Abstract
Pack ice around Svalbard was sampled during the expedition ARK XIX/1 of RV “Polarstern” (March–April 2003) in order to determine environmental conditions, species composition and abundances of sea-ice algae and heterotrophic protists during late winter. As compared to other seasons, species diversity of algae (total 40 taxa) was not low, but abundances (5,000–448,000 cells l−1) were lower by one to two orders of magnitude. Layers of high algal abundances were observed both at the bottom and in the ice interior. Inorganic nutrient concentrations (NO2, NO3, PO4, Si(OH)4) within the ice were mostly higher than during other seasons, and enriched compared to seawater by enrichment indices of 1.6–24.6 (corrected for losses through the desalination process). Thus, the survival of algae in Arctic pack ice was not limited by nutrients at the beginning of the productive season. Based on less-detailed physical data, light was considered as the most probable factor controlling the onset of the spring ice-algal bloom in the lower part of the ice, while low temperatures and salinities inhibit algal growth in the upper part of the ice at the end of the winter. Incorporation of ice algae probably took place during the entire freezing period. Possible overwintering strategies during the dark period, such as facultative heterotrophy, energy reserves, and resting spores are discussed.




Similar content being viewed by others
Explore related subjects
Discover the latest articles and news from researchers in related subjects, suggested using machine learning.References
Arrigo KR (2003) Primary production in sea ice. In: Thomas DN, Dieckmann GS (eds) Sea ice—an introduction to its physics, chemistry, biology and geology. Blackwell Science Ltd, Oxford, pp 143–183
Assur A (1958) Composition of sea ice and its tensile strength. Nat Res Council Publ 598:106–138
Bartsch A (1989) Die Eisalgenflora des Weddellmeeres (Antarktis): Artenzusammensetzung und Biomasse sowie Ökophysiologie ausgewählter Arten. Ber Polarforsch 63:1–110
Booth BC, Horner RA (1997) Microalgae on the Arctic Ocean Section, 1994: species abundance and biomass. Deep-Sea Res II 44:1607–1622
Comiso JC (2003) Large-scale characteristics and variability of the global sea ice cover. In: Thomas DN, Dieckmann GS (eds) Sea ice—an introduction to its physics, chemistry, biology and geology. Blackwell Science Ltd, Oxford, pp 112–142
Cota GF, Legendre L, Gosselin M, Ingram RG (1991) Ecology of bottom ice algae: I. Environmental controls and variability. J Mar Syst 2:257–277
Druzhkov NV, Druzhkova EI, Kuznetsov LL (2001) The sea-ice algal community of seasonal pack ice in the southwestern Kara Sea in late winter. Polar Biol 24:70–72
Eicken H, Lensu M, Leppäranta M, Tucker III WB, Gow AJ, Salmela O (1995) Thickness, structure, and properties of level summer multiyear ice in the Eurasian sector of the Arctic Ocean. J Geophys Res 100(C11):22697–22710
Fahl K, Kattner G (1993) Lipid content and fatty acid composition of algal communities in sea-ice and water from the Weddell Sea (Antarctica). Polar Biol 13:405–409
Fehling J (2000) Sympagische Protistengemeinschaften im arktischen Packeis der Framstraßenregion. Unpubl Diploma thesis, Univ Kiel, pp 86
Frankenstein G, Garner R (1967) Equations for determining the brine volume of sea ice from -0.5° to -22.9°C. J Glaciol l6:943–944
Garrison DL, Buck KR (1986) Organism losses during ice melting: a serious bias in sea ice community studies. Polar Biol 6:237–239
Golden KM, Ackley SF, Lytle VI (1998) The percolation phase transition in sea ice. Science 282:2238–2241
Gosselin M, Legendre L, Therriault JC, Demers S, Rochet M (1986) Physical control of the horizontal patchiness of sea-ice microalgae. Mar Ecol Prog Ser 29:289–298
Gosselin M, Legendre L, Therriault JC, Demers S (1990) Light and nutrient limitation of sea-ice microalgae (Hudson Bay, Canadian Arctic). J Phycol 26:220–232
Gosselin M, Levasseur M, Wheeler PA, Horner RA, Booth BC (1997) New measurements of phytoplankton and ice algal production in the Arctic Ocean. Deep-Sea Res II 44:1623–1644
Gradinger R (1998) Environmental controls of Arctic pack ice algal composition and development—a synopsis. Habilitation thesis. University of Kiel, Kiel, 82 pp
