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.     

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.     

Species of Thaumatomastix (Thaumatomastigidae, Protista incertae sedis) from the Arctic sea ice biota (North-East Water Polynya, NE Greenland)

The sea ice biota of polar regions contains numerous heterotrophic flagellates very few of which have been properly identified. The whole mount technique for transmission electron microscopy enables the identification of loricate and scaly forms. A survey of Arctic ice samples (North-East Water Polynya, NE Greenland) revealed the presence of ca. 12 taxa belonging to the phagotrophic genus Thaumatomastix (Protista incertae sedis). Species of Thaumatomastix possess siliceous body scales and one naked and one scale-covered flagellum. The presence in both Arctic samples and sea ice material previously examined from the Antarctic indicates that this genus is most likely ubiquitous in polar sea ice and may be an important component in sea ice biota microbial activities.     

Biological effects of Mississippi River nitrogen on the northern gulf of Mexico—a review and synthesis

The Mississippi River currently delivers approximately 1.82 Tg N year⁻¹ (1.3×10¹¹ mol N year⁻¹) to the northern Gulf of Mexico. This large input dominates the biological processes of the region. The “new” nitrogen from the river stimulates high levels of phytoplankton production which in turn support high rates of bacterial production, protozoan and metazoan grazing, and fisheries production. A portion of the particulate organic matter produced in the pelagic food web sinks out of the euphotic zone where it contributes to high rates of oxygen consumption in the bottom waters of the inner shelf, resulting in the development of an extensive zone of hypoxia each summer. In spite of the significance of this river system to the coastal ocean of the northern gulf, we do not have an adequate understanding of the inputs, processing and ultimate fates of river nitrogen. Here we review available literature on this important system and propose a conceptual model showing how biological processes evolve in the river plume between the point of discharge and the point where plume waters are fully diluted by mixing with oceanic water.     

Microbial community structure in the marginal ice zone of the Bellingshausen Sea

Phytoplankton, bacteria and microzooplankton were investigated on a transect in the Bellingshausen Sea during the ice melt period in November–December 1992. The transect along the 85°W meridian comprised seven stations that progressed from solid pack-ice (70°S), through melting ice into open water (67°S). The abundance, biomass and taxonomic composition were determined for each component of the microbial community. The phytoplankton was mostly dominated by diatoms, particularly small (<20 μm) species. Diatom abundance ranged from 66 000 cells l⁻¹ under the ice to 410 000 cells l⁻¹ in open water. Phytoplankton biomass varied from <1 to 167 mg C m⁻³, with diatoms comprising 89–95% of the total biomass in open water and autotrophic nanoflagellates comprising 57% under the ice. The standing stocks of autotrophs in the mixed layer ranged from 95 mg C m⁻² under the pack-ice to 9478 mg C m⁻² in open waters. Bacterial abundance in ice-covered and open water stations varied from 1.1 to 5.5×10⁸ cells l⁻¹. Bacterial biomass ranged from 2.4 mg C m⁻³ under pack-ice to an average of 14 mg C m⁻³ in open water. The microzooplankton consisted mainly of aloricate oligotrich ciliates and heterotrophic dinoflagellates and these were most abundant in open waters. Their biomass varied between 0.2 and 54 mg C m⁻³ with a minimum at depth under the ice and maximum in open surface waters. Microheterotrophic standing stocks varied between 396 mg C m⁻² under pack-ice and 3677 mg C m⁻² in the open waters. The standing stocks of the total microbial community increased consistently from 491 mg C m⁻² at the ice station to 13 155 mg C m⁻² in open waters, reflecting the productive response of the community to ice-melt. The composition of the microbial community also shifted markedly from one dominated by heterotrophs (82% of microbial stocks) at the ice station to one dominated by autotrophs (73% of microbial stocks) in the open water. Our estimates suggest that the microbial community comprised >100% of the total particulate organic carbon (POC) under the ice and 62–66% of the measured POC in the open waters.     

