Trophic relationships of the sea ice meiofauna in Frobisher Bay,Arctic Canada

Summary A distinct fauna consisting mainly of nematodes, harpacticoid and cyclopoid copepods, rotifers, turbellarians and polychaete larvae, inhabits the lower levels of the sea ice in Frobisher Bay. Similar faunas are found throughout circumpolar regions. Thirteen taxa of the Frobisher Bay ice fauna were entirely herbivorous. Their food consisted of 26 genera of algae dominated by Chlamydomonas, Nitzschia, Navicula and Chaetoceros. There was a clear tendency to feed on the most abundant ice algae, hence little evidence of selective feeding. High algal food concentrations in the ice (estimated at 5000 g C/l) were in sharp contrast with the scant nourishment available from phytoplankton under the ice (8 g C/l) from mid-winter until the start of the summer bloom. Algal stocks and estimated productivity rates indicate that ice meiofaunal food requirements may be met by the ice algae. All the major ice meiofaunal species are well adapted to feeding within the ice. All are small enough to enter brine channels and secure particulate prey from surfaces within confined spaces. The ice meiofaunal species are major consumers of the ice algae and therefore important links in the transfer of energy from the ice to pelagic and benthic predators, including fishes, birds and mammals.     

Extracellular enzymes of cold-adapted bacteria from Arctic sea ice,Canada Basin

A total of 338 aerobic heterotrophic bacterial strains were isolated from Arctic sea ice, Canada Basin (77°30′N–80°12′N). The capability of the isolates to produce protease, lipase, amylase, chitinase, β-galactosidase, cellulase and/or agarase was investigated. Isolates that were able to degrade tributyrin, skim milk, starch, lactose and chitin accounted for 71.6, 65.7, 38.5, 31.6 and 16.9% of sea ice strains, respectively. Lipase producers and/or protease producers were phylogenetically widespread among the isolated strains. Starch and/or lactose hydrolytic strains were mainly distributed among Colwellia, Marinomonas, Pseudoalteromonas, Pseudomonas and Shewanella isolates. Pseudoalteromonas tetraodonis, Pseudoalteromonas elyakovii, Bacillus firmus and Janibacter melonis isolates all have the ability to degrade chitin. Only some strains belonging to Pseudoalteromonas genus scored positive for agarase (6) and cellulose (9). The temperature dependences for lipase activities were determined for five psychrophilic and six psychrotolerant bacteria. At low temperatures, the psychrophilic bacterial lipase activity was not significantly higher than psychrotolerant bacterial lipase, though all lipases showed remarkably high activity with 10–36% residual activity at 0°C.     

Denitrification activity and oxygen dynamics in Arctic sea ice

Denitrification and oxygen dynamics were investigated in the sea ice of Franklin Bay (70°N), Canada. These investigations were complemented with measurements of denitrification rates in sea ice from different parts of the Arctic (69°N–85°N). Potential for bacterial denitrification activity (5–194 μmol N m⁻² day⁻¹) and anammox activity (3–5 μmol N m⁻² day⁻¹) in melt water from both first-year and multi-year sea ice was found. These values correspond to 27 and 7%, respectively, of the benthic denitrification and anammox activities in Arctic sediments. Although we report only potential denitrification and anammox rates, we present several indications that active denitrification in sea ice may occur in Franklin Bay (and elsewhere): (1) despite sea ice-algal primary production in the lower sea ice layers, heterotrophic activity resulted in net oxygen consumption in the sea ice of 1–3 μmol l⁻¹ sea ice per day at in situ light conditions, suggesting that O₂ depletion may occur prior to the spring bloom. (2) The ample organic carbon (DOC) and NO₃ ⁻ present in sea ice may support an active denitrification population. (3) Measurements of O₂ conditions in melted sea ice cores showed very low bulk concentrations, and in some cases anoxic conditions prevailed. (4) Laboratory studies using planar optodes for measuring the high-resolution two-dimensional O₂ distributions in sea ice confirmed the very dynamic and heterogeneous O₂ distribution in sea ice, displaying a mosaic of microsites of high and low O₂ concentrations. Brine enclosures and channels were strongly O₂ depleted in actively melting sea ice, and anoxic conditions in parts of the brine system would favour anaerobic processes.     

