An ice-binding protein from an Antarctic sea ice bacterium

An Antarctic sea ice bacterium of the Gram-negative genus Colwellia, strain SLW05, produces an extracellular substance that changes the morphology of growing ice. The active substance was identified as a approximately 25-kDa protein that was purified through its affinity for ice. The full gene sequence was determined and was found to encode a 253-amino acid protein with a calculated molecular mass of 26,350 Da. The predicted amino acid sequence is similar to predicted sequences of ice-binding proteins recently found in two species of sea ice diatoms and a species of snow mold. A recombinant ice-binding protein showed ice-binding activity and ice recrystallization inhibition activity. The protein is much smaller than bacterial ice-nucleating proteins and antifreeze proteins that have been previously described. The function of the protein is unknown but it may act as an ice recrystallization inhibitor to protect membranes in the frozen state.     

Biodiversity of gas vacuolate bacteria from Antarctic sea ice and water.

Psychrophilic, gas vacuolate, heterotrophic bacteria indigenous to sea ice communities in Antarctica have been isolated. Phylogenetic analysis of representative members of these bacteria shows that they belong to the alpha, beta, and gamma Proteobacteria and the Flavobacteria-Cytophaga group. This is the first report of gas vacuolate bacteria from the beta Proteobacteria and the Flavobacteria-Cytophaga groups.     

In situ primary production in young Antarctic sea ice

An in situ incubation technique used successfully to measure the photosynthetic carbon assimilation of internal algal assemblages within thick multiyear Arctic ice was developed and improved to measure the photosynthetic carbon assimilation within young sea ice only 50 cm thick (Eastern Weddell Sea, Antarctica). The light transmission was improved by the construction of a cylindrical frame instead of using a transparent acrylic-glass barrel. The new device enabled some of the first precise measurements of in situ photosynthetic carbon assimilation in newly formed Antarctic sea ice, which is an important component in the sea ice ecosystem of the Antarctic Ocean. The rates of carbon assimilation of the interior algal assemblage (top to 5 cm from bottom) was 0.25 mg C m^–2 d^–1 whereas the bottom algal community (lowest 5 cm) attained only 0.02 mg C m^–2 d^–1. Chl a specific production rates (P^Chl) for bottom algae (0.020 – 0.056 g C g chl a ^–1 h^–1) revealed strong light limitation, whereas the interior algae (P^Chl = 0.7 – 1.2 g C g chl a ^–1 h^–1) were probably more limited by low temperatures (< –5 °C) and high brine salinities.     

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     

Antarctic sea ice viral dynamics over an annual cycle

Viral abundance, burst sizes, lytic production and temperate phage were investigated in land-fast ice at two sites in Prydz Bay Antarctica (68°S, 77°E) between April and November 2008. Both ice cores and brine were collected. There was no seasonal pattern in viral or bacterial numbers. Across the two sites virus abundances ranged between 0.5 × 10⁵ and 5.1 × 10⁵ viruses ml⁻¹ in melted ice cores and 0.6 × 10⁵–3.5 × 10⁵ viruses ml⁻¹ in brine, and bacterial abundances between 2.7 × 10⁴ and 17.3 × 10⁴ cells ml⁻¹ in melted ice cores and 3.9 × 10⁴–32.5 × 10⁴ cells ml⁻¹ in brine. Virus to bacterium ratios (VBR) showed a clear seasonal pattern in ice cores with lowest values in winter (range 1.2–20.8), while VBRs in brine were lower (0.2–4.9). Lytic viral production range from undetectable to 2.0 × 10⁴ viruses ml⁻¹ h⁻¹ in ice cores with maximum rates in September and November. In brine maximum, lytic viral production occurred in November (1.18 × 10⁴ viruses ml⁻¹ h⁻¹). Low burst sizes were typical (3.94–4.03 viruses per bacterium in ice cores and 3.16–4.0 viruses per bacterium in brine) with unusually high levels of visibly infected cells—range 40–50%. This long-term investigation revealed that viral activity was apparent within the sea ice throughout its annual cycle. The findings are discussed within the context of limited data available on viruses in sea ice.     

