Past, present and future state of the biogeochemical Si cycle in the Baltic Sea

The Baltic Sea is one of many aquatic ecosystems that show long-term declines in dissolved silicate (DSi) concentrations due to anthropogenic alteration of the biogeochemical Si cycle. Reductions in DSi in aquatic ecosystems have been coupled to hydrological regulation reducing inputs, but also with eutrophication, although the relative significance of both processes remains unknown for the observed reductions in DSi concentrations. Here we combine present and historical data on water column DSi concentrations, together with estimates of present river DSi loads to the Baltic, the load prior to damming together with estimates of the long-term accumulation of BSi in sediments. In addition, a model has been used to evaluate the past, present and future state of the biogeochemical Si cycle in the Baltic Sea. The present day DSi load to the Baltic Sea is 855 ktons y^− 1. Hydrological regulation and eutrophication of inland waters can account for a reduction of 420 ktons y^− 1 less riverine DSi entering the Baltic Sea today. Using published data on basin-wide accumulation rates we estimate that 1074 ktons y^− 1 of biogenic silica (BSi) is accumulating in the sediments, which is 36% higher than earlier estimates from the literature (791 ktons y^− 1). The difference is largely due to the high reported sedimentation rates in the Bothnian Sea and the Bothnian Bay. Using river DSi loads and estimated BSi accumulation, our model was not able to estimate water column DSi concentrations as burial estimates exceeded DSi inputs. The model was then used to estimate the BSi burial from measured DSi concentrations and DSi load. The model estimate for the total burial of BSi in all three basins was 620 ktons y^− 1, 74% less than estimated from sedimentation rates and sediment BSi concentrations. The model predicted 20% less BSi accumulation in the Baltic Proper and 10% less in the Bothnian Bay than estimated, but with significantly less BSi accumulation in the Bothnian Sea by a factor of 3. The model suggests there is an overestimation of basin-wide sedimentation rates in the Bothnian Bay and the Bothnian Sea. In the Baltic Proper, modelling shows that historical DSi concentrations were 2.6 times higher at the turn of the last century (ca. 1900) than at present. Although the DSi decrease has leveled out and at present there are only restricted areas of the Baltic Sea with limiting DSi concentrations, further declines in DSi concentrations will lead to widespread DSi limitation of diatoms with severe implications for the food web.     

Diatom stratigraphy and long-term dissolved silica concentrations in the Baltic Sea

In many parts of the world coastal waters with anthropogenic eutrophication have experienced a gradual depletion of dissolved silica (DSi) stocks. This could put pressure on spring bloom diatom populations, e.g. by limiting the intensity of blooms or by causing shifts in species composition. In addition, eutrophication driven enhanced diatom growth is responsible for the redistribution of DSi from the water phase to the sediments, and changes in the growth conditions may be reflected in the sediment diatom stratigraphy.To test for changes in diatom communities we have analyzed four sediment cores from the Baltic Sea covering approximately the last 100 years. The sediment cores originate from the western Gulf of Finland, the Kattegat, the Baltic Proper and the Gulf of Riga. Three out of the four cores reveal only minor changes in composition of diatom assemblages, while the Gulf of Riga core contains major changes, occurring after the second World War. This area is set apart from the other Baltic Sea basins by a high frequency of low after spring bloom DSi concentrations (< 2 µmol L^− 1) during a relatively well defined time period from 1991–1998. In 1991 to 1993 a rapid decline of DSi spring concentrations and winter stocks (down to 5 µmol L^− 1) in the Gulf was preceded by exceptionally intense diatom spring blooms dominated by the heavily silicified species Thalassiosira baltica (1991–1992; up to 5.5 mg ww L^− 1). T. baltica has been the principal spring bloom diatom in the Gulf of Riga since records began in 1975. DSi consumption and biomass yield experiments with cultured T. baltica suggest that intense blooms can potentially exhaust the DSi stock of the water column and exceed the annual Si dissolution in the Gulf of Riga. The phytoplankton time series reveals another exceptional T. baltica bloom period in 1981–1983 (up to 8 mg L^− 1), which, however, took place before the regular DSi measurements. These periods may be reflected in the conspicuous accumulation of T. baltica frustules in the sediment core corresponding to ca. 1975–1985.     

