Abundant dissolved genetic material in Arctic sea ice Part II: Viral dynamics during autumn freeze-up

Viruses play a significant role in nutrient cycling within the world’s oceans and are important agents of horizontal gene transfer, but little is know about their entrainment into sea ice or their temporal dynamics once entrained. Nilas, grease ice, pancake ice, first-year sea ice floes up to 78 cm in thickness, and under-ice seawater were sampled widely across Amundsen Gulf (ca. 71^° N, 125^° W71^\circ \hbox{N}, 125^\circ \hbox{W}) for concentrations of viruses and bacteria. Here, we report exceptionally high virus-to-bacteria ratios in seawater (45–340) and sea ice (93–2,820) during the autumn freeze-up. Virus concentrations ranged from 4.8 to 27 × 10⁶  ml⁻¹ in seawater and, scaled to brine volume, 5.5 to 170 × 10⁷ ml⁻¹ in sea ice. Large enrichment indices indicated processes of active entrainment from source seawater, or viral production within the ice, which was observed in 2 of 3 bottle incubations of sea ice brine at a temperature (-7^°C-7^\circ\hbox{C}) and salinity ( 110 \permille110 \permille) approximating that in situ. Median predicted virus-to-bacteria contact rates (relative to underlying seawater) were greatest in the top of thick sea ice (66–78 cm: 130×) and lowest in the bottom of medium-thickness ice (33–37 cm: 23×). The great abundance of viruses and more frequent interactions between bacteria and viruses predicted in sea ice relative to underlying seawater suggest that sea ice may be a hot spot for virally mediated horizontal gene transfer in the polar marine environment.     

Biogeochemical conditions and ice algal photosynthetic parameters in Weddell Sea ice during early spring

Physical, biogeochemical and photosynthetic parameters were measured in sea ice brine and ice core bottom samples in the north-western Weddell Sea during early spring 2006. Sea ice brines collected from sackholes were characterised by cold temperatures (range −7.4 to −3.8°C), high salinities (range 61.4–118.0), and partly elevated dissolved oxygen concentrations (range 159–413 μmol kg⁻¹) when compared to surface seawater. Nitrate (range 0.5–76.3 μmol kg⁻¹), dissolved inorganic phosphate (range 0.2–7.0 μmol kg⁻¹) and silicic acid (range 74–285 μmol kg⁻¹) concentrations in sea ice brines were depleted when compared to surface seawater. In contrast, NH₄ ⁺ (range 0.3–23.0 μmol kg⁻¹) and dissolved organic carbon (range 140–707 μmol kg⁻¹) were enriched in the sea ice brines. Ice core bottom samples exhibited moderate temperatures and brine salinities, but high algal biomass (4.9–435.5 μg Chl a l⁻¹ brine) and silicic acid depletion. Pulse amplitude modulated fluorometry was used for the determination of the photosynthetic parameters F _v/F _m, α, rETR_max and E _k. The maximum quantum yield of photosystem II, F _v/F _m, ranged from 0.101 to 0.500 (average 0.284 ± 0.132) and 0.235 to 0.595 (average 0.368 ± 0.127) in the sea ice internal and bottom communities, respectively. The fluorometric measurements indicated medium ice algal photosynthetic activity both in the internal and bottom communities of the sea ice. An observed lack of correlation between biogeochemical and photosynthetic parameters was most likely due to temporally and spatially decoupled physical and biological processes in the sea ice brine channel system, and was also influenced by the temporal and spatial resolution of applied sampling techniques.     

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.     

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.     

Microbial loop malfunctioning in the annual sea ice at Terra Nova Bay (Antarctica)

We investigated organic carbon quantity and biochemical composition, prokaryotic abundance, biomass and carbon production in the annual and platelet sea ice of Terra Nova Bay (Antarctica), as well as the downward fluxes of organic matter released by melting ice during early spring. Huge amounts of biopolymeric C accumulated in the bottom layer of the ice column concomitantly with the early spring increase in sympagic algal biomass. Such organic material, mostly accounted for by autotrophic biomass, was characterised by a high food quality and was rapidly exported to the sea bottom during sea ice melting. Prokaryote abundance (up to 1.3 × 10⁹ cells L⁻¹) and extracellular enzymatic activities (up to 24.3 μM h⁻¹ for amino-peptidase activity) were extremely high, indicating high rates of organic C degradation in the bottom sea ice. Despite this, prokaryote C production values were very low (range 5–30 ng C L⁻¹ h⁻¹), suggesting that most of the degraded organic C was not channelled into prokaryote biomass. In the platelet ice, we found similar organic C concentrations, prokaryote abundance and biomass values and even higher extracellular enzymatic activities, but values of prokaryote C production (range 800–4,200 ng C L⁻¹ h⁻¹) were up to three orders of magnitude higher than in the intact bottom sea ice. Additional field and laboratory experiments revealed that the dissolved organic material derived from algae accumulating in the bottom sea ice significantly reduced prokaryote C production, suggesting the presence of a potential allopathic control of sympagic algae on prokaryote growth. This article belongs to a special topic: Five articles on Sea-ice communities in Terra Nova Bay (Ross Sea), coordinated by L. Guglielmo and V. Saggiomo, appear in this issue of Polar Biology. The studies were conducted in the frame of the National Program of Research in Antarctica (PNRA) of Italy.     

