Recent advances in sea-ice microbiology

Over the past 50 years there has been much effort invested in the investigation of the ecology of sea ice. Sea ice is an ephemeral feature of the Arctic and Southern Oceans and smaller water bodies such as the Baltic and Caspian Seas. The semisolid ice matrix provides a range of habitats in which a diverse range of microbial organisms thrive. In the past 5 years there has been considerable steps forward in sea-ice research, in particular regarding the analysis of sea-ice microstructure and the investigation of the diversity and adaptation of microbial communities. These studies include: (i) controlled simulated and in situ studies on a micrometer scale to unravel the dynamic of the microhabitat with consequences for the organisms; (ii) the introduction of molecular approaches to uncover the diversity of uncultured still unknown microorganisms; and (iii) studies into the molecular adaptation of selected model organisms to the extreme environment. This minireview presents some of the most recent findings from sea-ice studies within the framework of these aims.     

Evidence for active microbial nitrogen transformations in sea ice (Gulf of Bothnia, Baltic Sea) in midwinter

Nutrient concentrations, chlorophyll-a, bacterial biomass and relative activity of denitrifying organisms were investigated from ice-core, brine and underlying water samples in February 1998 in the Gulf of Bothnia, Baltic Sea. Examined sea ice was typical for the Baltic Sea; ice bulk salinity varied from 0.1 to 1.6 psu, and in underlying water salinity was from 4.2 to 4.7 psu. In 2- to 3-months-old sea ice (thickness 0.4–0.6 m), sea-ice communities were at the winter stage; chl-a concentrations were generally below 1 mg m⁻³ and heterotrophic organisms composed 7–20% of organism assemblage. In 1-month-old ice (thickness 0.2–0.25 m), an ice spring bloom was already developing and chl-a concentrations were up to 5.6 mg m⁻³. In relation to low salinity, high concentrations of NH⁺ ₄, NO⁻ ₂, PO³⁺ ₄ and SiOH₄ were found in the ice column. The results suggest that the upper part of ice accumulates atmospheric nutrient load during the ice season, and nutrients in the upper 10–20 cm of ice are mainly of atmospheric origin. The most important biological processes controlling the sea-ice nutrient status are nutrient regeneration, nutrient uptake and nitrogen transformations. Nutrient regeneration is specially active in the middle parts of the 50- to 60-cm-thick ice and subsequent accumulation of nutrients probably enhances the ice spring bloom. Nitrite accumulation and denitrifying activity were located in the same ice layers with nutrient regeneration, which together with the observed significant correlation between the concentrations of nitrogenous nutrients points to active nitrogen transformations occurring in the interior layers of sea ice in the Baltic Sea. Accepted: 12 June 2000     

Succession of sea-ice bacterial communities in the Baltic Sea fast ice

Coastal fast ice and underlying water of the northern Baltic Sea were sampled throughout the entire ice winter from January to late March in 2002 to study the succession of bacterial biomass, secondary production and community structure. Temperature gradient gel electrophoresis (TGGE) and sequencing of TGGE fragments were applied in the community structure analysis. Chlorophyll-a and composition of autotrophic and heterotrophic assemblages were also examined. Overall succession of ice organism assemblages consisted of a low-productive stage, the main algal bloom, and a heterotrophic post-bloom situation, as typical for the study area. The most important groups of organisms in ice in terms of biomass were dinoflagellates, plasticidic flagellates, rotifers and ciliates. Ice bacteria showed a specific succession not directly dependent on the overall succession events of ice organisms. Sequenced 16S rDNA fragments were mainly affiliated to α-, β-, and γ-proteobacterial phyla and Cytophaga–Flavobacterium–Bacteroides-group, and related to sequences from cold environments, also from the Baltic Sea. Temporal clustering of the TGGE fingerprints was stronger than spatial, although lower ice and underlying water communities always clustered together, pointing to the importance of ice maturity and ice–water interactions in shaping the bacterial communities.     

