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     

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.     

Abundant dissolved genetic material in Arctic sea ice Part I: Extracellular DNA

The porous medium of sea ice, a surface-rich environment characterized by low temperature and high salinity, has been proposed as a favorable site for horizontal gene transfer, but few measurements are available to assess the possibility of this mode of evolution in ice. Here, we report the first measurements of dissolved DNA in sea ice, measured by fluorescent dye staining of centrifugal-filter-concentrated samples of melted ice. Newly formed landfast and pack ice on the Canadian Arctic Shelf (ca. 71°N, 125°W) contained higher concentrations (scaled to volume of brine) of the major components of dissolved DNA—extracellular DNA and viruses—than the underlying seawater. Dissolved DNA was dominated by extracellular DNA in surface seawater (up to 95%), with viruses making up relatively larger fractions at depths below 100 m (up to 27%) and in thick sea ice (66–78 cm; up to 100%). Extracellular DNA was heterogeneously distributed, with concentrations up to 135 μg DNA L⁻¹ brine detected in landfast sea ice, higher than previously reported from any marine environment. Additionally, extracellular DNA was significantly highly enriched at the base of ice of medium thickness (33–37 cm), suggestive of in situ production. Relative to underlying seawater, higher concentrations of extracellular DNA, viruses, and bacteria, and the availability of numerous surfaces for attachment within the ice matrix suggest that sea ice may be a hotspot for HGT in the marine environment.     

Marine bacterioplankton at the Weddell Sea ice edge,distribution of psychrophilic and psychrotrophic populations

Summary In the eastern Weddell Sea on several transects from ice-covered, through ice melt, to open-ocean stations, total and heterotrophic bacteria were estimated to document an enhanced bacteriological biomass expected near the ice edge. The highest numbers of bacteria were found in melted ice cores, with 4.2·10³ CFUml^–1 and 1.1·10⁷ Cells ml^–1. Although brine from pore water samples average more than one order of magnitude less cells per ml, the highest bacterial production, 2.2·10⁷ cells l^–1 day^–1, was recorded in brine samples. All quantitatively studied bacterial parameters were lower under the ice than in the ice samples but there were no clear vertical gradients in the water column. In the studied spring situation, sea ice occurrence seems to play only a minor role in the general distribution of the seawater bacterioplankton. The bacterial community structure was investigated by carrying out 29 morphological and biochemical tests on 118 isolated strains. The bacterial communities inhabiting Antarctic pack ice differ from those found in underlying seawater. Although non fermentative Gram-negative rods were always dominant in seawater, Vibrio sp. represented more than 25% of the strains isolated from some ice samples. The results clearly indicated that a large majority of the bacteria isolated from seawater must be considered psychrotrophic but that truly psychrophilic strains occurred in melted ice and brine samples.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

The timing of sea ice formation and exposure to photosynthetically active radiation along the Western Antarctic Peninsula

Understanding the flow of solar energy into ecosystems is fundamental to understanding ecosystem productivity and dynamics. To gain a better understanding of this fundamental process in the Antarctic winter sea ice, we produced a model that estimates the time-integrated exposure of seasonal Antarctic sea ice to PAR through the use of remotely sensed sea ice concentrations, sea ice movement and spatially distributed PAR calculations that account for cloud cover and have applied this model over the past three decades. The resulting spatially distributed estimates of sea ice exposure to PAR by mid-winter are evaluated in context of changes in the timing of sea ice formation that have been documented along the Western Antarctic Peninsula (WAP) region and its potential effects on the variation (seasonal and inter-annual) in the accumulation of sea ice algae in this region. The analysis shows the ice pack is likely to have large inter-annual variations (10–100 fold) in productivity throughout the autumn to winter transition in the sea ice along the WAP. Moreover, the pack ice is likely to have spatial structure in regards to biological processes that cannot be determined from analysis of sea ice concentration information alone. The resulting inter-annual variations in winter processes are likely to affect the dynamics of Antarctic krill (Euphausia superba).     

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.     

