High concentrations of exopolymeric substances in Arctic winter sea ice: Implications for the polar ocean carbon cycle and cryoprotection of diatoms

Deep-Sea Research Part I: Oceanographic Research Papers

Volumn 49, Issue 12, 2002, Pages 2163-2181

  Krembs, C ^a   Eicken, H ^b   Junge, K ^a   Deming, J W ^a  

^a UNIVERSITY OF WASHINGTON   (United States)

^b UNIVERSITY OF ALASKA FAIRBANKS   (United States)

Author keywords

Arctic; Carbon; Diatom; Exopolymers; Polysaccharides; Sea ice

Indexed keywords

ECOLOGY; MICROORGANISMS; MICROSCOPIC EXAMINATION; POLYMERS;

EXOPOLYMERIC SUBSTANCES (EPS);

OCEANOGRAPHY;

CARBON CYCLE; DIATOM; EXOPOLYMER; MICROORGANISM; SEA ICE;

UNITED STATES;

BACILLARIOPHYTA; BACTERIA (MICROORGANISMS);

EID: 0346037119     PISSN: 09670637     EISSN: None     Source Type: Journal    
DOI: 10.1016/S0967-0637(02)00122-X     Document Type: Article

References (81)

1. Ackley, S.F., Dieckmann, G., Shen, H., 1987. Algal and foram incorporation into new sea ice. EOS 68, 1736.
2. Aletsee, L., Jahnke, J., 1992. Growth and productivity of the psychrophilic marine diatoms Thalassiosira antarctica Comber and Nitzschia frigida Grunow in batch cultures at temperatures below the freezing point of sea water. Polar Biology 11, 643-647.
3. Alexander, V., Horner, R., Clasby, C., 1973. Metabolism of Arctic sea ice organisms. Institute of Marine Science, University of Alaska, Fairbanks, Alaska. IMS report R74-4, p. 120.
4. Alldredge, A.L., Crocker, K.M., 1995. Why do sinking mucilage aggregates accumulate in the water column? The Science of the Total Environment 165, 15-22.
5. Alldredge, A.L., Passow, U., Logan, B.E., 1993. The abundance and significance of a class of large, transparent organic particles in the ocean. Deep-Sea Research I 40, 1131-1140.
6. Bronstein, V.L., Itkin, Y.A., Ishkov, G.S., 1981. Rejection and capture of cells by ice crystals on freezing aqueous solutions. Journal of Crystal Growth 52, 345-349.
7. Carpenter, E.J., Senjie, L., Capone, D.G., 2000. Bacterial activity in South Pole snow. Applied Environmental Microbiology 66, 4514-4517.
8. Chin, W.C., Orellana, M.V., Verdugo, P., 1998. Spontaneous assembly of marine dissolved organic matter into polymer gels. Nature 391, 568-572.
9. Cooksey, K.E., Wigglesworth-Cooksey, B., 1995. Adhesion of bacteria and diatom to surfaces in the sea: a review. Aquatic Microbial Ecology 9, 87-96.
10. Costerton, J.W., Lewandowski, Z., Caldwell, D.E., Korber, D.R., Lappin-Scott, H.M., 1995. Microbial biofilms. Annual Reviews of Microbiology 49, 711-745.
11. Cox, G.F.N., Weeks, W.F., 1983. Equations for determining the gas and brine volumes in sea-ice samples. Journal of Glaciology 29, 306-316.
12. Decho, A.W., 1990. Microbial exopolymer secretions in ocean environments: their role(s) in food webs and marine processes. Oceanographic Marine Biological Annual Reviews 28, 73-153.
13. Decho, A.W., 2000. Microbial biofilms in intertidal systems. Continental Shelf Research 20, 1257-1273.
14. Deming, J., Baross, J., 2001. Search and discovery of microbial enzymes from thermally extreme environments in the ocean. In: Dick, R.P., Burns, R.G. (Eds.), Enzymes in the Environment. Marcel Dekker, New York, pp. 327-362.
15. Eicken, H., Deming, J., Krembs, C., Stierle, A., Junge, K., Bock, C., Miller, H., 1999. Morphology and microphysics of sea-ice brine inclusions and their importance for fluid transport and microbial activity. Abstract, American Geophysical Union Annual Meeting, December 1999, San Francisco, USA.
16. Eicken, H., Bock, C., Wittig, R., Miller, H., Poertner, H.-O., 2000. Nuclear magnetic resonance imaging of sea ice pore fluids: methods and thermal evolution of pore microstructure. Cold Regions Science and Technology 31, 207-225.
17. Engel, A., 1998. Bildung, Zusammensetzung und Sinkgeschwindigkeit mariner Aggregate. Ph.D. Thesis, Berichte aus dem Institut für Meereskunde an der Christian-Albrechts-Universität, Kiel, p. 145.