Seasonal changes in substrate utilization patterns by bacterioplankton in the Amundsen Gulf (western Arctic)

Due to logistic difficulties, biological processes along the Arctic winter remain poorly known. In particular, carbon sources used by bacterioplankton have not been identified. A previous study in Franklin Bay suggested that polymers were one of the main substrates used by bacteria. During the Circumpolar Flaw Lead System Study, we analyzed metabolic capabilities of the heterotrophic bacterioplankton using Biolog MT2 MicroPlates^® amended with custom-selected substrates. Our purpose was to test whether the use of polymers was a peculiarity of Franklin Bay or a robust feature of the Arctic winter community. Seventeen stations were sampled in the Amundsen Gulf (western Arctic), a very dynamic area with heterogeneous ice conditions, from February to July 2008, at the surface (0–12 m), intermediate depths (20–70 m), and near the bottom (200 m). In winter, when nutrients and chlorophyll a concentrations were low, the number of substrates used was close to zero. In early spring, when the levels of chlorophyll a increased, so did the number of substrates used. This was followed by a 1-month period with no substrates used in April and May. Finally, the activity recovered toward the summer. Amino acids were occasionally used. However, the group of substrates most commonly used at all depths was carbohydrates, especially cellobiose, maltose, N-acetyl-d-glucosamine and glycogen. All these are either polymers or monomers derived from polymers. These results confirm that the heterotrophic bacterial assemblage relies heavily on polysaccharides for subsistence during the Arctic winter.     

Respiration and bacterial carbon dynamics in Arctic sea ice

Bacterial carbon demand, an important component of ecosystem dynamics in polar waters and sea ice, is a function of both bacterial production (BP) and respiration (BR). BP has been found to be generally higher in sea ice than underlying waters, but rates of BR and bacterial growth efficiency (BGE) are poorly characterized in sea ice. Using melted ice core incubations, community respiration (CR), BP, and bacterial abundance (BA) were studied in sea ice and at the ice–water interface (IWI) in the Western Canadian Arctic during the spring and summer 2008. CR was converted to BR empirically. BP increased over the season and was on average 22 times higher in sea ice as compared with the IWI. Rates in ice samples were highly variable ranging from 0.2 to 18.3 μg C l⁻¹ d⁻¹. BR was also higher in ice and on average ~10 times higher than BP but was less variable ranging from 2.39 to 22.5 μg C l⁻¹ d⁻¹. Given the high variability in BP and the relatively more stable rates of BR, BP was the main driver of estimated BGE (r ² = 0.97, P < 0.0001). We conclude that microbial respiration can consume a significant proportion of primary production in sea ice and may play an important role in biogenic CO₂ fluxes between the sea ice and atmosphere.     

Population dynamics of bacteria in Arctic sea ice

The dynamics of bacterial populations in annual sea ice were measured throughout the vernal bloom of ice algae near Resolute in the Canadian Arctic. The maximum concentration of bacteria was 6.0·10¹¹ cells·m^–2 (about 2.0·10¹⁰ cells·l^–1) and average cell volume was 0.473 m³ in the lower 4 cm of the ice sheet. On average, 37% of the bacteria were epiphytic and were most commonly attached (70%) to the dominant alga,Nitzschia frigida (58% of total algal numbers). Bacterial population dynamics appeared exponential, and specific growth rates were higher in the early season (0.058 day^–1), when algal biomass was increasing, than in the later season (0.0247 day^–1), when algal biomass was declining. The proportion of epiphytes and the average number of epiphytes per alga increased significantly (P<0.05) through the course of the algal bloom. The net production of bacteria was 67.1 mgC·m^–2 throughout the algal bloom period, of which 45.5 mgC·m^–2 occurred during the phase of declining algal biomass. Net algal production was 1942 mgC·m^–2. Sea ice bacteria (both arctic and antarctic) are more abundant than expected on the basis of relationships between bacterioplankton and chlorophyll concentrations in temperate waters, but ice bacteria biomass and net production are nonetheless small compared with the ice algal blooms that presumably support them.     