Phylogeographical structure, distribution and genetic variation of the green algae Ulva intestinalis and U. compressa (Chlorophyta) in the Baltic Sea area

The marine algae Ulva intestinalis and U. compressa are morphologically plastic with many overlapping characters and are therefore difficult to distinguish from each other. The present distribution of U. intestinalis and U. compressa is investigated along the salinity gradient in the Baltic Sea area through analyses of internal transcribed spacer (ITS) sequence data. Also, the amount and distribution of intraspecific genetic polymorphism in the ITS region is studied allowing inferences on the phylogeographical pattern and postglacial recolonization of the Baltic Sea area. The data show that of the two species only U. intestinalis occurs in the Baltic Sea. The distribution of U. compressa is more restricted than previously reported, and it was not found in salinities lower than 15 ppt. All of Scandinavia and the Baltic Sea were covered with ice during the last ice age and the organisms in the Baltic Sea must have colonized the area after the ice had started to melt. The genetic diversity of U. intestinalis and U. compressa in the Baltic Sea and the neighbouring area was found to be reduced compared to that in the British Isles. This reduction may be the result of either a historical reduction of diversity or an adaptation of specific clones to the northern environmental conditions.     

Experimental Evidence on Nutrient and Substrate Limitation of Baltic Sea sea-ice Algae and Bacteria

The effect of nutrient limitation on Baltic Sea ice algae, and substrate and nutrient limitation on ice bacteria, was studied in a series of in situ -experiments conducted during the winter of 2002 in northern Baltic Sea. Community level changes in algal biomass (chlorophyll a) and productivity, and bacterial thymidine and leucine incorporation were followed for one week after the addition of nutrient and/or organic carbon rich filtered seawater to the experimental units. The results showed the major contribution of snow cover to the algal responses during the beginning of the ice-covered season. Algal communities were able to grow even in January if no snow was present. Nutrient addition did occasionally have an effect on algal biomass and productivity in the ice. Surprisingly, seeding effect from the ice to the underlying water was negatively affected by the nutrient availability in March. Bacterial limitation varied between nutrient (phosphorus) and substrate limitations. The results showed, that limitation in both algal and bacterial communities changed periodically in the northern Baltic Sea ice.     

Biomass,composition and activity of organism assemblages along a salinity gradient in sea ice subjected to river discharge in the Baltic Sea

A study was undertaken to examine the activity and composition of the seasonal Baltic Sea land-fast sea-ice biota along a salinity gradient in March 2003 in a coastal location in the SW coast of Finland. Using a multi-variable data set, the less well-known algal and protozoan communities, and algal and bacterial production in relation to the physical and chemical environment were investigated. Also, the first coincident measurements of bacterial production and dissolved organic matter (DOM) in a sea-ice system are reported. Communities in sea ice were clearly autotrophy-dominated with algal biomass representing 79% of the total biomass. Protozoa and rotifers made up 18% of biomass in the ice and bacteria only 3%. Highest biomasses were found in mid-transect bottom ice. Water column assemblages were clearly more heterotrophic: 39% algae, 12% bacteria and 49% for rotifers and protozoa. Few significant correlations existed between DOM and bacterial variables, reflecting the complex origin of ice DOM. Dynamics of dissolved organic carbon, nitrogen and phosphorus (DOC, DON and DOP) were also uncoupled. A functional microbial loop is likely to be present in the studied ice. Existence of an under-ice freshwater plume affects the ecosystem functioning: Under-ice water communities are influenced directly by river-water mixing, whereas the ice system seems to be more independent—the interaction mainly taking place through the formation of active bottom communities.     

