Ice binding,recrystallization inhibition,and cryoprotective properties of ice-active substances associated with Antarctic sea ice diatoms

Extracellular macromolecules associated with Antarctic sea ice diatoms were previously shown to have ice-binding activities. The function of these ice-active substances (IASs) has not been identified. Here we show that two of the IASs have a strong ability to inhibit the recrystallization of ice, possibly signifying a cryoprotectant function. To test this possibility, two species of marine diatom (one Antarctic and one temperate) were subjected to a single freeze-thaw cycle (approximately 20h at -4 or -5 degrees C) in the presence or absence of IAS. Viability, based on a double staining technique, was 15-29% higher in the presence of IAS. Etching of single crystal ice hemispheres grown from dilute IAS solutions indicated that the IASs bind to specific faces of ice and are incorporated into the ice lattice. Together, these results suggest that the IASs acts as a cryoprotectant, probably through some ice-binding mechanism.     

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     

Semipurification and ice recrystallization inhibition activity of ice-active substances associated with Antarctic photosynthetic organisms.

Ice-active substances (IASs), i.e., macromolecular substances that modify the shape of growing ice crystals, were previously found to be associated with various terrestrial and aquatic photosynthetic organisms from Antarctica, but their chemical nature and function are unknown. In this study, we used the ice-binding properties of the IASs to semipurify IASs from a cyanobacterial mat, a eukaryotic green alga (Prasiola sp.), and a moss (Bryum sp.) and examined the ice recrystallization inhibition (RI) activities of the semipure materials. The semipure materials contain both protein and carbohydrate in which the carbohydrate accounted for 73, 52, and 37%, respectively, of the total carbohydrate + protein. The IASs had RI activity at concentrations of 1.4, 0.05, and 0.01 microg ml-1, respectively. RI activity was greatly reduced by heat treatment, suggesting that the IASs inhibit recrystallization through a specific interaction with ice. These results raise the possibility that the IASs increase freezing tolerance of their respective organisms by preventing the recrystallization of ice.     

Increased Land Use by Chukchi Sea Polar Bears in Relation to Changing Sea Ice Conditions

Recent observations suggest that polar bears (Ursus maritimus) are increasingly using land habitats in some parts of their range, where they have minimal access to their preferred prey, likely in response to loss of their sea ice habitat associated with climatic warming. We used location data from female polar bears fit with satellite radio collars to compare land use patterns in the Chukchi Sea between two periods (1986–1995 and 2008–2013) when substantial summer sea-ice loss occurred. In both time periods, polar bears predominantly occupied sea-ice, although land was used during the summer sea-ice retreat and during the winter for maternal denning. However, the proportion of bears on land for > 7 days between August and October increased between the two periods from 20.0% to 38.9%, and the average duration on land increased by 30 days. The majority of bears that used land in the summer and for denning came to Wrangel and Herald Islands (Russia), highlighting the importance of these northernmost land habitats to Chukchi Sea polar bears. Where bears summered and denned, and how long they spent there, was related to the timing and duration of sea ice retreat. Our results are consistent with other studies supporting increased land use as a common response of polar bears to sea-ice loss. Implications of increased land use for Chukchi Sea polar bears are unclear, because a recent study observed no change in body condition or reproductive indices between the two periods considered here. This result suggests that the ecology of this region may provide a degree of resilience to sea ice loss. However, projections of continued sea ice loss suggest that polar bears in the Chukchi Sea and other parts of the Arctic may increasingly use land habitats in the future, which has the potential to increase nutritional stress and human-polar bear interactions.     

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     

The impact of sea ice on the initiation of the spring bloom on the Newfoundland and Labrador Shelves

The relationship between sea ice and the phytoplankton springbloom over the Newfoundland and Labrador shelves is examinedusing remotely-sensed chlorophyll data and sea-ice data forthe period 1998–2004. A regression analysis between thetwo data sets shows that the retreat of sea ice precedes thespring bloom, and the inter-annual variation of the spring bloomis closely correlated with the start time of ice retreat. Thespring bloom off Canada's east coast usually starts on the easternGrand Banks. Here, the water properties are strongly influencedby sea ice on the Newfoundland shelves in early spring whenaccelerated ice melting causes the ice edge to retreat northand the melt water is advected south by the Labrador Current.After the ice retreat, the water on the eastern Grand Banksis rapidly stratified and the mixed layer shallows as a resultof surface freshening. The shallow mixed layer promotes phytoplanktongrowth. The regression analysis also reveals that an early springbloom or ice retreat tends to prolong the duration of the springbloom.     

