for example, Pseudomonas sp.) and photoheterotrophic bacteria (for example, unicellular diazotrophic cyanobacteria group A). In the GB, the first of two N₂ fixation peaks coincided with a similar nifH-expressing community as in RF, whereas the second peak was synchronous with increased nifH expression by an array of diazotrophs, including heterotrophic organisms as well as the heterocystous cyanobacterium Anabaena. Thus, we show for the first time that significant planktonic N₂ fixation takes place in mesohaline, temperate estuaries and that the importance of heterotrophic, photoheterotrophic and photosynthetic diazotrophs is clearly variable in space and time.

Loss of water clarity is one of the consequences of coastal eutrophication. Efforts have therefore been made to reduce external nutrient loadings of coastal waters. This paper documents improvements to water clarity between 1985 and 2008–2009 at four stations in the microtidal estuary Roskilde Fjord and find significant relationships to freshwater nutrient loadings. The paper then investigates to which extent changes in phytoplankton biomass (chlorophyll a (Chl a)), non-algal particulate organic matter (POM*), and residual attenuation in the water (Kb), respectively, can account for this optical improvement. Vertical light attenuation (Kd) declined, on average, by 34 %, accompanying a 71 % reduction of Chl a and an 80 % reduction of POM*. Residual attenuation declined by 26 % over the period in accordance with a measured 34 % decline of dissolved organic nitrogen. Analysis of simultaneous changes in light attenuation and Secchi depth also suggested a reduction of the scatter-to-absorption ratio over time. Considering the stronger reductions of particle concentrations than dissolved organic matter, the contribution of residual attenuation to vertical attenuation increased from 54 to 74 % in 1985 to 78 to 85 % in 2008–2009. Overall, efforts to reduce nutrient loading and improve water clarity appeared to have had a larger impact on POM* than on Chl a and colored dissolved organic matter concentrations in the estuary, which can account for the decrease in the scatter-to-absorption ratio. These optical changes lead to larger improvements of Secchi depth than of vertical light attenuation. The consequence of this is an overestimation (0.45–1.48 m) of the predicted increase of potential seagrass depth limits when based on Secchi depth rather than Kd.

The Western Baltic Sea is affected by eutrophication and receives nutrients from local land-based sources, atmospheric deposition and by advection from the neighbouring North Sea and Baltic Sea. In the present study, we evaluated the importance of local (Danish) versus external (Baltic Sea–North Sea) nutrient loads for surface Chl a- concentrations, total areal primary production and the vertical distributions of primary production in the W. Baltic Sea using the 3D coupled HBM-ERGOM model. This was assessed by improving descriptions of nutrient loads and primary production in the model and by conducting scenarios of different nutrient loads according to (i) the eutrophication level in 1990, (ii) the new Danish Water Plan (DWP) and iii) DWP+ the Baltic Sea Action Plan (BSAP2). The model results showed that local nutrient reductions have a profound effect on Chl a concentrations and primary production in the coastal areas of the Western Kattegat and the Belt Sea with the highest impact

In the 1980s, Danish coastal waters suffered from eutrophication and several nutrient management plans have been implemented during the years to improve ecological status. This study aims at giving a holistic ecosystem perspective on 25 years of mitigation measures. We report trends of nutrient inputs and the responses to these in various chemical and biological components. Nutrient inputs from land were reduced by ~50 % for nitrogen (N) and 56 % for phosphorus (P) since 1990. These reductions resulted in significant and parallel declines in nutrient concentrations, and initiated a shift in the dominance of primary producers towards reduced phytoplankton biomass (chlorophyll a concentration) and increased cover of macroalgae in deeper waters. In the last 5 years, eelgrass meadows have also expanded towards deeper waters, in response to improving water clarity. An expected improvement of bottom water oxygen conditions has not been observed, presumably because more frequent stratification and higher water temperatures have counteracted the expected positive effects of reduced nutrient inputs. The biomass of the benthic macrofauna decreased as expected, but it was composed of a drastic decline of filter feeders paralleled by a more moderate increase of deposit feeders. This shift was most likely induced by increasing stratification. The reduced benthic filtration along with the limited eelgrass cover probably kept relatively more particles in suspension, which can explain why improvements in the Secchi depths were modest. Overall, several ecosystem components demonstrated clear signs of improvement, suggesting that at least partial recovery is attainable. On this basis, we propose a conceptual scheme for recovery of shallow coastal ecosystems following marked reductions in nutrient inputs.

