Estimates of Southern Ocean primary production—constraints from predator carbon demand and nutrient drawdown

In view of the wide range of estimates for the total primary production for the Southern Ocean south of the Subantarctic Front—current estimates range from 1.2 to 3.5 Gtonne C year⁻¹—we have examined two indirect methods for assessing primary production. First, we have estimated the primary production needed to sustain the carbon requirements of the endotherm top predators in the ecosystem. Estimation of the carbon requirements for crabeater seals of about 7 Mtonne C year⁻¹ is extrapolated to a value for all endotherm predators of 15–30 Mtonne C year⁻¹. Current data indicate that 70–80% of the diet of this suite of predators is zooplankton (predominantly the euphausiid krill), making for highly efficient transfer from primary production to top predators. Our best estimate of Southern Ocean primary production by this method is of the order of 1.7 Gtonne C year⁻¹, or an averaged areal primary production of about 30–40 g C m⁻² year⁻¹. Our second approach is to estimate primary production from the drawdown of inorganic nutrients, based on the limited suite of studies from which an annual nutrient deficit can be calculated. Again, this indicates annual primary production of the order of 1.5 Gtonne. Although both methods have inherent uncertainties, taken together they provide a relatively robust constraint on annual primary production. For both methods to underestimate primary production by the 1–1.5 Gtonne C implied by the higher current estimates, carbon export from the Southern Ocean pelagic ecosystem would need to be much higher than is normally found in other oceans.     

Siliceous plankton dominate primary and new productivity during the onset of El Niño conditions in the Santa Barbara Basin, California

The potential for carbon export and the role of siliceous plankton in the cycling of C and N was assessed in natural plankton assemblages in the Santa Barbara Basin, California, by examining uptake rates of inorganic carbon, nitrate and silicic acid. In April–August 1997, the concentrations of chlorophyll a, particulate organic carbon, particulate organic nitrogen and biogenic silica were measured twice monthly, and results revealed the occurrence of at least three blooms, the largest in June. Particulate elemental ratios of C, N and Si were similar to ratios of nutrient-replete diatoms, suggesting that they dominated this bloom. Mean integrated rates of carbon, nitrate and silicon uptake during the 4-month study period are similar to other productive coastal and upwelling regions (103, 8.3 and 13 mmol m⁻² day⁻¹, respectively). New production rates were twice as high as previously reported in this region and indicate that high rates of new production along eastern boundary currents are not confined to the major coastal upwelling regions. C/NO₃⁻, Si/NO₃⁻ and Si/C uptake ratios varied widely, and mean integrated ratios were 14±5.4, 1.6±1.0 and 0.12±0.07 (S.D.), respectively. That mean C/NO₃⁻ uptake ratio corresponds to an f-ratio of about 0.5 indicating a large potential for particulate export. Based on the average Si/NO₃⁻ and Si/C uptake ratios, diatoms could perform all of the primary production and nitrate uptake that occurred during the study; these rates also suggest that export is controlled by diatoms in this system. The mean Si/C biomass ratio was lower than the mean Si/C uptake ratio, consistent with the preferential export of Si relative to C observed in sediment traps in the basin. The study took place during a period of surface-water warming, with nitrate and silicic acid concentrations decreasing throughout the onset of the 1997–1998 El Niño conditions. Although diatoms contributed less to particulate biomass during the low nutrient conditions, high f-ratios (0.33–0.66) were maintained.     

