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.     

Spatial and temporal variability of size-fractionated biomass and primary production in the Ross Sea (Antarctica) during austral spring and summer

The results of a study on the spatial and temporal dynamics of size-fractionated biomass and production of phytoplankton in the Ross Sea during the austral spring and summer are reported. The spring cruise took place in the offshore Ross Sea from 14 November to 14 December 1994. Sampling was carried out on a transect of 27 stations distributed from 76.5 to 72.0°S along 175°E, and covered the three main Antarctic environments of the polynya open waters, the marginal ice zone and the pack ice area. Three subsystems were identified. The subsystem of the polynya was characterised by the predominance of the micro- and nano-planktonic fractions, chlorophyll (Chl a) concentrations from 69.6 to 164.7 mg m⁻² and production rates from 0.68 to 1.14 g C m⁻² day⁻¹. The second subsystem, the marginal ice zone, showed a relative increase of the micro-planktonic fraction, high biomass levels (from 99.64 to 220 mg Chl m⁻²) and production rates from 0.99 to 2.7 g C m⁻² day⁻¹. The subsystem of the pack ice area had a phytoplankton community dominated by the pico-planktonic fraction and showed low biomasses (from 19.4 to 37.7 mg Chl m⁻²) and production rates (0.28 to 0.60 g C m⁻² day⁻¹). Selective grazing by krill is considered an important factor in determining the size structure of the phytoplankton communities. The summer study consisted of a time series carried out in inshore waters of Terra Nova Bay from 12 January to 8 February 1990. In a well stabilised water column and with high levels of PAR always available, the primary production rates of a community dominated by micro-plankton varied from 0.52 to 1.2 g C m⁻² day⁻¹ (average 0.84). A high P/B ratio, up to 3, and a remarkably elevated mean phaeopigment (Phaeo)/Chl a ratio of 2.4 indicated an active removal of biomass by grazing, confirmed by the presence of faecal pellets in quantities reaching 6000 m⁻³ in the upper 50 m. The peculiarities of the inshore versus offshore environments in terms of community size structure, production processes and their implications as regards the food web are discussed.     

Sediment community biomass and respiration in the Northeast water polynya, Greenland: a numerical simulation of benthic lander and spade core data

Sediment community metabolism (oxygen demand) was measured in the Northeast Water (NEW) polynya off Greenland employing two methods: in situ benthic chambers deployed with a benthic (GOMEX) lander and shipboard laboratory Batch Micro-Incubation Chambers (BMICs) utilizing ‘cores’ recovered from USNEL box cores. The mean benthic respiration rate measured with the lander was 0.057 mM O₂ m⁻² h⁻¹ (n = 5); whereas the mean measured with the BMICs was 0.11 mM O₂ m⁻² h⁻¹ (n = 21; p < 0.01 that the means were the same). In terms of carbon fluxes (14 and 27 mg C m⁻² d⁻¹), these respiration rates represent ca. 5–15% of the average net primary production measured in the euphotic zone in 1992. The biomass of the bacteria, meiofauna and macrofauna were measured at each location to quantify the relationship between total community respiration and total community biomass (mean 1.42 g C m⁻²). Average carbon residence time in the biota, calculated by dividing the biomass by the respiration, was on the order of 50–100 days, which is comparable to relatively oligotrophic continental margins at temperate latitudes.The biomass and respiration data for the aerobic heterotrophic bacteria, the infaunal invertebrates (meiofauna and macrofauna), and the epifaunal megabenthos (two species of brittle stars) are summarized in a ‘steady-state’ solution of a sediment food chain model, in terms of carbon. This carbon budget illustrates the relative importance of the sediment-dwelling invertebrates in the benthic subsystem, compared to the bacteria and the epibenthos, during the summer open-water period in mud-lined troughs at depths of about 300 m. The input needed to drive heterotrophic respiratory processes was within the range of the input of organic matter recorded in moored, time-sequencing sediment traps.A time-dependent numerical simulation of the model was run to investigate the potential responses of the three size groups of benthos to abrupt seasonal pulses of particulate organic matter. The model suggests that there is a time lag in the increase in bottom community biomass and respiration following the POC pulse, and provides hypothetical estimates for the potential carbon storage in the summer (open water), followed by catabolic losses during each ensuing winter (ice covered).This sequence of storage and respiration may contribute to the process of seasonal CO₂ ‘rectification’ (sensu Yager et al., 1995) in some Arctic ecosystems.     