Changes in dissolved silicate loads to the Baltic Sea — The effects of lakes and reservoirs

We tested the hypothesis that dissolved silicate (DSi) yields [kg km^− 2 yr^− 1] of 82 major watersheds of the Baltic Sea can be expressed as a function of the hydraulic load (HL) as a measure of water residence time and the total organic carbon (TOC) concentration, both variables potentially increasing the DSi yield. Most boreal rivers fitted a linear regression model using HL as an independent variable to explain the DSi yield. Rivers with high HL, i.e., shortest residence times, showed highest DSi yields up to 2300 kg km^− 2 yr^− 1. This is most likely caused by an excess supply of DSi, i.e., the geochemical sources prevail over biological sinks in these boreal watersheds. The DSi yield for regulated and unregulated larger rivers of the boreal watersheds constituting about 40% of the total water discharge and of the total DSi load to the Baltic Sea, respectively, can be expressed as: DSi yield = 190 + 49.5 HL[m yr^− 1] + 0.346 TOC [µM] (R² = 0.80). Since both HL and TOC concentrations have decreased after damming, the DSi yields have decreased significantly in the regulated boreal watersheds, for the River Luleälven we estimated more than 30%. The larger eutrophic watersheds draining cultivated landscape of the southern catchment of the Baltic Sea and representing about 50% of the annual water discharge to the Baltic Sea, deviated from this pattern and showed lower DSi yields between 60–580 kg km^− 2 yr^− 1. DSi yields showed saturation curve like relationship to HL and it appears that DSi is retained in the watersheds efficiently through biogenic silica (BSi) production and subsequent sedimentation along the entire river network. The relationship between HL and DSi yields for all larger cultivated watersheds was best fitted by a Freundlich isotherm (DSi = 115.7HL¹⁰⁹; R² = 0.73), because once lake and reservoir area exceeds 10% of the watershed area, minimum DSi yields were reached. To estimate an uperturbed DSi yield for the larger eutrophic southeastern watersheds is still difficult, since no unperturbed watersheds for comparison were available. However, a rough estimate indicate that the DSi flux from the cultivated watersheds to the Baltic Sea is nowadays only half the uperturbed flux. Overall, the riverine DSi loads to the Baltic Sea might have dropped with 30–40% during the last century.     

Pore water phosphate and ammonia below the permanent halocline in the south-eastern Baltic Sea and their benthic fluxes under anoxic conditions

In this paper the results of a study on the distribution of pore water phosphates and ammonia, and their fluxes under anoxic condition in a deep (> 70 m) accumulation-type bottom of the south-eastern Baltic Sea, namely in the Gdańsk Deep and the adjacent areas, are presented. All measurements were taken during the growth period, i.e. in September 2000, April 2001 and June 2002. Benthic phosphate and ammonia fluxes were estimated using Fick's First Law. Phosphate and ammonia concentrations ranged from 7.5 to 266.3 μmol dm^− 3 and from 53.6 to 1248.3 μmol dm^− 3, respectively. The values recorded in the central part of the Gdańsk Deep were lower than those found both on its slopes and on the SW slope of the Gotland Deep. The lowest phosphate contents were typical of the Oblique Sill which separates the Gdańsk and Gotland Deeps.In 1993–2002, as a result of anoxia the sediments in the Gdańsk Deep released about 5.1 × 10³ t P and 22.8 × 10³ t N. These loads supplied on average 1.5% and 0.9% of phytoplankton's demand for P and N, respectively. In comparison to the total external load of nutrients discharged to the Gulf of Gdańsk (i.e. 8.79 × 10³ t year^− 1 P_tot and 130.79 × 10³ t year^− 1 N_tot; [Witek, Z., Humborg, Ch., Savchuk, O., Grelowski, A. and Łysiak-Pastuszak, E., 2003. Nitrogen and phosphorus budgets of the Gulf of Gdańsk (Baltic Sea). Est. Coast. Shelf Sci., 57:239–248.]), the return flux of P and N from the anoxic sediments to the water column in the Gdańsk Deep was a minor source of these elements.     

Alterations in nutrient limitations — Scenarios of a changing Baltic Sea

Previous trend studies have shown increasing nitrogen and phosphorus as well as decreasing silica concentrations in the water mass of the Baltic Sea. This has had an impact on the amount of primary production, but also on the quality and succession of plankton species. Present study examines the spatial and temporal patterns of potential nutrient limitations in the Baltic Sea for the time period 1970–2000. Generally, low concentrations of DSi can limit the diatom blooms and such conditions are found in the Gulf of Riga and Gulf of Finland during spring and summer. Nutrient ratios, DSi:DIN, DSi:DIP and DIN:DIP, are often used to determine which nutrient may limit the primary production. Annual long-term temporal trends of silica to inorganic nitrogen and phosphorus respectively show consistent decreasing patterns. The largest slopes are detected during spring and summer for DSi:DIN and during spring for DSi:DIP ratios. For the DIN:DIP ratio significant slopes are only found in a few locations despite increasing levels for both nutrients, displaying a large variation in trends. In the open Baltic Proper the present trends are positive during winter and negative during spring and autumn. Gulf of Finland and Gulf of Riga are areas where both DSi:DIP and DSi:DIN ratios are found close to the Redfield ratios for diatoms. Together with the evaluated trends these suggest that the Gulfs may become silica limited in a relatively near future. These findings give some implications on the development and impact of changing nutrient concentrations.     

