On the structure and development of Arctic pack ice communities in Fram Strait: a multivariate approach

Summary The distribution of ice organisms was investigated in Fram Strait in May 1988 during the ARK V/1 expedition on RV Polarstern. Over a 3 week period the abundances of bacteria, diatoms, auto- and heterotrophic flagellates as well as various groups of meiofauna organisms were observed in the lowermost 30 cm of an ice floe. Data were obtained from three experimental fields under three different light regimes as a result of manipulations of the snow cover. The application of multivariate factor analysis on this time series data set resulted in the characterization of four succession stages of an Arctic sea ice community: 1) the diatom bottom assemblage, 2) the mixed autotrophic assemblage, 3) the mixed auto- and heterotrophic supra-bottom assemblage, and 4) the heterotrophic supra-bottom assemblage. The two most abundant meiofauna groups (Turbellaria, Ciliata) showed different preferences according to algal distribution. While turbellarians were most abundant in samples with mixed populations of diatoms and flagellates, ciliates reached their abundance maxima in samples dominated by diatoms, suggesting different prey selections. We have developed a model for the explanation of the spatial separation of auto- and heterotrophic organisms, highlighting the possible role of DOC production by ice algae and DOC transport with brine flow.     

Symbiosis between the stramenopile protist Solenicola setigera and the diatom Leptocylindrus mediterraneus in the North Eastern Arabian Sea

Symbiotic associations between the diatoms and other organisms like cyanobacteria, flagellates etc. is less studied in Indian waters. Here, we report the occurrences of Solenicola setigera, a colonial protist found only in association with centric chain forming diatom Leptocylindrus mediterraneus. We observed these consortia in the surface waters (~80?C100 colonies L-1) of North Eastern Arabian Sea (NEAS) during the spring inter monsoon season of 2011 and were also observed in the water column (<20 colonies L-1) up to 120?m depth. The consortium observed in the surface waters comprised single or few frustule with the protozoan colonies spread along the entire frustule, but those in the water column with the protozoan colony restricted to the central section of the frustule. Our observations suggest that Solenicola setigera is a heterotroph that grazes on pico and nanoplanktonic organisms thereby play an important ecological role in the North Eastern Arabian Sea ecosystem.     

The nutrient status in sea ice of the Weddell Sea during winter: effects of sea ice texture and algae

Summary Cores and brine samples from sea ice of the Weddell Sea were analyzed for nutrients (phosphate, nitrate and silicate), salinity and chlorophyll a during winter. Stratigraphic analyses of the cores were also carried out. Bulk nutrient concentrations in the sea ice fluctuated widely and did not correlate with salinity. Nutrient concentrations in cores were normalized to sea-water salinity to facilitate comparison. They varied between zero and two or three times those measured in the water column. Differentiation into young and old sea ice, however, revealed that nutrient concentrations in the young ice in many cases corresponded to those in surface seawater. In older ice, nutrients showed signs of increase as well as depletion or exhaustion relative to the water column. Differentiation of core sections according to ice textural classes and analyses of brine samples clarified some relationships between nutrients, salinity and algal biomass. Most of the changes in the nutrient concentrations are attributed to an increase in biological activity as the seasons progress. Silicate is expected to become the first nutrient likely to limit growth of diatoms in the ice which is ascribed to slower regeneration or dissolution of this nutrient relative to phosphate and nitrate. A consequence of silicate exhaustion may be the succession of different algal assemblages, from a diatom dominated community to one in which autotrophic flagellates form the largest component.     