The impact of ice melting on bacterioplankton in the Arctic Ocean

Global warming and the associated ice melt are leading to an increase in the organic carbon in the Arctic Ocean. We evaluated the effects of ice melt on bacterioplankton at 21 stations in the Greenland Sea and Arctic Ocean in the summer of 2007, when a historical minimum of Arctic ice coverage was measured. Polar Surface Waters, which have a low temperature and low salinity and originate mainly from melted ice, contained a very low abundance of bacteria (7.01 × 10⁵ ± 2.20 × 10⁵ cells ml⁻¹); however, these bacteria had high specific bacterial production (2.40 ± 1.61 fmol C bac⁻¹ d⁻¹) compared to those in Atlantic Waters. Specifically, bacterioplankton in Polar Surface Waters showed a preference for utilizing carbohydrates and had significantly higher specific activities of the glycosidases assayed, i.e. β-glucosidase, xylosidase, arabinosidase and cellobiosidase. Furthermore, bacterioplankton in Polar Sea Waters showed preferential growth on some of the carbohydrates in the Biolog Ecoplate, such as d-cellobiose and N-acetyl-d-glucosamine. Our results suggest that climate change and the associated melting of Arctic ice might induce changes in bacterioplankton functional diversity by enhancing the turnover of carbohydrates. Since organic aggregates are largely composed of polysaccharides, higher solubilization of aggregates might modify the carbon cycle, weaken the biological pump and have biogeochemical and ecological implications for the future Arctic Ocean.     

Notes on the biology of sea ice in the Arctic and Antarctic

The sea ice which covers large areas of the polar regions plays a major role in the marine ecosystem of both the Arctic and Southern Oceans. Not only do warmblooded animals depend on sea ice as a platform, but the sympagic organisms living internally within the sea ice or at the interfaces ice/snow and ice/water provide a substantial part of the total primary production of the ice covered regions. In addition sea ice organisms are an important food source for a variety of pelagic animals and may initiate phytoplankton spring blooms after ice melt by seeding effects.Sea ice organisms often are enriched by some orders of magnitude if the same volume of melted ice is compared to that of the underlying water column. Three hypotheses try to explain this discrepancy and are discussed. Investigations on the nutrient chemistry within the sea ice system and in-situ observations still are rare. Intense growth of sympagic organisms can result in nutrient deficiencies, at least in selected habitats. Advances in endoscopie methods may lead to a better understanding of the life within the sea ice.Paper presented at the Symposium on Polar regions: the challenge for biological and ecological research organised by the Swiss Committee for Polar Research, Basel on 2 October 1992     

Identification of the sea ice diatom biomarker IP25 in Arctic benthic macrofauna: direct evidence for a sea ice diatom diet in Arctic heterotrophs

Currently, the impact of declining seasonal sea ice extent in the Arctic on polar food webs remains uncertain. Previously, a range of proxy techniques has been employed to determine links between sea ice or phytoplankton primary production and the Arctic marine food web, although it is accepted that such approaches have their limitations. Here, we propose a novel approach to tracing sea ice primary production through Arctic food webs using the sea ice diatom biomarker, IP₂₅. Various benthic macrofaunal specimens were collected between March and May 2008 from Franklin Bay in the Amundsen Gulf, Arctic Canada, as part of the International Polar Year–Circumpolar Flaw Lead system study. Each specimen was analysed for the presence of the sea ice diatom biomarker IP₂₅ in order to provide evidence for feeding by benthic organisms on sea ice algae. IP₂₅ was found in nineteen out of the twenty-one specimens analysed, often as the most abundant of the highly branched isoprenoid biomarkers detected. The stable isotope composition of IP₂₅ (δ¹³C = −17.1 ± 0.5‰) in the sea urchin (Strongylocentrotus sp.) specimens was similar to that reported previously for this biomarker in Arctic sea ice, sedimenting particles and sediments. It is concluded that detection of IP₂₅ in Arctic benthic macrofauna represents a novel approach to providing convincing evidence for feeding on sea ice algae. It is also proposed that analysis of IP₂₅ may be used to trace trophic transfer of sea ice algal-derived organic matter through Arctic food webs in the future.     