Inter-annual variability of hypoxic conditions in a shallow estuary

Water quality data from two monitoring programs in the Pamlico River Estuary (PRE) were analyzed for dissolved oxygen (DO), salinity, temperature, and nutrient concentrations. Data were collected bi-weekly at 8 stations from 1997 to 2003 by East Carolina University and continuously at three stations from 1999 to 2003 by the U.S. Geological Survey. Hypoxic conditions were observed mostly in the upper to middle estuary, but the frequency of hypoxic events varied between years. During June to October in 1997–1999 (referred to as the oxic summers) bottom water hypoxia (DO < 2 mg l^− 1) was found in 8.7% of the observations. By contrast, during June to October in 2001–2003 (referred to as the hypoxic summers), 37.9% of the total measurements had DO concentrations less than 2 mg l^− 1. The more frequent and/or prolonged hypoxic conditions during the hypoxic summers were closely associated with stronger salinity stratification and greater loadings of nutrient and particulate matter.Salinity stratification appeared to be governed by patterns of freshwater discharge, and frequency of wind mixing events. The “oxic” summers were characterized by continuous low freshwater inflow (except one extremely high flow event due to hurricanes), stronger northeastward wind, and more frequent wind mixing events. In contrast, the hypoxic summers were characterized by frequent moderate freshwater inflow events, and fewer wind mixing events.The greater loadings of nutrient (nitrate, ammonium, and phosphate) and particulate matter during the hypoxic summers were primarily due to higher river discharges. At the head of the PRE, no significant differences were found in concentrations of nutrient and particulate nitrogen between the oxic and the hypoxic summers. In addition, chlorophyll a concentrations were averaged above 30 μg l^− 1 (maximum 167 μg l^− 1) during the hypoxic summers, significantly higher than those during the oxic summers (averaged around 15 μg l^− 1).     

Organic carbon budget for the Gulf of Bothnia

We calculated input of organic carbon to the unproductive, brackish water basin of the Gulf of Bothnia from rivers, point sources and the atmosphere. We also calculated the net exchange of organic carbon between the Gulf of Bothnia and the adjacent marine system, the Baltic Proper. We compared the input with sinks for organic carbon; permanent incorporation in sediments and mineralization and subsequent evasion of CO₂ to the atmosphere. The major fluxes were riverine input (1500 Gg C year^− 1), exchange with the Baltic Proper (depending on which of several possible DOC concentration differences between the basins that was used in the calculation, the flux varied between an outflow of 466 and an input of 950 Gg C year^− 1), sediment burial (1100 Gg C year^− 1) and evasion to the atmosphere (3610 Gg C year^− 1). The largest single net flux was the emission of CO₂ to the atmosphere, mainly caused by bacterial mineralization of organic carbon. Input and output did not match in our budget which we ascribe uncertainties in the calculation of the exchange of organic carbon between the Gulf of Bothnia and the Baltic Proper, and the fact that CO₂ emission, which in our calculation represented 1 year (2002) may have been overestimated in comparison with long-term means. We conclude that net heterotrophy of the Gulf of Bothnia was due to input of organic carbon from both the catchment and from the Baltic Proper and that the future degree of net heterotrophy will be sensible to both catchment export of organic carbon and to the ongoing eutrophication of the Baltic Proper.     