Movements and behavior of satellite-tagged spotted seals (Phoca largha) in the Bering and Chukchi Seas

Satellite-linked tags were attached to 12 spotted seals (Phoca largha) captured at a coastal lagoon in the eastern Chukchi Sea during August 1991–1993. Movements of seals were tracked for 32–298 days using the Argos system. Of 9,651 total location records obtained, 7,268 were usable. Individual seals were located on 41–96% of the days that tags were operational. During August–November, tagged seals alternated haul-outs at coastal sites lasting 1–304 h with trips to sea of 14–901 h. Coastal haul-outs occurred at 14 sites in western Alaska and eastern Russia. On several trips to sea, seals covered distances of more than 1,000 km. Movement southward from the Chukchi Sea generally began in October, with most of the seals passing through the Bering Strait during November. Seals first hauled out on sea ice in October (Chukchi Sea) or November (Bering Sea), and generally moved southward during October–December as sea-ice coverage increased. Seven seals, whose transmitters were still operating, spent December to June in the Bering Sea region between Kuskokwim Bay and Anadyr Gulf, which corresponded to the location of the ice front. The seals made active east-west movements within the ice front. Spotted seals are unlike other ice-breeding seals in that they regularly use coastal haul-outs during summer and autumn. Compared to the closely related Pacific harbor seal (Phoca vitulina richardsi), spotted seals make much longer trips to sea and spend longer continuous periods at their haul-outs during summer and autumn. Received: 9 April 1997 / Accepted: 30 September 1997     

Infiltration phyto- and protozooplankton assemblages in the annual sea ice of Disko Island, West Greenland, spring 1996

During the spring of 1996 we occupied a station on annual sea ice located several kilometers from Disko Island, West Greenland in water depths greater than 200 m. The goal of this 3-week field season was to characterize sea-ice communities and the underlying water column prior to, and during, ice break-up. A heavier than usual snow load depressed the sea ice below sea level and the snow-ice interface became flooded. Some of this flooded region subsequently refroze and the whole process repeated itself when additional snow accumulated. The infiltration phytoplankton and protozooplankton assemblages that developed in this region were abundant and diverse. Algal biomass in the infiltration layer was approximately an order of magnitude greater than in the underlying water column but an order of magnitude less than in the well-developed bottom ice community. The infiltration autotrophic assemblage resembled the bottom-ice assemblage while the protozooplankton assemblage was more similar to the water column assemblage. Received: 13 February 1998 / Accepted: 30 May 1998     

Comparison of wintertime eukaryotic community from sea ice and open water in the Baltic Sea, based on sequencing of the 18S rRNA gene

The Baltic Sea is one of the world’s largest brackish water basins and is traditionally considered to be species poor. Here, we assessed the diversity of the nano-sized eukaryotic microbial wintertime community, using molecular ecological methods based on sequencing of small-subunit ribosomal RNA gene clone libraries. The results demonstrate that a rich community of small eukaryotes inhabits the Baltic Sea ice and water during winter. The community was dominated by alveolates and stramenopiles. Ciliates and cercozoans were the richest groups present, while in contrast to previous studies, diatoms showed a lower richness than expected. Furthermore, fungi and parasitic Syndiniales were present both in the water and in the sea ice. Some of the organisms in the sea-ice community were active, based on the RNA data, but a number of organisms were inactive or remnants from the freezing process. The results demonstrate that the sea-ice communities in the Baltic Sea are highly diverse and that water and ice of different ages include different protistan assemblages. Our study emphasizes the potential loss in biodiversity through diminishing ice cover as a result of climate change.     

Acquisition of freeze protection in a sea-ice crustacean through horizontal gene transfer?

Sea ice is permeated by small brine channels, which are characterised by sub-zero temperatures and varying salinities. Despite sometimes extreme conditions a diverse fauna and flora thrives within the brine channels. The dominant calanoid copepods of Antarctic sea ice are Stephos longipes and Paralabidocera antarctica. Here, I report for the first time thermal hysteresis (TH) in the haemolymph of a crustacean, S. longipes, whereas P. antarctica has no such activity. TH, the non-colligative prevention of ice growth, seems to enable S. longipes to exploit all available microhabitats within sea ice, especially the surface layer, in which strong temperature fluctuations can occur. In contrast, P. antarctica only thrives within the lowermost centimetres of sea ice, where temperature fluctuations are moderate. S. longipes possesses two isoforms of a protein with TH activity. A high homology to a group of (putative) antifreeze proteins from diatoms, bacteria and a snow mold and, in contrast, no homologs in any metazoan lineage suggest that this protein was obtained through horizontal gene transfer (HGT). Further analysis of available sequence data from sea-ice organisms indicates that these antifreeze proteins were probably transferred horizontally several times. Temperature and salinity fluctuations within the brine channel system are proposed to provide “natural transformation” conditions enabling HGT and thus making this habitat a potential “hot spot” for HGT.     