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     

Bacterial activity in sea ice and open water of the Weddell Sea,Antarctica: A microautoradiographic study

Metabolic activity of bacteria was investigated in open water, newly forming sea ice, and successive stages of pack ice in the Weddell Sea. Microautoradiography, using [³H]leucine as substrate, was compared with incorporation rates of [³H]leucine into proteins. Relation of [³H]leucine incorporation to the biomass of active bacteria provides information about changes of specific metabolic activity of cells. During a phytoplankton bloom in an ice-free, stratified water column, total numbers of bacteria in the euphotic zone averaged 2.3 × 10⁵ ml^–1, but only about 13% showed activity via leucine uptake. Growth rate of the active bacteria was estimated as 0.3–0.4 days^–1. Total cell concentration of bacteria in 400 m depth was 6.6 × 10⁴ ml^–1. Nearly 50% of these cells were active, although biomass production and specific growth rate were only about one-tenth that of the surface populations. When sea ice was forming in high concentrations of phytoplankton, bacterial biomass in the newly formed ice was 49.1 ng C ml^–1, exceeding that in open water by about one order of magnitude. Attachment of large bacteria to algal cells seems to cause their enrichment in the new ice, since specific bacterial activity was reduced during ice formation, and enrichment of bacteria was not observed when ice formed at low algal concentration. During growth of pack ice, biomass of bacteria increased within the brine channel system. Specific activity was still reduced at these later stages of ice development, and percentages of active cells were as low as 3–5%. In old, thick pack ice, bacterial activity was high and about 30% of cells were active. However, biomass-specific activity of bacteria remained significantly lower than that in open water. It is concluded that bacterial assemblages different to those of open water developed within the ice and were dominated by bacteria with lower average metabolic activity than those of ice-free water.     

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.     

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.     

Seasonal variation in marine phytoplankton and ice algae at a shallow antarctic coastal site

The phytoplankton population near Davis, Vestfold Hills, Antarctica was monitored throughout 1982. Chlorophyll-a determinations and counts of living cells in both the water column and sea ice demonstrated a marked seasonality in phytoplankton abundane and species composition. From April to October nanoplanktonic organisms contributed most of the chlorophyll-a in both the sea ice and water column. Blooms of diatoms occurred in May, November and December in the bottom of the sea-ice and in January and February in the water column. Phaeocystis pouchetii was dominant during December in the water column. Large numbers of dead diatoms were found in winter. The concentrations of nitrate, dissolved inorganic phosphate and dissolved silicate increased throughout the year until December, when the concentrations of nitrate and silicate fell sharply, followed a month later by a reduction in phosphate concentration. The diversity of phytoplankton was greatest during the summer months.     

POPULATION DYNAMICS OF MICROALGAE IN THE UPPER LAND-FAST SEA ICE AT A SNOW-FREE LOCATION

The population dynamics of interior ice microalgae were investigated at a snow-free site on annual land-fast sea ice in McMurdo Sound, Antarctica, during the austral spring and summer of 1995 – 96. A dynamic successional sequence was observed with life history transformations playing an important role. During late November and early December (austral spring), cryo- and halotolerant dinoflagellates and chrysophytes bloomed in brine channels within the upper ice. At this time, competition and grazing pressure are low because of the inability of most marine species to grow under the extreme environmental conditions found in the upper ice during the austral spring. In November and December, dinoflagellates, chrysophytes, and prasinophytes contributed an average of 66%, 44%, and < 1% of the phytoflagellate biomass, respectively. Both the dinoflagellates and the chrysophytes encysted in December, with cyst formation most intense just prior to surface melt and flushing of the ice. The cysts appear to be an adaptation for survival and dispersal in the plankton during ice decay and/or overwintering in the sea ice. In January (austral summer), when ice temperatures were similar to those in the water column, pennate diatoms replaced flagellates as the photosynthetic dominants in the upper sea ice. The upper land-fast sea ice undergoes dramatic seasonal changes in light availability, temperature, brine salinity, and inorganic nutrient availability. Ephemeral blooms of cyst-forming phytoflagellates exploit this habitat in the austral spring, when both inorganic nutrients and light are available but temperatures <− 2° C and brine salinities elevated.     

