Polar Biology

, Volume 12, Issue 3–4, pp 429–444 | Cite as

Ecology of sea ice biota

2. Global significance
  • Louis Legendre
  • Stephen F. Ackley
  • Gerhard S. Dieckmann
  • Bjørn Gulliksen
  • Rita Horner
  • Takao Hoshiai
  • Igor A. Melnikov
  • William S. Reeburgh
  • Michael Spindler
  • Cornelius W. Sullivan


The sea ice does not only determine the ecology of ice biota, but it also influences the pelagic systems under the ice cover and at ice edges. In this paper, new estimates of Arctic and Antarctic production of biogenic carbon are derived, and differences as well as similarities between the two oceans are examined. In ice-covered seas, high algal concentrations (blooms) occur in association with several types of conditions. Blooms often lead to high sedimentation of intact cells and faecal pellets. In addition to ice-related blooms, there is progressive accumulation of organic matter in Arctic multi-year ice, whose fate may potentially be similar to that of blooms. A fraction of the carbon fixed by microalgae that grow in sea ice or in relation to it is exported out of the production zone. This includes particulate material sinking out of the euphotic zone, and also material passed on to the food web. Pathways through which ice algal production does reach various components of the pelagic and benthic food webs, and through them such top predators as marine mammals and birds, are discussed. Concerning global climate change and biogeochemical fluxes of carbon, not all export pathways from the euphotic zone result in the sequestration of carbon for periods of hundreds of years or more. This is because various processes, that take place in both the ice and the water column, contribute to mineralize organic carbon into CO2 before it becomes sequestered. Processes that favour the production and accumulation of biogenic carbon as well as its export to deep waters and sequestration are discussed, together with those that influence mineralization in the upper ice-covered ocean.


Microalgae Faecal Pellet Euphotic Zone Benthic Food Biogenic Carbon 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Ackley SF (1982) Ice scavenging and nucleation: two mechanisms for the incorporation of algae into newly forming sea ice. Eos Trans Am Geophys Union 63:65Google Scholar
  2. Ackley SF, Dieckmann GS, Shen HT (1987) Algal and foram incorporation into new sea ice. Eos Trans Am Geophys Union 68:1736Google Scholar
  3. Ackley SF, Sullivan CW (in press) Physical controls on the development and characteristics of the Antarctic sea ice ecosystemGoogle Scholar
  4. Ackley SF, Lange M, Wadhams P (1990) Snow cover effects on Antarctic sea ice thickness. In: Ackley SF, Weeks WF (eds) Sea ice properties and processes, CRREL Monogr 90-1, pp 16–21Google Scholar
  5. Agatha S, Wilbert N, Spindler M, Elbrächter M (1990) Euplotide ciliates in sea ice of the Weddell Sea (Antarctica). Acta Protozool 29:221–228Google Scholar
  6. Alexander V (1980) Interrelationships between the seasonal sea ice and biological regimes. Cold Regions Sci Tech Rep 2:157–178Google Scholar
  7. Alexander V, Horner R, Clasby RC (1974) Metabolism of Arctic sea ice organisms. Inst Mar Sci Univ Alaska Rep R74-4, 120 ppGoogle Scholar
  8. Alldredge AL (1984) The quantitative significance of gelatinous zooplankton as pelagic consumers. In: Fasham MJR (ed) Flows of energy and materials in marine ecosystems: Theory and practice. Plenum, New York, pp 407–433Google Scholar
  9. Anderson JB (1975) Factors controlling CaCO3 dissolution in the Weddell Sea from foraminiferal distribution patterns. Mar Geol 19:315–332Google Scholar
  10. Anderson LG, Dyrssen D, Jones EP (1990) An assessment of transport of atmospheric CO2 into the Arctic Ocean. J Geophys Res 95:1703–1711Google Scholar
  11. Andriashev AP (1954) Fishes of the northern seas of the U.S.S.R. Israel Progr Sci Transl, Jerusalem 164, 617 pp (transl. from Russian)Google Scholar
  12. Apollonio S (1965) Chlorophyll in Arctic sea ice. Arctic 18:118–122Google Scholar
  13. Arrigo KR, Dieckmann GS, Gosselin M, Sullivan CW (1990) Studies on the nutrient status in sea ice and underlying platelet layer. Antarct J US 25:185–188Google Scholar