Temporal and vertical variations of lipid biomarkers during a bottom ice diatom bloom in the Canadian Beaufort Sea: further evidence for the use of the IP25 biomarker as a proxy for spring Arctic sea ice

Variations in the concentrations of the sea ice diatom biomarker, IP₂₅ (Ice Proxy with 25 carbon atoms), were measured in the bottom 10 cm of sea ice collected from the eastern Beaufort Sea and Amundsen Gulf from January to June 2008, as part of the International Polar Year–Circumpolar Flaw Lead system study. Temporal and vertical changes in IP₂₅ concentrations were compared against other biomarkers and indicators of ice algal production. IP₂₅ was not detected in sea ice samples collected from mid-winter to early spring, likely as a result of light-limiting conditions for algal growth and accumulation. From early March to mid-June, IP₂₅ concentrations correlated well with those of fatty acids (r = 0.79; P < 0.001), less so with total sterols (r = 0.63; P < 0.001) and qualitatively with chlorophyll a concentrations and diatom cell abundances from adjacent sea ice cores. Approximately 90% of the total sea ice IP₂₅ accumulation occurred from mid-March to late-May, coincident with the ice algal bloom period. The majority (ca. 87–93%) of IP₂₅ was biosynthesised within the lower 5 cm of the sea ice where brine volume fractions were >5% which is consistent with the hypothesis that brine channel connectivity limits the internal colonisation of sea ice by diatoms. Maximum IP₂₅ concentrations occurred at 1–3 cm from the ice–water interface providing further evidence for a selective sea ice diatom origin for this biomarker. In contrast, vertical concentration profiles for fatty acids and sterols indicated mixed sources for these biomarkers.     

Sea ice meiofauna distribution on local to pan‐Arctic scales

Arctic sea ice provides microhabitats for biota that inhabit the liquid‐filled network of brine channels and the ice–water interface. We used meta‐analysis of 23 published and unpublished datasets comprising 721 ice cores to synthesize the variability in composition and abundance of sea ice meiofauna at spatial scales ranging from within a single ice core to pan‐Arctic and seasonal scales. Two‐thirds of meiofauna individuals occurred in the bottom 10 cm of the ice. Locally, replicate cores taken within meters of each other were broadly similar in meiofauna composition and abundance, while those a few km apart varied more; 75% of variation was explained by station. At the regional scale (Bering Sea first‐year ice), meiofauna abundance varied over two orders of magnitude. At the pan‐Arctic scale, the same phyla were found across the region, with taxa that have resting stages or tolerance to extreme conditions (e.g., nematodes and rotifers) dominating abundances. Meroplankton, however, was restricted to nearshore locations and landfast sea ice. Light availability, ice thickness, and distance from land were significant predictor variables for community composition on different scales. On a seasonal scale, abundances varied broadly for all taxa and in relation to the annual ice algal bloom cycle in both landfast and pack ice. Documentation of ice biota composition, abundance, and natural variability is critical for evaluating responses to decline in Arctic sea ice. Consistent methodology and protocols must be established for comparability of meiofauna monitoring across the Arctic. We recommend to (1) increase taxonomic resolution of sea ice meiofauna, (2) focus sampling on times of peak abundance when seasonal sampling is impossible, (3) include the bottom 30 cm of ice cores rather than only bottom 10 cm, (4) preserve specimens for molecular analysis to improve taxonomic resolution, and (5) formulate a trait‐based framework that relates to ecosystem functioning.     

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.     

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.     