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.     

Size-fractionated phytoplankton biomass and species composition in the Indian sector of the Southern Ocean during austral summer

During the late austral summer of 1994, Antarctic waters were characterized by low phytoplankton biomass. Along the 62°E meridian transect, between 49°S and 67°S, chlorophyll (Chl.) a concentration in the upper 150 m was on average 0.2 mg m⁻³. However, in the Seasonal Ice Zone (SIZ) chlorophyll a concentrations were higher, with a characteristic deep chlorophyll maximum. The highest value (0.6 mg Chl. a m⁻³) was measured at the Antarctic Divergence, 64°S, corresponding to the depth of the temperature minimum (100 m). This deep biomass maximum decreased from South to North, disappeared in the Permanently Open Ocean Zone (POOZ) and reappeared with less vigour in the vicinity of the Polar Front Zone (PFZ). In the SIZ, the upper mixed layer was shallow, biomass was higher and the >10 μm fraction was predominant. In this zone the >10 μm, 2–10 μm and <2 μm size fractions represented on the average 46%, 25.1% and 28.9% of the total integrated Chl. a stock in the upper 100 m, respectively. The phytoplankton assemblage was diverse, mainly composed of large diatoms and dinoflagellate cells which contributed 42.7% and 33.1% of the autotrophic carbon biomass, respectively. Moving northwards, in parallel with the decrease in biomass, the biomass of autotrophic pico- and nanoflagellates (mainly Cryptophytes) increased steadily. In the POOZ, the picoplanktonic size fraction contributed 47.4% of the total integrated Chl. a stock. A phytoplankton community structure with low biomass and picoplankton-dominated assemblage in the POOZ contrasted with the relatively rich, diverse and diatom-dominated assemblage in the SIZ. These differences reflect the spatial and temporal variations prevailing in the Southern Ocean pelagic ecosystem.     

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.     

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.     

Estimation of copepod production in a flooded savanna of Venezuela

This is the first contribution to the copepod production in Venezuelan tropical savannas. Total abundance, biomass, production and mean P/B ratio of nauplii, copepodids and adults were determined in interdaily samples from a flooded, embanked savanna during February and March (end of dry season), and May (beginning of rainy season). Highest values of biomass and total production were recorded during dry season (61.5 mg m⁻³, 153.8 mg m⁻³ day⁻¹, respectively), compared to the rainy season (5.6 mg m⁻³, 45.9 mg m⁻³ day⁻¹). The last values are related to a low population density found during rainy season. Highest production were observed in copepodids at the end of the dry season. Significant differences of production between nauplii and copepodid stages, as well as between nauplii and adults, were also found.     

Influence of the Mackenzie River plume on the sinking export of particulate material on the shelf

We examined the influence of the Mackenzie River plume on sinking fluxes of particulate organic and inorganic material on the Mackenzie Shelf, Canadian Arctic. Short-term particle interceptor traps were deployed under the halocline at 3 stations across the shelf during fall 2002 and at 3 stations along the shelf edge during summer 2004. During the two sampling periods, the horizontal patterns in sinking fluxes of particulate organic carbon (POC) and chlorophyll a (chl a) paralleled those in chl a biomass within the plume. Highest sinking fluxes of particulate organic material occurred at stations strongly influenced by the river plume (maximum POC sinking fluxes at 25 m of 98 mg C m^− 2 d^− 1 and 197 mg C m^− 2 d^− 1 in 2002 and 2004, respectively). The biogeochemical composition of the sinking material varied seasonally with phytoplankton and fecal pellets contributing considerably to the sinking flux in summer, while amorphous detritus dominated in the fall. Also, the sinking phytoplankton assemblage showed a seasonal succession from a dominance of diatoms in summer to flagellates and dinoflagellates in the fall. The presence of the freshwater diatom Eunotia sp. in the sinking assemblage directly underneath the river plume indicates the contribution of a phytoplankton community carried by the plume to the sinking export of organic material. Yet, increasing chl a and BioSi sinking fluxes with depth indicated an export of phytoplankton from the water column below the river plume during summer and fall. Grazing activity, mostly by copepods, and to a lesser extent by appendicularians, appeared to occur in a well-defined stratum underneath the river plume, particularly during summer. These results show that the Mackenzie River influences the magnitude and composition of the sinking material on the shelf in summer and fall, but does not constitute the only source of material sinking to depth at stations influenced by the river plume.     