Dynamics of inorganic nutrient species in the Bohai seawaters

Within the frame of a Sino-German Joint Research Program, two cruises of “R/V Dong Fang Hong 2” were carried out in September–October 1998 and April–May 1999, respectively, to understand the dynamics of nutrients in the Bohai. Nutrient species (NO₃⁻, NO₂⁻, NH₄⁺, PO₄³⁻ and SiO₃²⁻) are determined colorimetrically on board for five anchor and 30 grid stations. In situ incubation experiments are performed to determine planktonic nutrient uptake and benthic exchange flux. Nutrient concentrations display short-term variability and seasonal change in the Bohai, with higher levels in shallow coastal waters than in the Central Bohai. The influence of riverine discharge on nutrient levels can be seen from salinity isopleths, nutrient distribution and species ratios. Near-bottom (nb) waters have similar nutrient concentrations as to the surface waters in the Central Bohai, whereas stratification takes place in the Bohai Strait and North Yellow Sea. In situ incubation experiments provide evidence that the uptake ratio (i.e. N, P) by phytoplankton is proportional to the ratios among nutrient species in ambient waters. Based on the data of this study and previously publications, a preliminary estimate of nutrient budgets via riverine input and atmospheric deposition is established. The results indicate that atmospheric deposition gains importance over rivers in delivering nutrients into the Bohai and sustain the new production, following recent decrease in riverine inflow caused by drought periods in North China and damming practices. A historical review of nutrient data indicates that concentrations of nitrogen increase and phosphorus and silica decrease in the Central Bohai over last 40 years. This potentially has an important influence on the health of ecosystem in Bohai (e.g. food web and community structure), though further study is needed to examine the scenario in more detail.     

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.     

Carbon and nutrient dynamics in a hypertrophic lagoon in southwestern Taiwan

Dissolved and particulate phases of carbon (DIC, DOC, POC) and nutrients (DIN, DIP, DSi, DON, DOP, PN) were investigated bimonthly from August 1999 to August 2000 to study biogeochemical dynamics of carbon and nutrients in Tapong Bay, a small semi-enclosed and hypertrophic lagoon in southwestern Taiwan. The lagoon has only a tidal inlet for exchanging water between Tapong Bay and Taiwan Strait, which may result in low water exchange rates and various oxygen-deficient conditions in bottom water of the inner bay during warm seasons. The water exchange time of Tapong Bay ranges from 7 days (summer) to 13 days (winter) with a mean of 10 days. Nutrient dynamics were largely ascribed to allochthonous inputs, biological and exported removals in the lagoon. Diffusion fluxes from sediments to overlying water accounted for only about 7.6% of annual DIN inputs and 1.0% of annual DIP inputs. High primary productivity (89 mol C m⁻² year⁻¹) supported by abundant nutrients primarily drove the lagoon into a hypertrophic condition as particulate organic matter was derived mainly from biological production. Excess of DIP appeared to occur throughout the study period in the lagoon. Temperature, solar radiation and turbidity, rather than nutrients, perhaps controlled seasonal variations of primary productivity. The net ecosystem production (NEP) derived from daily changes of DOC and POC inventories was about 6.3 mmol C m⁻² day⁻¹ that was close to 6.7 mmol C m⁻² day⁻¹ simulated from the biogeochemical modeling. Therefore, the net ecosystem production (NEP) rate of organic carbon estimated from the biogeochemical model was reliable, and the NEP was temporally variable with an annual mean of 5.8 mol C m⁻² year⁻¹, implying that Tapong Bay was an autotrophic system. Although calcification proceeded pronouncedly in warm seasons, an invasion of CO₂ was significant in this system. In terms of nitrogen budget, the annual nitrogen fixation exceeded the annual denitrification with a magnitude of 1.30 mol N m⁻² year⁻¹, which may be supported by the abundance of nitrogen fixation microplanktons in the lagoon.     

Pb and Pu in the Northeast Water Polynya, Greenland: particle dynamics and sediment mixing rates