Distribution patterns of epilithic diatoms along climatic,spatial and physicochemical variables in the Baltic Sea

The species richness and community composition of the diatom communities were studied in the Baltic Sea, Northern Europe, to enhance knowledge about the diversity of these organisms in a brackish water ecosystem. Many organisms in the Baltic Sea have been studied extensively, but studies investigating littoral diatoms are scarce. The goal of this study was to examine the importance of climatic, spatial and water physicochemical variables as drivers of epilithic diatoms in the Gulf of Finland and the Gulf of Bothnia. The variation in species richness was best explained by pH, total phosphorus and total nitrogen. Redundancy Analysis indicated that the most important factors correlating with species composition were air temperature, silicon, total phosphorus, water temperature, salinity and pH. Variation Partitioning showed that the species composition was mostly affected by climatic and spatial variables, whereas physicochemical variables had little impact. However, the strongest factor was the combined influence of climatic, spatial and physicochemical variables. The results suggest that diatom species richness in the northern Baltic Sea is primarily regulated by local factors, while climatic and spatial variables have little impact on richness. Species composition is mostly affected by climatic and spatial variables. We conclude that understanding the distribution patterns of Baltic Sea diatoms requires the inclusion of climatic, spatial and water chemistry variables.     

The Native Bacterioplankton Community in the Central Baltic Sea Is Influenced by Freshwater Bacterial Species

The Baltic Sea is one of the largest brackish environments on Earth. Despite extensive knowledge about food web interactions and pelagic ecosystem functioning, information about the bacterial community composition in the Baltic Sea is scarce. We hypothesized that due to the eutrophic low-salinity environment and the long water residence time (>5 years), the bacterioplankton community from the Baltic proper shows a native “brackish” composition influenced by both freshwater and marine phylotypes. The bacterial community composition in surface water (3-m depth) was examined at a single station throughout a full year. Denaturing gradient gel electrophoresis (DGGE) showed that the community composition changed over the year. Further, it indicated that at the four extensive samplings (16S rRNA gene clone libraries and bacterial isolates from low- and high-nutrient agar plates and seawater cultures), different bacterial assemblages associated with different environmental conditions were present. Overall, the sequencing of 26 DGGE bands, 160 clones, 209 plate isolates, and 9 dilution culture isolates showed that the bacterial assemblage in surface waters of the central Baltic Sea was dominated by Bacteroidetes but exhibited a pronounced influence of typical freshwater phylogenetic groups within Actinobacteria, Verrucomicrobia, and Betaproteobacteria and a lack of typical marine taxa. This first comprehensive analysis of bacterial community composition in the central Baltic Sea points to the existence of an autochthonous estuarine community uniquely adapted to the environmental conditions prevailing in this brackish environment.     

Long-term dynamics of zooplankton in the southeastern Baltic Sea

Long-term research in the Baltic Sea revealed the basic trends of zooplankton community variations depending on oceanographic processes. Alternation of the periods of increase and decrease in salinity of the Baltic Sea against the background of climate changes (temperature increase) and eutrophication affect the state of the entire Baltic ecosystem, including zooplankton. For these periods, the dynamics of zooplankton in the Baltic Sea were analyzed based on literature data and results of regular research in the southeastern Baltic Sea during 1998–2007. The changes in the hydrological situation were accompanied by significant changes in the zooplankton community. In the 1990s–2000s, the abundance and biomass of brackish-water and thermophilous species primarily of Cladocera and Copepoda increased markedly. The role of the previously dominant marine copepod Pseudocalanus elongatus decreased due to salinity reduction in the deep-water part of the Baltic Sea. Maximum development of zooplankton occurred in years of the greatest warming-up of the water (2001, 2005–2007) against the background of a general positive trend of zooplankton abundance in the last decade.     