Dissolved extracellular polymeric substances (dEPS) dynamics and bacterial growth during sea ice formation in an ice tank study

Extracellular polymeric substances (EPS) are known to help microorganisms to survive under extreme conditions in sea ice. High concentrations of EPS are reported in sea ice from both poles; however, production and dynamics of EPS during sea ice formation have been little studied to date. This investigation followed the production and partitioning of existing and newly formed dissolved organic matter (DOM) including dissolved carbohydrates (dCHO), dissolved uronic acids (dUA) and dissolved EPS (dEPS), along with bacterial abundances during early stages of ice formation. Sea ice was formed from North Sea water with (A) ambient DOM (NSW) and (B) with additional algal-derived DOM (ADOM) in a 6d experiment in replicated mesocosms. In ADOM seawater, total bacterial numbers (TBN) increased throughout the experiment, whereas bacterial growth occurred for 5d only in the NSW seawater. TBN progressively decreased within developing sea ice but with a 2-fold greater decline in NSW compared to ADOM ice. There were significant increases in the concentrations of dCHO in ice. Percentage contribution of dEPS was highest (63%) in the colder, uppermost parts in ADOM ice suggesting the development of a cold-adapted community, producing dEPS possibly for cryo-protection and/or protection from high salinity brines. We conclude that in the early stages of ice formation, allochthonous organic matter was incorporated from parent seawater into sea ice and that once ice formation had established, there were significant changes in the concentrations and composition of dissolved organic carbon pool, resulting mainly from the production of autochthonous DOM by the bacteria.     

Aerial surveys for Antarctic minke whales (Balaenoptera bonaerensis) reveal sea ice dependent distribution patterns

This study investigates the distribution of Antarctic minke whales (AMW) in relation to sea ice concentration and variations therein. Information on AMW densities in the sea ice‐covered parts of the Southern Ocean is required to contextualize abundance estimates obtained from circumpolar shipboard surveys in open waters, suggesting a 30% decline in AMW abundance. Conventional line‐transect shipboard surveys for density estimation are impossible in ice‐covered regions, therefore we used icebreaker‐supported helicopter surveys to obtain information on AMW densities along gradients of 0%–100% of ice concentration. We conducted five helicopter surveys in the Southern Ocean, between 2006 and 2013. Distance sampling data, satellite‐derived sea‐ice data, and bathymetric parameters were used in generalized additive models (GAMs) to produce predictions on how the density of AMWs varied over space and time, and with environmental covariates. Ice concentration, distance to the ice edge and distance from the shelf break were found to describe the distribution of AMWs. Highest densities were predicted at the ice edge and through to medium ice concentrations. Medium densities were found up to 500 km into the ice edge in all concentrations of ice. Very low numbers of AMWs were found in the ice‐free waters of the West Antarctic Peninsula (WAP). A consistent relationship between AMW distribution and sea ice concentration weakens the support for the hypothesis that varying numbers of AMWs in ice‐covered waters were responsible for observed changes in estimated abundance. The potential decline in AMW abundance stresses the need for conservation measures and further studies into the AMW population status. Very low numbers of AMWs recorded in the ice‐free waters along the WAP support the hypothesis that this species is strongly dependent on sea ice and that forecasted sea ice changes have the potential of heavily impacting AMWs.     

Dynamics of ice algae and phytoplankton in Frobisher Bay

Summary Vertical and seasonal variations of ice algae and phytoplankton were studied in relation to their physico-chemical environments in Frobisher Bay from 1979 to 1986. The biomass, estimated by both chlorophyll a concentrations and cell counts, was greater in the ice algae than in the phytoplankton in the underlying sea-water during winter and spring. Algal distribution in the sea ice varied vertically and seasonally, while in the underlying water column the phytoplankton distribution was much less variable. The ice algal bloom occurred at the bottom of the ice, particularly in the lower 5 cm during late spring, while the phytoplankton bloom took place at depths between 1 and 10 m during early summer after the ice bloom was over. The community structure of the ice algae changed from pennate to centric diatoms as the ice melted. The centrics dominated through the fall, and then decreased as the pennates increased in dominance when the ice formed again in winter. Species diversity and number were greater in the sea ice than in the seawater, but they were similar vertically within each habitat. The evenness of the species distribution did not vary with ice thickness or water depth. Species composition, abundance and dominance of ice algae and phytoplankton continually change both vertically and seasonally. The differential abilities of the species to attain maximal growth rates under various environmental conditions may result in species succession. Evidence is given for the major role of environmental factors regulating the dynamics of ice algae and phytoplankton.     