18. Engel, A., Passow, U., 2001. The carbon and nitrogen content of transparent exopolymer particles (TEP) derived from exudates of marine diatoms. Marine Ecology Progress Series 219, 1-10.
19. Evans, C.A., O'Reilly, J.E., Thomas, J.P., 1987. A handbook for the measurement of chlorophyll a and primary production. SCAR Biomass Research Series Handbook 8, 1-114.
20. Franks, F., Asquith, M.H., Hammond, C.C., Skaer, H.B., Echlin, P., 1977. Polymeric cryoprotectants in the preservation of biological ultrastructure. I. Low temperature states of aqueous solutions of hydrophilic polymers. Journal of Microscopy 110, 223-238.
21. Freitag, J., 1999. The hydraulic properties of Arctic sea-ice implications for the small scale particle transport. Berichte zur Polarforschung, 325, 150pp.
22. Giannelli, V., Thomas, D.N., Kennedy, H.A., Dieckmann, G.S., Kattner, G., Haas, C., 2001. The behavior of dissolved organic matter during sea ice formation. Annals of Glaciology 33, 317-321.
23. Gleitz, M., Vonderloeff, M.R., Thomas, D.N., Dieckmann, G.S., Millero, F.J., 1995. Comparison of summer and winter in organic carbon, oxygen and nutrient concentrations in Antarctic sea ice brine. Marine Chemistry 51, 81-91.
24. Golden, K.M., Ackley, S.F., Lytle, V.I., 1998. The percolation phase transition in sea ice. Science 282, 2238-2241.
25. Gosselin, M., Levasseur, M., Wheeler, P.A., Horner, R.A., Booth, B.C., 1997. New measurements of phytoplankton and ice algal production in the Arctic ocean. Deep-Sea Research II 44, 1623-1644.
26. Gradinger, R., Spindler, M., Weissenberger, J., 1992. On the structure and development of Arctic pack ice communities in Fram Strait: a multivariate approach. Polar Biology 12, 727-733.
27. Grimm, K.A., Lange, C.B., Gill, A.S., 1997. Self-sedimentation of phytoplankton blooms in the geologic record. Sedimentary Geology 110 (3-4), 151-161.
28. Günther, S., Gleitz, M., Dieckmann, G.S., 1999. Biogeochemistry of Antarctic Sea ice: a case study on platelet layers at Drescher Inlet, Weddell Sea. Marine Ecological Progress Series 177, 1-13.
29. Hansell, D.A., Whitledge, T.E., Goering, J.J., 1993. Patterns of nitrate utilization and new production over the Bering-Chukchi shelf. Continental-Shelf-Research 13 (5-6), 601-627.
30. Hargens, A.R., Shabica, S.V., 1973. Protection against lethal freezing temperatures by mucus in an Antarctic limpet. Cryobiology 10, 331-337.
31. Heissenberger, A., Herndl, G.J., 1994. Formation of high molecular weight material by free living marine bacteria. Marine Ecological Progress Series 57, 207-217.
32. Helmke, E., Weyland, H., 1995. Bacteria in sea ice and underlying water of the eastern Weddell Sea in midwinter. Marine Ecological Progress Series 117, 269-287.
33. Hillman, S.R., 1984. Near-shore chlorophyll concentrations in the Chukchi Sea. Polar Record 22 (137), 182-186.
34. Hoagland, K.D., Rosowski, J.R., Gretz, M.R., Roemer, S.R., 1993. Diatom extracellular polymeric substances: function, fine structure, chemistry, and physiology. Journal of Phycology 29, 537-566.
35. Hong, Y., Smith, W.O., White, A.M., 1997. Studies on transparent exopolymer particles (TEP) produced in the Ross Sea (Antartica) and by Phaeocystis antarctica (Prymnesiophyceae). Journal of Phycology 33, 368-379.
36. Horner, R.A., 1996. Ice algal investigations: historical perspective. Proceedings of the NIPR Symposium. Polar Biology 9, 1-12.
37. Horner, R.A., Ackley, S.F., Dieckmann, G.S., Gulliksen, B., Hoshiai, T., Legendre, L., Melnikov, I.A., Reeburgh, W.S., Spindler, M., Sullivan, C.W., 1992. Ecology of sea ice biota. 1. Habitat, terminology, and methodology. Polar Biology 12, 417-427.
38. Hung, W.T., Chang, I.L., Hong, S.G., Young, J.D., Chen, G.W., Lee, D.L., 1996. Floc migration and chemical composition change in a freezing chamber. Journal of Environmental Science and Health Part A: Environmental Science and Engineering and Toxic and Hazardous Substance Control 31, 1053-1066.