  14. Atlas RM, Griffiths RP (1984) Bacterial populations of the Beaufort Sea. In: Barnes PW, Schell DM, Reimnitz E (eds) The Alaskan Beaufort Sea: ecosystems and environments. Academic Press, Orlando, pp 327–345Google Scholar
  15. Azam F, Fenchel T, Field JG, Gray JS, Meyer-Reil LA, Thingstad F (1983) The ecological role of water column microbes in the sea. Mar Ecol Prog Ser 10:257–263Google Scholar
  16. Bain H, Sekerak AD (1978) Aspects of biology of Arctic cod, Boreogadus saida, in the central Canadian Arctic. LGL ltd, Toronto, Ontario, 104 ppGoogle Scholar
  17. Bé AWH (1977) An ecological, zoogeographic and taxonomic review of recent planktonic foraminifera. In: Ramsey ATS (ed) Oceanic micropaleontology, vol. 1, Academic Press, London, pp 1–100Google Scholar
  18. Berger WH (1978) Deep-sea carbonate: pteropod distribution and the aragonite compensation depth. Deep-Sea Res 25:447–452Google Scholar
  19. Berger WH (1982) Deglacial CO2 buildup: constraints on the coral reef model. Paleogeog Paleoclim Paleoecol 40:235–253Google Scholar
  20. Bergmann MA, Welch HE, Butler-Walker JE, Sifred TD (1991) Ice algal photosynthesis at Resolute and Saqvaqjuac in the Candian Arctic. J Mar Syst 2:43–52Google Scholar
  21. Berner RA, Honjo S (1981) Pelagic sedimentation of aragonite: its geochemical significance. Science 211:940–942Google Scholar
  22. Betzer PR, Byrne RH, Acker JG, Lewis CS, Jolley RR, Feeley RA (1984) The oceanic carbonate system: a reassessment of biogenic controls. Science 226:1074–1077Google Scholar
  23. Blaxter JHS (1975) Fish vision and applied research. In: Ali MA (ed) Vision in fishes. Plenum, New York, pp 757–773Google Scholar
  24. Bradstreet MSW, Cross WE (1982) Trophic relationships at high Arctic ice edges. Arctic 35:1–12Google Scholar
  25. Broecker WS (1991) Keeping global change honest. Global Biogeochem Cycles 5:191–192Google Scholar
  26. Brown RGB (1980) Seabirds as marine animals. In: Burger J, Olla BL, Winn HE (eds) Behaviour of marine animals, vol 4. Plenum, New York, pp 1–39Google Scholar
  27. Bunch JN, Harland RC (1990) Bacterial production in the bottom surface of sea ice in the Canadian subarctic. Can J Fish Aquat Sci 47:1986–1995Google Scholar
  28. Burkholder PR, Mandelli EF (1965) Productivity of microalgae in Antarctic sea ice. Science 33:177–184Google Scholar
  29. Burns JJ (1981) Bearded seal Erignathus barbatus Erxleben, 1777. In: Ridgway SH, Harrison RJ (eds) Handbook of marine mammals, vol 2. Seals. Academic Press, London, pp 145–170Google Scholar
  30. Bustnes JO, Erikstad KE (1988) The diets of sympatric wintering populations of common eider Somateria mollissima and king eider S. spectabillis in northern Norway. Omis Fenn 65:163–168Google Scholar
  31. Byrne RH, Acker JG, Betzer PR, Feely RA, Cates MH (1984) Water column dissolution of aragonite in the Pacific Ocean, Nature 312:321–326Google Scholar
  32. Carey AG Jr (1985) Marine ice fauna: Arctic In: Horner RA (ed) Sea ice biota. CRC Press, Boca Raton, pp 173–190Google Scholar
  33. Carey AG Jr (1987) Particle flux beneath fast ice in the shallow southwestern Beaufort Sea, Arctic Ocean. Mar Ecol Prog Ser 40:247–257Google Scholar
  34. Cosmiso JC and Sullivan CW (1986) Satellite microwave and in situ observations of the Weddell Sea ice cover and its marginal ice zone. J Geophys Res 91:9663–9681Google Scholar
  35. Cosmiso JC, Maynard NG, Smith WO Jr, Sullivan CW (1990) Satellite ocean color studies of Antarctic ice edge in summer/autumn. J Geophys Res 95:9481–9486Google Scholar
  36. Cota GF, Kottmeier ST, Robinson DH, Smith WO Jr, Sullivan CW (1990) Bacterioplankton in the marginal ice zone of the Weddell Sea: biomass, production and metabolic activities during austral autumn. Deep-Sea Res 37:1145–1167Google Scholar
  37. Cota GF, Legendre L, Gosselin M, Ingram RG. 1991. Ecology of bottom ice algae. I. Environmental controls and variability. J Mar Syst 2:257–277Google Scholar
  38. Cota GF, Smith WO Jr (1989) AMERIEZ 1988: Phytoplankton biomass and productivity in the marginal ice zone of the Weddell-Scotia Sea during austral winter. Antarct J US 24:152–153Google Scholar
  39. Craig PC, Griffiths W, Haldorsen L, McElderry L (1982) Ecological studies of arctic cod (Boreogadus saida) in Beaufort Sea coastal waters. Alaska, Can J Fish Aquat Sci 39:395–406Google Scholar