The topography of the ice-water interface – its influence on the colonization of sea ice by algae

The ice-water interface constitutes an important habitat for polar organisms, characterized by extreme variability in physical and biological properties, which can range over an order of magnitude on decimeter scales. The porous nature of sea ice allows vertical fluid exchange within the ice and across the ice-water interface in much the same way as the sediment-water interface. The present paper reports on experiments examining the effect of irregular undersides of sea ice on the variability of algal biomass, using flume tanks with constant unidirectional flow. Bulges and depressions at the ice-water interface altered the interfacial pore water flux and affected the spatial distribution and abundance of ice-associated algae substantially. Dye tracer experiments demonstrated that interfacial water fluxes around ice bulges are by a factor of 10-100 higher than published indirect estimates based on algal nutrient demands in flat fast ice. Further, underside relief structures migrated downstream, illustrating their dynamic and transient nature. The presence of a relief fostered algal accumulation compared to a flat underside. Algal growth occurred at bulge sites facing the current, while particles accumulated in the wake further downstream. We infer from our experiments that sea ice is not only a source for algae production, but can also serve as sink for organic material from the water column. We propose that local formation and ablation of ice around underside features is an important process which induces high variations in the sea-ice habitat structure and in growth of sympagic organisms, and could partially explain the high natural variability observed in the abundance and colonization patterns of sea-ice organisms.     

Responses of Baltic Sea Ice and Open-Water Natural Bacterial Communities to Salinity Change

To investigate the responses of Baltic Sea wintertime bacterial communities to changing salinity (5 to 26 practical salinity units), an experimental study was conducted. Bacterial communities of Baltic seawater and sea ice from a coastal site in southwest Finland were used in two batch culture experiments run for 17 or 18 days at 0°C. Bacterial abundance, cell volume, and leucine and thymidine incorporation were measured during the experiments. The bacterial community structure was assessed using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified partial 16S rRNA genes with sequencing of DGGE bands from initial communities and communities of day 10 or 13 of the experiment. The sea ice-derived bacterial community was metabolically more active than the open-water community at the start of the experiment. Ice-derived bacterial communities were able to adapt to salinity change with smaller effects on physiology and community structure, whereas in the open-water bacterial communities, the bacterial cell volume evolution, bacterial abundance, and community structure responses indicated the presence of salinity stress. The closest relatives for all eight partial 16S rRNA gene sequences obtained were either organisms found in polar sea ice and other cold habitats or those found in summertime Baltic seawater. All sequences except one were associated with the α- and γ-proteobacteria or the Cytophaga-Flavobacterium-Bacteroides group. The overall physiological and community structure responses were parallel in ice-derived and open-water bacterial assemblages, which points to a linkage between community structure and physiology. These results support previous assumptions of the role of salinity fluctuation as a major selective factor shaping the sea ice bacterial community structure.     

Further observations on flagellates within sea ice in northern Bothnian Bay, the Baltic Sea

To improve our knowledge of flagellates inhabiting the Baltic Sea ice and water column during the winter, material was obtained from northern Bothnian Bay in March/April 1995. Light microscopical observations on live and fixed material and further transmission electron microscopy of whole mounts revealed 47 nanoflagellate taxa. In addition, detached scales of eight taxa were encountered. It is now evident that nearly all nanoflagellate classes are present within Bothnian Bay sea ice. The most common groups were cryptomonads, dinoflagellates, chrysophytes, prasinophytes, choanoflagellates and heterotrophic flagellates of unknown systematic position (Protista incertae sedis). Most flagellates in Baltic Sea ice biota apparently thrive in both the water column and the sea ice, while some, e.g. Paraphysomonas spp. (Chrysophyceae), heterotrophic euglenids, volvocalean chlorophytes, and some taxa of uncertain systematic affinity, are more frequently found within the sea ice. Received: 9 February 1997 / Accepted: 23 September 1997     

Evidence for a pan-Arctic sea-ice diatom diet in Strongylocentrotus spp

The Arctic marine food web is based on organic matter produced by both phytoplankton and sea-ice algae. With the decline of Arctic sea ice, the sustained availability of organic carbon of sea-ice origin is unclear. Recently, the detection of the sea-ice diatom biomarker IP₂₅ in a range of Arctic benthic macrofauna indicated that this is a highly suitable biomarker for the identification of organic carbon derived from sea-ice primary production in Arctic food webs. However, the data presented previously were restricted to a single geographical region in the Canadian Arctic. Here, we show that IP₂₅ is present in sea urchins of the genus Strongylocentrotus collected from ten locations with seasonal sea-ice cover from the Canadian Archipelago, Greenland and Spitsbergen. In contrast, IP₂₅ was not found in specimens of Echinus esculentus collected from the southwest UK, where sea ice is absent. Our findings provide evidence that the presence of IP₂₅ in macrobenthic organisms can be used across different Arctic regions as a versatile indicator of a diet containing carbon of sea-ice origin.     