Distributions of the radionuclides ²¹⁰Pb and ^239,240Pu in sediment cores from the Northeast Water Polynya, Greenland, showed that these nuclides reached depths of 5–15 cm by particle mixing and sediment accumulation. End-member average values of the particle mixing coefficient and sediment accumulation rate were 0.13 cm² y⁻¹ and 0.06 cm y⁻¹, obtained from the ²¹⁰Pb profiles by assuming that each process is dominant relative to the other. Both ²¹⁰Pb and ^239,240Pu were measured on four cores; using the Pu data to constrain mixing rates produced corrected sediment accumulation rates that were 20–80% of the values calculated by neglecting mixing. Organic carbon burial in the polynya sediments was ≤0.4 mmol m⁻² d⁻¹, based on measured POC values at depth in the sediments and sediment accumulation rates corrected for mixing. This value is about 1% of the independently measured POC flux leaving the euphotic zone and compares with benthic carbon remineralization rates of 7% calculated by others from O₂ uptake in the sediments.The inventories of excess ²¹⁰Pb in the sediments ranged from 6 to 28 dpm cm⁻². Relative to the atmospheric input of ²¹⁰Pb and in situ production from decay of ²²⁶Ra, approximately 5 dpm cm⁻² of ²¹⁰Pb was being removed from the water column. The difference between the removal from the water column and sediment inventories suggests a net import of ²¹⁰Pb to the polynya. This may occur by input of dissolved ²¹⁰Pb from offshore waters or by input of ²¹⁰Pb carried by sea ice. Particulate matter in land-derived fast ice adjacent to the polynya contained 330 ± 14 dpm of excess ²¹⁰Pb g⁻¹. If particles transported in sea ice are comparable to those extracted from fast ice, then sea ice transport into the polynya followed by melting may be an important source of excess ²¹⁰Pb to the area. Fast ice also may contribute ²¹⁰Pb if portions break off and melt within the polynya, as occurred in 1993.     

Microphytobenthos biomass at Kerguelen's Land (Subantarctic Indian Ocean): repartition and variability during austral summers

Microphytobenthos biomass has been measured at several coastal sites on the SE of the main island of the Kerguelen Archipelago (Indian Ocean), during several austral summers (1985–1992), using a conventional fluorometric method. Heterogeneity tests, conducted on two different intertidal sites (Port-Aux-Français, PAF; and Port-Raymond, PRA), showed low standard deviations, whereas the mean concentrations were highly different. Pigment concentrations showed a high variability related to the characteristics of the sediments: from low biomass in coarse intertidal sand, submitted to intense scouring (0.32±0.31 μg Chl a g⁻¹ dw, 0.29±0.14 μg Phaeo g⁻¹ dw) to high biomass in intertidal muddy sand in sheltered areas, particularly along estuaries (54 μg Chl a, 15 μg Phaeo g⁻¹ dw at Korrigan). The subtidal muddy sediments under a Macrocystis pyrifera (Linné) and Durvillaea antarctica (Chamisso in Choris) belt exhibited high concentrations in phaeopigment (Phaeo) (up to 136±83 μg g⁻¹ dw; PRA), while the concentration of chlorophyll a (Chl a) was relatively low. The dense macroalgal canopy supports an important epiphytic diatom biomass (mainly the genera Cocconeis Ehrenberg and Grammatophora Ehrenberg), which is sedimenting after degradation and is in part responsible of the high levels of Phaeo in all sediments. Macroalgal debris were observed, but diatom frustules were dominant in most surficial subtidal sediments. A circatidal mud, in the Morbihan Bay, made of a sponge spicule mat (50 m deep; 4.96 μg Chl a g⁻¹ dw), showed a very low Chl a/Phaeo ratio (0.1), while it reached up to 6 in intertidal sand. Surprisingly, a penguin rookery beach, at the east side of Courbet Peninsula, was characterized by a very low biomass (0.07±0.04 μg Chl a g⁻¹ dw), while it was nutrient enriched, particularly with nitrates.In comparison with the data at the similar latitude, but in temperate regions from the Northern Hemisphere, the microphytobenthos biomass, recorded at Kerguelen's Land, exhibited relatively high pigment concentrations, particularly the Phaeo, and supported a dense and diversified subtidal macrofauna composed of polychaetes (particularly Thelepus extensus Hutchings and Glasby), sea urchins, mytillids and gammarids. The exuberant macroalgal canopy, coastal indentations and low tidal amplitude must be in part responsible of these large benthic primary and secondary biomasses.     