The coastal edge of the Northeast Water polynya in spring 1993

Multidisciplinary, marine ecological observations were conducted at the shallow water edge of the Northeast Water in June, 1993. Although variable in size and shape, a small polynya was constantly present at Eskimonaes, at the fast-ice edge of Ingolfsfjord. A shallow stratified layer developed at the water sufface at negative water and air temperatures—an effect of sea ice melting in cold water early in the season. Nutrients were recorded in considerable quantities, although by mid July NO₃ had become depleted. The chlorophyll and phytoplankton maxima at 8–12 m depth had peak values of 2 mg chl a m⁻³, typical for Arctic algal blooms. The phytoplankton included over 90 species and was dominated by the Fragillariopsis group. Zooplankton was poor in biomass and density, but over 23 taxa were found, with the copepods Oithona similis and Pseudocalanus acuspes being numerically dominant. Sedimentation was approximately 0.2 g dry weight m⁻² d⁻¹ and suspended matter concentrations ranged from 4 to 19 mg l⁻¹. The benthos was represented by hard bottom forms only, with a surprisingly dense cover of macrophytes. Juvenile sea urchins (Strongylocentrotus droebachiensis), brittle stars (Ophiocten sericeum) and amphipods were dominant. Higher trophic levels were represented by benthic feeders, such as eiders and walruses. The area observed was more similar to high Arctic fjord ecosystems than to the offshore central Northeast Water polynya.     

Seasonal carbon distribution of copepods in the eastern Weddell Sea, Antarctica

Copepods were sampled by a multiple opening-closing net in the eastern Weddell Sea during various seasons (late winter/early spring, summer, autumn). Total copepod biomass integrated over the upper 1000 m varied seasonally between 1.7 mg C m⁻³ in late winter/early spring and 3.7 mg C m⁻³ in autumn. After the dark season the copepods were rather evenly distributed vertically and highest biomass levels were found in the mid-water layers between about 200 m and 500 m. By contrast, especially in summer but also in autumn copepod biomass concentrated in the uppermost water layer. A total of 64 calanoid species were identified in the upper 1000 m with maximum species numbers in the deepest layer. The large calanoids Calanus propinquus, Calanoides acutus, Metridia gerlachei, Euchaeta antarctica and the small calanoid Microcalanus pygmaeus prevailed and accounted for 60–70% of total copepod biomass, while the small poecilostomatoid Oncaea and the cyclopoid Oithona species comprised about 20%. Hence, the distribution pattern of the entire copepod biomass is strongly influenced by the life cycles of a few dominant species.     

Role of filtering and biodeposition by Adamussium colbecki in circulation of organic matter in Terra Nova Bay (Ross Sea, Antarctica)

At Terra Nova Bay, the scallop Adamussium colbecki (Smith, 1902) characterises the soft and hard bottoms from 20 to 80 m depth, constituting large beds and reaching high values of density (50–60 individuals/m²) and biomass (120 g/m² DW soft tissues). To assess its role in the organic matter recycling in the coastal ecosystem, its filtering and biodeposition rates were evaluated in laboratory experiments during the austral summer 1993/94. Filtration rates, measured in a flow-through system, were calculated from the difference in particulate organic carbon (POC), nitrogen (PON) and chlorophyll-a (Chl-a) concentration in inflow and outflow water. Experiments were performed using natural sea water with POC, PON and Chl-a concentrations of about 450 μg/l, 90 μg/l and 2 μg/l, respectively. The biodeposition rate and the biochemical composition of the biodeposits were studied in order to detect how the organic matter is transformed through feeding activity of A. colbecki. At +1°C temperature, the average filtering rate was about 1 l h⁻¹ g⁻¹ (DW soft tissues) in specimens ranging in body mass from 2 to 3 g (DW soft tissues) and 6–7 cm long. The biodeposition rate in 3–8 cm long specimens, ranging from 0.4 to 5.7 g (DW soft tissues), was about 5.65 mg DW/g DW/day, leading to an estimate of Corg flux, through biodeposition by A. colbecki, of about 21 mg C m⁻² day⁻¹ at in situ conditions. Comparison between the biochemical composition of seston and biodeposits shows a decrease of the labile compounds, of the Chl-a/phaeopigments ratio in the biodeposits. The recorded C/N ratio decrease suggests a microbial colonisation in the biodeposits. This study suggests that Adamussium colbecki plays an important role in coupling the material fluxes from the water column to the sea bed, processing about 14% of total Carbon flux from the water column to the sediments, with an assimilation efficiency of 36%.     