Th-derived particulate organic carbon fluxes in the northern Barents Sea with comparison to drifting sediment trap fluxes

Large-volume sampling of ²³⁴Th was conducted to estimate particulate organic carbon (POC) export in conjunction with drifting sediment trap deployments in the northern Barents Sea in July 2003 and May 2005. ²³⁴Th-derived POC fluxes averaged 42.3 ± 39.7 mmol C m^− 2 d^− 1 in 2003 and 47.1 ± 30.6 mmol C m^− 2 d^− 1 in 2005. Sediment trap POC fluxes averaged 13.1 ± 8.2 mmol C m^− 2 d^− 1 in 2003 and 17.3 ± 11.4 mmol C m^− 2 d^− 1 in 2005, but better reflected the transient bloom conditions that were observed at each station within a season. Although ²³⁴Th fluxes agreed within a factor 2 at most stations and depths sampled, sediment trap POC fluxes were lower than large-volume POC flux estimates at almost every station. This may represent an under-collection of POC by the drifting sediment traps or, conversely, an over-collection of POC by the large-volume sampling of ²³⁴Th. It is hypothesized that the offset between the two methods is partly due to the presence of the prymnesiophyte Phaeocystis pouchetii, which potentially causes a large variation in > 53-μm POC/²³⁴Th ratios. Due to the large proportion of dissolved carbon or mucilage released by P. pouchetii, and because it is thought that P. pouchetii does not contribute significantly to the vertical export of biogenic matter in the Barents Sea, the application of large-volume sampling of ²³⁴Th may yield relatively high, and possibly inaccurate POC/²³⁴Th ratios. Hence, POC fluxes derived from ²³⁴Th sampling may be inappropriate and drifting sediment traps might be a more reliable method to measure the vertical export of biogenic matter in regions that have recurrent P. pouchetii blooms, such as the Barents Sea.     

Thorium-234 derived information on particle residence times and sediment deposition in shallow waters of the south-western Baltic Sea

Activities of the naturally occurring, short-lived and highly particle-reactive radionuclide tracer ²³⁴Th in the dissolved and particulate phase were measured at three shallow-water stations (maximum water depths: 15.6, 22.7 and 30.1 m) in Mecklenburg Bay (south-western Baltic Sea) to constrain the time scales of the dynamics and the depositional fate of particulate matter. Activities of particle-associated (> 0.4 μm) and total (particulate + dissolved) ²³⁴Th were in the range of 0.08–0.11 dpm L^− 1 and 0.11–0.20 dpm L^− 1, respectively. The activity ratio of total ²³⁴Th and its long-lived and conservative parent nuclide ²³⁸U was well below unity (range: 0.09–0.19) indicating substantial radioactive disequilibria throughout the water column, very dynamic trace-metal scavenging and particle export from the water column at all three stations. For the discussion the ²³⁴Th data of this study were combined with previously published water-column ²³⁴Th and particulate-matter data from Mecklenburg Bay (Kersten et al., 1998. Applied Geochemistry 13, 339–347). The resulting average vertical distribution of total ²³⁴Th/²³⁸U disequilibria was used to estimate the depositional ²³⁴Th flux to the sediment. There was a virtually constant net downward flux of ²³⁴Th of about 28 dpm m^− 2 d^− 1 leaving each water layer of one meter thickness. Thorium-234-derived net residence times of particulate material regarding settling from a given layer in the water column were typically on the order of days, but with maximum values of up to a couple of weeks. Based on an average ratio of particulate matter (PM) to particle-associated ²³⁴Th a net flux of about 145 mg PM m^− 2 d^− 1 was estimated to leave each water layer of one meter thickness. The estimated cumulative water-column-derived particulate-matter fluxes at the seafloor are higher by a factor of about 2 than previously published sediment-derived estimates for Mecklenburg Bay. This suggests that about half of the settling particulate material is exported from the study area and/or subject to processes such as mechanical breakdown, remineralisation and dissolution. Lateral particulate-matter redistribution and particle breakdown in the water column (as opposed to the sediment) seem to be favoured by (repeated) particle resuspension from and resettling to the seafloor before ultimate sedimentary burial. The importance of net lateral redistribution of particulate material seems to increase towards the seafloor and be particularly high within the bottommost few meters of the water column.     

Seasonality of picophytoplankton chlorophyll a and biomass in the central Cantabrian Sea, southern Bay of Biscay