  40. Cramp S, Simmons KEL (1977) Handbook of the birds of Europe, the Middle East and North Africa. The birds of the Western Palearctic, vol 1. Oxford Univ Press, Oxford, 722 ppGoogle Scholar
  41. Cushing DH (1972) The production cycle and the numbers of marine fish. Symp Zool Soc London 29:213–232Google Scholar
  42. Dahms HU, Dieckmann GS (1987) Drescheriella glacialis gen. nov., sp. nov. (Copepoda, Harpacticoida) from Antarctic sea ice. Polar Biol 7:329–337Google Scholar
  43. Daly KL (1990) Overwintering development, growth, and feeding of larval Euphausia superba in the Antarctic marginal ice zone. Limnol Oceanogr 35:1564–1576Google Scholar
  44. Daly KL, Macaulay MC (1988) Abundance and distribution of krill in the ice edge zone of the Weddell Sea, austral spring 1983. Deep-Sea Res 35:21–41Google Scholar
  45. Davis RA, Finley KJ, Richardson WJ (1980) The present status and future management of Arctic marine mammals in Canada. Sci Adv Board NW Territories, Rep No 3, Dept Information, Gov NWT, 93 ppGoogle Scholar
  46. Demers S, Legendre L, Maestrini SY, Rochet M, Ingram RG (1989) Nitrogenous nutrition of sea-ice microalgae. Polar Biol 9:377–383Google Scholar
  47. DeWitt HH (1971) Coastal and deep-water benthic fishes of the Antarctic. Antarctic map folio ser, folio 15. Am Geogr Soc, New York, pp 1–10Google Scholar
  48. Dieckmann G, Sullivan CW, Garrison D (1990) Seasonal standing crop of ice algae in pack ice of the Weddell Sea, Antarctica. Eos Trans Am Geophys Union 71:79Google Scholar
  49. Dieckmann GS, Spindler M, Lange MA, Ackley SF, Eicken H (1991) Antractic sea ice: A habitat for the foraminifer Neogloboquadrina pachyderma. J Foram Res 21:184–191Google Scholar
  50. Drolet R, Fortier L, Ponton D, Gilbert M (1991) Production of fish larvae and their prey in subarctic southeastern Hudson Bay Mar Ecol Prog Ser 77:105–118Google Scholar
  51. Dunbar MJ (1957) The determinants of production in northern seas: A study of the biology of Themisto libellula Mandt. Can J Zool 35:797–819Google Scholar
  52. Dymond J, Lyle M (1985) Flux comparisons between sediments and sediment traps in the eastern tropical Pacific: Implications for atmospheric CO2 variations during the Pleistocene. Limnol Oceanogr 30:699–712Google Scholar
  53. Eastman JT, DeVries AL (1985) Adaptations for cryopelagic life in the Antarctic notothenoid fish Pagothenia borchgrevinki. Polar Biol 4:45–52Google Scholar
  54. Eilertsen HC, Tande KS, Hegseth EN (1989) Potential of herbivorous copepods for regulating the spring phytoplankton bloom in the Barents Sea. Rapp P-V Reun Cons Int Explor Mer 188:154–163Google Scholar
  55. Fabry VJ (1990) Shell growth rates of pteropod and heteropod molluscs and aragonite production in the open ocean: Implications for the marine carbonate system. J Mar Res 48:209–222Google Scholar
  56. Fay FH (1981) Walrus Odobenus rosmarus (Linnaeus, 1758). In: Ridgway SH, Harrison RJ (eds) Handbook of marine mammals, vol 1. The walrus, sea lions, fur seals and sea otter. Academic Press, London, pp 1–23Google Scholar
  57. Fenchel T (1984) Suspended marine bacteria as a food source. In: Fasham MJR (ed) Flows of energy and materials in marine ecosystems. Plenum, New York, pp 301–315Google Scholar
  58. Fenchel T, Lee CC (1972) Studies on alliates associated with sea ice from Antarctica. 1. The nature of the fauna. Arch Protistenk 114:231–236Google Scholar
  59. Finley KJ, Miller GW, Davis RA, Koski WR (1983) A distinctive large breeding population of ringed seals (Phoca hispida) inhabiting the Baffin Bay pack ice. Arctic 36:162–173Google Scholar
  60. Fischer G, Fütterer D, Gersond R, Honjo S, Ostermann D, Wefer G (1988) Seasonal variability of particle flux in the Weddell Sea and its relation to ice cover. Nature 335:426–428Google Scholar
  61. Garrison DL, Ackley SF, Buck KR (1983) A physical mechanism for establishing algal populations in frazil ice. Nature 306:363–65Google Scholar
  62. Garrison DL, Buck KR (1989) The biota of Antarctic pack ice in the Weddell Sea and Antarctic Peninsula regions. Polar Biol 10:211–219Google Scholar
  63. Garrison DL, Buck KR (1991) Surface-layer sea ice assemblages in Antarctic pack ice during the austral spring: environmental conditions, primary production and community structure. Mar Ecol Prog Ser 75:161–172Google Scholar