The biota of Antarctic pack ice in the Weddell sea and Antarctic Peninsula regions

Summary Pack ice surrounding Antarctica supports rich and varied populations of microbial organisms. As part of the Antarctic Marine Ecosystem Research in the Ice Edge Zone (AMERIEZ) studies, we have examined this community during the late spring, autumn, and winter. Although organisms are found throughout the ice, the richest concentrations often occur in the surface layer. The ice flora consists of diatoms and flagellates. Chrysophyte cysts (archaeomonads) of unknown affinity and dinoflagellate cysts are abundant and may serve as overwintering stages in ice. The ice fauna includes a variety of heterotrophic flagellates, ciliates, and micrometazoa. The abundance of heterotrophs indicates an active food web within the ice community. Ice may serve as a temporary habitat or refuge for many of the microbial forms and some of these appear to provide an inoculum for planktonic populations when ice melts. Larger consumers, such as copepods and the Antarctic krill, Euphausia superba are often found on the underside of ice floes and within weathered floes. The importance of the ice biota as a food resource for these pelagic consumers is unknown.     

Release of an ice-active substance by Antarctic sea ice diatoms

Interstitial water from the diatom-rich ice platelet layer in McMurdo Sound, Antarctica contains a macromolecular, ice-active substance (IAS) that, at in situ concentrations, causes dense pitting on the basal surfaces of growing ice platelets. In this respect, it resembles several fish antifreezes that also cause pitting on ice surfaces, but unlike the antifreezes, it does not lower the freezing point. The IAS appeared to be released by diatoms, as extracts from the diatoms contained IAS, while seawater from a diatom-free area did not. No evidence of IAS was found in several species of temperate water diatoms. The ice-pitting activity of the IAS was destroyed by proteases and by incubation at 40° C, but not by periodate oxidation, or by incubation with galactosidase or endonuclease. Thus, activity appears to arise from a protein or protein component, and not from carbohydrate or nucleic acids. Potential roles of the IAS in the sea ice community are discussed.     

Seasonal changes of Antarctic marine bacterioplankton and sea ice bacterial assemblages

The distributions of bacterial populations in sea ice and underlying seawater were investigated on the continental shelf of the “Terre Adélie” area. A reference station was sampled weekly from January 1991 to January 1992. In winter, the survey included a minimum of six sampling layers: surface and bottom ice, brine, seawater from the interface, and at 0.5 and 2 m depth. In seawater, the total bacterial abundance ranged from 0.5 × 10⁵ cells ml⁻¹ in July to 6.0 × 10⁵ cells ml⁻¹ after ice break. Values reaching 2.5 × 10⁶ cells ml⁻¹ were recorded in the overlying ice cover. Mean cell volumes were twice as high in brine as in seawater. The saprophytic bacterial abundance ranged from 5.0 × 10⁴ CFU (colony-forming units) ml⁻¹ in some winter interface samples to less than 1.0 × 10³ CFU ml⁻¹ in most of the summer seawater samples. In sea ice a clear decreasing gradient for most of the studied bacterial parameters from the surface layers towards the bottom layer was found. The ice cover had a discernible impact on underlying seawater, but its influence was restricted to a limited interface layer.     

Physical parameters influencing diatom community structure in eastern Antarctic sea ice

Diatom assemblages obtained from fast ice around Prydz Bay, Antarctica, are distinctly different from those obtained from pack ice in the same area. The dominant species in all ice cores were Fragilariopsis curta, F. cylindrus, Nitzschia stellata and Pseudonitzschia turgiduloides. Entomoneis kjellmanii and Cocconeis spp. were more characteristic of fast ice samples and F. curta of pack ice samples. Ice crystal type (i.e. whether frazil or congelation crystal) is an important factor determining the algal composition of the ice. Other significant influences include the time of year the ice forms and the salinity of the ice.     

Diversity and association of psychrophilic bacteria in Antarctic sea ice.