Eelgrass depth limits and water clarity in the Skive Fjord estuarine system have not improved despite nutrient input reductions of 30%. Long-term monitoring data (1989–2010) were used to investigate the underlying causes. Dissolved inorganic and organic nitrogen concentrations decreased significantly over time, whereas particulate organic nitrogen concentration, assumed to consist primarily of phytoplankton and phytoplankton detritus and calculated as a proportional factor to chlorophyll a, did not change. Total organic carbon, mostly of autochthonous origin, remained constant despite reduced nitrogen concentrations, resulting in an increasing C:N ratio of the organic material in the water column. Phytoplankton primary production also remained constant suggesting that phytoplankton growth was only limited by nitrogen to a minor degree. Alleviated grazing pressure caused by a reduction in the blue mussel standing stock and a pelagic food web dominated by jellyfish may have contributed to the constantly high phytoplankton levels. Particulate inorganic matter, likely reflecting sediment resuspension, increased over time, most probably in response to removal of blue mussels and declining eelgrass cover. The Skive Fjord estuarine system is affected by multiple pressures—nutrient enrichment, mussel dredging and climate change that must be addressed together for water clarity to improve and eelgrass to recover.

The effects of temperature and food availability on feeding and egg production of the Arctic copepod Calanus hyperboreus were investigated in Disko Bay, western Greenland, from winter to spring 2009. The abundance of females in the near bottom layer and the egg production of C. hyperboreus prior to the spring bloom document that reproduction relies on lipid stores. The maximum in situ egg production (± SE) of 54 ± 8 eggs female −1 d −1 was recorded in mid-February at chlorophyll a concentrations below 0.1 µg l −1 , whereas no egg production was observed in mid-April when the spring bloom developed. After reproduction, the females migrated to the surface layer to exploit the bloom and refill their lipid stores. In 2 laboratory experiments, initiated before and during the spring bloom, mature females were kept with and without food at 5 different temperatures ranging from 0 to 10°C and the fecal pellet and egg production were monitored. Food had a clear effect on fecal pellet production but no effect on egg production, while temperature did not have an effect on egg or fecal pellet production in any of the experiments. Analyses of carbon and lipid content of the females before and after the experiments did not reflect any effect of food or temperature in the pre-bloom experiment, whereas in the bloom experiment a clear positive effect of food was detected in female biochemical profiles. The lack of a temperature response suggests a future warmer ocean could be unfavorable for C. hyperboreus compared to smaller Calanus spp. which are reported to exploit minor temperature elevations for increased egg production.

Benthic flora can contribute significantly to gross primary production (GPP) of shallow coastal waters where light reaches the sea bottom. We quantified and compared benthic and pelagic GPP along nutrient gradients in time and space in the shallow estuary, Limfjorden, Denmark, based on monitoring data combined with historical information. Limfjorden experienced a shift from a pristine, benthic-dominated clear water regime with high total GPP in the early twentieth century to a eutrophic, plankton-dominated regime still with high total GPP in the 1980s when nutrient loadings peaked. Recent reductions in nutrient loadings reduced pelagic GPP, particularly in spring, but water clarity and benthic GPP did not increase correspondingly, so total GPP declined. The most nutrient-rich basins have remained plankton-dominated, with benthic vegetation constrained to shallow waters. The results support existing evidence that total GPP of shallow coastal areas does not increase systematically with eutrophication. Furthermore, the results suggest that total GPP may decline temporarily during oligotrophication as pelagic GPP declines, while feedback mechanisms delay or prevent restoration to a state with benthic dominance of GPP.

The relative importance of nutrient inputs from streams and the open sea for the water quality is a key issue for an effective management of eutrophication problems in estuaries. In order to resolve this question we have developed a coupled 3D hydrodynamic–ecological model with numerical nitrogen tracking for Horsens estuary, which is a typical Danish estuary that is shallow and well ventilated with water from the adjacent open sea. The nitrogen tracking technique revealed that nitrogen discharges from local streams accounted for the major part (40%–90%) of the nitrogen in NO3, NH4 and detritus whereas only 15% to 50% of the nitrogen in phytoplankton originated from streams. Effects of nitrogen loading reductions were most pronounced for nutrient concentrations and Chl a concentrations whereas the light condition was only slightly improved. The impact of stream nitrogen reductions on nitrate concentrations, Chl a concentrations and light attenuation was respectively 7, 2 and 3 times higher than the impact of open sea nitrogen reductions. Hence, even in estuaries that are well ventilated with water from the adjacent sea significant environmental improvements can be expected after reduction in local nutrient discharges.