  64. Garrison DL, Sullivan CW, Ackley SF (1986) Sea ice microbial communities in Antarctica. BioScience 36:243–250Google Scholar
  65. Gilbert M, Fortier L, Ponton D, Drolet R (1992) Feeding ecology of marine fish larvae across the Great Whale River plume in seasonally ice-covered southeastern Hudson Bay. Mar Ecol Prog Ser 19 (in press)Google Scholar
  66. Goldman JC (1988) Spatial and temporal discontinuities of biological processes in pelagic surface waters. In Rothschild BJ (ed) Toward a theory on biological-physical interactions in the World Ocean. Kluwer Acad Publ, Dordrecht, pp 273–296Google Scholar
  67. Gradinger R, Spindler M, Henschel D (1992) Development of Arctic sea ice organisms under graded snow cover. Polar Res (in press)Google Scholar
  68. Grainger EH, Mohammed AA, Lovrity JE (1985) The sea ice fauna of Frobisher Bay, Arctic Canada. Arctic 38:23–30Google Scholar
  69. 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 35:153–164Google Scholar
  70. Gulliksen B, Lønne OJ (1989) Distribution, abundance, and ecological importance of marine sympagic fauna in the Arctic. Rapp P-V Reun Cons Int Explor Mer 188:133–138Google Scholar
  71. Hamner WM, Hamner PP, Strand SW, Gilmer RW (1983) Behavior of antarctic krill, Euphausia superba: chemoreception, feeding, schooling, and molting. Science 220:433–435Google Scholar
  72. Harbison GR (1988) Observations on the epipelagic gelatinous fauna of McMurdo Sound. Ant J US 23:135–136Google Scholar
  73. Haug T, Gulliksen B (1982) Size, age, occurrence, growth and food of Greenland halibut Reinhardtius hippoglossoides (Walbaum) in coastal waters of western Spitzbergen. Sarsia 68:293–297Google Scholar
  74. Hemleben C, Spindler M, Anderson OR (1989) Modern planktonic foraminifera. Springer, New York, 363 ppGoogle Scholar
  75. Herman Y, Andersen OGN (1989) Foraminifera and pteropoda beneath the Arctic sea ice: new distributions. In: Herman Y (ed) The Arctic seas climatology, oceanography, geology, and biology. Van Nostrand Reinhold Co, New York, pp 223–234Google Scholar
  76. Hjort J (1914) Fluctuations in the great fisheries of northern Europe viewed in light of biological research. Rapp P-V Reun Cons Perm Int Explor Mer 20:1–228Google Scholar
  77. Hognestad PT (1968) 1. Observations on polar cod in the Barents Sea. Rapp P-V Reun Cons Int Explor Mer 158:126–130Google Scholar
  78. Honjo S (1980) Material fluxes and modes of sedimentation in the mesopelagic and bathypelagic zones. J Mar Res 38:53–97Google Scholar
  79. Horner R, Ackley SF, Dieckmann GS, Gulliksen B, Horner R, Hoshiai T, Melnikov IA, Reeburgh WS, Spindler M, Sullivan CW (1992) Ecology of sea ice biota. 1. Habitat and terminology. Polar Biol 12 (in press)Google Scholar
  80. Horner R, Alaxander V (1972) Algal populations in Arctic sea ice: an investigation of heterotrophy. Limnol Oceanogr 17:454–458Google Scholar
  81. Horner R, Schrader GC (1982) Relative contributions of ice algae, phytoplankton, and benthic microalgae to primary production in nearshore regions of the Beaufort Sea. Arctic 35:485–503Google Scholar
  82. Hoshiai T, Tanimura A (1986) Sea ice meiofauna at Syowa station, Antarctica. Mem Natl Inst Polar Res, Spec Issue 44:118–124Google Scholar
  83. Hoshiai T, Tanimura A, Fukuchi M, Watanabe K (1989) Feeding by the Nototheniid fish, Pagothenia borchgrevinki on the ice-associated copepod, Paralabidocera antarctica. Proc NIPR Symp Polar Biol 2:61–64Google Scholar
  84. Hoshiai T, Tanimura A, Watanabe K (1987) Ice algae as food of an Antarctic ice-associated copepod, Paralabidocera antarctica (I.C. Thompson). Proc NIPR Symp Polar Biol 1:105–111Google Scholar
  85. Hubold G (1985) Stomach contents of the Antarctic silverfish Pleuragramma antarcticum from the southern and eastern Weddell Sea (Antarctica). Polar Biol 5:43–48Google Scholar
  86. Jennings JC Jr, Gordon LI, Nelson DM (1984) Nutrient depletion indicates high primary productivity in the Weddell Sea. Nature 308:51–54Google Scholar
  87. Johnsen G, Hegseth EN (1991) Photoadaptation of sea-ice microalgae in the Barents Sea. Polar Biol 11:179–184Google Scholar