The bacterial populations associated with sea ice sampled from Antarctic coastal areas were investigated by use of a phenotypic approach and a phylogenetic approach based on genes encoding 16S rRNA (16S rDNA). The diversity of bacteria associated with sea ice was also compared with the bacterial diversity of seawater underlying sea ice. Psychrophilic (optimal growth temperature, < or = 15 degrees C; no growth occurring at 20 degrees C) bacterial diversity was found to be significantly enriched in sea ice samples possessing platelet and bottom ice diatom assemblages, with 2 to 9 distinct (average, 5.6 +/- 1.8) psychrophilic taxa isolated per sample. Substantially fewer psychrophilic isolates were recovered from ice cores with a low or negligible population of ice diatoms or from under-ice seawater samples (less than one distinct taxon isolated per sample). In addition, psychrophilic taxa that were isolated from under-ice seawater samples were in general phylogenetically distinct from psychrophilic taxa isolated from sea ice cores. The taxonomic distributions of psychrotrophic bacterial isolates (optimal growth temperature, > 20 degrees C; growth can occur at approximately 4 degrees C) isolated from sea ice cores and under-ice seawater were quite similar. Overall, bacterial isolates from Antarctic sea ice were found to belong to four phylogenetic groups, the alpha and gamma subdivisions of the Proteobacteria, the gram-positive branch, and the Flexibacter-Bacteroides-Cytophaga phylum. Most of the sea ice strains examined appeared to be novel taxa based on phylogenetic comparisons, with 45% of the strains being psychrophilic. 16S rDNA sequence analysis revealed that psychrophilic strains belonged to the genera Colwellia, Shewanella, Marinobacter, Planococcus, and novel phylogenetic lineages adjacent to Colwellia and Alteromonas and within the Flexibacter-Bacteroides-Cytophaga phylum. Psychrotrophic strains were found to be members of the genera Pseudoalteromonas, Psychrobacter, Halomonas, Pseudomonas, Hyphomonas, Sphingomonas, Arthrobacter, Planococcus, and Halobacillus. From this survey, it is proposed that ice diatom assemblages provide niches conducive to the proliferation of a diverse array of psychrophilic bacterial species.     

Contribution of sea ice microbial production to Antarctic benthic communities is driven by sea ice dynamics and composition of functional guilds

Organic matter produced by the sea ice microbial community (SIMCo) is an important link between sea ice dynamics and secondary production in near‐shore food webs of Antarctica. Sea ice conditions in McMurdo Sound were quantified from time series of MODIS satellite images for Sept. 1 through Feb. 28 of 2007–2015. A predictable sea ice persistence gradient along the length of the Sound and evidence for a distinct change in sea ice dynamics in 2011 were observed. We used stable isotope analysis (δ¹³C and δ¹⁵N) of SIMCo, suspended particulate organic matter (SPOM) and shallow water (10–20 m) macroinvertebrates to reveal patterns in trophic structure of, and incorporation of organic matter from SIMCo into, benthic communities at eight sites distributed along the sea ice persistence gradient. Mass‐balance analysis revealed distinct trophic architecture among communities and large fluxes of SIMCo into the near‐shore food web, with the estimates ranging from 2 to 84% of organic matter derived from SIMCo for individual species. Analysis of patterns in density, and biomass of macroinvertebrate communities among sites allowed us to model net incorporation of organic matter from SIMCo, in terms of biomass per unit area (g/m²), into benthic communities. Here, organic matter derived from SIMCo supported 39 to 71 per cent of total biomass. Furthermore, for six species, we observed declines in contribution of SIMCo between years with persistent sea ice (2008–2009) and years with extensive sea ice breakout (2012–2015). Our data demonstrate the vital role of SIMCo in ecosystem function in Antarctica and strong linkages between sea ice dynamics and near‐shore secondary productivity. These results have important implications for our understanding of how benthic communities will respond to changes in sea ice dynamics associated with climate change and highlight the important role of shallow water macroinvertebrate communities as sentinels of change for the Antarctic marine ecosystem.     

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     

Turbellaria (Archoophora: Acoela) from Antarctic sea ice endofauna – examination of their micromorphology

Acoel Turbellaria constitute a regular component of the metazoa populating Antarctic sea ice (sea-ice endofauna). Two species were collected, which differ in colour, size, shape and egg spawning season. They do not resemble any known pelagic species. Their small body diameter of less than 300 μm allows them to penetrate deeply into the network of brine channels. Their vertical distribution within one ice floe was positively correlated with the accumulation of algal biomass; maxima for both parameters were found in the bottom 5 cm of the floe. The method by which the Turbellaria invade the sea ice is not clear. At present we have no indication that they pass through a pelagic or benthic stage in their life-cycle. As the Turbellaria were found to populate sea ice in areas with water depths ranging from 370 to 4450 m, the presence of benthic phases in their life-cycle, either free-living or epizooic, is not very probable. We suggest that the Turbellaria either use migrating invertebrates as a vector for their propagation or pass through a pelagic stage in their life-cycle. Accepted: 14 December 1998     