Composition and distribution of the pelagic and sympagic algal assemblages in the Laptev Sea during autumnal freeze-up

The phytoplankton and ice algal assemblages in the SiberianLaptev Sea during the autumnal freeze-up period of 1995 aredescribed. The spatial distribution of algal taxa (diatoms,dinoflagellates, chrysophytes, chlorophytes) in the newly formedice and waters at the surface and at 5 m depth differed considerablybetween regions. This was also true for algal biomass measuredby in situ fluorescence, chlorophyll (Chl) a and taxon-specificcarbon content. Highest in situ fluorescence and Chl a concentrations(ranging from 0.1 to 3.2 µg l^–1) occurred in surfacewaters with maxima in Buor Khaya Bay east of Lena Delta. Thealgal standing stock on the shelf consisted mainly of diatoms,dinoflagellates, chrysophytes and chlorophytes with a totalabundance (excluding unidentified flagellates <10 µm)in surface waters of 351–33 660 cells l^–1. Highestalgal abundance occurred close to the Lena Delta. Phytoplanktonbiomass (phytoplankton carbon; PPC) ranged from 0.1 to 5.3 µgC l^–1 in surface waters and from 0.3 to 2.1 µg Cl^–1 at 5 m depth, and followed the distribution patternof abundances. However, the distribution of Chl a differed considerablyfrom the distribution pattern shown by PPC. The algal assemblagein the sea ice, which could not be quantified due to high sedimentload, was dominated by diatom species, accompanied by dinoflagellates.Thus, already during the early stage of autumnal freeze-up,incorporation processes, selective enrichment and subsequentgrowth lead to differences between surface water and sea icealgal assemblages.     

Bacterial Standing Stock, Activity, and Carbon Production during Formation and Growth of Sea Ice in the Weddell Sea, Antarctica

Bacterial response to formation and growth of sea ice was investigated during autumn in the northeastern Weddell Sea. Changes in standing stock, activity, and carbon production of bacteria were determined in successive stages of ice development. During initial ice formation, concentrations of bacterial cells, in the order of 1 × 10⁸ to 3 × 10⁸ liter^-1, were not enhanced within the ice matrix. This suggests that physical enrichment of bacteria by ice crystals is not effective. Due to low concentrations of phytoplankton in the water column during freezing, incorporation of bacteria into newly formed ice via attachment to algal cells or aggregates was not recorded in this study. As soon as the ice had formed, the general metabolic activity of bacterial populations was strongly suppressed. Furthermore, the ratio of [³H]leucine incorporation into proteins to [³H]thymidine incorporation into DNA changed during ice growth. In thick pack ice, bacterial activity recovered and growth rates up to 0.6 day^-1 indicated actively dividing populations. However, biomass-specific utilization of organic compounds remained lower than in open water. Bacterial concentrations of up to 2.8 × 10⁹ cells liter^-1 along with considerably enlarged cell volumes accumulated within thick pack ice, suggesting reduced mortality rates of bacteria within the small brine pores. In the course of ice development, bacterial carbon production increased from about 0.01 to 0.4 μg of C liter^-1 h^-1. In thick ice, bacterial secondary production exceeded primary production of microalgae.     

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     

Protists in the marine ice of the Amery Ice Shelf, East Antarctica

Samples of marine ice were collected from the Amery Ice Shelf, a large embayed ice shelf in East Antarctica, during the Austral summer of 2001–2002. The samples came from a site ∼90 km from the iceberg calving front of the shelf, where the ice is 479 m thick and the lower 203 m is composed of accreted marine ice. Protists identified within the marine ice layer of the Amery Ice Shelf include diatoms, chrysophytes, silicoflagellates and dinoflagellates. The numerical dominance of sea ice indicator diatoms such as Fragilariopsis curta, Fragilariopsis cylindrus, Fragilariopsis rhombica and Chaetoceros resting spores, and the presence of cold open water diatoms such as Fragilariopsis kerguelensis and species of Thalassiosira suggest the protist composition of the Amery marine ice is attributable to seeding from melting pack and/or fast ice protist communities in the highly productive waters of Prydz Bay to the north.     

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.     