Responses of Baltic Sea ice and open-water natural bacterial communities to salinity change

To investigate the responses of Baltic Sea wintertime bacterial communities to changing salinity (5 to 26 practical salinity units), an experimental study was conducted. Bacterial communities of Baltic seawater and sea ice from a coastal site in southwest Finland were used in two batch culture experiments run for 17 or 18 days at 0 degrees C. Bacterial abundance, cell volume, and leucine and thymidine incorporation were measured during the experiments. The bacterial community structure was assessed using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified partial 16S rRNA genes with sequencing of DGGE bands from initial communities and communities of day 10 or 13 of the experiment. The sea ice-derived bacterial community was metabolically more active than the open-water community at the start of the experiment. Ice-derived bacterial communities were able to adapt to salinity change with smaller effects on physiology and community structure, whereas in the open-water bacterial communities, the bacterial cell volume evolution, bacterial abundance, and community structure responses indicated the presence of salinity stress. The closest relatives for all eight partial 16S rRNA gene sequences obtained were either organisms found in polar sea ice and other cold habitats or those found in summertime Baltic seawater. All sequences except one were associated with the alpha- and gamma-proteobacteria or the Cytophaga-Flavobacterium-Bacteroides group. The overall physiological and community structure responses were parallel in ice-derived and open-water bacterial assemblages, which points to a linkage between community structure and physiology. These results support previous assumptions of the role of salinity fluctuation as a major selective factor shaping the sea ice bacterial community structure.     

Study of bacterial communities in Antarctic coastal waters by a combination of 16S rRNA and 16S rDNA sequencing

An ecological study on distribution of Antarctic bacterial communities was determined by 16S-based phylogenetic analyses of clone libraries derived from RNA and DNA extracted from two different marine areas and compared between each other. Superficial seawater samples were collected from four stations in Ross Sea, three of them located in Rod Bay and one in Evans Cove; for each station two clone libraries (16S rDNA and 16S rRNA) were prepared and evident divergences between DNA and RNA libraries of each site were obtained. Of all phylotypes 93.6% were found in RNA libraries; in contrast, only 31 phylotypes (70.5%) were retrieved from total microbial community (DNA libraries). DNA and RNA sequences related to gamma-Proteobacteria and Bacteroidetes groups, typical for Antarctic sea-ice bacterial communities, were detected in analysed sites. 16S rDNA and rRNA libraries derived from the two different areas were enriched by picophytoplanktonic 16S sequences of plastid and mitochondrion origins, reflecting that the algal blooms occurred during sampling (Antarctic summer 2003). The finding in Rod Bay libraries of high percentage of DNA clones apparently affiliated with beta-Proteobacteria typical for activated sludges and well water could be explained by the presence of a sewage depuration system at this site. Obtained results clearly demonstrate that combination of 16S rDNA and 16S rRNA gene sequencing is preferred approach to have a more reliable vision on the composition of microbial communities.     

Vertical distribution of sympagic meiofauna in sea ice in the Canadian Beaufort Sea