Gradinger R (1999) Vertical fine structure of the biomass and composition of algal communities in Arctic pack ice. Mar Biol 133:745–754
Gradinger R, Friedrich C, Spindler M (1999) Abundance, biomass and composition of the sea ice biota of the Greenland Sea pack ice. Deep-Sea Res II 46:1457–1472
Gradinger R, Ikävalko J (1998) Organism incorporation into newly forming Arctic sea ice in the Greenland Sea. J Plankt Res 20:871–886
Gradinger R, Spindler M, Henschel D (1991) Development of Arctic sea-ice organisms under graded snow cover. Polar Res 10:295–307
Grossi SM, Kottmeier ST, Moe RL, Taylor GT, Sullivan CW (1987) Sea ice microbial communities. VI. Growth and primary production in bottom ice under graded snow cover. Mar Ecol Prog Ser 35:153–164
Haecky P, Jonsson S, Andersson A (1998) Influence of sea ice on the composition of the spring phytoplankton bloom in the Northern Baltic Sea. Polar Biol 20:1–8
Hegseth EN (1992) Sub-ice algal assemblages of the Barents Sea: species composition, chemical composition, and growth rates. Polar Biol 12:485–496
Hegseth EN (1997) Phytoplankton of the Barents Sea—the end of a growth season. Polar Biol 17:235–241
Heimdal BR (1983) Phytoplankton and nutrients in the waters North-west of Spitsbergen in the autumn of 1979. J Plankton Res 5:901–918
Horner RA, Ackley SF, Dieckmann GS, Gulliksen B, Hoshiai T, Legendre L, Melnikov IA, Reeburgh WS, Spindler M, Sullivan CW (1992) Ecology of sea ice biota. 1. Habitat, terminology, and methodology. Polar Biol 12:417–427
Horner R, Schrader GC (1982) Relative contribution of ice algae, phytoplankton, and benthic microalgae to primary production in nearshore regions of the Beaufort Sea. Arctic 35:485–503
Ikävalko J (2004) Checklist of unicellular and invertebrate organisms within and closely associated with sea ice in the Arctic regions. Meri Series 52:1–41
Junge K, Eicken H, Deming JW (2004) Bacterial activity at -2 to -20°C in Arctic wintertime sea ice. Appl Environ Microbiol 70:550–557
Junge K, Eicken H, Swanson BD, Deming JW (2006) Bacterial incorporation of leucine into protein down to -20°C with evidence for potential activity in sub-eutectic saline ice formations. Cryobiology 52:417–429
Kaartokallio H (2001) Evidence for active microbial nitrogen transformations in sea ice (Gulf of Bothnia, Baltic Sea) in midwinter. Polar Biol 24:21–28
Kirst GO, Wiencke C (1995) Ecophysiology of polar algae. J Phycol 31:181–199
Kottmeier ST, Sullivan CW (1988) Sea ice microbial communities (SIMCO). 9. Effects of temperature and salinity on rates of metabolism and growth of autotrophs and heterotrophs. Polar Biol 8:293–304
Krembs C, Eicken H, Junge K, Deming JW (2002) High concentrations of exopolymeric substances in Arctic winter sea ice: implication for the polar ocean carbon cycle and cryoprotection of diatoms. Deep-Sea Res I 49:2163–2181
Leppäranta M, Manninen T (1988) The brine and gas content of sea ice with attention to low salinities and high temperatures. Finn Inst Mar Res Intern Rep 2:1–14
Lieser J (2005) Sea ice conditions in the northern North Atlantic in 2003 and 2004. Ber Polarforsch 504:1–197
Lizotte MP (2003) The microbiology of sea ice. In: Thomas DN, Dieckmann GS (eds) Sea ice—an introduction to its physics, chemistry, biology and geology. Blackwell Science Ltd, Oxford, pp 184–210
Lozan JL, Kausch H (2004) Angewandte Statistik für Naturwissenschaftler. Wissenschaftliche Auswertungen, Hamburg, p 300
Lüpkes C, Hartmann J, Birnbaum G, Cohrs W, Yelland M, Pascal R, Spieß T, Buschmann M (2004) Convection over Arctic leads. Ber Polarforsch 481:47–55
Maykut GA (1985) The ice environment. In: Horner RA (ed) Sea ice biota. CRC Press, Boca Raton, pp 21–82
Melnikov IA, Kolosova EG, Welch HE, Zhitnia LS (2002) Sea ice biological communities and nutrient dynamics in the Canada basin of the Arctic Ocean. Deep-Sea Res I 49:1623–1249
Meiners K, Fehling J, Granskog M, Spindler M (2002) Abundance, biomass and composition of biota in Baltic sea ice and underlying water (March 2000). Polar Biol 25:761–770
Meiners K, Gradinger R, Fehling J, Civitarese G, Spindler M (2003) Vertical distribution of exopolymer particles in sea ice of the Fram Strait (Arctic) during autumn. Mar Ecol Prog Ser 248:1–13