A diatom based transfer function for reconstructing sea ice concentrations in the North Atlantic

A diatom based sea ice transfer function is developed using 99 surface sediment samples from the North Atlantic and the associated modern sea ice concentrations. Canonical correspondence analysis (CCA) is applied to the species assemblages of the surface sediment samples and the association of the species with two environmental parameters, August sea surface temperature and May sea ice concentration, is assessed. The results of this analysis indicate negative correlation between sea ice and sea surface temperature and that a group of diatom species is strongly associated with sea ice, especially May sea ice concentration. The results of the CCA legitimate the development of a diatom based sea ice transfer function. The maximum likelihood method has been applied as the transfer function method, as it has been proven most suitable with this particular data set. The newly developed transfer function is then used to reconstruct May sea ice concentration in three cases, each focusing on a different time period: the Last Glacial Maximum, the Younger Dryas and the Holocene. In all three cases the transfer function produces reasonable results when compared to other paleoclimatic proxy results. This suggests that the sea ice concentration reconstructed by the diatom based sea ice transfer function is a valid and reliable method, which can be applied as a valid proxy for May sea ice concentration.     

Vertical distribution of bacteria in arctic sea ice

Abstract Heterotrophic bacteria were enumerated in north polar sea ice cores obtained near Point Barrow, Alaska. Highest concentrations of total and viable bacteria were found in the layer containing the sea ice microbial community identified by the maximum chlorophyll a content. Gas vacuolate bacteria were also found in the sea ice, a discovery which is consistent with their recent report from antarctic sea ice microbial communities. The gas vacuolate bacteria comprised 0.2% or less of the viable bacteria isolated from sea ice cores, lower than concentrations reported for most antarctic samples. Most gas vacuolate isolates from the sea ice cores were pigmented pink, orange, or yellow. An ice core from nearby saline Elson Lagoon contained an inverted sea ice microbial community with highest chlorophyll a concentrations and bacterial counts found in the top 0–20 cm of the ice. This surface layer also contained high numbers (up to 186 bacteria/ml) of a nonpigmented, gas vacuolate, elongated rod-shaped bacterium.     

Sea ice bacterial growth rate, growth efficiency and preference for inorganic nitrogen sources in the Baltic Sea

Seasonal Baltic Sea ice is structurally similar to polar sea ice and provides habitats for diverse ice organism assemblages that are integral to the biogeochemistry and ecology of the sea during winter. Temperature and inorganic nitrogen sources have been suggested to control bacterial growth, with increasing dependence on ammonium at low temperatures. To study the bacterial growth and preference for the nitrogen source, we conducted experiments at 0 and 4°C, using ammonium and nitrate as nitrogen sources at two coastal fast-ice stations in the Gulf of Finland and in the Gulf of Bothnia during three successive winters. The two study sites differ markedly in relation to the allochthonous dissolved organic matter supply from the catchment area. High levels of bacterial growth were recorded at both study sites, with community generation times of 15–37 h. The measured bacterial growth efficiencies of 20–58% suggest that the Baltic sea ice brines provide a rich medium for bacterial growth and efficient functioning of bacteria-based food webs. Our experiments with sea ice samples showed a preference for ammonium at both temperatures and high potential growth in both types of nitrogen supplies. No major differences in phosphorus depletion rates were found at the two temperatures, but rates were always highest when ammonium was added to the experiments. These experiments point out that ice maturity, presumably through changes in bacterial community structure, impacts nitrogen processes and that these processes are pronounced prior to melting of the ice.     