This is the first study to determine vertical distribution patterns of sympagic meiofauna, including metazoans, protozoans and eggs >20 μm, in the Amundsen Gulf (southeastern Beaufort Sea, Arctic). Full sea-ice cores were sampled from mid of March to end of May 2008 (Circumpolar Flaw Lead system study). Investigations were performed on first-year ice from three pack- and three fast-ice stations. Additionally, 5-cm bottom-ice sections were sampled at 13 pack-ice and 5 fast-ice stations. The metazoan community was composed of nematodes, rotifers, copepods, copepod nauplii, platyhelminthes and a few rare taxa such as mollusks, cnidarians and nemerteans. High numbers of eggs, between 50 and 2,188 eggs L⁻¹, particularly of nematodes and copepods, were present in the ice. Investigations revealed also eggs of the pelagic species Calanus hyperboreus and Sagitta spp. within the ice, so that further research is needed to clarify whether more organisms than expected might use this habitat as a reproduction ground. Many different morphotypes of protozoans were observed in the samples, especially ciliates of the order Euplotida. The highest abundance was always found in the lowermost 5 cm of the ice cores, nevertheless sympagic meiofauna was not restricted to that part of the ice. Integrated meiofauna abundance ranged between 41 and 4,738 × 10² Ind. m⁻² and was highest in the fast ice in early May. Differences between pack and fast ice in terms of integrated meiofauna communities and vertical distribution were not significant, while the analysis of the bottom-ice sections indicated both a temporal development and ice-type-specific differences.     

The role of sea ice in structuring Antarctic ecosystems

Summary This paper focusses on the links between growth, persistence and decay of sea ice and the structure of Antarctic marine ecosystems on different spatial and temporal scales. Sea-ice growth may divide an oceanic ecosystem into two dissimilar compartments: (1) the water column, with primary production controlled by the reduction of irradiative fluxes due to the snow-laden sea-ice cover and thermo-haline convection, and (2) the pore space within the ice with incorporated organisms switching from a planktonic to a kryohaline mode of life. In the ice, physical boundary conditions are set by (1) the irradiance which is controlled by the optical properties of snow and ice and (2) the ambient temperature which controls salinity and brine volume. Partly due to the high levels of biomass within the sea-ice system, interaction between different groups of organisms concentrates on the planar environment predefined by the ice cover. As a result of regional structuring of ecosystems, four sea-ice regimes may be recognized: seasonal pack ice, coastal zone, perennial pack ice, and marginal ice zone. These regimes are interwoven through the temporal structuring of ecosystems brought about by ice-cover seasonality and ice drift. In comparison with open-water pelagic ecosystems, sea ice appears of particular importance as it partly inverts the ecosystem structure and enhances the degree of ecological variability.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Lipid Content and fatty acid composition of algal communities in sea-ice and water from the Weddell Sea (Antarctica)

The lipid and fatty acid compositions of microalgae were investigated in sea-ice and water samples from six different habitats of the Weddell Sea (Antarctica). All sea-ice samples and ice-associated water contained high algal biomass dominated by centric and pennate diatoms. Cells partially filled with oil droplets and resting spores were found. In the cells from the ice platelet layer triacylglycerols formed the largest component of the lipids. The fatty acid composition of sea-ice microalgae was dominated by the 16:1(n-7), 16:0, 18:1(n-9) and 20:5 (n-3) fatty acids. Except 18:1, they are typical for diatom fatty acids. These fatty acids were most abundant in pieces of first year ice with a brown colouration (brown-ice) and in the water column directly below sea-ice (sub-ice water). The small amounts of non-diatom acids, as 22:6 (n-3) and 18:4 (n-3), clearly showed that the sea-ice communities were not purely composed of diatoms. The most striking difference, in comparison to the general fatty acid composition of diatoms, was the high proportion of the 18:1 fatty acid in all samples, which might be caused by detrital material or lipid accumulation within cells and resting spores. In general, no clear adaptation of the fatty acid composition to the Antarctic and sea-ice environment was found. The fatty acid composition of the particulate matter from the water column was totally different from all other samples dominated by the saturated fatty acids 16:0 and 18:0.     