  88. Joos F, Sarmiento JL, Siegenthaler U (1991) Estimates of the effect of Southern Ocean iron fertilization on atmospheric CO2 concentrations. Nature 349:772–775Google Scholar
  89. Kennett JP (1970) Comparison of Globigerina pachyderma (Ehrenberg) in Arctic and Antarctic areas. Contrib Cushman Found Foram Res 21:47–49Google Scholar
  90. Klumov SK (1937) Polar cod and their importance for certain life processes in the Arctic. Izv Akad Nauk USSR, Ser Biol 1:175 (in Russian)Google Scholar
  91. Kobayashi HA (1974) Growth cycle and related vertical distribution of the thecosomatous pteropod Spiratella (“Limacina”) helicina in the central Arctic Ocean. Mar Biol 26:295–301Google Scholar
  92. Kottmeier ST, Grossi SM, Sullivan SW (1987) Sea ice microbial communities. VIII. Bacterial production in annual sea ice of McMurdo Sound, Antarctica. Mar Ecol Prog Ser 35:175–186Google Scholar
  93. Kottmeier ST, Miller MA, Lizotte MP, Craft LL, Gulliksen B, Sullivan CW (1985) Ecology of sea ice microbial communities (SIMCO) during the 1984 winter to summer transition in McMurdo Sound, Antarctica. Antarct J US 20:128–130Google Scholar
  94. Kottmeier ST, Sullivan CW (1987) Late winter primary production and bacterial production in sea ice and seawater west of the Antarctic Peninsula. Mar Ecol Prog Ser 36:287–298Google Scholar
  95. Kottmeier ST, Sullivan CW (1990) Bacterial biomass and production in pack ice of Antarctic marginal ice edge zones. Deep-Sea Res 37:1311–1330Google Scholar
  96. Larssen BB, Elverhøi A, Aagaard P (1987) Study of particulate material in sea ice in the Farm Strait — a contribution to paleoclimatic research? Polar Res 5:313–315Google Scholar
  97. Last JM (1980) The food of twenty species of fish larvae in the west-central North sea. Fish Res Tech Rep MAFF Direct Fish Res, Lowestoft 60:1–44Google Scholar
  98. Legendre L (1990) The significance of microalgal blooms for fisheries and for the export of particulate organic carbon in oceans. J Plankton Res 12:681–699Google Scholar
  99. Legendre L, Demers S, Gosselin M (1987) Chlorophyll and photosynthetic efficiency of size-fractionated sea-ice microalgae (Hudson Bay, Canadian Arctic). Mar Ecol Prog Ser 40:199–203Google Scholar
  100. Legendre L, Gosselin M (1989) New production and export of organic matter to the deep ocean: consequences of some recent discoveries. Limnol Oceanogr 34:1374–1380Google Scholar
  101. Legendre L, Ingram RG, Poulin M (1981) Physical control of phytoplankton production under sea ice (Manitounuk Sound, Hudson Bay). Can J Fish Aquat Sci 38:1385–1392Google Scholar
  102. Legendre L, Le Fèvre J (1989) Hydrodynamical singularities as controls of recycled versus export production in oceans. In: Berger WH, Smetacek VS, Wefer G (eds) Productivity of the ocean: present and past. Wiley, Chichester, pp 49–63Google Scholar
  103. Legendre L, Le Fèvre J (1991) From individual plankton cells to pelagic marine ecosystems and to global biogeochemical cycles. In: Demers S (ed) Particle analysis in oceanography. Springer, Berlin, pp 261–299Google Scholar
  104. Leventer A, Dunbar RB (1987) Diatom flux in McMurdo Sound, Antarctica. Mar Micropaleontol 12:49–64Google Scholar
  105. Leventer A, Dunbar RB (1988) Recent diatom record of McMurdo Sound, Antarctica: implications for history of sea ice extent. Paleoceanography 3:259–274Google Scholar
  106. Longhorst AR (1991) A reply to Broecker's charges. Global Biogeochem Cycles 5:315–316Google Scholar
  107. Longhurst AR, Bedo AW, Harrison WG, Head EJH, Sameoto DD (1990) Vertical flux of respiratory carbon by oceanic diel migrant biota. Deep-Sea Res 37:685–694Google Scholar
  108. Lønne OJ, Gulliksen B (1989) Size, age and diet of polar cod, Boreogadus saida (Lepechin 1773), in the ice covered waters. Polar Biol 9:187–191Google Scholar
  109. Lowry LF, Frost KJ (1981) Distribution, growth and foods of arctic cod (Boreogadus saida) in the Bering, Chuckchi, and Beaufort Seas. Can Field Nat 95:186–191Google Scholar
  110. Lowry LF, Frost KJ, Burns JJ (1979) Potential resource competition in the southeastern Bering Sea: fisheries and phocid seals. Proc 29th Alaska Sci Conf, Fairbanks, 15–17 August 1978, pp 287–296Google Scholar
  111. Lowry LF, Frost KJ, Burns JJ (1980) Variability in the diet of ringed seals (Phoca hispida) in Alaska. Can J Fish Aquat Sci 37:2254–2261Google Scholar