Surface-associated diatoms from marine habitats at Cape Evans,Antarctica, including the first record of living <Emphasis Type="Italic">Eunotogramma marginopunctatum</Emphasis>

The diversity of diatom communities found epiphytically on red macroalgae collected under the sea ice at Cape Evans, Antarctica, was studied microscopically. Comparison of communities along a depth profile showed that species diversity decreased as depth below sea ice increased. Dominant taxa also changed with depth, with Cocconeis fasciolata dominant at 10 and 15 m, Porosira glacialis at 20 m and Eunotogramma marginopunctatum at 25 m. Epiphytic communities were also compared to sympagic and sediment-associated communities collected from a single depth. Species richness was greatest for communities associated with surface sediments while the poorest was found in sympagic, or sea-ice, communities. E. marginopunctatum, previously only described from fossil material, was found associated epiphytically on macroalgae from 20 to 25 m below sea ice.     

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.     

Diving behaviour of Adélie penguins (Pygoscelis adeliae) at Dumont D'Urville, Antarctica: nocturnal patterns of diving and rapid adaptations to changes in sea-ice condition

The diving behaviour of Adélie penguins (Pygoscelis adeliae) was studied with time-depth recorders at Dumont D'Urville, Antarctica, during the breeding seasons in 1995/1996 and 1996/1997. We studied penguins foraging at all breeding stages, in various sea-ice conditions. For the first time in this species we observed nocturnal patterns of diving as the penguins dived more frequently and spent more time underwater around midnight than around noon. This behaviour may be related to the abundance of neritic krill, Euphausia crystallorophias, in the diet. Dive depth and duration varied extensively over the cycle, and appeared related to sea ice conditions rather than representative of the locality (22 m/78 s and 40 m/102 s with and without sea-ice, respectively). Comparisons with other studies showed that different diving behaviour previously observed in different localities can also occur at the same locality, and in some cases over a single foraging trip of a single penguin when short-term variation of external conditions occurred. Accepted: 27 September 1999     

Long-term increases in young-of-the-year growth of Arctic cisco Coregonus autumnalis and environmental influences

Arctic cisco Coregonus autumnalis young-of-year (YOY) growth was used as a proxy to examine the long-term response of a high-latitude fish population to changing climate from 1978 to 2004. YOY growth increased over time (r2 = 0·29) and was correlated with monthly averages of the Arctic oscillation index, air temperature, east wind speed, sea-ice concentration and river discharge with and without time lags. Overall, the most prevalent correlates to YOY growth were sea-ice concentration lagged 1 year (significant correlations in 7 months; r2 = 0·14-0·31) and Mackenzie River discharge lagged 2 years (significant correlations in 8 months; r2 = 0·13-0·50). The results suggest that decreased sea-ice concentrations and increased river discharge fuel primary production and that life cycles of prey species linking increased primary production to fish growth are responsible for the time lag. Oceanographic studies also suggest that sea ice concentration and fluvial inputs from the Mackenzie River are key factors influencing productivity in the Beaufort Sea. Future research should assess the possible mechanism relating sea ice concentration and river discharge to productivity at upper trophic levels.     

Protist assemblages in winter sea ice: setting the stage for the spring ice algal bloom

This study documents, for the first time, the abundance and species composition of protist assemblages in Arctic sea ice during the dark winter period. Lack of knowledge of sea-ice assemblages during the dark period has left questions about the retention and survival of protist species that initiate the ice algal bloom. Sea-ice and surface water samples were collected between December 27, 2007 and January 31, 2008 within the Cape Bathurst flaw lead, Canadian Beaufort Sea. Samples were analyzed for protist identification and counts, chlorophyll (chl) a, and total particulate carbon and nitrogen concentrations. Sea-ice chl a concentrations (max. 0.27 μg l⁻¹) and total protist abundances (max. 4 × 10³ cells l⁻¹) were very low, indicating minimal retention of protists in the ice during winter. The diversity of winter ice protists (134 taxa) was comparable to spring ice assemblages. Pennate diatoms dominated the winter protist assemblage numerically (averaging 77% of total protist abundances), with Nitzschia frigida being the most abundant species. Only 56 taxa were identified in surface waters, where dinoflagellates were the dominant group. Our results indicate that differences in the timing of ice formation may have a greater impact on the abundance than structure of protist assemblages present in winter sea ice and at the onset of the spring ice algal bloom.     