Biogenic silica recycling in sea ice inferred from Si-isotopes: constraints from Arctic winter first-year sea ice

We report silicon isotopic composition (δ³⁰Si vs. NBS28) in Arctic sea ice, based on sampling of silicic acid from both brine and seawater in a small Greenlandic bay in March 2010. Our measurements show that just before the productive period, δ³⁰Si of sea-ice brine similar to δ³⁰Si of the underlying seawater. Hence, there is no Si isotopic fractionation during sea-ice growth by physical processes such as brine convection. This finding brings credit and support to the conclusions of previous work on the impact of biogenic processes on sea ice δ³⁰Si: any δ³⁰Si change results from a combination of biogenic silica production and dissolution. We use this insight to interpret data from an earlier study of sea-ice δ³⁰Si in Antarctic pack ice that show a large accumulation of biogenic silica. Based on these data, we estimate a significant contribution of biogenic silica dissolution (D) to production (P), with a D:P ratio between 0.4 and 0.9. This finding has significant implications for the understanding and parameterization of the sea ice Si-biogeochemical cycle, i.e. previous studies assumed little or no biogenic silica dissolution in sea ice.     

In-situ observations on the distribution and behavior of amphipods and Arctic cod (Boreogadus saida) under the sea ice of the High Arctic Canada Basin

The occurrence and behavior of sympagic amphipods and Arctic cod (Boreogadus saida) were studied in the High Arctic Canada Basin by diving under the ice at seven stations in summer 2002. Still images of video-transects were used to obtain animal abundances and information on the structure of the ice environment. Mean amphipod abundances for the stations varied between 1 and 23 individuals m^–2, with an increase towards the western part of the basin. The standard deviation within the 31–51 images analyzed per station was small (<1 individual m^–2). Gammarus wilkitzkii was found in low abundances, often hiding in small ice gaps. Small amphipods (Onisimus spp., Apherusa glacialis, and juveniles of all species) tended to move freely along the bottom of the floes. B. saida occurred in narrow wedges of seawater along the edges of melting ice floes at three stations in water depths of 10–50 cm and was never found under the ice. The fish occurred in schools of 1–28 per wedge. Fish were inactive and did not escape the approaching diver. Resting in the wedges may be a strategy to reduce energetic requirements and avoid predators.We dedicate this publication to Professor Dr. J. Lenz (Kiel University).     

Protists in Arctic drift and land‐fast sea ice

Global climate change is having profound impacts on polar ice with changes in the duration and extent of both land‐fast ice and drift ice, which is part of the polar ice pack. Sea ice is a distinct habitat and the morphologically identifiable sympagic community living within sea ice can be readily distinguished from pelagic species. Sympagic metazoa and diatoms have been studied extensively since they can be identified using microscopy techniques. However, non‐diatom eukaryotic cells living in ice have received much less attention despite taxa such as the dinoflagellate Polarella and the cercozoan Cryothecomonas being isolated from sea ice. Other small flagellates have also been reported, suggesting complex microbial food webs. Since smaller flagellates are fragile, often poorly preserved, and are difficult for non‐experts to identify, we applied high throughput tag sequencing of the V4 region of the 18S rRNA gene to investigate the eukaryotic microbiome within the ice. The sea ice communities were diverse (190 taxa) and included many heterotrophic and mixotrophic species. Dinoflagellates (43 taxa), diatoms (29 taxa) and cercozoans (12 taxa) accounted for ~80% of the sequences. The sympagic communities living within drift ice and land‐fast ice harbored taxonomically distinct communities and we highlight specific taxa of dinoflagellates and diatoms that may be indicators of land‐fast and drift ice.     