  112. Manak DK, Mysak L (1989) On the relationship between Arctic sea ice anomalies and fluctuations in northern Canadian air temperature and river discharge. Atoms Ocean 27:682–691Google Scholar
  113. Marr JWS (1962) The natural history and geography of the Antarctic krill (Euphausia superba). Discovery Rep 32:33–464Google Scholar
  114. Marra J, Boardman DC (1984) Late winter chlorophyll a distributions in the Weddell Sea. Mar Ecol Prog Ser 19:197–205Google Scholar
  115. Marschall HP (1988) The overwintering strategy of Antarctic krill under the pack ice of the Weddell Sea. Polar Biol 9:129–135Google Scholar
  116. McLaren IA (1958) The biology of the ringed seal (Phoca hispida Schreber) in the eastern Canadian Arctic. Bull Fish Res Board Can 118:1–97Google Scholar
  117. Mehlum F, Giertz I (1984) Feeding ecology of seabirds in the Svalbard area-a preliminary report. Norsk Polarinst, Rapp Ser 16:1–41Google Scholar
  118. Melnikov IA (1989) Ecosystem of the Arctic sea ice. Inst Oceanol, Acad Sci USSR, Moscow (In Russian)Google Scholar
  119. Melnikov IA, Bondarchuk LL (1987) Ecology of mass accumulations of colonial diatom algae under drifting Arctic ice. Oceanology 27:233–236Google Scholar
  120. Melnikov IA, Pavlov GL (1978) Peculiarities of organic carbon distribution in the waters and ice of the Arctic basin. Oceanology 18:248–253Google Scholar
  121. Moritz RE, Aagaard K, Baker DJ, Codispoti LA, Smith SL, Smith WO, Tipper RC, Walsh JE (1990) Arctic system science. Oceanatmosphere-ice interactions. Joint Oceanogr Inst Inc, Washington DCGoogle Scholar
  122. Mucci A (1983) The solubility of calcite and aragonite in sewater at various salinites, temperatures and one atmospheric pressure. Am J Sci 283:780–799Google Scholar
  123. Nelson DM, Smith WO Jr (1986) Phytoplankton bloom dynamics of the western Ross Sea ice edge. II. Mesoscale cycling of nitrogen and silicon. Deep-Sea Res 33:1389–1412Google Scholar
  124. Nelson DM, Smith WO Jr, Gordon LI, Huber BA (1987) Spring distributions of density, nutrients and phytoplankton biomass in the ice edge zone of the Weddell-Scotia Sea. J Geophys Res 92:7181–7190Google Scholar
  125. Nelson DM, Smith WO Jr, Muench RD, Gordon LI, Sullivan CW, Husby DM (1989) Particulate matter and nutrient distributions in the ice-edge zone of the Weddell Sea: Relationship to hydrography during late summer. Deep-Sea Res 36:191–209Google Scholar
  126. Niebauer HJ, Alexander V, Henrichs S (1990) Physical and biological oceanographic interaction in the spring bloom at the Bering Sea marginal ice edge zone. J Geophys Res 95:22239–22241Google Scholar
  127. Nöthig EM, Bodungen B von (1989) Occurrence and vertical flux of faecal pellets of probably protozoan origin in the southeastern Weddell Sea (Antarctica). Mar Ecol Prog Ser 56:281–289Google Scholar
  128. Olson RJ (1980) Nitrate and ammonium uptake in Antarctic waters. Limnol Oceanogr 25:1064–1074Google Scholar
  129. Palmisano AC, Moe RL, Sullivan CW (1987) Sea ice microbial communities. VII. Changes in under-ice spectral irradiance during the development of Antarctic sea ice microbial communities. Mar Ecol Prog Ser 35:165–173Google Scholar
  130. Peng TH, Broecker WS (1991) Dynamic limitations on the Antarctic iron limitation strategy. Nature 349:227–229Google Scholar
  131. Pfirman S, Gasard JC, Wollenburg I, Mudie P, Abelmann A (1989) Particle-laden Eurasian Arctic sea ice: observations from July and August 1987. Polar Res 7:59–66Google Scholar
  132. Platt T, Subba Rao DV (1975) Primary production of marine microphytes. In: Cooper JP (ed) Photosynthesis and productivity in different environments. Cambridge Univ Press, Cambridge, pp 249–280Google Scholar
  133. Plötz J (1986) Summer diet of Weddell seals (Leptonychotes weddellii) in the eastern and southern Weddell sea, Antarctica. Polar Biol 6:97–102Google Scholar
  134. Palmisano AC, Sullivan CW (1983) Sea Ice Microbial Communities (SIMCO). I. Distribution, abundance and primary production of ice microalgae in McMurdo Sound, Antarctica in 1980. Polar Biol 2:171–177Google Scholar
  135. Pomeroy LR, Diebel D (1986) Temperature regulation of bacterial activity during the spring bloom in Newfoundland coastal waters. Science 233:359–361Google Scholar