Picophytoplankton and nanophytoplankton abundance and distribution in the southeastern Beaufort Sea (Mackenzie Shelf and Amundsen Gulf) during Fall 2002

The distribution of picophytoplankton (0.2–2 µm) and nanophytoplankton (2–20 µm) in the Beaufort Sea–Mackenzie Shelf and Amundsen Gulf regions during autumn, 2002 is examined relative to their ambient water mass properties (salinity, temperature and nutrients: nitrate + nitrite, phosphate, and silicate) and to the ratio of variable to maximum fluorescence, Fv/Fm. Total phytoplankton and cell abundances (< 20 µm) were mainly correlated with salinity. Significant differences in picophytoplankton cell numbers were found among waters near the mouth of the Mackenzie River, ice melt waters and the underlying halocline water masses of Pacific origin. Picophytoplankton was the most abundant phytoplankton fraction during the autumnal season, probably reflecting low nitrate concentrations (surface waters average ~ 0.65 µM). The ratio Fv/Fm averaged 0.44, indicating that cells were still physiologically active, even though their concentrations were low (max Chl a = 0.9 mg m^− 3). No significant differences in Fv/Fm were evident in the different water masses, indicating that rate limiting conditions for photosynthesis and growth were uniform across the whole system, which was in a pre-winter stage, and was probably already experiencing light limitation as a result of shortening day lengths.     

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.     

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.     

Large-scale physical controls on phytoplankton growth in the Irminger Sea, Part II: Model study of the physical and meteorological preconditioning

The upper water column in the Irminger Sea is characterized by cold fresh arctic and subarctic waters and warm saline North Atlantic waters. In this study the local physical and meteorological preconditioning of the phytoplankton development over an annual cycle in the upper water column in four physical zones of the Irminger Sea is investigated. Data from four cruises of the UK's Marine Productivity programme are combined with results from a coupled biological–physical nitrogen–phytoplankton–zooplankton–detritus model run using realistic forcing. The observations and model predictions are compared and analyzed to identify the key parameters and processes which determine the observed heterogeneity in biological production in the Irminger Sea. The simulations show differences in the onset of the bloom, in the time of the occurrence of the maximum phytoplankton biomass and in the length of the bloom between the zones. The longest phytoplankton bloom of 90 days duration was predicted for the East Greenland Current of Atlantic origin zone. In contrast, for the Central Irminger Sea zone a phytoplankton bloom with a start at the beginning of May and the shortest duration of only 70 days was simulated. The latest onset of the phytoplankton bloom in mid May and the latest occurrence of the maximum biomass (end of July) were predicted for the Northern Irminger Current zone. Here the bloom lasted for 80 days. In contrast the phytoplankton bloom in the Southern Irminger Current zone started at the same time as in Central Irminger Sea, but peaked end of June and lasted for 80 days. For all four zones relatively low daily (0.3–0.5 g C m^− 2d^− 1) and annual primary production was simulated, ranging between 35.6 g C m^− 2y^− 1 in the East Greenland Current of Atlantic origin zone and 45.6 g C m^− 2y^− 1 in the Northern Irminger Current zone. The model successfully simulated the observed regional and spatial differences in terms of the maximum depth of winter mixing, the onset of stratification and the development of the seasonal thermocline, and the differences in biological characteristics between the zones. The initial properties of the water column and the seasonal cycle of physical and meteorological forcing in each of the zones are responsible for the observed differences during the Marine Productivity cruises. The timing of the transition from mixing to stratification regime, and the different prevailing light levels in each zone are identified as the crucial processes/parameters for the understanding of the dynamics of the pelagic ecosystem in the Irminger Sea.     