Modelling the silica pump in the Permanently Open Ocean Zone of the Southern Ocean

A coupled 1D physical–biogeochemical model has been built to simulate the cycles of silicon and of nitrogen in the Indian sector of the Permanently Open Ocean Zone of the Southern Ocean. Based on a simplified trophic network, that includes two size classes of phytoplankton and of zooplankton, and a microbial loop, it has been calibrated by reference to surface physical, chemical and biological data sets collected at the KERFIX time-series station (50°40′S–68°25′E). The model correctly reproduces the high nutrient low chlorophyll features typical of the studied area. In a region where the spring–summer mixed layer depth is usually deeper than 60 m, the maximum of chlorophyll never exceeds 1.5 mg m⁻³, and the annual primary production is only 68 g C m⁻² year⁻¹. In the surface layer nitrate is never exhausted (range 27–23.5 mmoles m⁻³) while silicic acid shows strong seasonal variations (range 5–20 mmoles m⁻³). On an annual basis 71% of the primary production sustained by nanophytoplankton is grazed by microzooplankton. Compared to North Atlantic, siliceous microphytoplankton is mainly prevented from blooming because of an unfavourable spring–summer light-mixing regime. Silicic acid limitation (high half saturation constant for Si uptake: 8 mmoles m⁻³) also plays a major role on diatom growth. Mesozooplankton grazing pressure excerpts its influence especially in late spring. The model illustrates the efficiency of the silica pump in the Southern Ocean: up to 63% of the biogenic silica that has been synthetized in the photic layer is exported towards the deep ocean, while only 11% of the particulate organic nitrogen escapes recycling in the surface layer.     

Determination of anthropogenic CO2 in the North Atlantic Ocean using water mass ages and CO2 equilibrium chemistry

A new method to calculate the anthropogenic CO₂ (ΔDIC_ant) within the water column of the North Atlantic Ocean is presented. The method exploits the equilibrium chemistry of the carbonate system with reference to temperature, salinity and the partial pressure of atmospheric CO₂ (pCO_2,atm). ΔDIC_ant is calculated with reference to the ventilation ages of water masses derived from tracer data and to the time history of pCO_2,atm. The method is applied to data recorded during the WOCE program on the WHP A1/E transect in the North Atlantic Ocean, where we characterise six key water masses by their relationships of dissolved inorganic carbon (DIC) and apparent oxygen utilisation (AOU). The error in determining ΔDIC_ant is reduced significantly by minimising the number of values referred to, especially by avoiding any use of remineralisation ratios of particulate organic matter. The distribution of ΔDIC_ant shows highest values of up to 45 μmol kg⁻¹ in the surface waters falling to 28–33 μmol kg⁻¹ in the Irminger Sea west of the Mid-Atlantic Ridge. The eastern basin is imprinted by older water masses revealing decreasing values down to 10 μmol kg⁻¹ ΔDIC_ant in the Antarctic Bottom Water. These findings indicate the penetration of the whole water column of the North Atlantic Ocean by anthropogenic CO₂.     

Nutrient cycling and primary production in the marine systems of the Arctic and Antarctic

Primary production events in both the Arctic and the Antarctic are highly localized. Carbon-14 incubations that did not account for this caused antarctic primary production estimates to be revised too far downwards from the historic view of high productivity. The primary production regime in the Arctic is even more heterogeneous than in the Antarctic. Arctic primary production rates are in the process of being revised upwards because of a better spatial and temporal distribution of incubation experiments and a re-awakening of interest in estimating new production from the distribution of chemical variables. Similarly, recent examination of temporal changes in nitrate concentrations and recognition of the importance of ice-edge blooms has caused antarctic primary productivity to be revised upwards. In both the Arctic and the Antarctic, the ratio of “new” to total primary production is high, and neglect of this fact can lead to an underestimation of the potential that these regions have for influencing global cycles of bioactive chemicals. Some recent data on temporal changes in nitrate from Fram Strait emphasize the poor state of our knowledge by suggesting an unexpectedly high “new” production rate of 1 g C m⁻² d⁻¹ for a 35 day experiment that encountered an early Phaeocystis bloom. Chemical distributions suggest that new production over the shelf seas that border the Polar Basin is about 50 g Cm⁻² yr⁻¹.The shelves in the Arctic Ocean's marginal and adjacent seas comprise 25% of the total global continental shelf. These extensive shallow regions have much higher rates of primary production than the Polar Basin and may be globally significant sites of denitrification. Globally significant silica deposition could occur on these shelves or on the adjacent slopes.Because of the differences in geomorphology and stratification, global warming is likely to increase primary production in the Arctic and will probably decrease antarctic primary production.In addition to sharing high ratios of “new” to total primary production, high ammonium concentrations occur in the Arctic and Antarctic. It is possible that these accumulations arise from a strong repression of nitrification at low temperatures.     