The onset of spring bloom in temperate areas is a transition period where the low productive, winter phytoplankton community is transformed into a high productive spring community. Downwelling irradiance, mixing depth and the ability of the phytoplankton community to utilize the light, are key parameters determining the timing of the onset of the spring bloom. Knowing these parameters would thus provide tools for modeling the spring bloom and enhance our knowledge of ecophysiological processes during this period. Our main objective with this study was to provide data for the growth characteristics of some key species forming the spring bloom in the Gulf of Finland, and to apply those results in a simple dynamic model for the onset of the spring bloom, in order to test if the timing of the spring bloom predicted by the models corresponds to field observations. We investigated the photosynthetic characteristics of three diatoms and two dinoflagellates ( Chaetoceros wighamii , Melosira arctica , Thalassiosira baltica , Scrippsiella hangoei and Woloszynskia halophila ), at low temperatures (4–5 °C). All of these species are common during spring bloom in the Baltic Sea. Cultures of these species were acclimated to different irradiance regimes prior to measurements of photosynthesis, respiration, pigment concentration and light absorption. We did not find a positive relationship between respiration and growth rate, and we hypothesize that this relationship, which is well established at higher temperatures, is negligible or absent at low temperatures ( P m ), and maximum light utilization coefficient ( α ) was lowest and respiration ( R ) highest in the dinoflagellates. We made a model of the onset of the spring bloom in the western part of Gulf of Finland, using the obtained data together with monitoring data of mixing depth and water transparency from this area. Model results were compared to field observations of chlorophyll- a (Chl- a ) concentration. There was a good agreement between the model predictions and the observed onset of the spring bloom for the diatoms. S. hangoei , however, was not able to reach positive production in the model, and W. halophila had the similar growth characteristics as S. hangoei . Consequently, these species must have other competition strategies enabling them to exist and grow during spring bloom.

Accurate quantification of pelagic primary production is essential for quantifying the marine carbon turnover and the energy supply to the food web. Knowing the electron requirement (Κ) for carbon (C) fixation (Κ C) and oxygen (O2) production (Κ O2), variable fluorescence has the potential to quantify primary production in microalgae, and hereby increasing spatial and temporal resolution of measurements compared to traditional methods. Here we quantify Κ C and Κ O2 through measures of Pulse Amplitude Modulated (PAM) fluorometry, C fixation and O2 production in an Arctic fjord (Godthåbsfjorden, W Greenland). Through short- (2h) and long-term (24h) experiments, rates of electron transfer (ETRPSII), C fixation and/or O2 production were quantified and compared. Absolute rates of ETR were derived by accounting for Photosystem II light absorption and spectral light composition. Two-hour incubations revealed a linear relationship between ETRPSII and gross 14C fixation (R2 = 0.81) during light-limited photosynthesis, giving a Κ C of 7.6 ± 0.6 (mean ± S.E.) mol é (mol C)−1. Diel net rates also demonstrated a linear relationship between ETRPSII and C fixation giving a Κ C of 11.2 ± 1.3 mol é (mol C)−1 (R2 = 0.86). For net O2 production the electron requirement was lower than for net C fixation giving 6.5 ± 0.9 mol é (mol O2)−1 (R2 = 0.94). This, however, still is an electron requirement 1.6 times higher than the theoretical minimum for O2 production [i.e. 4 mol é (mol O2)−1]. The discrepancy is explained by respiratory activity and non-photochemical electron requirements and the variability is discussed. In conclusion, the bio-optical method and derived electron requirement support conversion of ETR to units of C or O2, paving the road for improved spatial and temporal resolution of primary production estimates.

