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     

Influence of sea ice on the composition of the spring phytoplankton bloom in the northern Baltic Sea

During the late winter and spring of 1994, the influence of sea ice on phytoplankton succession in the water was studied at a coastal station in the northern Baltic Sea. Ice cores were taken together with water samples from the underlying water and analysed for algal composition, chlorophyll a and nutrients. Sediment traps were placed under the ice and near the bottom, and the sedimented material was analysed for algal composition. The highest concentration of ice algae (4.1 mmol C m⁻²) was found shortly before ice break-up in the middle of April, coincidental with the onset of an under-ice phytoplankton bloom. The ice algae were dominated by the diatoms Chaetoceros wighamii Brightwell, Melosira arctica (Ehrenberg) Dickie and Nitzschia frigida Grunow. Under the ice the diatom Achnanthes taeniata Grunow and the dinoflagellate Peridiniella catenata (Levander) Balech were dominant. Calculations of sinking rates and residence times of the dominant ice algal species in the photic water column indicated that only one ice algal species (Chaetoceros wighamii) had a seeding effect on the water column: this diatom dominated the spring phytoplankton bloom in the water together with Achnanthes taeniata and Peridiniella catenata. Received: 9 May 1997 / Accepted: 15 February 1998     

Nitrogen uptake in the infiltration community, an ice algal community in Antarctic pack-ice

An infiltration community was the dominating ice algal community in pack-ice off Queen Maud Land, Southern Ocean, in January 1993. The community was dominated by autotrophic processes, and the most common species were the prymnesiophyte Phaeocystis antarctica and the diatoms Chaetoceros neglectus and Fragilariopsis cylindrus. The concentration of chlorophyll a was 1.3–47.9 μg l⁻¹, and the inner part of the community was nitrate depleted. Uptake rates of nitrate, nitrite, ammonium, urea and amino acids were measured using ¹⁵N. Nitrate was the major nitrogen source for ice algal growth (67 ± 6% nitrate uptake). It is suggested that % nitrate uptake in the infiltration community decreases during the growth season, from 92% during spring (literature data) to 67% during summer. Scalar irradiance in the infiltration community was high and variable. It reached ca. 2000 μmol m⁻² s⁻¹ at some locations, and nitrate uptake rate was potentially photoinhibited at irradiances >500 μmol m⁻² s⁻¹. Nitrate uptake rate in an average infiltration community (0.6 m of snow cover) was lowered by 13% over a 2-week period due to photoinhibition. Received: 16 December 1996 / Accepted: 5 January 1998     

Phytoplankton biomass and primary production in the marginal ice zone of the northwestern Weddell Sea during austral summer

During the austral summer of 1995, distributions of phytoplankton biomass (as chlorophyll a), primary production, and nutrient concentrations along two north-south transects in the marginal ice zone of the northwestern Weddell Sea were examined as part of the 8th Korean Antarctic Research Program. An extensive phytoplankton bloom, ranging from 1.6 to 11.2 mg m⁻³ in surface chlorophyll a concentration, was encountered along the eastern transect and extended ca. 180 km north of the ice edge. The spatial extent of the bloom was closely related to the density field induced by the input of meltwater from the retreating sea ice. However, the extent (ca. 200 km) of the phytoplankton bloom along the western transect exceeded the meltwater-influenced zone (ca. 18 km). The extensive bloom along the western transect was more closely related to local hydrography than to the proximity of the ice edge and the resulting meltwater-induced stability of the upper water column. In addition, the marginal ice zone on the western transect was characterized by a deep, high phytoplankton biomass (up to 8 mg Chl a m⁻³) extending to 100-m depth, and the decreased nutrient concentration, which was probably caused by passive sinking from the upper euphotic zone and in situ growth. Despite the low bloom intensity relative to the marginal ice zone in both of the transects, mean primary productivity (2.6 g C m⁻² day⁻¹) in shelf waters corresponding to the northern side of the western transect was as high as in the marginal ice zone (2.1 g C m⁻² day⁻¹), and was 4.8 times greater than that in open waters, suggesting that shelf waters are as highly productive as the marginal ice zone. A comparison between the historical productivity data and our data also shows that the most productive regions in the Southern Ocean are shelf waters and the marginal ice zone, with emerging evidence of frontal regions as another major productive site. Accepted: 27 September 1998     

Water level and currents of tidal and infragravity periods at Tague Reef, St. Croix (USVI)