The trophic role of the tunicate Salpa thompsoni in the Antarctic marine ecosystem

During a repeat grid survey and drogue study carried out in austral summer 1994/95, the abundance and feeding activity of salps were estimated in the Lazarev Sea region from net tows and in situ measurements of gut fluorescence. Throughout the survey area, Salpa thompsoni accounted for >95% of the total salp stock while Ihlea racovitzai was consistently represented in very low abundances. Maximum densities of S. thompsoni, with ≈4000 ind. 1000 m⁻³, were recorded in the Marginal Ice Zone (MIZ) in December when chlorophyll-a concentrations were well below 1 mg m⁻³. A dramatic decrease in salp stock was observed at the beginning of January, when S. thompsoni virtually disappeared from the most productive area of the MIZ where chlorophyll-a concentrations had by then reached bloom levels of 1.5–3 mg (Chl-a) m⁻³. In situ grazing measurements showed that throughout the cruise S. thompsoni exhibited the highest ingestion rates per individual of any of the most abundant components of the grazing pelagic community, with maxima of ≈160 μg (pigm) ind. ⁻¹ d⁻¹. These feeding rates are 3 to 5 times higher than those previously obtained using in vitro incubations. The total daily consumption of the population of S. thompsoni varied from 0.3 to 108% of daily primary production. We suggest that competitive removal of food by S. thompsoni, rather than direct predation, is responsible for the low krill abundances generally associated with salp swarms.     

Nutrient status of the Northeast Water Polynya

The nutrient distribution in the Northeast Water Polynya (NEW) was investigated intensively between the end of May and the beginning of August 1993 during the R/V Polarstern cruise ARK IX. The major characteristics were low initial nitrate concentrations (ca. 4 μM) in the surface mixed layer of the East Greenland Shelf Water, accompanied by high silicate values (ca. 10–14 μM). These concentrations were not reduced by phytoplankton growth. Silicate was rather homogeneously distributed in the entire water column, whereas nitrate increased continuously with depth to about 13 μM. Phosphate concentrations were about 1.1 μM and had a similar distribution to that of silicate. During the course of the summer, nutrients became depleted, and nitrate was exhausted in large parts of the NEW. Silicate was reduced to values of less than 2 μM at some stations which implies that diatom growth continued despite nitrate depletion, ammonium serving as a nitrogen source. The polynya is fertilised by water with the initial nutrient concentrations downstream of the Norske Øer Ice Shelf. This process continuously supplies nutrients to the surface throughout the year and these are transported northward by the anticyclonic surface circulation following the topography of the trough system. The northern boundary of this tongue of relatively nutrient-rich water is controlled by the uptake of nutrients by phytoplankton in summer. Its extemsion is variable due to interactions between biological processes, circulation and ice cover. In the Ob Bank region the nutrient distribution can be altered by the inflow of Polar Water from the north when strong northerly winds prevail as happened during the first part of the study.     

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.     

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.     

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).     