Ecology of bottom ice algae: II. Dynamics, distributions and productivity

Spring blooms of bottom ice algae are a common feature of landfast congelation ice in polar regions. Because ice algae are usually associated with a substrate, their population dynamics can be followed with considerable confidence. Although ice algal dynamics are closely related to irradiance, their dynamics and distributions are influenced by other abiotic and biotic factors. Ice algal abundance varies horizontally over all scales examined. Factors such as grazing and nutrient availability may contribute to local and geographic differences. Loss terms for most sea ice assemblages are largely unquantified. Ice algal biomass is most concentrated near the ice-water interface in spring.Environmental factors affecting ice algal abundance and productivity are considered here, emphasizing recent results from several well-studied sites. Biomass accumulation, growth rates and productivity have been documented for spring blooms of bottom interstitial and sub-ice assemblages. On an areal basis biomass accumulation in bottom ice assemblages can be comparable with planktonic systems. At low ambient temperatures and irradiances average specific growth rates (≤ 0.25 d⁻¹) and production rates (≤ 1.0 mg C mg Chl⁻¹ h⁻¹) for ice algae are low. Current methods of measuring productivity are compared. Results are consistently low but variable with little systematic difference among them. At present, apparent differences in productivity between bottom ice assemblages in the Arctic and Antarctic, or among different antarctic assemblages, are so confounded by methodological and other sources of variability that no firm differences can be detected.     

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.     

100-years-changes in the phytoplankton community of Kiel Bight (Baltic Sea)

Literature data from 1905/06, 1912/13 and 1949/50 were compared with recent data (2001–2003) from Kiel Bight in order to investigate changes in phytoplankton composition and biomass, which may serve as indicators of environmental changes. In terms of biomass, diatomophyceae and dinophyceae are by far the most important groups. Their ratio is still close to unity. The share of diatomophyceae increased strongly in years with exceptionally high summer blooms (2001) or exceptionally early spring blooms (2003). The summer and autumn blooms of Chaetoceros and Skeletonema, detected in the early 20th century, are replaced by other diatoms (Cerataulina pelagica, Dactyliosolen fragilissimus, Proboscia alata, Pseudo-nitzschia spp.). Chaetoceros and Skeletonema are still important components of the spring blooms. Now as before, the autumn blooms are dominated by Ceratium spp., sometimes also by diatoms. Newly appearing bloom-forming species are mostly potentially toxic (Dictyocha speculum, Prorocentrum minimum, Pseudo-nitzschia spp.). The total phytoplankton biomass has roughly doubled in the course of the last century. The reference condition for phytoplankton biomass in Kiel Bight in the sense of the Water Framework Directive was defined at 55 mg C m^− 3 (± 10%, annual mean). The mean annual biomass of diatomophyceae and dinophyceae was 25 mg C m^− 3 (± 40%) for each, indicating that the sum of their carbon biomass amounted to 90% (± 10%) of the total phytoplankton biomass on an annual average. Diatomophyceae represented at least 80% of carbon biomass in the spring bloom peak at the beginning of the 20th century.     

Variability of mesozooplankton spatial distribution in the North Aegean Sea, as influenced by the Black Sea waters outflow

The North Aegean Sea constitutes an important region of the Mediterranean Sea since in its eastern part the mesotrophic, low salinity and relatively cold water from the Black Sea (outflowing from the Dardanelles strait) meets the oligotrophic, warm and very saline water of Levantine origin, thus forming a thermohaline front. Mesozooplankton samples were collected at discrete layers according to the hydrology of the upper 100 m, during May 1997 and September 1998. In May highest biomass and abundance values (up to 66.82 mg m^− 3 and 14,157 ind m^− 3) were detected in the 10–20 m layer (within the halocline) of the stations positioned close to the Dardanelles strait. The front moved slightly southwards in September, characterized by high biomass and abundance values within the halocline layer. The areas moderately or non influenced by Black Sea water revealed lower standing stock values than the frontal area in both cruises and maxima were detected in the uppermost low salinity layer. Samples collected at the stations and/or layers more influenced by Black Sea water were distinguished from those collected at layers and/or stations more affected by Levantine waters in both periods. In May the former samples were characterized by the copepods Acartia clausi, Centropages typicus, Paracalanus parvus. The abundance of the above species decreased gradually with increasing salinity, in the horizontal and/or in the vertical dimension, with a parallel increase of the copepods Oithona plumifera, Oithona copepodites, Oncaea media, Ctenocalanus vanus, Farranula rostrata. During September the frontal area as well as that covered by the modified Black Sea water, were highly dominated by the cladoceran Penilia avirostris and doliolids. For both seasons, MDS plots, issued from the combination of mesozooplankton and water-type data, revealed the gradual differentiation of zooplankton composition from the frontal area towards the area covered by Levantine water, following the spreading and mixing of the Black sea water. The observed temporal and spatial variability in the distribution pattern of mesozooplankton standing stock and species composition seems to depend considerably on the variability of circulation and frontal flows.