Seasonal changes in the abundance and biomass of cyanobacteria (Synechococcus and Prochlorococcus) and picoeukaryotes were studied by flow cytometry in the upper layers of the central Cantabrian Sea continental shelf, from April 2002 to April 2006. The study area displayed the typical hydrographic conditions of temperate coastal zones. A marked seasonality of the relative contribution of prokaryotes and eukaryotes was found. While cyanobacteria were generally more abundant for most of the year (up to 2.4 10⁵ cells mL^− 1), picoeukaryotes dominated the community (up to 10⁴ cells mL^− 1) from February to May. The disappearance of Prochlorococcus from spring through summer is likely related to shifts in the prevailing current regime. The maximum total abundance of picophytoplankton was consistently found in late summer–early autumn. Mean photic-layer picoplanktonic chlorophyll a ranged from 0.06 to 0.53 µg L^− 1 with a relatively high mean contribution to total values (33 ± 2% SE), showing maxima around autumn and minima in spring. Biomass (range 0.58–40.16 mg C m^− 3) was generally dominated by picoeukaryotes (mean ± SE, 4.28 ± 0.27 mg C m^− 3) with an average contribution of cyanobacteria of 30 ± 2%. Different seasonality of pigment and biomass values resulted in a clear temporal pattern of picophytoplanktonic carbon to chlorophyll a ratio, which ranged from 10 (winter) to 140 (summer). This study highlights the important contribution of picoplanktonic chlorophyll a and carbon biomass in this coastal ecosystem.     

Time series for dissolved cadmium at a coastal station in the Western Baltic Sea

Concentrations of dissolved Cd, Cd(diss), were measured weekly from June 1991 to June 1994 at a coastal station in the western Baltic Sea. The mean concentration of 204 pmol/dm³ is about 50% higher than in open Baltic Sea surface waters. A distinct seasonal cycle was observed with elevated concentrations in winter and spring (272 pmol/dm³) and lower values in summer and autumn (131 pmol/dm³). Relating the seasonal changes in Cd(diss) to the nutrient cycle revealed ΔCd(diss)/ΔNO₃ and ΔCd(diss)/ΔPO₄ ratios which are consistent with other measurements and seem to confirm the concept of a nutrient-like biogeochemistry of Cd. However, a time shift of two to three months exists between the depletion of nutrients in spring and the depletion of Cd(diss). Possibly, this indicates a decoupling of Cd(diss) from nutrients during the spring plankton bloom. However, no final conclusions can be drawn yet.Cd(diss) concentrations decreased significantly during the three year measurement period, whereas nitrate concentrations increased. A possible linkage between eutrophication and the Cd budget of the Baltic Sea is discussed.     

Impact of macrozoobenthic structures on near-bed sediment fluxes

Sandy sediments in shallow coastal waters of the Baltic Sea are often characterised by large numbers of biogenic structures which are produced by macrozoobenthos species. A series of experiments was devised to quantify how the interaction of such structures with the near-bed flow regime affects the sediment flux. Most experiments were done with simplified replicates of structures generated by typical species commonly found in the Mecklenburg Bight, starting with solitary structures and regularly-spaced arrays in a range of characteristic population densities, followed by a complex benthic macrofauna community, both artificial and alive. A laboratory flume channel, equipped with an acoustic Doppler flow sensor and a topography scanning laser, was used for high-resolution measurements (2 mm horizontal step size and 0.3 mm vertical resolution) of sand erosion (220 µm median grain size, at 20 cm s^− 1) and fine particle deposition (8 µm grain size, at 5 cm s^− 1). Sediment transport threshold values were measured for each layout. As a rule-of-thumb, both the erosion fluxes and the deposition of suspended matter increased considerably at low population densities (below 2%, expressed as percent of the sediment surface covered, i.e. roughness density RD). Above densities of 4%, erosion almost stopped inside the test arrays, and deposition remained well below the level of unpopulated areas. An attempt to extrapolate these findings to field conditions (using field current velocity data from 2001) showed that the net flux switched from erosion to deposition for densities above 5%. These parameters can now be integrated into a numerical sediment transport model coupling waves, currents, sediment dynamics and biological processes, which is currently under construction at the Baltic Sea Research Institute (IOW), Rostock, Germany.     

Seasonal study of dissolved CH4, CO2 and N2O in a shallow tidal system of the bay of Cádiz (SW Spain)