Appendicularian ecophysiology I: Food concentration dependent clearance rate, assimilation efficiency, growth and reproduction of Oikopleura dioica

Three aspects of the appendicularian O. dioica's ecophysiology were measured here: 1) morphological parameters over a wide range of appendicularian sizes, including mature animals in order to document the morphological characteristics inducing reproduction; 2) clearance rate and assimilation efficiency using feeding incubations with different algal concentrations and 3) the effect of food concentration on growth, mortality and reproduction.The relationship between the body carbon weight and the clearance rate follows a power function, with an exponent of 0.91 (± 0.07). The rate of particles retention increases with the food concentration following a Michaelis–Menten relationship (half-saturation constant = 151 ± 22 µg C l^− 1, maximum clearance rate = 12 ± 1 µg C µg C^− 1 d^− 1). The carbon assimilation efficiency decreases with the increasing food concentration. As a result, appendicularian growth which is limited in concentrations lower than 50 µg C l^− 1 is saturated above 100 µg C l^− 1.In immature animals the gonad represents less than 30% of the body volume whereas in mature individuals, its volume varies between 50% and 87% (mean 63%) suggesting that gonad/total volume ratio can be used as indicator of the maturation stages. The gonad weight in mature animals represents 70.3 (± 4.6)% of the total body carbon weight. Two major maturity stages can explain the changes in energy allocation: i) the somatic growth, when less energy is invested in gonad growth when compared to the rest of the body and ii) the maturation phase where most of the assimilated matter is invested in gonad maturation. This process is rapid, lasting only few hours. For this reason we measured completely mature organisms that are generally not measured during the experimental work with appendicularians. In food-limited conditions, the gonad maturation process starts with smaller individuals and ends with smaller reproductive animals having the same gonad to total volume ratio than in unlimited food conditions. The results obtained in this study were used to model the life cycle of O. dioica (see Lombard, F., Sciandra, A. and Gorsky, G., 2009-this volume. Appendicularian ecophysiology. II. Modeling nutrition, metabolism, growth and reproduction of the appendicularian Oikopleura dioica.).     

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.     

Turbulence in coastal fronts near the mouths of Block Island and Long Island Sounds

Measurements of turbulence were performed in four frontal locations near the mouths of Block Island Sound (BIS) and Long Island Sound (LIS). These measurements extend from the offshore front associated with BIS and Mid-Atlantic Bight Shelf water, to the onshore fronts near the Montauk Point (MK) headland, and the Connecticut River plume front. The latter feature is closely associated with the major fresh water input to LIS. Turbulent kinetic energy (TKE) dissipation rate, ε, was obtained using shear probes mounted on an autonomous underwater vehicle. Offshore, the BIS estuarine outflow front showed, during spring season and ebb tide, maximum TKE dissipation rate, ε, estimates of order 10^− 5 W/kg, with background values of order 10^− 6 to 10^− 9 W/kg. Edwards et al. [Edwards, C.A., Fake, T.A., and Bogden, P.S., 2004a. Spring–summer frontogenesis at the mouth of Block Island Sound: 1. A numerical investigation into tidal and buoyancy-forced motion. Journal of Geophysical Research 109 (C12021), doi:10.1029/2003JC002132.] model this front as the boundary of a tidally driven, baroclinically adjusted BIS flow around the MK headland eddy. At the entrance to BIS, near MK, two additional fronts are observed, one of which was over sand waves. For the headland site front east of MK, without sand waves, during ebb tide, ε estimates of 10^− 5 to 10^− 6 W/kg were observed. The model shows that this front is at the northern end of an anti-cyclonic headland eddy, and within a region of strong tidal mixing. For the headland site front further northeast over sand waves, maximum ε estimates were of order 10^− 4 W/kg within a background of order 10^− 7–10^− 6 W/kg. From the model, this front is at the northeastern edge of the anti-cyclonic headland eddy and within the tidal mixing zone. For the Connecticut River plume front, a surface trapped plume, during ebb tide, maximum ε estimates of 10^− 5 W/kg were obtained, within a background of 10^− 6 to 10^− 8 W/kg. Of all four fronts, the river plume front has the largest finescale mean-square shear, S² ~ 0.15 s^− 2. All of the frontal locations had local values of the buoyancy Reynolds number indicating strong isotropic turbulence at the dissipation scales. Local values of the Froude number indicated shear instability in all of the fronts.     