conference abstract. EGU2015-11364; The greenhouse gas nitrous oxide (N2O) mainly originates in direct emissions from agricultural soils due to microbial reactions stimulated by the use of nitrogen fertilisers. Indirect N2O emissions from water systems due to nitrogen leaching and deposition from crop fields range between 26 and 37% of direct agricultural emissions, indicating their potential importance and uncertainty (Reay et al. 2012). The study presented here couples a top-down approach with eddy covariance (EC) and a bottom-up approach using different models and measurements. A QCL sensor at 96-m height on a tall tower measures the emissions of N2O from 1100 ha of crop fields and from the south part of the Roskilde fjord, in a 5-km radius area around the tall tower at Roskilde, Denmark. The bottom-up approach includes ecosystem modelling with CERES-EGC for the crops and PaSIM for the grasslands, and the N2O fluxes from the Roskilde fjord are derived from N2O sea water concentration measurements. EC measurements are now available from July to December 2014, and indicate a magnitude of the emissions from the crop fields around 0.2 mg N2O-N m-2 day-1 (range -9 to 5) which is consistent with the CERES-EGC simulations and calculations using IPCC emission factors. N2O fluxes from the Roskilde fjord in May and July indicated quite constant N2O concentrations around 0.1 µg N L-1 despite variations of nitrate and ammonium in the fjord. The calculated fluxes from these concentrations and the tall tower measurements consistently ranged between -7 and 6 mg N2O-N m-2 day-1. The study site also contains a waste water treatment plant, whose direct emissions will be measured in early 2015 using a dynamic plume tracer dispersion method (Mønster et al. 2014). A refined source attribution methodology together with more measurements and simulations of the N2O fluxes from the different land uses in this study site will provide a clearer view of the dynamics and budgets of N2O at the regional scale. The complementarity between these bottom-up and top-down approaches and their usefulness to disentangle direct and indirect N2O fluxes will also be discussed. Acknowledgements: This work was funded by the EU-FP7 InGOS project. References: Mønster JG, Samuelsson J, Kjeldsen P, Rella CW, Scheutz C. Quantifying methane emission from fugitive sources by combining tracer release and downwind measurements - A sensitivity analysis based on multiple field surveys. Waste Management. 2014 Aug;34(8):1416-28. Reay DS, Davidson EA, Smith KA, Smith P, Melillo JM, Dentener F, et al. Global agriculture and nitrous oxide emissions. Nature Clim Change. 2012 Jun;2(6):410-6

Scattering alters the path of photons, but ultimately their removal from the water column occurs by absorption by one of four components: water itself, coloured dissolved organic matter (CDOM), phytoplankton pigments or non-algal particulate matter (NAP). We calculated absorption budgets for two fjordal systems, Godthabsfjord (SW Greenland) and Young Sound (NE Greenland), based on these components and evaluated the fate of solar radiation in each system. Absorption by phytoplankton pigment accounted for 15–32% of photons in Godthabsfjord whilst in Young Sound the corresponding fraction was only 5–8%. NAP accounted for 13–25% of absorption in Young Sound and only 7–8% in Godthabsfjord whilst fractions of absorption by CDOM were more similar: 6–13% in Godthabsfjord and 6–18% in Young Sound. In typical temperate estuarine systems, nutrients, CDOM and particulate matter are supplied by riverine sources. Increased nutrient supply will tend to increase productivity whilst increased concentrations of CDOM and particles will increase light attenuation, thereby reducing productivity. The two Greenlandic fjords differ from typical estuarine systems in that their supply of nutrients and particles is decoupled. Freshwater feeding them comes from glacial melt and contains particles but low concentrations of nutrients. New nutrients are supplied by entrainment of oceanic water at the mouths of the fjords and sediment–water exchange of remineralized nutrients. Attenuation and thereby light availability in the two fjords is strongly correlated with turbidity (Godthabsfjord: r2 = 0.90 p