Winter–spring dynamics in sea-ice carbon cycling in the coastal Arctic Ocean

An understanding of microbial interactions in first-year sea ice on Arctic shelves is essential for identifying potential responses of the Arctic Ocean carbon cycle to changing sea-ice conditions. This study assessed dissolved and particulate organic carbon (DOC, POC), exopolymeric substances (EPS), chlorophyll a, bacteria and protists, in a seasonal (24 February to 20 June 2004) investigation of first-year sea ice and associated surface waters on the Mackenzie Shelf. The dynamics of and relationships between different sea-ice carbon pools were investigated for the periods prior to, during and following the sea-ice-algal bloom, under high and low snow cover. A predominantly heterotrophic sea-ice community was observed prior to the ice-algal bloom under high snow cover only. However, the heterotrophic community persisted throughout the study with bacteria accounting for, on average, 44% of the non-diatom particulate carbon biomass overall the study period. There was an extensive accumulation of sea-ice organic carbon following the onset of the ice-algal bloom, with diatoms driving seasonal and spatial trends in particulate sea-ice biomass. DOC and EPS were also significant sea-ice carbon contributors such that sea-ice DOC concentrations were higher than, or equivalent to, sea-ice-algal carbon concentrations prior to and following the algal bloom, respectively. Sea-ice-algal carbon, DOC and EPS-carbon concentrations were significantly interrelated under high and low snow cover during the algal bloom (r values ≥ 0.74, p < 0.01). These relationships suggest that algae are primarily responsible for the large pools of DOC and EPS-carbon and that similar stressors and/or processes could be involved in regulating their release. This study demonstrates that DOC can play a major role in organic carbon cycling on Arctic shelves.     

Crystallographic structure of sea ice along a salinity gradient and environmental control of microalgae in the brine cells

Duplicate ice cores were taken in southeastern Hudson Bay (Canadian Arctic), at 8 stations along a transect, from the mouth of the Great Whale River to saline waters 25 km offshore. One core from each station was cut into 10-cm sections, which were melted at room temperature for determinations of salinity, nutrients, algal pigments and taxonomic composition of the microalgal assemblages. On the second core, thin sections were cut every 2 cm to optically determine the ratio of brine/gas pockets to the total sections. Relative volumes of brine in melted samples were computed from the observed ratios and calculated values, and used to convert per unit brine volume the concentrations of pigments, nutrients and algal cells, initially measured per unit volume of melted sample.Salinity of the melted samples indicate the presence of freshwater beneath the sea ice close to shore throughout the growth season, and the vertical salinity profiles suggest intermittent intrusions of freshwater. Nutrient concentrations in the brine were high, suggesting that microalgae in the brine cells were generally not nutrient limited. Concentrations of ice algae in the brine cells reached relatively high values, that are often of the same order as those reported for eutrophic marine waters. Statistical analyses identified the rate of ice growth as the most important factor controlling the vertical and horizontal distributions of algal biomass and taxonomic composition in the sea ice, along the salinity gradient. Higher biomasses were generally associated with slower ice growth and also possibly lower grazing activity. In addition, the rate of ice growth influenced the distribution of microalgal assemblages, with nitrogen limitation potentially playing a secondary role in some instances.     

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.     

Fluxes of biogenic carbon in the Southern Ocean: roles of large microphagous zooplankton

The Southern Ocean is an extreme environment, where waters are permanently cold, a seasonal ice cover extends over large areas, and the solar energy available for photosynthesis is severely restricted, either by vertical mixing to considerable depths or, especially south of the Antarctic Circle, by prolonged seasonal periods of low or no irradiance. Such conditions would normally lead to low productivity and a water column dominated by recycling processes involving microbial components of pelagic communities but this does not seem to be the case in the Southern Ocean, where there is efficient export to large apex predators and deep waters. This paper investigates the role of large microphagous zooplankton (salps, krill, and some large copepods) in the partitioning of biogenic carbon among the pools of short- and long-lived organic carbon and sequestered biogenic carbon. Large microphagous zooplankton are able to ingest microbial-sized particles and thus repackage small, non-sinking particles into both metazoan biomass and large, rapidly sinking faeces. Given the wide spatio-temporal extent of microbial trophic pathways in the Southern Ocean, large zooplankton that are omnivorous or able to ingest small food particles have a competitive advantage over herbivorous zooplankton. Krill efficiently transfer carbon to a wide array of apex predators and their faecal pellets are exported to depth during occasional brief sedimentation episodes in spring time. Salps may be a significant link towards some fish (directly) and other apex predators (indirectly) and, at some locations (especially in offshore waters) and time, they may account for most of the downward flux of biogenic carbon. Large copepods are a trophic link towards fish and at least one whale species, and their grazing activity generally impedes the export of organic particles to depth. As a result, biogenic carbon is channelled mainly towards apex predators and episodically into the deep ocean. Without these original interactions, Antarctic waters might well be dominated by microbial components and recycling processes instead of active export from the generally small primary producers towards large apex predators.     