Organism losses during ice melting: A serious bias in sea ice community studies

Summary When ice samples are melted, microorganisms living within the brine inclusions are subjected to rapid and extreme changes in salinities. This procedure results in substantial losses of flagellates and ciliates. Most of these losses can be prevented if ice samples are melted in larger volumes of sterile sea water to buffer salinity and osmotic changes. Since most studies on the ice biota have ignored, or have been unable to avoid this bias, current views of the composition and activity of sea ice communities are based on assemblages over-representing organisms with rigid cell material.     

The role of sea ice for vascular plant dispersal in the Arctic

Sea ice has been suggested to be an important factor for dispersal of vascular plants in the Arctic. To assess its role for postglacial colonization in the North Atlantic region, we compiled data on the first Late Glacial to Holocene occurrence of vascular plant species in East Greenland, Iceland, the Faroe Islands and Svalbard. For each record, we reconstructed likely past dispersal events using data on species distributions and genetics. We compared these data to sea-ice reconstructions to evaluate the potential role of sea ice in these past colonization events and finally evaluated these results using a compilation of driftwood records as an independent source of evidence that sea ice can disperse biological material. Our results show that sea ice was, in general, more prevalent along the most likely dispersal routes at times of assumed first colonization than along other possible routes. Also, driftwood is frequently dispersed in regions that have sea ice today. Thus, sea ice may act as an important dispersal agent. Melting sea ice may hamper future dispersal of Arctic plants and thereby cause more genetic differentiation. It may also limit the northwards expansion of competing boreal species, and hence favour the persistence of Arctic species.     

Organism incorporation into newly forming Arctic sea ice in the Greenland Sea

New ice formation, protist incorporation and enrichment in differentstages of young Arctic sea ice (grease,nilas and pancake ice)were studied in the Greenland Sea in autumn 1995. Nutrients(nitrite, nitrate, phosphateand silicate), salinity and abundanceestimates of organisms were analysed from surface water andnew ice samples. The abundances of bacteria, diatoms, and photo-and hetero trophic flagellates in the ice and water column weredetermined using epifluorescence microscopy. An enrichment indexwas calculated to compare the abundance of organisms in thewater column with different stages of young sea ice. The resultsclearly show that (i) protist incorporation already begins duringthe first stages of new sea ice formation, (ii) incorporationof protists is selective, showing preference for diatoms witha relatively large cell size and (iii) enrichment of organisms,in particular diatoms, takes place in young sea ice in the GreenlandSea. The selectivity of the incorpor ation process and the evidentpreference for diatoms are presumably a result of the largercell size and/orcertain properties of the cell surface (e.g.stickiness) that enhance their incorporation. The calculatedenrichment indices were relatively low for bacteria and flagellates.     

Vertical distribution of bacteria in Arctic sea ice from the Barents and Laptev Seas

The vertical distribution of bacterial abundance and biomass was investigated in relation to algal biomass in ice cores taken from drifting ice floes in two Arctic shelf areas: the Barents Sea and the Laptev Sea. Bacteria were not homogeneously distributed throughout the cores but occurred in dense layers. Different types of distribution patterns were found: either a single maximum occurred inside or at the bottom of the ice floe or maxima were found in different parts of the floes. Bacterial concentrations ranged from 0.4 to 36.7 · 10⁵ cells ml⁻¹. The size spectra of sea-ice bacteria were determined by image analysis. Cell sizes showed considerable variation between the ice floes. In multi-year sea ice, the largest bacteria were observed in the area of an internal chlorophyll a maximum. No specific vertical distribution patterns were found in first-year ice floes. Bacterial biomass for the ice cores ranged from 19.2 to 79.2 mg C m⁻², and the ratio of bacterial:ice algal biomass ranged from 0.43 to 10.42. A comparison with data collected from fast ice revealed large differences in terms of cell size, abundance and biomass. Received: 7 September 1995 / Accepted: 10 September 1996     