During 2004, 10 samplings were performed in order to measure dissolved methane (CH₄), carbon dioxide (CO₂) and nitrous oxide (N₂O) in the surface waters of Río San Pedro, a tidal creek in the salt marsh area of the Bay of Cádiz (SW Spain). The inner partvs of the creek is affected by the inputs coming from an intensive fish farm and the drainage of an extensive salt marsh area.Dissolved CH₄, CO₂ and N₂O concentrations ranged from 11 to 88 nM, 36 to 108 μM and 14 to 50 nM, respectively. Surface waters were in all cases oversaturated with respect to the atmosphere, reaching values of up to 5000% for CH₄, 1240% for CO₂ and 840% for N₂O. Dissolved CH₄, CO₂ and N₂O showed a significant tidal and seasonal variability. Over a tidal cycle, concentrations were always highest during low tide, which points to the influence of the inputs from the fish farm effluent and the drainage of the adjacent salt marsh area, as well as in situ production within the system. Dissolved CH₄, CO₂ and N₂O seasonal patterns were similar and showed maximum concentrations in summer conditions. Using four different parameterizations to calculate the gas transfer coefficients [Liss, P.S. and Merlivat, L., 1986. Air-sea exchange rates: introduction and synthesis. In P. Buat-Ménard (Ed.), The Role of Air-Sea Exchanges in Geochemical Cycling. Reidel, Dordrecht, The Netherlands, p. 113–127.; Clark, J.F., Schlosser, P., Simpson, H.J., Stute, M., Wanninkhof, R., and Ho, D.T., 1995. Relationship between gas transfer velocities and wind speeds in the tidal Hudson River determined by the dual tracer technique. In: B. Jähne and E. Monahan (Eds.), Air-Water Gas Transfer: AEON Verlag and Studio, Hanau, Germany, pp. 785–800.; Carini, S., Weston, N., Hopkinson, G., Tucker, J., Giblin, A. and Vallino, J., 1996. Gas exchanges rates in the Parker River estuary, Massachusetts. Biol. Bull., 191: 333–334.; Kremer, J.N., Reischauer, A. and D'Avanzo, C., 2003. Estuary-specific variation in the air-water gas exchange coefficient for oxygen. Estuaries, 26: 829–836.], the averaged air–water fluxes of CH₄, CO₂ and N₂O from the creek to the atmosphere ranged between 34 and 150 μmol CH₄ m^− 2 day^− 1, 73 and 177 mmol CO₂ m^− 2 day^− 1 and 24 and 62 μmol N₂O m⁻² day⁻¹, respectively.     

Production of Acartia omorii (Copepoda: Calanoida) in Ilkwang Bay, southeastern coast of Korea

Production of the marine calanoid copepod Acartia omorii was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay on the southeastern coast of Korea. A. omorii (nauplii + copepodites + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. omorii was averaged at 0.44 mgC m^− 3, with peaks in February and July, and relatively low biomass in late summer and fall. Egg production rate ranged from 2.4 to 151.9 μgC m^− 3 day^− 1, which was equivalent to 95–6075 eggs m^− 3 day^− 1. Fecundity of an adult female was averaged at 38 eggs female^− 1 day^− 1. Instantaneous growth rates of copepodites were higher than those of nauplii stages. Annual production of A. omorii ranged from 33.5 mgC m^− 3 year^− 1 to 221 mgC m^− 2 year^− 1, showing a seasonal variation of daily production rate with peaks in February and July. The daily production rate of A. omorii was significantly correlated with chlorophyll a concentration. These results suggest that standing stocks and/or productivity of phytoplankton are the major influencing factors, rather than water temperature for the seasonal variation of production of A. omorii in Ilkwang Bay.     

Distribution of phytoplankton in the southern Black Sea in summer 1996, spring and autumn 1998

The species composition, abundance, and biomass of micro- (>15 μm) and nano- (<15 μm) phytoplankton were studied along the southern Black Sea during June–July 1996 and March–April and September 1998. A total of 150 species were identified, 50% of them being dinoflagellates. The average total phytoplankton abundance changed from 77×10³ cells l⁻¹ in spring to 110×10³ cells l⁻¹ in autumn and biomass from 250 μg l⁻¹ in summer to 1370 μg l⁻¹ in spring. Based on the extensive sampling grid from June–July 1996, phytoplankton seemed to have a rather homogeneous biomass distribution in the southern Black Sea. In all periods, the coccolithophorid Emiliania huxleyi was the most abundant species, its contribution to the total abundance ranging from 73% in autumn to 43% in spring. However, in terms of biomass, diatoms made up the bulk of phytoplankton in spring (97%, majority being Proboscia alata) and autumn (73%, majority being Pseudosolenia calcar-avis), and dinoflagellates in summer (74%, Gymnodinium sp.). There was a remarkable similarity in the dominant species between the western and eastern regions of the southern Black Sea, indicating transport of phytoplankton within the basin.     