  136. Pomeroy LR, Wiebe WJ (1988) Energetics of microbial food webs. Hydrobiologia 159:7–18Google Scholar
  137. Quast JC (1974) Density distribution of juvenile arctic cod, Boreogadus saida in the eastern Chukchi Sea in the fall of 1970. Fish Bull 72:1094–1105Google Scholar
  138. Rass T (1968) Spawning and development of polar cod. Rapp P-V Reun Cons Perm Int Explor Mer 158:135–137Google Scholar
  139. Redfield AC (1958) The biological control of chemical factors in the environment. Am Sci 46:205–222Google Scholar
  140. Redfield AC, Ketchum BH, Richards FA (1963) The influence of organisms on the composition of sea water. In: Hill MN (ed) The sea, vol 2. Interscience, New York, pp 26–77Google Scholar
  141. Reynolds LA, Thunell RC (1985) Seasonal succession of planktonic foraminifera in the subpolar North Pacific. J Foram Res 15:282–301Google Scholar
  142. Runge JA, Ingram RG (1988) Underice grazing by planktonic, calanoid copepods in relation to a bloom of ice microalgae in southeastern Hudson Bay. Limnol Oceanogr 33:280–286Google Scholar
  143. Runge JA, Ingram RG (1991) Under ice feeding and diel migration by the planktonic copepods Calanus glacialis and Pseudocalanus minutus in relation to the ice algal production cycle in southeastern Hudson Bay. Mar Biol 108:217–225Google Scholar
  144. Sarmiento JL, Toggweiler JR (1984) A new model for the role of the ocean in determining atmospheric pCO2. Nature 308:621–624Google Scholar
  145. Sarmiento JL, Toggweiler JR, Najjar R (1988) Ocean carbon-cycle dynamics and atmospheric pCO2. Philos Trans R Soc A 325:3–21Google Scholar
  146. Sekerak AD (1982) Young-of-the-year cod (Boreogadus) in Lancaster Sound and Western Baffin Bay. Arctic 35:75–87Google Scholar
  147. Siegenthaler U, Wenk T (1984) Rapid atmospheric CO2 variations and ocean circulation. Nature 308:624–626Google Scholar
  148. Smith REH, Clement P, Cota GF, Li WKW (1987) Intracellular photosynthate allocation and the control of Arctic marine ice algal production. J Phycol 23:124–132Google Scholar
  149. Smith REH, Clement P, Head E (1989) Biosynthesis and photosynthate allocation patterns of arctic ice algae. Limnol Oceanogr 34:591–605Google Scholar
  150. Smith SL, Smith WO, Codispoti LA, Wilson DL (1985) Biological observations in the marginal ice zone of the East Greenland Sea. J Mar Res 43:693–717Google Scholar
  151. Smith WO, Nelson DM (1985) Phytoplankton bloom produced by a receding ice edge in the Ross Sea: Spatial coherence with the density field. Science 227:163–166Google Scholar
  152. Smith WO, Nelson DM (1986) Importance of ice edge phytoplankton production in the Southern Ocean. BioScience 36:251–257Google Scholar
  153. Spindler M (1990) A comparison of arctic and antarctic sea ice and the effects of different properties on sea ice biota. In: Bliel U, Thiede J (eds) Geological history of the polar oceans: Arctic versus Antarctic. Kluwer, Dordrecht, pp 173–186Google Scholar
  154. Spindler M., Dieckmann GS (1986) Distribution and abundance of the planktic foraminifer Neogloboquadrina pachyderma in sea ice of the Weddell Sea. Polar Biol 5:185–191Google Scholar
  155. Spindler M, Dieckmann GS, Lange MA (1990) Seasonal and geographic variations in sea ice community structure of the Weddell Sea, Antarctica. In: Kerry KR, Hempel G (eds) Antarctic ecosystems. Ecological change and conservation. Springer, Berlin, pp 129–135Google Scholar
  156. Spoel S van der, Boltovskoy D (1981) Pteropoda. In: Boltovskoy D (ed) Atlas del zooplankton del Atlántico suboccidental y métodos de trabajo con el zooplankton marino. Publ Espc INIDEP, Mar del Plata, Argentina, pp 493–531Google Scholar
  157. Stehman CF (1972) Planktonic foraminifera in Baffin Bay, Davis Strait and the Labrador Sea. Mar Sed 8:13–19Google Scholar
  158. Stretch JJ, Hamner PP, Hamner WM, Michel WC, Cook J, Sullivan CW (1988) Foraging behavior of antarctic krill Euphausia superba on sea ice microalgae. Mar Ecol Prog Ser 44:131–139Google Scholar
  159. Subba Rao DV, Platt T (1984) Primary production of Arctic waters. Polar Biol 3:191–201Google Scholar
  160. Suess E (1973) Interaction of organic compounds with calcium carbonate. II. Organo-carbonate association in recent sediments. Cheochim Cosmochim Acta 37:2435–2447Google Scholar