Persistence of bacterial and archaeal communities in sea ice through an Arctic winter

The structure of bacterial communities in first‐year spring and summer sea ice differs from that in source seawaters, suggesting selection during ice formation in autumn or taxon‐specific mortality in the ice during winter. We tested these hypotheses by weekly sampling (January–March 2004) of first‐year winter sea ice (Franklin Bay, Western Arctic) that experienced temperatures from ?9°C to ?26°C, generating community fingerprints and clone libraries for Bacteria and Archaea. Despite severe conditions and significant decreases in microbial abundance, no significant changes in richness or community structure were detected in the ice. Communities of Bacteria and Archaea in the ice, as in under‐ice seawater, were dominated by SAR11 clade Alphaproteobacteria and Marine Group I Crenarchaeota, neither of which is known from later season sea ice. The bacterial ice library contained clones of Gammaproteobacteria from oligotrophic seawater clades (e.g. OM60, OM182) but no clones from gammaproteobacterial genera commonly detected in later season sea ice by similar methods (e.g. Colwellia, Psychrobacter). The only common sea ice bacterial genus detected in winter ice was Polaribacter. Overall, selection during ice formation and mortality during winter appear to play minor roles in the process of microbial succession that leads to distinctive spring and summer sea ice communities.     

Isolation and characterization of marine psychrophilic phage-host systems from Arctic sea ice

Phage-host systems from extreme cold environments have rarely been surveyed. This study is concerned with the isolation and characterization of three different phage-host systems from Arctic sea ice and melt pond samples collected north-west of Svalbard (Arctic). On the basis of 16S rDNA sequences, the three bacterial phage hosts exhibited the greatest similarity to the species Shewanella frigidimarina (96.0%), Flavobacterium hibernum (94.0%), and Colwellia psychrerythraea (98.4%), respectively. The host bacteria are psychrophilic with good growth at 0°C, resulting in a rapid formation of visible colonies at this temperature. The phages showed an even more pronounced adaptation to cold temperatures than the bacteria, with growth maxima below 14°C and good plaque formation at 0°C. Transmission electron microscopy (TEM) examinations revealed that the bacteriophages belonged to the tailed, double-stranded DNA phage families Siphoviridae and Myoviridae. All three phages were host-specific.Communicated by K. Horikoshi     

Diversity and cold-active hydrolytic enzymes of culturable bacteria associated with Arctic sea ice,Spitzbergen

The diversity of culturable bacteria associated with sea ice from four permanently cold fjords of Spitzbergen, Arctic Ocean, was investigated. A total of 116 psychrophilic and psychrotolerant strains were isolated under aerobic conditions at 4°C. The isolates were grouped using amplified rDNA restriction analysis fingerprinting and identified by partial sequencing of 16S rRNA gene. The bacterial isolates fell in five phylogenetic groups: subclasses and of Proteobacteria, the Bacillus–Clostridium group, the order Actinomycetales, and the Cytophaga–Flexibacter–Bacteroides (CFB) phylum. Over 70% of the isolates were affiliated with the Proteobacteria subclass. Based on phylogenetic analysis (<98% sequence similarity), over 40% of Arctic isolates represent potentially novel species or genera. Most of the isolates were psychrotolerant and grew optimally between 20 and 25°C. Only a few strains were psychrophilic, with an optimal growth at 10–15°C. The majority of the bacterial strains were able to secrete a broad range of cold-active hydrolytic enzymes into the medium at a cultivation temperature of 4°C. The isolates that are able to degrade proteins (skim milk, casein), lipids (olive oil), and polysaccharides (starch, pectin) account for, respectively, 56, 31, and 21% of sea-ice and seawater strains. The temperature dependences for enzyme production during growth and enzymatic activity were determined for two selected enzymes, -amylase and -galactosidase. Interestingly, high levels of enzyme productions were measured at growth temperatures between 4 and 10°C, and almost no production was detected at higher temperatures (20–30°C). Catalytic activity was detected even below the freezing point of water (at –5°C), demonstrating the unique properties of these enzymes.