39. Jetzek, K.C., Stanton, T.K., Gow, A.J., Lange, M.J., 1990. Acoustical and morphological properties of undeformed sea ice, laboratory and field results. In: Ackley, S.F., Weeks, W.F. (Eds.), Sea Ice Properties and Processes. Proceedings of the W.F. Weeks Sea Ice Symposium, San Francisco, December 1988. US Army Cold Regions Research and Engineering Laboratory, Monograph, pp. 67-75.
40. Junge, K., Krembs, C., Deming, J., Stierle, A., Eicken, H., 2001a. A microscopic approach to investigate bacteria under in-situ conditions in sea-ice samples. Annals of Glaciology 33, 304-310.
41. Junge, K., Eicken, H., Deming, J.W., 2001b. An Arctic wintertime study of sea-ice bacteria: abundance, activity and diversity at in-situ temperatures of -2 to -20°C. Abstract, ASM Annual Meeting, Orlando, Florida, May.
42. Kawahara, H., Ikugawa, H., Obata, H., 1996. Isolation and characterization of a marine ice-nucleating bacterium, Pseudomonas sp. KUIN-5, which produces cellulose and secretes it into the culture broth. Bioscience, Biotechnology Biochemistry 60 (9), 1474-1478.
43. Kirst, G.O., Wiencke, C., 1995. Ecophysiology of polar algae. Journal of Phycology 31, 181-199.
44. Knight, C.A., DeVries, A.L., 1994. Effects of a polymeric, nonequilibrium "antifreese" upon ice growth from water. Journal of Crystal Growth 143, 301-310.
45. Kottmeier, S.T., Sullivan, C.W., 1990. Bacterial biomass and production in pack ice of Antarctic marginal ice edge zones. Deep Sea Research I 37 (8A), 1311-1330.
46. Krembs, C., Engel, A., 2001a. Abundance and variability of microorganisms and TEP across the ice-water interface of melting first-year sea-ice in the Laptev Sea (Arctic). Marine Biology 138, 173-185.
47. Krembs, C., Gradinger, R., Spindler, M., 2000b. Implications of brine channel geometry and surface area for the interaction of sympagic organisms in Arctic Sea ice. Journal of Experimental Marine Biology and Ecology 243, 55-80.
48. Krembs, C., Junge, K., Deming, J.W., Eicken, H., 2000b. First observations on concentration and potential production and fate of organic polymers in winter sea ice from the Chukchi Sea. Abstract, International Symposium on Sea Ice and its Interactions with the Ocean Atmosphere and Biosphere, Fairbanks, Alaska, USA, 19-23 June 2000.
49. Marion, G.M., Grant, S.A., 1994. FREZCHEM: a chemical-thermodynamic model for aqueous solutions at subzero temperatures. CRREL Special Report 94-18, 35pp.
50. Maykut, G.A., 1985. The ice environment. In: Horner, R.A. (Ed.), Sea Ice Biota. CRC Press, Boca Raton, FL, pp. 21-82.
51. McConville, M.J., 1985. Chemical composition and biochemistry of sea ice microalgae. In: Horner, R.A. (Ed.), Sea Ice Biota. CRC Press, Boca Raton, FL, pp. 105-129.
52. McConville, M.J., Wetherbee, R., Bacic, A., 1999. Subcellular location and composition of the wall and secreted extracellular sulphated polysaccharides/proteoglycans of diatom Stauroneis amphioxys Gregory. Protoplasma 206, 188-200.
53. Mock, T., Gradinger, R., 2000. Changes in the photosynthetic carbon allocation in algal assemblages of Arctic sea ice with decreasing nutrient concentrations and irradiance. Marine Ecological Progress Series 202, 1-11.
54. Murase, N., 1992. Ice crystallization in biological systems. In: Maeno, N., Hondoh, T. (Eds.), Physics and Chemistry of ice. Hokkaido University Press, Sapporo, pp. 245-248.
55. Okutani, K., 1985. Isolation and fractionation of an extracellular polysaccharide from marine vibrio. Bulletin of the Japanese Society of Fish Nissuishi 51 (3), 493-496.
56. Palmisano, A.C., Sullivan, C.W., 1985. Growth, metabolism and dark survival in sea ice microalgae. In: Horner, R.A. (Ed.), Sea Ice Biota. CRC Press, Boca Raton, FL, pp. 131-146.
57. Passow, U., Alldredge, A.L., 1995. A dye-binding assay for the spectrophotometric measurement of transparent exopolymer particles (TEP). Limnology and Oceanography 40 (7), 1326-1335.
58. Passow, U., Kozlowski, W., Vernet, M., 1995. Palmer LTER: temporal variability of transparent exopolymer particles in Arthur Harbor during the 1994-1995 growth season. Antarctic Journal of the United States 30 (5), 265-266.