  161. Sugimura Y, Suzuki Y (1988) A high temperature catalytic oxydation method of non-volatile dissolved organic carbon in seawater by direct injection of liquid samples. Mar Chem 24:105–131Google Scholar
  162. Sullivan CW, Palmisano AC (1984) Sea ice microbial communities: distribution, abundance, and diversity of ice bacteria in McMurdo Sound, Antarctica, in 1980. Appl Environ Microbiol 47:788–795Google Scholar
  163. Sullivan CW, McClain CR, Comiso JC, Smith WO Jr (1988) Phytoplankton standing crops within an Antarctic ice edge assessed by satellite remote sensing. J Geophys Res 93:12487–12498Google Scholar
  164. Sullivan CW, Cota GF, Krempin DW, Smith WO Jr (1990) Distribution and activity of bacterioplankton in the marginal ice zone of the Weddell-Scotia Sea during austral spring. Mar Ecol Prog Ser 63:239–252Google Scholar
  165. Takahashi K, Bé AWH (1984) Planktonic foraminifera: factors controlling sinking speeds. Deep-Sea Res 31:1477–1500Google Scholar
  166. Toggweiler JR (1989) Is the downward dissolved organic matter (DOM) flux important in carbon transport? In: Berger WH, Smetacek VS, Wefer G (eds) Productivity of the ocean: present and past. Wiley and Sons, Chichester, pp 65–83Google Scholar
  167. Tourangeau S, Runge JA (1991) Reproduction of Calanus glacialis in relation to an ice microalgal bloom in southeastern Hudson Bay. Mar Biol 108:227–233Google Scholar
  168. Tremblay C, Runge JA, Legendre L (1989) Grazing and sedimentation of ice algae during and immediately after a bloom at the ice water interface. Mar Ecol Prog Ser 56:291–300Google Scholar
  169. Volk T, Hoffert MI (1985) Ocean carbon pumps: analysis of relative strengths and efficiencies in ocean-driven atmospheric CO2 changes. In: Sundquist ET, Broecker WS (eds) The carbon cycle and atmospheric CO2: natural variations archean to present. AGU Monogr 32, pp 99–110Google Scholar
  170. Volk T (1989) Sensitivity of climate and atmospheric CO2 to deepocean and shallow-ocean carbonate burial. Nature 337:637–638Google Scholar
  171. Wallace DW, Moore RM, Jones EP (1987) Ventilation of the Arctic Ocean cold halocline: Rates of diapycnal ans isopycnal transport, oxygen utilization and primary production inferred using chlorofluoromethane distributions. Deep-Sea Res 34:1957–1979Google Scholar
  172. Walsh JJ (1990) Arctic carbon sinks; present and future. Global Biogeochem Cycles 3:393–411Google Scholar
  173. Wassmann P, Vernet M, Mitchell GB, Rey F (1990) Mass sedimentation of Phaeocystis pouchetii in the Barents Sea. Mar Ecol Prog Ser 66:183–195Google Scholar
  174. Wefer G, Fischer G, Fütterer DK, Gersonde R, Honjo S, Ostermann D (1990) Particle sedimentation and productivity in Antarctic waters of the Atlantic sector. In: Bleil U, Thiede J (eds) Geological history of the polar oceans: Arctic versus Antarctic. Kluwer, Dordrecht, pp 363–379Google Scholar
  175. Wilson DL, Smith WO Jr, Nelson DM (1986) Phytoplankton bloom dynamics of the western Ross Sea ice edge. I. Primary productivity and species-specific production. Deep-Sea Res 33:1375–1387Google Scholar

Copyright information

© Springer-Verlag GmbH & Co. KG 1992

Authors and Affiliations

  • Louis Legendre
    • 1
  • Stephen F. Ackley
    • 2
  • Gerhard S. Dieckmann
    • 3
  • Bjørn Gulliksen
    • 4
  • Rita Horner
    • 5
  • Takao Hoshiai
    • 6
  • Igor A. Melnikov
    • 7
  • William S. Reeburgh
    • 8
  • Michael Spindler
    • 9
  • Cornelius W. Sullivan
    • 10
  1. 1.Département de biologieUniversité LavaiQuébecCanada
  2. 2.U.S. Army Cold Regions Research and Engineering LaboratoryHanoverUSA
  3. 3.Alfred-Wegener-Institut für Polar- und MeeresforschungBremerhavenFederal Republic of Germany
  4. 4.University/Tromsø (NFH)TromsøNorway
  5. 5.School of Oceanography WB-10University of WashingtonSeattleUSA
  6. 6.National Institute of Polar ResearchTokyoJapan
  7. 7.Institute of OceanologyAcademy of SciencesMoscowRussia
  8. 8.Institute of Marine ScienceUniversity of AlaskaFairbanksUSA
  9. 9.Institute for Polar EcologyKiel UniversityKielFederal Republic of Germany
  10. 10.Department of Biological SciencesUniversity of Southern CaliforniaLos AngelesUSA

Personalised recommendations