A two-week study, at Tague Reef, St. Croix, USVI investigated the magnitude and spatial variation of tides, sea level differences, infragravity waves, and unidirectional cross-reef currents on a modern coral reef. Infragravity oscillations of water level (∼ 27 min period) of 1–2 cm height correlate with a quarter wavelength resonance over the shelf. Particle displacements associated with these waves may be important to the dispersive characteristics of the reef environment. Estimates of cross-reef mass transport per unit width ranged from 0.058 to 0.032 m²s ^-1. Sea level differences across the reef (1–4 cm) varied at diurnal and infragravity periods with contributions from wave set-up, and a small contribution from cross-shelf wind stress to the observed sea level differences. The quadratic bottom friction coefficient over the reef was estimated at 0.06–0.2, 20–70 times greater than on open shelves, reflecting the reef’s extreme bottom roughness. Accepted: 28 February 1998     

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     

Summer plankton beneath the McMurdo Ice Shelf at white Island,McMurdo Sound,Antarctica

 The zooplankton of the under-shelf-ice ecosystem at White Island (78^°10′ S, 167^°30′ E), McMurdo Sound, Antarctica was investigated during December 1976 and January 1977. The water column was sampled through a hole in the McMurdo Ice Shelf over a water depth of 67 m. Seawater temperatures under the ice shelf ranged from −1.91 to 1.96^°C. Dissolved oxygen levels ranged from 5.0–6.05 ml l^-1 in early December to 4.65–4.8 ml l^-1 in late January. Current speeds of up to 0.13 m s^-1 were recorded at a depth of 50 m and a predominantly northward flow was detected. Light levels under the shelf ice were low with less than 1% of the incident light being transmitted to a depth of 3 m. No chlorophyll a was detected within the water column throughout the investigation. Mean zooplankton biomass values in the water column ranged from 12 to 447 mg wet weight m^-3 and were similar to values recorded elsewhere from Antarctic inshore waters, but were very much higher than those recorded from under seasonal sea ice in McMurdo Sound. Thirty-two zooplankton species were recorded including 1 ostracod, 21 copepods (10 calanoids, 3 cyclopoids and 8 harpacticoids), 4 amphipods, 2 euphausiids, a chaetognath and 3 pteropods. Larvae of polychaetes and fish were found on some occasions. The species composition in general was similar to that recorded from McMurdo Sound and other Antarctic inshore localities. Among the Copepoda, however, there were a number of species, especially among the Harpacticoidea, that have not been found previously in McMurdo Sound and the Ross Sea, but that are known to be associated with ice in other localities in Antarctica. Two recently described species are known only from White Island. They were present in the water column but were most abundant in the surface water of the tide crack where they were the most abundant zooplankters. The tide crack, which probably is an extension of the under-ice habitat, is apparently a significant nursery area for amphipods and copepod species. Received: 23 November 1994/Accepted 7 May 1995     

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     

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     

Abundance, biomass and small-scale distribution of cryopelagic amphipods in the Franz Josef Land area (Arctic)

Arctic sea ice is inhabited by several amphipod species. Abundance, biomass and small-scale distribution of these cryopelagic (=ice associated) amphipods were investigated near Franz Josef Land in summer 1994. The mean abundance of all species was 420 ind./m²; the mean biomass was 10.61 g ww/m². Gammarus wilkitzkii was the dominant species, whereas Apherusa glacialis, Onisimus nanseni and O. glacialis were only scarcely found. Amphipods were concentrated at the edges of ice floes and were less frequent in areas further away under the ice. The relationship between the distribution and ecological/physiological requirements of cryopelagic amphipods, as well as the small-scale morphology of Arctic sea ice, are discussed. Received: 14 January 1998 / Accepted 14 April 1998     

Pelagic occurrence of the sympagic amphipod Gammarus wilkitzkii in ice-free waters of the Greenland Sea – dead end or part of life-cycle?

The sympagic (=ice-associated) amphipod Gammarus wilkitzkii usually lives attached to the underside of Arctic sea ice. During an expedition to the Greenland Sea in May/June 1997, high numbers of this species were found in pelagic Rectangular Midwater Trawl catches (0–500 m water depth) in an ice-free area, 35–42 km away from the ice edge. The amphipods seemed to have maintained position in the water column for at least 4 days. Mean biomass data (length: 2.9 cm, organic content: 73% dry mass), gut fullness (>50% in 85% of specimens) and sex ratio (females:males = 1:1.5) of these amphipods were very similar to values for under-ice populations. Due to their relatively high body density (mean: 1.134 g cm⁻³), the energy demand for swimming was assumed to be high. Measurements of oxygen consumption of swimming and resting amphipods (8.8 and 4.0 J g wet mass⁻¹ day⁻¹, respectively) suggested that, from an energetic point of view, G. wilkitzkii would maintain position in an ice-free water column for the time period. Accepted: 11 January 1999     

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     

Effects of snow removal and algal photoacclimation on growth and export of ice algae