Unusual narwhal sea ice entrapments and delayed autumn freeze-up trends

Sea ice entrapments of narwhals (Monodon monoceros) occur when rapid changes in weather and wind conditions create a formation of fast ice in bays or passages used by whales. Between 2008 and 2010, four entrapments of narwhals were reported in Canada and Greenland. In each case, large groups (40–600 individuals) succumbed in the sea ice at three separate summering localities, two of these where entrapments had never before been reported. We examined long-term trends in autumn freeze-up timing (date when sea ice concentration rises above some threshold) on the 6 largest narwhal summering areas using sea ice concentration from satellite passive microwave data (1979–2009). We found strongly positive and significant trends (P < 0.001) in progressively later dates of autumn freeze-up in all summering areas. Autumn freeze-up occurs between 0.5 and 1 day later per year, or roughly 2–4 weeks later, over the 31-year time series. This indicates that sea ice conditions on narwhal summering areas are changing rapidly. The question remains whether entrapment events on summering areas are random or whether narwhals are adapting to changes in sea ice freeze-up by prolonging their summer residence time.     

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.     

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.     

Polar zoobenthos blue carbon storage increases with sea ice losses,because across‐shelf growth gains from longer algal blooms outweigh ice scour mortality in the shallows

One of the major climate‐forced global changes has been white to blue to green; losses of sea ice extent in time and space around Arctic and West Antarctic seas has increased open water and the duration (though not magnitude) of phytoplankton blooms. Blueing of the poles has increases potential for heat absorption for positive feedback but conversely the longer phytoplankton blooms have increased carbon export to storage and sequestration by shelf benthos. However, ice shelf collapses and glacier retreat can calve more icebergs, and the increased open water allows icebergs more opportunities to scour the seabed, reducing zoobenthic blue carbon capture and storage. Here the size and variability in benthic blue carbon in mega and macrobenthos was assessed in time and space at Ryder and Marguerite bays of the West Antarctic Peninsula (WAP). In particular the influence of the duration of primary productivity and ice scour are investigated from the shallows to typical shelf depths of 500 m. Ice scour frequency dominated influence on benthic blue carbon at 5 m, to comparable with phytoplankton duration by 25 m depth. At 500 m only phytoplankton duration was significant and influential. WAP zoobenthos was calculated to generate ~10⁷, 4.5 × 10⁶ and 1.6 × 10⁶ tonnes per year (between 2002 and 2015) in terms of production, immobilization and sequestration of carbon respectively. Thus about 1% of annual primary productivity has sequestration potential at the end of the trophic cascade. Polar zoobenthic blue carbon capture and storage responses to sea ice losses, the largest negative feedback on climate change, has been underestimated despite some offsetting of gain by increased ice scouring with more open water. Equivalent survey of Arctic and sub‐Antarctic shelves, for which new projects have started, should reveal the true extent of this feedback and how much its variability contributes to uncertainty in climate models.     

Distribution and abundance of the planktic foraminifer Neogloboquadrina pachyderma in sea ice of the Weddell Sea (Antarctica)

Summary Sea ice cores were obtained from eleven fast ice stations and one floe in the Weddell Sea, Antarctica in January–February 1985. All cores from the north eastern part of the Weddell Sea contained numerous living and dead planktic foraminifers of the species Neogloboquadrina pachyderma (Ehrenberg), while cores drilled in southern parts were barren of foraminifers with one exception. Foraminiferal abundances were variable, with numbers up to 320 individuals per liter melted sea ice. Distribution of foraminifers appears to be patchy, parallel cores taken less than 30 cm apart contained numbers which varied considerably. On the other hand, three cores taken on a transect each more than 3 km apart showed striking similarities. In general, small dead tests were found in the upper parts of the sea ice cores while large living individuals mainly occurred in lower sections. Abundant diatoms probably serve as a food source for the foraminifers. Correlation of foraminiferal abundance with salinity, chlorophyll and nutrient profiles are inconsistent. The possible mechanism of incorporation of N. pachyderma into the ice is discussed.     

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.     

北极浮冰生态学研究进展

近年来随着北极地区的开放和全球变化对北极地区生态环境和海冰现存量的影响日益显现,北极浮冰生态学研究得到了广泛的重视和实质性的进展.最新研究结果显示,浮冰本身包含了一个复杂的生物群落,高纬度浮冰生物群落的初级产量远高于原先的估算,浮冰生物群落在北极海洋生态系统中的作用被进一步确认.但由于对浮冰生物群落的研究受后勤保障条件的制约,目前尚有大量科学问题有待今后进一步深入研究,预期我国科学家将在其中做出贡献.