Variability of snow and ice thermal, physical and optical properties pertinent to sea ice algae biomass during spring

Changes in the thermal, physical and optical properties of the snow–sea ice system and feedbacks between various temporal and spatial scales affect accumulation of microalgae at the sea ice bottom and are the focus of this research. During the spring transition period, May 4 to June 9, 2002, we closely monitored atmospheric conditions and properties of the snow–sea ice system, including thermal, physical and optical properties of the snow cover (e.g., temperature, grain size, light attenuation), ice thickness and salinity, and biomass of bottom ice algae. Results show that snowdrift size averaged 31.2 by 10.6 m with a depth range of 2 to 45 cm. Snowpacks also varied in age, distinguished by coincident peaks of snow salinity and grain size and a lower PAR extinction coefficient. Spatial variability of the snowpack was superimposed by temporal variability associated with seasonal snow–ice melt and wind redistribution of snow. Maximum biomass of ice algae was observed under intermediate snow covers. Under thin snow covers, algae biomass declined steadily coincident with seasonal warming and desalination of the ice cover. Under thick snow covers, algae biomass was negatively correlated with snow depth. These results suggest that thin snow covers were associated with a thermal effect causing sloughing of algae, whereas under deep snow, algae were still light limited and thermally insulated from the warming atmosphere. Our results highlight the importance of snow cover history on the sea ice system operating below. Furthermore, in the context of current climate change scenarios, shifts in snow depth would result in decreases of ice algae biomass.     

Benthic community and food web structure on the continental shelf of the Bay of Biscay (North Eastern Atlantic) revealed by stable isotopes analysis

The North Bay of Biscay continental shelf is a major French demersal fishery, but little was known on the trophic food web of its benthic communities. In order to determine the benthic trophic web, the objectives of this study are to describe the macro- and megafaunal benthic community structure (species richness, abundance and biomass) and to establish the trophic pathways (food sources and trophic levels) by applying carbon and nitrogen stable isotopic analysis to the main benthic and demersal species (invertebrates and fish). Two distinct benthic communities have been identified: a muddy sand community within the central part of the bay, and an outer Bay of Biscay Ditrupa sand community of higher species richness, abundance and biomass than the muddy sand community. Deposit-feeders, suspension feeders and predators, distributed in three main trophic levels, dominate both communities. Large differences in stable carbon ratio values within the primary consumers provide evidence of two different food sources: i) a pelagic food source made up of recent sedimenting particulate organic matter on which zooplankton and suprabenthos feed and ii) a benthic detrital food source supplying deposit feeders and partly benthic suspension feeders. Differences in isotopic signatures were also observed within the upper trophic levels that allowed estimation of the contribution of each food source component to the diet of the upper consumers. Finally, the use of stable isotopic composition together with the species' feeding strategy allow identification of the main differences between the trophic functioning of the two benthic communities and highlight the importance of the role of detrital pathways in the carbon cycling within the continental shelf benthic trophic web.     

Characteristics and potential impacts of under-ice river plumes in the seasonally ice-covered Bothnian Bay (Baltic Sea)

The northernmost basin of the Baltic Sea, the Bothnian Bay, is ice-covered for about half the year. During this time, distinct under-ice river plumes develop, even seaward of the smallest rivers, that are substantially thicker and larger in extent than during the summer months. Wind mixing is negligible, and during late spring in April or May, the highest annual discharge occur while the sea is ice covered, thus providing conditions for the formation of extensive under-ice plumes. These plumes are characterised by high levels of trace elements (e.g., Al, Fe and Zn), organic matter (TOC and dissolved organic carbon [DOC]), nutrients and also optically active substances (colored dissolved organic matter, CDOM). The under-ice plumes provide an important pathway for undiluted transport of land-derived substances to the pelagic waters of the basin, affecting the salinity, chemistry and optical properties of coastal waters. Freshwater ice growth on the underside of an existing sea ice sheet also restricts the buildup of sea ice and under-ice algal communities, potentially in large areas along the coasts. Plume water influences the optical characteristics of coastal waters for a period of time after ice break-up, potentially affecting primary production in these areas. Furthermore, the formation of under-ice plumes potentially has a positive feedback on the ice season length due to freshening of the coastal waters (earlier freeze-up) and restricted oceanic heat flux (slower melting).     