Analysis of the hydrophysical structure of the Sea of Azov in the period of the bottom anoxia development

The hydrophysical and hydrochemical structure of the Sea of Azov, with developed bottom anoxia, was studied during the RV “Akvanavt” cruise from July 31 to August 03, 2001. The anoxic zone with a thickness from 0.5 to 4 m above the bottom was found in all deep regions of the Sea. Concentrations of hydrochemical parameters were similar to the pronounced anoxic conditions (about 90 mmol m^− 3 of hydrogen sulfide, 17 mmol m^− 3 of ammonia, 6 mmol m^− 3 of phosphate, 7 mmol m^− 3 of total manganese). The hydrophysical structure was characterized by the uniform distribution of temperature in the upper 6–7 m mixed layer (UML). Below this a thin (0.4–0.8 m) thermocline layer was observed, just above the anoxic waters. Formation of this phenomenon was connected with that summer weather conditions. Intensive rains led to increased influx of river waters in June. That resulted in large input of allochtonous organic matter (OM) and inorganic nutrients; the latter were consumed on the additional autochthonous organic matter production. In July the weather was characterized by a significant rise in the daily averaged air temperature and large oscillations of temperature during the day. In this period a wind of constant direction was absent, but wind bursts were observed. The completed analyses showed that the formation of such a structure could be connected with the following factors: (i) positive growth trends of the daily averaged temperature and the daily oscillations of temperature, (ii) presence of wind bursts. The joint action of these factors resulted in the formation of the UML. The amplitude of wind bursts determined the depth of UML, and the value of trend determined the value of the temperature change in the thermocline. An initial presence of bottom halocline (caused by the Black Sea water influx to the bottom of the Sea of Azov) prevented the heating of the bottom layer and therefore led to an increase of vertical gradient of temperature in the thermocline. The spatial distribution of the turbulent exchange coefficient confirmed the existence of a “stagnation” area located above the anoxia zone, which is also, apparently, the reason for its occurrence.     

Protist entrapment in newly formed sea ice in the Coastal Arctic Ocean

Protist abundance and taxonomic composition were determined in four development stages of newly formed sea ice (new ice, nilas, young ice and thin first-year ice) and in the underlying surface waters of the Canadian Beaufort Sea from 30 September to 19 November 2003. Pico- and nanoalgae were counted by flow cytometry whereas photosynthetic and heterotrophic protists ≥ 4 µm were identified and counted by inverted microscopy. Protists were always present in sea ice and surface water samples throughout the study period. The most abundant protists in sea ice and surface waters were cells < 4 µm. They were less abundant in sea ice (418–3051 × 10³ cells L^− 1) than in surface waters (1393–5373 × 10³ cells L^− 1). In contrast, larger protists (≥ 4 µm) were more abundant in sea ice (59–821 × 10³ cells L^− 1) than in surface waters (22–256 × 10³ cells L^− 1). These results suggest a selective incorporation of larger cells into sea ice. The ≥ 4 µm protist assemblage was composed of a total number of 73 taxa, including 12 centric diatom species, 7 pennate diatoms, 11 dinoflagellates and 16 flagellates. The taxonomic composition in the early stage of ice formation (i.e., new ice) was very similar to that observed in surface waters and was composed of a mixed population of nanoflagellates (Prasinophyceae and Prymnesiophyceae), diatoms (mainly Chaetoceros species) and dinoflagellates. In older stages of sea ice (i.e., young ice and thin first-year ice), the taxonomic composition became markedly different from that of the surface waters. These older ice samples contained relatively fewer Prasinophyceae and more unidentified nanoflagellates than the younger ice. Diatom resting spores and dinoflagellate cysts were generally more abundant in sea ice than in surface waters. However, further studies are needed to determine the importance of this winter survival strategy in Arctic sea ice. This study clearly shows the selective incorporation of large cells (≥ 4 µm) in newly formed sea ice and the change in the taxonomic composition of protists between sea ice and surface waters as the fall season progresses.     

On the distribution of dissolved lead in the Loire estuary and the North Biscay continental shelf, France