Small scale vertical gradients of Arctic ice algal photophysiological properties

Photosynthetic parameters of phytoplankton and sea ice algae from landfast sea ice of the Chukchi Sea off Point Barrow, Alaska, were assessed in spring 2005 and winter through spring 2006 using Pulse Amplitude Modulated (PAM) fluorometry including estimates of maximum quantum efficiency (F _v/F _m), maximum relative electron transport rate (rETR_max), photosynthetic efficiency (α), and the photoadaptive index (E _k). The use of centrifuged brine samples allowed to document vertical gradients in ice algal acclimation with 5 cm vertical resolution for the first time. Bottom ice algae (0–5 cm from ice–water interface) expressed low F _v/F _m (0.331–0.426) and low α (0.098–0.130 (μmol photons m⁻²s⁻¹)⁻¹) in December. F _v/F _m and α increased in March and May (0.468–0.588 and 0.141–0.438 (μmol photons m⁻²s⁻¹)⁻¹, respectively) indicating increased photosynthetic activity. In addition, increases in rETR_max (3.3–16.4 a.u.) and E _k (20–88 μmol photons m⁻² s⁻¹) from December to May illustrates a higher potential for primary productivity as communities become better acclimated to under-ice light conditions. In conclusion, photosynthetic performance by ice algae (as assessed by PAM fluorometry) was tightly linked to sea ice salinity, temperature, and inorganic nutrient concentrations (mainly nitrogen).     

Zooplankton boom and ice amphipod bust below melting sea ice in the Amundsen Gulf, Arctic Canada

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m⁻³) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m⁻³) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind. m⁻²) and wet biomass (<0.2 g m⁻²). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.     

Effects of Incubation Temperature on Growth and Production of Exopolysaccharides by an Antarctic Sea Ice Bacterium Grown in Batch Culture

The sea ice microbial community plays a key role in the productivity of the Southern Ocean. Exopolysaccharide (EPS) is a major component of the exopolymer secreted by many marine bacteria to enhance survival and is abundant in sea ice brine channels, but little is known about its function there. This study investigated the effects of temperature on EPS production in batch culture by CAM025, a marine bacterium isolated from sea ice sampled from the Southern Ocean. Previous studies have shown that CAM025 is a member of the genus Pseudoalteromonas and therefore belongs to a group found to be abundant in sea ice by culture-dependent and -independent techniques. Batch cultures were grown at −2°C, 10°C, and 20°C, and cell number, optical density, pH, glucose concentration, and viscosity were monitored. The yield of EPS at −2°C and 10°C was 30 times higher than at 20°C, which is the optimum growth temperature for many psychrotolerant strains. EPS may have a cryoprotective role in brine channels of sea ice, where extremes of high salinity and low temperature impose pressures on microbial growth and survival. The EPS produced at −2°C and 10°C had a higher uronic acid content than that produced at 20°C. The availability of iron as a trace metal is of critical importance in the Southern Ocean, where it is known to limit primary production. EPS from strain CAM025 is polyanionic and may bind dissolved cations such at trace metals, and therefore the presence of bacterial EPS in the Antarctic marine environment may have important ecological implications.     

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.     

Decomposition ‘hotspots’ in a rewetted peatland: implications for water quality and carbon cycling

The hypothesis that specific components of seawater, such as particulate, dissolved and colloidal organic and inorganic material, render virions non-infective has long been postulated, but never rigorously tested. To address this hypothesis, the plaque assay method was used to derive infective decay rates, k, of two bacteriophages—P1 (marine host: PWH3a) and T4 (enteric host: E. coli B). We compared k values of bacteriophage suspended in serial filtrations of seawater, with and without autoclaving and UV oxidation. Both phages exhibited reduced decay rates in particle-free water (<0.2 μm) compared to <10 μm filtrate. The largest decrease in virion decay rates was achieved by autoclaving the 0.2 μm filtrate. UV oxidation of <0.2 μm filtrate, however, yielded higher decay rates than observed in autoclaved treatments. The lowest k values were seen in ultra-filtered seawater (<10 kDa). Exposure to a wide range of concentrations of Pronase E (a proteolytic enzyme), inorganic clay (kaolinite or montmorillonite), and organic particles (phytoplankton debris) did not promote phage inactivation. P1 infective titers were also not consistently reduced by exposures to axenic cultures of a resistant host mutant (PWH3a-R) and a non-host marine bacterium (MB-5). Finally, phage were exposed to a range of temperatures to derive activation energies required for phage inactivation. Application of the Arrhenius model to inactivation of T4 and P1 yielded activation energies (E _a) of 49 and 40 kJ mol⁻¹, respectively. This is the first comprehensive analysis in which specific seawater components were assayed for their ability to inactivate bacteriophage. Inactivation of these phage does not appear to depend on capsomere denaturation, proteolytic extracellular enzymes, sorption to non-host bacteria, clay particles or particulate organic debris, but is accelerated by naturally occurring particles, which include living organisms, and heat-labile colloids and macromolecules >10 kDa.     