Mesoscale distribution and functional diversity of picoeukaryotes in the first-year sea ice of the Canadian Arctic

Sea ice, a characteristic feature of polar waters, is home to diverse microbial communities. Sea-ice picoeukaryotes (unicellular eukaryotes with cell size <3 μm) have received little attention compared with diatoms that dominate the spring bloom in Arctic first-year sea ice. Here, we investigated the abundance of all picoeukaryotes, and of 11 groups (chlorophytes, cryptophytes, bolidophytes, haptophytes, Pavlovaphyceae, Phaeocystis spp., pedinellales, stramenopiles groups MAST-1, MAST-2 and MAST-6 and Syndiniales Group II) at 13 first-year sea-ice stations localized in Barrow Strait and in the vicinity of Cornwallis Island, Canadian Arctic Archipelago. We applied Catalyzed Reporter Deposition–Fluorescence In Situ Hybridization to identify selected groups at a single cell level. Pavlovaphyceae and stramenopiles from groups MAST-2 and MAST-6 were for the first time reported from sea ice. Total numbers of picoeukaryotes were significantly higher in the vicinity of Cornwallis Island than in Barrow Strait. Similar trend was observed for all the groups except for haptophytes. Chlorophytes and cryptophytes were the dominant plastidic, and MAST-2 most numerous aplastidic of all the groups investigated. Numbers of total picoeukaryotes, chlorophytes and MAST-2 stramenopiles were positively correlated with the thickness of snow cover. All studied algal and MAST groups fed on bacteria. Presence of picoeukaryotes from various trophic groups (mixotrophs, phagotrophic and parasitic heterotrophs) indicates the diverse ecological roles picoeukaryotes have in sea ice. Yet, >50% of total sea-ice picoeukaryote cells remained unidentified, highlighting the need for further study of functional and phylogenetic sea-ice diversity, to elucidate the risks posed by ongoing Arctic changes.     

The occurrence of <Emphasis Type="Italic">Eualus gaimardii gibba</Emphasis> Krøyer 1841 (Crustacea,Decapoda) in the sympagic habitat: an example of bentho-sympagic coupling

We report the first records of a decapod species (Eualus gaimardii gibba) in association with Arctic multiyear sea-ice. Seven specimens inhabiting brine channels in the ice were collected during two field campaigns (September 2003, March/April 2004) by diver operated suction samplers. Salinity tolerance experiments (14–50 PSU) suggest that the species is stenohaline. Gut examinations revealed remains of the sympagic amphipod Gammarus wilkitzkii. All collected individuals were found in ice above relatively shallow waters (60–100 m depth) suggesting bentho-sympagic coupling.     

A winter sighting of killer whales (Orcinus orca ) in Antarctic sea ice

 A group of killer whales was sighted in open leads well inside Antarctic sea ice during August 1995. This was the first winter sighting of killer whales in Antarctic waters since 1955, and contradicts the view that all killer whales migrate north prior to the winter. A small calf was observed, providing the first evidence of a cetacean species breeding in Antarctic waters. Several potential prey species were also present. The sighting highlights the importance of lead and polynya systems to marine mammals, which probably use them to disperse within the winter sea-ice zone. Received: 23 April 1996 / Accepted: 25 August 1996     

Modelled and measured dynamics of viruses in Arctic winter sea-ice brines

We describe a model based on diffusion theory and the temperature-dependent mechanism of brine concentration in sea ice to argue that, if viruses partition with bacteria into sea-ice brine inclusions, contact rates between the two can be higher in winter sea ice than in seawater, increasing the probability of infection and possible virus production. To examine this hypothesis, we determined viral and bacterial concentrations in select winter sea-ice horizons using epifluorescence microscopy. Viral concentrations ranged from 1.6 to 82 x 10(6) ml(-1) of brine volume of the ice, with highest values in brines from coldest (-24 to -31 degrees C) ice horizons. Calculated virus-bacteria contact rates in underlying -1 degrees C seawater were similar to those in brines of -11 degrees C ice but up to 600 times lower than those in ice brines at or below -24 degrees C. We then incubated native bacterial and viral assemblages from winter sea ice for 8 days in brine at a temperature (-12 degrees C) and salinity ( approximately 160 psu) near expected in situ values, monitoring their concentrations microscopically. While different cores yielded different results, consistent with known spatial heterogeneity in sea ice, these experiments provided unambiguous evidence for viral persistence and production, as well as for bacterial growth, in -12 degrees C brine.