Net growth of ice algae in response to changes in overlying snow cover was studied after manipulating snow thickness on land-fast, Arctic sea ice. Parallel laboratory experiments measured the effect of changing irradiance on growth rate of the ice diatom, Nitzschia frigida. After complete removal of thick snow (≥9 cm), in situ ice algae biomass declined (over 7–12 days), while removal of thin snow layers (4–5 cm), or partial snow removal, increased net algal growth. Ice bottom ablation sometimes followed snow removal, but did not always result in net loss of algae. Similarly, in laboratory experiments, small increases in irradiance increased algal growth rate, while greater light shifts suppressed growth for 3–6 days. However, N. frigida could acclimate to relatively high irradiance (110 μmol photons m² s⁻¹). The results suggest that algal loss following removal of a thick snow layer was due to the combination of photoinhibition and bottom ablation. The smaller relative increase in irradiance after removal of thin or partial snow layers allowed algae to maintain high specific-growth rates that compensated for loss from physical mechanisms. Thus, the response of ice algae to snow loss depends both on the amount of change in snow depth and algal photophysiology. The complex response of ice algae growth and export loss to frequently changing snow fields may contribute to horizontal and temporal patchiness of ecologically and biogeochemically important variables in sea ice and should be considered in predictions of how climate change will affect Arctic marine ecosystems.     

Microbiological Explorations in the Northern Part of the Barents Sea in Early Winter

The total number of microorganisms and rates of microbial processes of the carbon cycle were determined in snow, sea ice, water, and seafloor sediments of the northern part of the Barents Sea from September to October, 1998. The explorations were carried out in two areas: along the transection from Franz Josef Land to Victoria Island and along the continental slope region covered with solid ice at latitude 81°–82° N and longitude 37°–39° E. At the time of study, the ice cover was represented by thick one-year old ice (up to 1.2 m), perennial ice (up to 1.85 m), and pack ice. The number of bacteria in the snow cover, sea ice, and seawater was 12 to 14, 50 to 110, and 10 to 240 × 10³ cells/ml, respectively. Rates of dark CO₂ assimilation, glucose utilization, and methane oxidation by bacteria were determined. The highest rate of microbial processes was found in samples of the lowermost newly formed sea ice. The lowest level of activity for all processes was observed in melted snow water. A direct relation was shown between the concentration of C_org, the bacterial biomass, and the values of ¹³C_org in mixtures of melted snow and ice. The number of microorganisms and rates of microbial processes in seafloor sediments measured at the stations on the continental slope are comparable to those in the central part of the Barents Sea and the northern part of the Kara Sea.     

Seasonal variation in plankton, submicrometre particles and size-fractionated dissolved organic carbon in Antarctic coastal waters

Concentrations of plankton, suspended particles 0.74–87 μm equivalent spherical diameter and dissolved organic carbon (DOC) were measured from May to February at an Antarctic coastal site. Bacteria-sized particles 0.74–1 μm diameter, and bacterial cells and heterotrophic protists all exhibited a seasonal minimum during winter and maxima in summer. Bacteria composed <10% of the bacteria-sized particles. Release of autotrophic protists from the ice caused water column biomass of autotrophs to reach maximum concentrations in October and November, but maximum cell concentration in the water column was reached in January. Microheterotroph biomass weakly reflected the release of the ice algal community but reached maximum concentration during the water column bloom in January. Total DOC concentrations varied from 0.36 mg C l⁻¹ in July to 3.10 mg C l⁻¹ in October, with a yearly average of 1.51 mg C l⁻¹. Ultrafiltration of DOC revealed that the molecular weight composition of the DOC differed greatly through the year. DOC <5 kDa molecular weight reached a maximum of 1.25 mg C l⁻¹ in October and accounted for up to 60% of total DOC in July. Concentrations of high molecular weight DOC (>100 kDa) were highest in July and November, with the DOC (100 kDa–0.5 μm) fraction reaching a maximum of 1.22 mg C l⁻¹ in November and composing 82% of the total DOC in January. Wet chemical oxidation and high-temperature catalytic oxidation organic carbon analyses were compared. Good correlation was observed between methods during summer but no significant correlation existed in winter, indicating that winter DOC may be refractory. Accepted: 21 March 2000     

Seasonal distribution of sympagic amphipods near Chesterfield Inlet, N.W.T., Canada

The seasonal distribution of sympagic amphipods was investigated in the Chesterfield Inlet area of northwestern Hudson Bay (63°30′N). Amphipod abundance was measured by photographic samples and species composition was determined by sweep net samples. Twelve species of amphipods were collected, the most common being Ischyrocerus anguipes, Pontogeneia inermis, Apherusa megalops and Weyprechtia pinguis. The major environmental variable affecting amphipod distribution was water depth. Amphipod abundance was highest near 20 m and near zero past 50 m. The maximum recorded abundance was 1367 m⁻². A minor factor affecting the distribution of amphipods was snow depth, through its modifying effect on light and thereby the growth of ice algae. Amphipods began to inhabit the sea ice shortly after its formation. From the beginning of March, the number of amphipods on the ice increased steadily to about the 3rd week of April, after which numbers declined. This pattern coincided with the seasonal ice algae abundance. Amphipods reduced ice algal biomass over 20-m depth by 63%. No evidence of diurnal changes in abundance was observed. Received: 15 May 1996 / Accepted: 4 November 1996     