Ability of a “minimum” microbial food web model to reproduce response patterns observed in mesocosms manipulated with N and P, glucose, and Si

We compared an idealised mathematical model of the lower part of the pelagic food web to experimental data from a mesocosm experiment in which the supplies of mineral nutrients (nitrogen and phosphorous), bioavailable dissolved organic carbon (BDOC, as glucose), and silicate were manipulated. The central hypothesis of the experiment was that bacterial consumption of BDOC depends on whether the growth rate of heterotrophic bacteria is limited by organic-C or by mineral nutrients. In previous work, this hypothesis was examined qualitatively using a conceptual food web model. Here we explore the extent to which a “simplest possible” mathematical version of this conceptual model can reproduce the observed dynamics. The model combines algal–bacterial competition for mineral nutrients (phosphorous) and accounts for alternative limitation of bacterial and diatom growth rates by organic carbon and by silicate, respectively. Due to a slower succession in the diatom–copepod, compared to the flagellate–ciliate link, silicate availability increases the magnitude and extends the duration of phytoplankton blooms induced by mineral nutrient addition. As a result, Si interferes negatively with bacterial consumption of BDOC consumption by increasing and prolonging algal–bacterial competition for mineral nutrients. In order to reproduce the difference in primary production between Si and non-Si amended treatments, we had to assume a carbon overflow mechanism in diatom C-fixation. This model satisfactorily reproduced central features observed in the mesocosm experiment, including the dynamics of glucose consumption, algal, bacterial, and mesozooplankton biomass. While the parameter set chosen allows the model to reproduce the pattern seen in bacterial production, we were not able to find a single set of parameters that simultaneously reproduces both the level and the pattern observed for bacterial production. Profound changes in bacterial morphology and stoichiometry were reported in glucose-amended mesocosms. Our “simplest possible” model with one bacterial population with fixed stoichiometry cannot reproduce this, and we suggest that a more elaborate representation of the bacterial community is required for more accurate reproduction of bacterial production.     

Study of the seasonal cycle of the biogeochemical processes in the Ligurian Sea using a 1D interdisciplinary model

A one-dimensional coupled physical–biogeochemical model has been built to study the pelagic food web of the Ligurian Sea (NW Mediterranean Sea). The physical model is the turbulent closure model (version 1D) developed at the GeoHydrodynamics and Environmental Laboratory (GHER) of the University of Liège. The ecosystem model contains 19 state variables describing the carbon and nitrogen cycles of the pelagic food web. Phytoplankton and zooplankton are both divided in three size-based compartments and the model includes an explicit representation of the microbial loop including bacteria, dissolved organic matter, nano-, and microzooplankton. The internal carbon/nitrogen ratio is assumed variable for phytoplankton and detritus, and constant for zooplankton and bacteria. Silicate is considered as a potential limiting nutrient of phytoplankton's growth. The aggregation model described by Kriest and Evans in (Proc. Ind. Acad. Sci., Earth Planet. Sci. 109 (4) (2000) 453) is used to evaluate the sinking rate of particulate detritus. The model is forced at the air–sea interface by meteorological data coming from the “Côte d'Azur” Meteorological Buoy. The dynamics of atmospheric fluxes in the Mediterranean Sea (DYFAMED) time-series data obtained during the year 2000 are used to calibrate and validate the biological model. The comparison of model results within in situ DYFAMED data shows that although some processes are not represented by the model, such as horizontal and vertical advections, model results are overall in agreement with observations and differences observed can be explained with environmental conditions.     

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.     

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.     

Analysis of the contribution of ice algae to the ice-covered ecosystem in Lake Saroma by means of a coupled ice–ocean ecosystem model

A new coupled ice–ocean ecosystem model that links the pelagic and ice ecosystems was used to clarify the role of ice algae in ice-covered ocean ecosystems. The model was applied to Lake Saroma (Hokkaido, Japan) in 1992. Comparison of the model's results with observational data confirmed that the model reproduced the behavior of the ecosystem with acceptable accuracy during the period from winter to spring. The primary production of the ice algae is effectively transported into the pelagic system by means of physical releasing effects: brine convection, ice melting and freezing, and diffusion generated at the bottom of the ice. Ice algae released from the ice are rapidly exported because of their high sinking speed and the shallow depth of Lake Saroma. For this reason, the zooplankton in Lake Saroma cannot graze these released algae. However, zooplankton actively graze the ice algae living along the bottom of the ice. These results show that, before their release, ice algae play an important role as a food source for overwintering zooplankton. A sensitivity analysis revealed a positive correlation between the sinking speed of the released ice algae and the magnitude of the spring bloom by pelagic phytoplankton, and that the time when secondary production becomes active is an important factor in the linkage between these two algal populations.     