The gulf of Bothnia: The northernmost part of the Baltic Sea

Summary The Baltic Sea, one of the largest brackish water areas in the world, can be characterized as a young, cold sea containing an impoverished ecosystem due to salinity stress. The present Baltic Sea was formed as late as 2000 to 2500 years ago when the Danish sounds became more narrow and shallow. The inflow of freshwater from the surrounding land areas caused the Baltic to gradually attain its brackish character. Today the Baltic covers an area of some 366,000 km² as a series of basins separated by shallower areas and filled with about 22,000 km³ of brackish water. These basins are, from north to south, the Gulf of Bothnia, the Gulf of Finland, the Gotland Sea and the Bornholm Sea. The climate gradient ranges from almost arctic conditions in the extreme north to a more maritime climate in the southern parts. The North Sea salt water is connected to the Baltic through the shallow Kattegat and the sills in the Danish sounds. The inflow of salt water occurs in two different ways,viz. as a continuous flow along the bottom due to the salinity gradient and as pulses of salt water generated by the distribution of air pressure and the direction of the wind. The freshwater input (500 km³) from mainly the large rivers equals roughly the net outflow and stresses the south-bound current along the Swedish coast that also compensates for the salt water inflow. Tidal movements can be seen in the southern Baltic, but are of minor importance for the system. The residence time of the total water mass is 25 years and the hydrographical conditions within the different basins are stable and dominated by a permanent halocline, and a thermocline developing every spring. The salinity ranges from about 1–2 per mille in the innermost part of the Gulf of Bothnia to 10–15 per mille in the Bornholm Sea. Total vertical mixing takes place during winter in at least the northern parts of the sea. Due to the climate-gradient, the ice condition differs from about four months of total ice-cover in the inner parts of the Gulf of Bothnia to one month or less of coastal ice in the southern part of the Baltic. Thus, the seasonal effect is more pronounced in the northern parts.The living systems of the Baltic are reduced and adapted to these varying conditions. When comparing the deeper soft bottoms of the Gulf of Bothnia to the rest of the Baltic, the following pattern can be seen. The pelagic primary productivity increases by a factor 6 from north to south. The southern parts of the sea show a pronounced spring peak, while in the north the spring development is delayed or replaced by a summer maximum. The total increase of the macrofauna biomass is striking, from about 1 g.m^–2 (w.wt) in the north to 100 g.m^–2 (w.wt) or more in the south. The meiofauna and the zooplankton biomasses show less variability. The meiofauna increases by a factor of 2–4, giving a biomass of about twice that of the macrofauna in the northernmost part. The extremely low salinity of this area causes the exclusion of bivalves (filter-feeders) from the fauna. Available data, pooled with the high metabolic rate of the meiofauna, roughly follow the changes in primary productivity within the Baltic Sea. The changing ratio of macro- to meiofauna, as well as results from intensive studies of the macrobenthic amphipodPontoporeia affinis (Lindström), suggest that the macrofauna is regulated mainly by food limitation and that the benthic and pelagic systems are closely coupled.     

A nursery area for the Antarctic silverfish Pleuragramma antarcticum at Terra Nova Bay (Ross Sea): first estimate of distribution and abundance of eggs and larvae under the seasonal sea-ice

Pleuragramma antarcticum is the dominant pelagic fish in the waters of the continental shelf in high Antarctic regions, where it plays a key role in the food web. A nursery ground for eggs of this species was first identified in 2002 in Terra Nova Bay (Ross Sea), where eggs were found trapped in ice platelets under the sea-ice during the spring. As part of a monitoring program aimed at understanding the geographic and temporal characteristics of this nursery ground, the present study reports on surveys carried out in the austral springs of 2005 and 2006 using a simple and effective method for sampling from the sea-ice. These surveys enabled the evaluation of the spatial range of the nursery area of the Antarctic silverfish in the sea-ice of the coastal area of Victoria Land between the Coulman Island and the Drygalski Glacier Tongue. P. antarcticum eggs were concentrated in an area of Terra Nova Bay of about 270?km², encompassing two adjacent sites, Gerlache Inlet and Silverfish Bay. The present results add information on life cycle and hatching period of the Antarctic silverfish and confirm the importance of the Terra Nova Bay as a nursery area for this important species. Moreover, the survey points to the sea-ice cover and platelet ice as important environmental features of the nursery area.