Holocene paleoclimate change in the Antarctic Peninsula: evidence from the diatom,sedimentary and geochemical record

Holocene, marine deposition in Lallemand Fjord, Antarctic Peninsula, is reinterpreted using statistical analyses (cluster analysis, analysis of variance, nonmetric multidimensional scaling and multiple regression) to compare diatom assemblages and the primary sedimentological proxies. The assemblages have been deposited in a variable sea ice zone over the last ca. 10,500 yr BP in response to a climate change. In the Late Pleistocene/early Holocene (10,580–7890 yr BP), a sea ice diatom assemblage was deposited in the presence of a retreating ice shelf at the head of the fjord. In the mid Holocene (7890–3850 yr BP), an open water assemblage was deposited and sea ice cover was at a minimum. We associate the assemblage with climatic warming, which characterizes much of the Antarctic Peninsula during this time. A second sea ice assemblage, different from that deposited in the early Holocene, has been deposited in Lallemand Fjord since the late Holocene (<3850 yr BP). The assemblage reflects Neoglacial cooling, an increase in sea ice extent and/or an advance of the Müller Ice Shelf.     

Nutrient stress gradient in the bottom 5 cm of fast ice, McMurdo Sound, Antarctica

One-centimeter-scale vertical sampling of fast ice from McMurdo Sound, Antarctica reveals evidence of progressive nutrient limitation with distance above the ice/water interface. Over the bottom 6 cm photosynthetically active radiation increases by between 1.8 and 3 times, C:N increases from 6.8 to 19.8 and δ¹³C increases from −18 to −12. Fatty acid composition also changes with a consistent decline in polyunsaturated fatty acids and a rise in saturated fatty acids. These factors all suggest severe and progressive nutrient limitation with distance from the ice/water interface. Accepted: 5 September 1998     

The diversity,abundance and fate of ice algae and phytoplankton in the Bering Sea

Despite being an essential part of the marine food web during periods of ice cover, sea ice algae have not been studied in any detail in the Bering Sea. In this study, we investigated the diversity, abundance and ultimate fate of ice algae in the Bering Sea using sea ice, water and sub-ice sediment trap samples collected during two spring periods in 2008 and 2009: ice growth (March–mid-April) and ice melt (mid-April–May). The total ice algal species inventory included 68 species, dominated by typical Arctic ice algal diatom taxa. Only three species were determined from the water samples; we interpret the strong overlap in species as seeding of algal cells from the sea ice. Algal abundances in the ice exceeded 10⁷ cells l^?1 in the bottom 2-cm layer and were on average three orders of magnitude higher than in the water column. The vertical flux of algal cells beneath the ice during the period of ice melt (>10⁸ cells m^?2 day^?1) exceeded export during the ice growth period by one order of magnitude; the vertical flux during both periods can only be sustained by the release of algae from the ice. Differences in the relative species proportions of algae among sample types indicated that the fate of the released ice algae was species specific, with some taxa contributing to seeding in the water column, while other taxa were preferentially exported.     

Protozooplankton in the Weddell Sea,Antarctica: Abundance and distribution in the ice-edge zone

Summary Protozooplankton were sampled in the iceedge zone of the Weddell Sea during the austral spring of 1983 and the austral autumn of 1986. Protozooplankton biomass was dominated by flagellates and ciliates. Other protozoa and micrometazoa contributed a relatively small fraction to the heterotrophic biomass. During both cruises protozoan biomass, chlorophyll a concentrations, phytoplankton production and bacterial biomass and production were low at ice covered stations. During the spring cruise, protozooplankton, phytoplankton, and bacterioplankton reached high concentrations in a welldeveloped ice edge bloom 100 km north of the receding ice edge. During the autumn cruise, the highest concentrations of biomass were in open water well-separated from the ice edge. Integrated protozoan biomass was <12% of the biomass of phytoplankton during the spring cruise and in the autumn the percentages at some stations were >20%. Bacterial biomass exceeded protozooplankton biomass at ice covered stations but in open water stations during the fall cruise, protozooplankton biomass reached twice that of bacteria in the upper 100m of the water column. The biomass of different protozoan groups was positively correlated with primary production, chlorophyll a concentrations and bacterial production and biomass, suggesting that the protozoan abundances were largely controlled by prey availability and productivity. Population grazing rates calculated from clearance rates in the literature indicated that protozooplankton were capable of consuming significant portions of the daily phyto- and bacterioplankton production.