59. Porter, K.G., Feig, Y.S., 1980. The use of DAPI for identifying and counting aquatic microflora. Limnology and Oceanography 25, 943-948.
60. Rasband, W.S., Bright, D.S., 1995. NIH Image: a public domain image processing program for the macintosh. Journal of Microbeam Analysis 4, 137-149.
61. Raymond, J.A., Sullivan, C.W., DeVries, A.L., 1994. Release of an ice-active substance by Antarctic sea ice diatoms. Polar Biology 14 (1), 71-75.
62. Riebesell, U., Schloss, I., Smetacek, V., 1991. Aggregation of ice algae during sea ice melting in the NW Weddell Sea. Polar Biology 1, 239-248.
63. Roos, Y., 1995. Characterization of food polymers using state diagrams. Journal of Food Engineering 24, 339-360.
64. Smith, D.J., Underwood, C., 2000. The production of extracellular carbohydrates by estuarine benthic diatoms: the effect of growth phase and light and dark treatment. Journal of Phycology 36, 321-333.
65. Smith, D.J., Graham, J., Underwood, C., 1998. Exopolymer production by intertidal epipelic diatoms. Limnology and Oceanography 43 (7), 1578-1591.
66. Stierle, A., Eicken, H., Krembs, C., Junge, K., Deming, J.W., 2000. Characterization of sedimentary particles and salt precipitates in coastal sea ice from Barrow, Alaska. Abstract, International Symposium on Sea Ice and its Interactions with the Ocean Atmosphere and Biosphere, Fairbanks, Alaska, USA, 19-23 June 2000.
67. Stoderegger, K., Herndl, G., 1998. Production and release of bacterial capsular material and its subsequent utilization by marine bacterioplankton. Limnology and Oceanography 43, 877-884.
68. Stoecker, D.K., Gustafson, D.E., Merrell, J.R., Black, M.M.D., Baier, C.T., 1997. Excystment and growth of chrysophytes and dinoflagellates at low temperatures and high salinities in Antarctic sea-ice. Journal of Phycology 33, 585-595.
69. Sullivan, C.W., Palmisano, A.C., 1984. Sea ice microbial communities: distribution, abundance, and diversity of ice bacteria in McMurdo Sound, Antarctica, in 1980. Applied Environmental Microbiology 47, 788-795.
70. Sutton, R.L., 1991. Critical cooling rates to avoid ice crystalization in aqueous cryoprotectant solutions containing polymers. Chemical Society Faraday Transactions 87, 3747-3751.
71. Syvertsen, E.E., 1991. Ice algae in the Barents Sea: types of assemblages, origin, fate and role in the ice-edge phytoplankton bloom. Polar Research 10, 277-287.
72. Taylor, R., Fletcher, R.L., 1998. Cryopreservation of eukaryotic algae - a review of methodologies. Journal of Applied Phycology 10 (5), 481-501.
73. Thomas, D.N., Lara, R.J., 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 Biology 15, 477-483.
74. Thomas, D.N., Kattner, G., Engbrodt, R., Giannelli, V., Kennedy, H., Haas, C., Dieckmann, G.S., 2001. Dissolved organic matter in Antarctic sea ice. Annals of Glaciology 33, 297-303.
75. Underwood, G.J.C., Smith, D.J., 1998. Predicting epipelic diatom exopolymer concentrations in intertidal sediments from sediment chlorophyll a. Microbial Ecology 35, 116-125.
76. UNESCO, 1994. Protocols for the Joint Global Ocean Flux Study (JGOFS) core measurements, IOC Manual and Guides, 29pp.
77. Weeks, W.F., Ackley, S.F., 1986. The growth, structure and properties of sea ice. In: Untersteiner, N. (Ed.), The Geophysics of Sea Ice. Martinus Nijhoff Publishers, Dordrecht (NATO ASI B146), pp. 9-164.
78. Weissenberger, J., Dieckmann, G., Gradinger, R., Spindler, M., 1992. Sea ice: a cast technique to examine and analyze brine pockets and channel structure. Limnology and Oceanography 37, 179-183.
79. Yoshida, H., Hatakeyama, T., Hatakeyama, H., 1990. Phase transitions of the water-xanthan system. Polymer 31, 693-698.
80. Yoshida, H., Hatakeyama, T., Hatakeyama, H., 1992. Crystallization dynamics of water in polysaccharide hydrogels. In: Maeno, N., Hondoh, T. (Eds.), Physics and Chemistry of Ice. Hokkaido University Press, Sapporo, pp. 284-290.
81. Zhang, Q., Gradinger, R., Spindler, M., 1998. Dark survival of marine microalgae in the high Arctic (Greenland Sea). Polarforschung 65, 111-116.