High‐throughput sequencing reveals neustonic and planktonic microbial eukaryote diversity in coastal waters

Neustonic organisms inhabit the sea surface microlayer (SML) and have important roles in marine ecosystem functioning. Here, we use high‐throughput 18S rRNA gene sequencing to characterize protist and fungal diversity in the SML at a coastal time‐series station and compare with underlying plankton assemblages. Protist diversity was higher in February (pre‐bloom) compared to April (spring bloom), and was lower in the neuston than in the plankton. Major protist groups, including Stramenopiles and Alveolata, dominated both neuston and plankton assemblages. Chrysophytes and diatoms were enriched in the neuston in April, with diatoms showing distinct changes in community composition between the sampling periods. Pezizomycetes dominated planktonic fungi assemblages, whereas fungal diversity in the neuston was more varied. This is the first study to utilize a molecular‐based approach to characterize neustonic protist and fungal assemblages, and provides the most comprehensive diversity assessment to date of this ecosystem. Variability in the SML microeukaryote assemblage structure has potential implications for biogeochemical and food web processes at the air‐sea interface.     

Protozoan Bacterivory in the Ice and the Water Column of a Cold Temperate Lagoon

Abstract Bacterial abundance and bacterivorous protist abundance and activity were examined in ice-brine and water column communities of a cold temperate Japanese lagoon (Saroma-Ko Lagoon, Hokkaido, 44°N, 144°E), during the late winter phase of ice community development (February–March 1992). Bacterial abundance averaged 6 and 1 × 10⁵ cells ml⁻¹ in the ice-brine and plankton samples, respectively, and generally decreased during the sampling period. Bacterivorous protists, identified based on direct observation of short-term (<1 h) ingested fluorescently labeled bacteria (FLB) in their food vacuoles, were largely dominated by flagellates, mainly cryothecomonad-type and chrysomonad-like cells and small dinoflagellates of the genus Gymnodinium. Bacterivorous ciliates included mainly the prostomatid Urotricha sp., the scuticociliates Uronema and Cyclidium, the choreotrichs Lohmaniella oviformis and Strobilidium, and the hypotrich Euplotes sp. Protist abundance averaged 4 × 10³ and 8.1 cells ml⁻¹ in the ice-brine and 0.3 × 10³ and 1.2 cells ml⁻¹ in the plankton, for flagellates and ciliates, respectively. In contrast to bacteria, the abundance of protists generally increased throughout the sampling period, indicating predator–prey interactions. Protistan bacterivory, measured from the rate of FLB disappearance over 24 h, averaged 36% (ice) and 24% (plankton) of bacterial standing stock and exhibited the same seasonal pattern as for protist abundance. The calculated specific clearance (range, 2–67 nl protozoa⁻¹ h⁻¹) and ingestion (<1–26 particles protozoa⁻¹ h⁻¹) rates were likely to be minimal estimates and grazing impact may have been higher on occasion. Indications for the dependence of ``bacterivorous protists' on nonbacterial food items were also provided. Although alternative sources of bacterial loss are likely to be of importance, this study provides evidence for the potential of protozoan assemblages as bacterial grazers in both sea ice-brine biota and water column at the southern limit of sea ice in the northern hemisphere. Received: 30 July 1998; Accepted: 18 November 1998     

Experimental formation of sea ice: importance of water circulation and wave action for incorporation of phytoplankton and bacteria

The efficiency of physical concentration mechanisms for enrichment of algae and bacteria in newly formed sea-ice was investigated under defined conditions in the laboratory. Sea-ice formation was simulated in a 3,000 l tank under different patterns of water movement. When ice formed in an artificially generated current pattern, algal cells were substantially enriched within the ice matrix. Enrichment factors for chlorophyll a calculated from the ratio between the concentrations in ice and underlying water reached values of up to 53. Repeated mixing of ice crystals into the water column, as well as flow of water through the new ice layer, contributed to the enrichment of algae in the ice. Wave action during ice formation revealed lower phytoplankton enrichment factors of up to 9. Mixing of floating ice crystals with underlying water and pumping of water into the ice matrix by periodical expansion and compression of the slush ice layer were responsible for the wave-induced enrichment of algal cells. Physical enrichment of bacteria within the ice was negligible. Bacterial biomass within new ice was enhanced only when the concentration of algae was high. At low algal biomass, bacteria experienced substantial losses in the ice, most likely due to brine drainage, which were not observed for the microalgae. Bacterial cells are therefore not scavenged by ice crystals and the observed enrichment and sustainment of bacterial biomass within newly formed ice depend on their attachment to cells or aggregates of algae. Division rates of bacteria changed only slightly during ice formation. Received: 21 October 1997 / Accepted: 9 April 1998     