Distribution of diatoms in the Northeast Water Polynya, Greenland

This study formed part of the Northeast Water project (NEW project) which dealt with physical, geophysical and biological processes in the Northeast Water Polynya off Northeast Greenland. This was part of the International Arctic Polynya Programme (IAPP). The diatom composition of the water masses, sea ice and melt ponds was analysed to show the relationship between ice and the water column near the ice with regard to the origin and fate of the cells in the ice and melt ponds. Fragilariopsis oceanica, Fragiliria sp. I and Chaetoceros socialis usually dominated the phytoplankton, while the ice and melt pond samples showed a wide range of assemblages, with different single-celled pennates and two undescribed species, Navicula sp. 1 and Nitzschia sp. 1 often dominant. Planktonic algae in sea ice can be released into the water column during ice break-up and melt, thus contributing to the spring bloom in the water column, if the timing of the release and the species composition are correct. The number of different ice algal assemblages supports the theory that cells originated from the water column, the benthos and freshwater. In addition, differential growth in the sea ice or melt ponds often altered the relative abundance of species in comparison with what is usually found in their original habitat. However, many of the cells in the ice and melt ponds were dead (empty frustules), making it difficult to determine whether the cells had actually lived in these habitats.     

Primary productivity of the Palmer Long Term Ecological Research Area and the Southern Ocean

A major objective of the Palmer Long Term Ecological Research (Palmer LTER) project is to obtain a comprehensive understanding of the various components of the Antarctic marine ecosystem. Phytoplankton production plays a key role in this so-called high nutrient, low chlorophyll environment, and factors that regulate production include those that control cell growth (light, temperature, and nutrients) and those that control cell accumulation rate and hence population growth (water column stability, grazing, and sinking). Sea ice mediates several of these factors and frequently conditions the water column for a spring bloom which is characterized by a pulse of production restricted in both time and space. This study models the spatial and temporal variability of primary production within the Palmer LTER area west of the Antarctic Peninsula and discusses this production in the context of historical data for the Southern Ocean. Primary production for the Southern Ocean and the Palmer LTER area have been computed using both light-pigment production models [Smith, R.C., Bidigare, R.R., Prézelin, B.B., Baker, K.S., Brooks, J.M., 1987. Optical characterization of primary productivity across a coastal front. Mar. Biol. (96), 575–591; Bidigare, R.R., Smith, R.C., Baker, K.S., Marra, J., 1987. Oceanic primary production estimates from measurements of spectral irradiance and pigment concentrations. Global Biogeochem. Cycles (1), 171–186; Morel, A., Berthon, J.F., 1989. Surface pigments, algal biomass profiles and potential production of the euphotic layer—relationships reinvestigated in view of remote-sensing applications. Limnol. Oceanogr. (34), 1545–1562] and an ice edge production model [Nelson, D.M., Smith, W.O., 1986. Phytoplankton bloom dynamics of the western Ross Sea ice edge: II. Mesoscale cycling of nitrogen and silicon. Deep-Sea Res. (33), 1389–1412; Wilson, D.L., Smith, W.O., Nelson, D.M., 1986. Phytoplankton bloom dynamics of the Western Ross Sea ice edge: I. primary productivity and species-specific production. Deep-Sea Res., 33, 1375–1387; Smith, W.O., Nelson, D.M., 1986. Importance of ice edge phytoplankton production in the Southern Ocean. BioScience (36), 251–257]. Chlorophyll concentrations, total photosynthetically available radiation (PAR) and sea ice concentrations were derived from satellite data. These same parameters, in addition to hydrodynamic conditions, have also been determined from shipboard and Palmer Station observations during the LTER program. Model results are compared, sensitivity studies evaluated, and productivity of the Palmer LTER region is discussed in terms of its space time distribution, seasonal and interannual variability, and overall contribution to the marine ecology of the Southern Ocean.     

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.