Miljømålet for god økologisk tilstand i de marine vandområder er bestemmende for hvilken belastning områderne kan tåle (dvs. for målbelastningen). Det er derfor en forudsætning for vandplanerne, at man har fastsat grænsen mellem God og Moderat (G-M) tilstand.G-M grænserne for sommerklorofylkoncentrationer kan fastlægges ud fra kendskab til klorofylkoncentrationer i en referencesituation og en såkaldt interkalibreret EQR værdi, som definerer den tilladelige forskel mellem referencesituationen og G-M grænsen. For danske kystområder er referenceniveauet for klorofyl undersøgt ved at anvende de modeller, der er udviklet til den marine Vandplans-værktøjskasse. Undersøgelsen er baseret på estimater af referencebelastningen, der bygger på basis af data fra DCE/Aarhus Universitet og data fra Baltic Nest Institute. Hvor det er muligt er der anvendt en ensemble tilgang, hvor referencetilstandens klorofylniveau er beregnet med begge de 2 typer af modelværktøjer, statistiske modeller og mekanistiske modeller, der indgår i værktøjskassen (se abstract om disse). På basis af resultaterne og en typeopdeling af havområder er G-M grænsernefastlagt for de enkelte vandområder. Denne præsentation vil gennemgå de beregninger der er foretaget og diskutere resultater og usikkerheder. Miljømålet for god økologisk tilstand i de marine vandområder er bestemmende for hvilken belastning områderne kan tåle (dvs. for målbelastningen). Det er derfor en forudsætning for vandplanerne, at man har fastsat grænsen mellem God og Moderat (G-M) tilstand.G-M grænserne for sommerklorofylkoncentrationer kan fastlægges ud fra kendskab til klorofylkoncentrationer i en referencesituation og en såkaldt interkalibreret EQR værdi, som definerer den tilladelige forskel mellem referencesituationen og G-M grænsen. For danske kystområder er referenceniveauet for klorofyl undersøgt ved at anvende de modeller, der er udviklet til den marine Vandplans-værktøjskasse. Undersøgelsen er baseret på estimater af referencebelastningen, der bygger på basis af data fra DCE/Aarhus Universitet og data fra Baltic Nest Institute. Hvor det er muligt er der anvendt en ensemble tilgang, hvor referencetilstandens klorofylniveau er beregnet med begge de 2 typer af modelværktøjer, statistiske modeller og mekanistiske modeller, der indgår i værktøjskassen (se abstract om disse). På basis af resultaterne og en typeopdeling af havområder er G-M grænserne fastlagt for de enkelte vandområder. Denne præsentation vil gennemgå de beregninger der er foretaget og diskutere resultater og usikkerheder.

Production and bioavailability of dissolved organic matter was quantified on a time scale of two days from size fractions ranging from bacteria to zooplankton in the York River, Virginia. The goal was to find the main contributor to DOM. Batch incubation experiments were labeled with N15-ammonium and C13-bicarbonate, and degradation kinetics of dissolved organic matter was measured over 14 days. Chlorophyll a increased to a maximum of 14 ± 0.6 μg L-1 during the incubation. Inorganic nitrogen was taken up producing a maximum of 10 ± 0.3 μmol particulate nitrogen L-1. The increase in biomass was mainly in the phytoplankton size fraction, which on average contributed 62 % of total particulate nitrogen and 61 % of total particulate carbon. Up to 5 ± 0.4 μmol dissolved organic nitrogen L-1 and 33 ± 6.2 μmol dissolved organic carbon L-1 was produced during the incubation. Bioavailability of phytoplankton produced dissolved organic carbon was 12 ± 1 % and higher than in the presence of bacteria, microzooplankton, or copepods (7 ± 3 %). The pattern for bioavailability of dissolved organic nitrogen was less clear and ranged from 4 – 7 %. This study revealed that phytoplankton was the main contributor to the production of dissolved organic matter.

Processes governing productivity at the base of the pelagic food web of the southern Indian Ocean are influenced primarily by physical‐chemical conditions with implications for the structure and function of the entire pelagic food web. Here, we report observations along a great circle transect from Cape Town, South Africa, to Broome in north western Australia. Primary production was tightly linked to water column stability and nutrient availability, with high productivity (1144 mg C m 22 day 21 ) in the sub-tropical convergence zone, and falling off by an order of magnitude in the sub-tropical gyre and tropical waters off northwest Australia. Primary production was largely confined to the GF/F fraction (GF/F .75% of total production and usually much higher) and the photosynthetic parameters showed adaption to the prevailing light levels. Bacterial production ranged from 19 to 155 mg C m 22 day 21 . No relationships were found between bacterial production and primary production or phytoplankton biomass and bacterial production seem more related to the state of the phytoplankton community as high bacterial production was observed in a post-bloom situation. The average BP:PP ratio was 31% (range 3.5‐71%).