Anchor ice and benthic disturbance in shallow Antarctic waters: interspecific variation in initiation and propagation of ice crystals

Sea ice typically forms at the ocean's surface, but given a source of supercooled water, an unusual form of ice--anchor ice--can grow on objects in the water column or at the seafloor. For several decades, ecologists have considered anchor ice to be an important agent of disturbance in the shallow-water benthic communities of McMurdo Sound, Antarctica, and potentially elsewhere in polar seas. Divers have documented anchor ice in the McMurdo communities, and its presence coincides with reduced abundance of the sponge Homaxinella balfourensis, which provides habitat for a diverse assemblage of benthic organisms. However, the mechanism of this disturbance has not been explored. Here we show interspecific differences in anchor-ice formation and propagation characteristics for Antarctic benthic organisms. The sponges H. balfourensis and Suberites caminatus show increased incidence of formation and accelerated spread of ice crystals compared to urchins and sea stars. Anchor ice also forms readily on sediments, from which it can grow and adhere to organisms. Our results are consistent with, and provide a potential first step toward, an explanation for disturbance patterns observed in shallow polar benthic communities. Interspecific differences in ice formation raise questions about how surface tissue characteristics such as surface area, rugosity, and mucus coating affect ice formation on invertebrates.     

Gas Vacuolate Bacteria from the Sea Ice of Antarctica

Gas-vacuolate heterotrophic bacteria from marine habitats are reported here for the first time. They have been isolated from Antarctic sea ice microbial communities and the underlying water column. The predominant gas-vacuolate bacterium from the sea ice is filamentous and pigmented, whereas those of the water column are unicellular and nonpigmented. The highest concentrations of bacteria in sea ice were found in conjunction with the highest algal (chlorophyll a) concentrations.     

PHOTOPROTECTION OF SEA‐ICE MICROALGAL COMMUNITIES FROM THE EAST ANTARCTIC PACK ICE1

All photosynthetic organisms endeavor to balance energy supply with demand. For sea‐ice diatoms, as with all marine photoautotrophs, light is the most important factor for determining growth and carbon‐fixation rates. Light varies from extremely low to often relatively high irradiances within the sea‐ice environment, meaning that sea‐ice algae require moderate physiological plasticity that is necessary for rapid light acclimation and photoprotection. This study investigated photoprotective mechanisms employed by bottom Antarctic sea‐ice algae in response to relatively high irradiances to understand how they acclimate to the environmental conditions presented during early spring, as the light climate begins to intensify and snow and sea‐ice thinning commences. The sea‐ice microalgae displayed high photosynthetic plasticity to increased irradiance, with a rapid decline in photochemical efficiency that was completely reversible when placed under low light. Similarly, the photoprotective xanthophyll pigment diatoxanthin (Dt) was immediately activated but reversed during recovery under low light. The xanthophyll inhibitor dithiothreitol (DTT) and state transition inhibitor sodium fluoride (NaF) were used in under‐ice in situ incubations and revealed that nonphotochemical quenching (NPQ) via xanthophyll‐cycle activation was the preferred method for light acclimation and photoprotection by bottom sea‐ice algae. This study showed that bottom sea‐ice algae from the east Antarctic possess a high level of plasticity in their light‐acclimation capabilities and identified the xanthophyll cycle as a critical mechanism in photoprotection and the preferred means by which sea‐ice diatoms regulate energy flow to PSII.