Nitrogenous discharges to the eastern Gulf of Finland, the Baltic Sea: Elemental flows, stable isotope signatures, and their estuarine modification

We studied the nutrient input to the Gulf of Finland via River Neva, the largest river discharging freshwater to the Baltic Sea, and characterised the isotopic signatures (¹⁵N, ¹⁸O, ¹³C) in dissolved and particulate substances (NO₃⁻, PON, POC, DIC) in the River Neva over two seasonal cycles, as well as in samples from St. Petersburg wastewater treatment plants (NO₃⁻, NH₄⁺, PON, POC). These riverine and municipal discharges account for 40% of terrestrial inorganic N loading to the Gulf of Finland, representing annually 7% of the total nitrogen pool in the water mass of the whole Gulf. To describe and evaluate the modification of these isotopic signals along a Gulf of Finland transect towards the Baltic Proper, two cruises were arranged, one in late spring after the annual maximum in River Neva runoff, and one in autumn, in the late phase of the annual growth season.River Neva nitrate signatures of ¹⁵N and ¹⁸O indicated major agricultural fertilizer origin of nitrogen, and the isotopic composition was clearly lighter (δ¹⁵N-NO₃⁻ mean of 2.4‰ air) than previously measured from more southern rivers discharging into the Baltic Sea. Because of the light composition of the River Neva N source, close to the ¹⁵N signatures of the open Gulf, as well as of the efficient depletion of the inorganic load already in the innermost estuary, straightforward end-member tracer analysis of the transport of N in the basin is problematic. St. Petersburg wastewater ammonium showed, however, high δ¹⁵N values (ca. 13‰), which gives a first estimate of 5.8‰ for δ¹⁵N of the easternmost estuarine total inorganic N source. The available sediment data from the basin (δ¹⁵N 6 to 8‰) somewhat exceeds the average source signature. This emphasizes the significance of biological transformation processes, most importantly assimilation of inorganic nitrogen, food web interactions and denitrification, which all involve isotopic fractionation, for the mass balance models describing the dynamics of the sources and sinks of the N cycle of the basin.     

Bacterial production and microbial food web structure in a large arctic river and the coastal Arctic Ocean

Globally significant quantities of organic carbon are stored in northern permafrost soils, but little is known about how this carbon is processed by microbial communities once it enters rivers and is transported to the coastal Arctic Ocean. As part of the Arctic River-Delta Experiment (ARDEX), we measured environmental and microbiological variables along a 300 km transect in the Mackenzie River and coastal Beaufort Sea, in July–August 2004. Surface bacterial concentrations averaged 6.7 × 10⁵ cells mL^− 1 with no significant differences between sampling zones. Picocyanobacteria were abundant in the river, and mostly observed as cell colonies. Their concentrations in the surface waters decreased across the salinity gradient, dropping from 51,000 (river) to 30 (sea) cells mL^− 1. There were accompanying shifts in protist community structure, from diatoms, cryptophytes, heterotrophic protists and chrysophytes in the river, to dinoflagellates, prymnesiophytes, chrysophytes, prasinophytes, diatoms and heterotrophic protists in the Beaufort Sea.Size-fractionated bacterial production, as measured by ³H–leucine uptake, varied from 76 to 416 ng C L^− 1 h^− 1. The contribution of particle-attached bacteria (> 3 µm fraction) to total bacterial production decreased from > 90% at the Mackenzie River stations to < 20% at an offshore marine site, and the relative importance of this particle-based fraction was inversely correlated with salinity and positively correlated with particulate organic carbon concentrations. Glucose enrichment experiments indicated that bacterial metabolism was carbon limited in the Mackenzie River but not in the coastal ocean. Prior exposure of water samples to full sunlight increased the biolability of dissolved organic carbon (DOC) in the Mackenzie River but decreased it in the Beaufort Sea.Estimated depth-integrated bacterial respiration rates in the Mackenzie River were higher than depth-integrated primary production rates, while at the marine stations bacterial respiration rates were near or below the integrated primary production rates. Consistent with these results, P_CO2 measurements showed surface water supersaturation in the river (mean of 146% of air equilibrium values) and subsaturation or near-saturation in the coastal sea. These results show a well-developed microbial food web in the Mackenzie River system that will likely convert tundra carbon to atmospheric CO₂ at increasing rates as the arctic climate continues to warm.