Mock T, Gradinger R (1999) Determination of Arctic ice algal production with a new in situ incubation technique. Mar Ecol Prog Ser 77:15–26
Okolodkov YB (1992) Cryopelagic flora of the Chukchi, East Siberian and Laptev Seas. Proc. NIPRI Symp. Polar Biol 5:28–43
Palmisano AC, Sullivan CW (1985) Growth, metabolism, and dark survival in sea ice microalgae. In: Horner RA (ed) Sea ice biota. CRC Press, Boca Raton, pp 131–146
Perovich DK, Andreas EL, Curry JA, Eicken H, Fairrall CW, Grenfell TC, Guest PS, Intrieri J, Kadko D, Lindsay RW, McPhee MG, Morison J, Moritz RE, Paulson CA, Pegau WS, Persson POG, Pinkel R, Richter-Menge JA, Stanton T, Stern HM, Tucker WB III, Uttal T (1999) Year on ice gives climate insigths. Eos 80:481–486
Perovich DK, Grenfell TC, Richter-Menge JA, Light B, Tucker III WB, Eicken H (2003) Thin and thinner: sea ice mass balance measurements during SHEBA. J Geophys Res 108(C3):8050
Redfield AC, Ketchum BH, Richards FA (1963) The influence of organisms on the composition of sea water. In: Hill MN (ed) The sea. Wiley, New York, pp 26–77
Rysgaard S, Glud RN (2004) Anaerobic N2 production in Arctic sea ice. Limnol Oceanogr 49:86–94
Rysgaard S, Kühl M, Glud RN, Hansen JW (2001) Biomass, production and horizontal patchiness of sea ice algae in a high-Arctic fjord (Young-Sound, NE Greenland). Mar Ecol Prog Ser 223:15–26
Schauer U, Kattner G (2004) The expedition ARKTIS XIX/1 a,b and XIX/2 of the Research Vessel “Polarstern” in 2003. Ber Polarforsch 481:1–190
Schnack-Schiel SB (2003) The macrobiology of sea ice. In: Thomas DN, Dieckmann GS (eds) Sea ice—an introduction to its physics, chemistry, biology and geology. Blackwell Science Ltd, Oxford, pp 211–239
Schünemann H, Werner I (2005) Seasonal variations in the distribution patterns of sympagic metazoans in Arctic pack ice. Mar Biol 146:1091–1102
Smith REH, Clément P, Head E (1989) Biosynthesis and photosynthate allocation patterns of arctic ice algae. Limnol Oceanogr 34:591–605
Smith REH, Gosselin M, Taguchi S (1997) The influence of major inorganic nutrients on the growth and physiology of high arctic ice algae. J Mar Syst 11:63–70
Spies A, Brockmann UH, Kattner G (1988) Nutrient regimes in the marginal ice zone of the Greenland Sea in summer. Mar Ecol Prog Ser 47:195–204
Syvertsen EE (1991) Ice algae in the Barents Sea: Types of assemblages, origin, fate and role in the ice-edge phytoplankton bloom. Polar Res 10:277–287
Thomas DN, Lara RJ, Eicken H, Kattner G, Skoog A (1995) Dissolved organic matter in Arctic multi-year sea ice during winter: major components and relationship to ice characteristics. Polar Biol 15:447–483
Tuschling K, Juterzenka Kv, Okolodkov YB, Anoshkin A (2000) Composition and distribution of the pelagic and sympagic algal assemblages in the Laptev Sea during autumnal freeze-up. J Plankton Res 22:843–864
Utermöhl H (1958) Zur Vervollkommnung der quantitativen Phytoplankton-Methodik. Mitt Int Ver Theor Angew Limnol 9:1–38
Werner I (1997) Grazing of arctic under-ice amphipods on sea-ice algae. Mar Ecol Prog Ser 160:93–99
Werner I, Auel H (2005) Seasonal variability in abundance, respiration and lipid composition of Arctic under-ice amphipods. Mar Ecol Prog Ser 292:251–262
Zhang Q, Gradinger R, Spindler M (1998) Dark survival of marine microalgae in the High Arctic (Greenland Sea). Polarforsch. 65:111–116
Acknowledgments
The support of the captain, the crew and the chief scientists of the RV “Polarstern” expedition ARK XIX/1 is gratefully acknowledged. All ice studies would have been impossible without the helpful cooperation of many colleagues from the ice working groups and numerous “polar bear watchers”. We thank A. Scheltz and R. Kiko for help in the field and H. Johannsen for conducting the nutrient measurements. K. Meiners and K. Bischof are acknowledged for constructive discussions and J. Schwarz for language corrections. The manuscript benefited substantially by the constructive comments made by three referees and the editor.
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
About this article
Cite this article
Werner, I., Ikävalko, J. & Schünemann, H. Sea-ice algae in Arctic pack ice during late winter. Polar Biol 30, 1493–1504 (2007). https://doi.org/10.1007/s00300-007-0310-2
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00300-007-0310-2