Physiological adaptations to low temperature and brine exposure in the circumpolar amphipod Gammarus wilkitzkii

Summary The amphipod Gammarus wilkitzkii does not survive being frozen totally into solid sea ice. When the animals are cooled in air or freezing seawater, they will freeze and die at a temperature of about-4° C. However, during sea ice growth, the amphipods may tolerate to stay in the vicinity of the ice by conforming to the ambient brine in a salinity range of 34 ppt to about 60 ppt. A passive relationship between the concentrations of the haemolymph and seawater Na⁺ and Cl^-, lowers the melting point of the body fluids of the animals, thus preventing internal ice formation at low temperatures.     

Effects of incubation temperature on growth and production of exopolysaccharides by an antarctic sea ice bacterium grown in batch culture

The sea ice microbial community plays a key role in the productivity of the Southern Ocean. Exopolysaccharide (EPS) is a major component of the exopolymer secreted by many marine bacteria to enhance survival and is abundant in sea ice brine channels, but little is known about its function there. This study investigated the effects of temperature on EPS production in batch culture by CAM025, a marine bacterium isolated from sea ice sampled from the Southern Ocean. Previous studies have shown that CAM025 is a member of the genus Pseudoalteromonas and therefore belongs to a group found to be abundant in sea ice by culture-dependent and -independent techniques. Batch cultures were grown at -2 degrees C, 10 degrees C, and 20 degrees C, and cell number, optical density, pH, glucose concentration, and viscosity were monitored. The yield of EPS at -2 degrees C and 10 degrees C was 30 times higher than at 20 degrees C, which is the optimum growth temperature for many psychrotolerant strains. EPS may have a cryoprotective role in brine channels of sea ice, where extremes of high salinity and low temperature impose pressures on microbial growth and survival. The EPS produced at -2 degrees C and 10 degrees C had a higher uronic acid content than that produced at 20 degrees C. The availability of iron as a trace metal is of critical importance in the Southern Ocean, where it is known to limit primary production. EPS from strain CAM025 is polyanionic and may bind dissolved cations such at trace metals, and therefore the presence of bacterial EPS in the Antarctic marine environment may have important ecological implications.     

Linking ice structure and microscale variability of algal biomass in Arctic first-year sea ice using an in situ photographic technique

Microscale photographs were taken of the ice bottom to examine linkages of algal chlorophyll a (chl a) biomass distribution with bottom ice features in thick Arctic first-year sea ice during a spring field program which took place from May 5 to 21, 2003. The photographic technique developed in this paper has resulted in the first in situ observations of microscale variability in bottom ice algae distribution in Arctic first-year sea ice in relation to ice morphology. Observations of brine channel diameter (1.65–2.68 mm) and number density (5.33–10.35 per 100 cm²) showed that the number of these channels at the bottom of thick first-year sea ice may be greater than previously measured on extracted ice samples. A variogram analysis showed that over areas of low chl a biomass (≤20.7 mg chl a m⁻²), patchiness in bottom ice chl a biomass was at the scale of brine layer spacing and small brine channels (∼1–3 mm). Over areas of high chl a biomass (≥34.6 mg chl a m⁻²), patchiness in biomass was related to the spacing of larger brine channels on the ice bottom (∼10–26 mm). Brine layers and channels are thought to provide microscale maxima of light, nutrient replenishment and space availability which would explain the small scale patchiness over areas of low algal biomass. However, ice melt and erosion near brine channels may play a more important role in areas with high algal biomass and low snow cover.     

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.     

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     

Fluxes associated with brine motion in growing sea ice

Summary Laboratory and field studies have demonstrated that fluid motion occurs at two locations in growing sea ice: in a network of brine channels and within the skeletal layer at the ice-water interface. Brine channel fluxes estimated using brine channel areal density from natural sea ice and channel velocities from laboratory studies are compared with recent measurements reported in the literature. Fluxes into the porous skeletal layer of sea ice may be estimated using rates of nutrient uptake by ice algae and adjacent seawater nutrient concentrations. Both approaches indicate fluxes of the order of 10^-6 cc cm^-2 s^-1 (l m^-2 h^-1), which are about equal to fluxes reported in bioirrigated sediments. Fluxes of this magnitude indicate a very short residence time for the liquid phase in the skeletal layer, suggesting that this fluid motion may be important in maintaining the ice algae community.