Effects of Antarctic sea ice biota on seeding as studied in aquarium experiments

Summary The potential seeding impact of sea ice microbial communities was studied during late austral winter early spring 1988 in the Weddell Sea, Antarctica. Experiments were performed in seawater aquariums with natural seawater and seawater enriched with crushed ice. Algal, protozoan and bacterial cell numbers were followed, as well as nutrients and DOC levels. The results showed a potential seeding effect of sea ice communities to the water column. However, the type of ice communities differed greatly from each other and the effect of such seeding will be patchy. In our experiments seeding of seawater by ice rich in algae, flagellates and/or particulate organic carbon lead to the development of communities dominated either by diatoms or bacteria.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

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.     

Purification and biochemical characterization of a cold-active lipase from Antarctic sea ice bacteria Pseudoalteromonas sp. NJ 70

An extracellular cold-active lipase from Antarctic sea ice bacteria Pseudoalteromonas sp. NJ 70 was purified and characterized. The overall purification based on lipase activity was 27.5-fold with a yield of 25.4 %. The purified lipase showed as a single band on SDS-PAGE with an apparent molecular weight of 37 kDa. The optimum temperature and pH were 35 °C and 7.0, respectively. The lipase activity was enhanced by Ca(2+) and Mg(2+), while was partially inhibited by other metals such as Cu(2+), Zn(2+), Ba(2+), Pb(2+), Fe(2+) and Mn(2+). The lipase had high tolerance to a wide range of NaCl concentrations (0-2 M NaCl). It exhibited high levels of activity in the presence of DTT, Thiourea, H(2)O(2) as well as in the presence of various detergents such as Span 60, Tween-80, Triton X-100. In addition, the lipase showed a preference for long-chain p-nitrophenyl esters (C(12)-C(18)). These results indicated that this lipase could be a novel cold-active lipase.     

Effect of melting Antarctic sea ice on the fate of microbial communities studied in microcosms

Although algal growth in the iron-deficient Southern Ocean surface waters is generally low, there is considerable evidence that winter sea ice contains high amounts of iron and organic matter leading to ice-edge blooms during austral spring. We used field observations and ship-based microcosm experiments to study the effect of the seeding by sea ice microorganisms, and the fertilization by organic matter and iron on the planktonic community at the onset of spring/summer in the Weddell Sea. Pack ice was a major source of autotrophs resulting in a ninefold to 27-fold increase in the sea ice-fertilized seawater microcosm compared to the ice-free seawater microcosm. However, heterotrophs were released in lower numbers (only a 2- to 6-fold increase). Pack ice was also an important source of dissolved organic matter for the planktonic community. Small algae (<10 μm) and bacteria released from melting sea ice were able to thrive in seawater. Field observations show that the supply of iron from melting sea ice had occurred well before our arrival onsite, and the supply of iron to the microcosms was therefore low. We finally ran a “sequential melting” experiment to monitor the release of ice constituents in seawater. Brine drainage occurred first and was associated with the release of dissolved elements (salts, dissolved organic carbon and dissolved iron). Particulate organic carbon and particulate iron were released with low-salinity waters at a later stage.     

Mycosporine-like amino acids in Antarctic sea ice algae,and their response to UVB radiation

Mycosporine like amino acids (MAAs) were detected in low concentration in sea ice algae growing in situ at Cape Evans, Antarctica. Four areas of sea ice were covered with plastics of different UV absorption exposing the bottom- ice algal community to a range of UV doses for a period of 15 days. Algae were exposed to visible radiation only; visible + UV radiation; and visible + enhanced UV radiation. MAA content per cell at the start of the experiment was low in snow-covered plots but higher in samples from ice with no snow cover. During the study period, the MAA content per cell reduced in all treatments, but the rate of this decline was less under both ambient UV and visible radiation than under snow covered plots. While low doses of UVB radiation may have stimulated some MAA production (or at least slowed its loss), relatively high doses of UVB radiation resulted in almost complete loss of MAAs from ice algal cells. Despite this reduction in MAA content per cell, the diatoms in all samples grew well, and there was no discernible effect on viability. This suggests that MAAs may play a minor role as photoprotectants in sea ice algae. The unique structure of the bottom ice algal community may provide a self-shading effect such that algal cells closest to the surface of the ice contain more MAAs than those below them and confer a degree of protection on the community as a whole.