The dissolved lead was studied in the whole salinity gradient of the system composed of the Loire estuary and the North Biscay continental shelf. About 130 samples were collected in winter 2001 and spring 2002 during Nutrigas and Gasprod campaigns (Programme PNEC-Golfe de Gascogne, RV Thalassa) and metal measurements were conducted on board by Potentiometric Stripping Analysis. In the Loire estuary, levels of dissolved lead ranged from 0.15 to 0.24 nM and from 0.04 to 0.26 nM in winter and spring, respectively. Compared to the concentrations reported in 1987 and 1990 (0.4–1.7 nM; Boutier, B., Chiffoleau, J.F., Auger, D., Truquet, I., 1993. Influence of the Loire river on dissolved lead and cadmium concentrations in coastal waters of Brittany. Estuar. Coast. Shelf S., 36:133–143, Estuarine, Coastal and Shelf Science 36, 133–143) our study indicated much lower values. The fall in concentration in the estuary could be attributed to the stopping of activity of Octel, a big manufacturer of tetra alkyl lead. Discharge in dissolved metal to the continental shelf by the Loire river was assessed as 7.5 and 1.9 kg day^− 1 for winter and spring, respectively. On the continental shelf, levels of dissolved lead varied within 0.06 and 0.27 nM in winter (0.15 ± 0.06 nM, sd = 1.96, n = 49), whereas concentrations measured in spring were in the range 0.06–0.17 nM (0.09 ± 0.03 nM, sd = 1.96, n = 60). This difference in metal concentration was related to the amounts of rainfall that have fallen over the continental shelf: estimations of inputs by this way (74 and 32 kg day^− 1 in winter and spring, respectively) appeared to be significantly higher than inputs from the Loire river (7.5 and 1.9 kg day^− 1 in winter and spring, respectively). The distributions of dissolved metal in the surface waters highlighted the role of suspended particular matter (SPM) for a rapid “trapping” of lead near the mouth of the estuary. The vertical distributions showed, in the stratified area, a biological transfer of lead between winter and spring from surface waters to the halocline.     

Microbial food web responses to light and nutrients beneath the coastal Arctic Ocean sea ice during the winter–spring transition

We measured the abundance and biomass of phototrophic and heterotrophic microbes in the upper mixed layer of the water column in ice-covered Franklin Bay, Beaufort Sea, Canada, from December 2003 to May 2004, and evaluated the influence of light and nutrients on these communities by way of a shipboard enrichment experiment. Bacterial cell concentrations showed no consistent trends throughout the sampling period, averaging (± SD) 2.4 (0.9) × 10⁸ cells L^− 1; integrated bacterial biomass for the upper mixed layer ranged from 1.33 mg C m^− 3 to 3.60 mg C m^− 3. Small cells numerically dominated the heterotrophic protist community in both winter and spring, but in terms of biomass, protists with a diameter > 10 µm generally dominated the standing stocks. Heterotrophic protist biomass integrated over the upper mixed layer ranged from 1.23 mg C m^− 3 to 6.56 mg C m^− 3. Phytoplankton biomass was low and variable, but persisted during the winter period. The standing stock of pigment-containing protists ranged from a minimum value of 0.38 mg C m^− 3 in winter to a maximal value of 6.09 mg C m^− 3 in spring and the most abundant taxa were Micromonas-like cells. These picoprasinophytes began to increase under the ice in February and their population size was positively correlated with surface irradiance. Despite the continuing presence of sea ice, phytoplankton biomass rose by more than an order of magnitude in the upper mixed layer by May. The shipboard experiment in April showed that this phototrophic increase in the community was not responsive to pulsed nutrient enrichment, with all treatments showing a strong growth response to improved irradiance conditions. Molecular (DGGE) and microscopic analyses indicated that most components of the eukaryotic community responded positively to the light treatment. These results show the persistence of a phototrophic inoculum throughout winter darkness, and the strong seasonal response by arctic microbial food webs to sub-ice irradiance in early spring.     

Variations of the abundance and nucleic acid content of heterotrophic bacteria in Beaufort Shelf waters during winter and spring

Depth profiles of heterotrophic bacteria abundance were measured weekly over a 6-month period from December to May in Franklin Bay, a 230 m-deep coastal Arctic Ocean site of the southeastern Beaufort Sea. Total bacteria, low nucleic acid (LNA) and high nucleic acid (HNA) bacteria abundances were measured using flow cytometry after SYBR Green I staining. The HNA bacteria abundance in surface waters started to increase 5–6 weeks after phytoplankton growth resumed in spring, increasing from 1 × 10⁵ to 3 × 10⁵ cells mL^− 1 over an 8-week period, with a net growth rate of 0.018 d^− 1. LNA bacteria response was delayed by more than two months relative to the beginning of the phytoplankton biomass accumulation and had a lower net growth rate of 0.013 d^− 1. The marked increase in bacterial abundance occurred before any significant increase in organic matter input from river discharge (as indicated by the unchanged surface water salinity and DOC concentrations), and in the absence of water temperature increase. The abundance of bacteria below the halocline was relatively high until January (up to 5 × 10⁵ cells mL^− 1) but then decreased to values close to 2 × 10⁵ cells mL^− 1. The three-fold bacterial abundance increase observed in surface waters in spring was mostly due to HNA bacteria, supporting the idea that these cells are the most active.