The oceanic gel phase: a bridge in the DOM–POM continuum

Marine Chemistry - Tập 92 Số 1-4 - Trang 67-85 - 2004
Pedro Verdugo1, Alice L. Alldredge2, Farooq Azam3, David L. Kirchman4, Uta Passow5, Peter H. Santschi6
1Department of Bioengineering, University of Washington 357962, Friday Harbor Laboratories, Friday Harbor, WA 98250, United States
2Department of Ecology, Evolution and Marine Biology, University of California, Santa Barbara, CA 93106, United States
3Scripps Institution of Oceanography, UCSD, 9500 Gilman Drive, La Jolla, CA 92093-0202, United States
4University of Delaware, Graduate College of Marine Studies, 700 Pilottown Road, Lewes, DE 19958, United States
5Alfred Wegener Institute for Polar and Marine Research, D‐27570 Bremerhaven (Germany)
6Department of Oceanography and Marine Sciences, Laboratory of Oceanographic and Environmental Research, Texas A&M University, Galveston, TX 77551, United States

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Alldredge, 1988, Characteristics, dynamics and significance of marine snow, Prog. Oceanogr., 20, 41, 10.1016/0079-6611(88)90053-5

Alldredge, 1989, Direct observations of the mass flocculation of diatom blooms: characteristics, settling velocities and formation of diatom aggregates, Deep-Sea Res., 36, 159, 10.1016/0198-0149(89)90131-3

Alldredge, 1995, Aggregation in marine systems, Deep-Sea Res., 42, 1, 10.1016/0967-0645(95)90003-9

Alldredge, 1993, The abundance and significance of a class of large, transparent organic particles in the ocean, Deep-Sea Res., Part 1, Oceanogr. Res. Pap., 40, 1131, 10.1016/0967-0637(93)90129-Q

Aluwihare, 1997, A major biopolymeric component to dissolved organic carbon in surface sea water, Nature, 387, 166, 10.1038/387166a0

Aluwihare, 2002, Chemical composition and cycling of dissolved organic matter in the Mid-Atlantic Bight, Deep-Sea Res., Part 2, Top. Stud. Oceanogr., 49, 4421, 10.1016/S0967-0645(02)00124-8

Amon, 1994, Rapid cycling of high-molecular weight dissolved organic matter in the ocean, Nature, 369, 549, 10.1038/369549a0

Azam, 1998, Microbial control of oceanic carbon flux: the plot thickens, Science, 280, 694, 10.1126/science.280.5364.694

Azam, 1993, Bacteria–organic matter coupling and its significance for oceanographic carbon cycling, Microb. Ecol., 28, 167, 10.1007/BF00166806

Balnois, 1999, Atomic force microscopy of humic substances: effects of pH and ionic strength, Environ. Sci. Technol., 33, 3911, 10.1021/es990365n

Balnois, 2000, Conformations of succinoglycan as observed by atomic force microscopy, Macromolecules, 33, 7440, 10.1021/ma0002951

Bauer, 2002, Carbon isotopic composition of DOM, 405

Benedetti, 1996, Humic substances considered as a heterogeneous Donnan gel phase, Environ. Sci. Technol., 30, 1805, 10.1021/es950012y

Benner, 2003, Abundance of amino sugars and peptidoglycan in marine particulate and dissolved organic matter, Limnol. Oceanogr., 48, 118, 10.4319/lo.2003.48.1.0118

Benner, 2002, Chemical composition and reactivity, 59

Benner, 1992, Bulk chemical characteristics of dissolved organic matter in the ocean, Science, 255, 1561, 10.1126/science.255.5051.1561

Bergamaschi, 1999, Distributions of uronic acids and O-methyl sugars in sinking and sedimentary particles in two coastal marine environments, Geochim. Cosmochim. Acta, 63, 413, 10.1016/S0016-7037(99)00075-7

Biddanda, 1997, Carbon, nitrogen, and carbohydrate fluxes during the production of particulate and dissolved organic matter by marine phytoplankton, Limnol. Oceanogr., 42, 506, 10.4319/lo.1997.42.3.0506

Biersmith, 1998, Carbohydrates in phytoplankton and freshly produced dissolved organic matter, Mar. Chem., 63, 131, 10.1016/S0304-4203(98)00057-7

Blackburn, 1998, Microscale nutrient patches in planktonic habitats shown by chemotactic bacteria, Science, 282, 2254, 10.1126/science.282.5397.2254

Borch, 1997, Concentration and composition of dissolved combined neutral sugars (polysaccharides) in seawater determined by HPLC-PAD, Mar. Chem., 57, 85, 10.1016/S0304-4203(97)00002-9

Borsheim, 1999, Monthly profiles of DOC, mono- and polysaccharides at two locations in the Trondheimsfjord (Norway) during two years, Mar. Chem., 63, 255, 10.1016/S0304-4203(98)00066-8

Buffle, 1993, vol. 1 and 2

Bundschuh, 2001, Quantification of aquatic nano particles after different steps of Bodensee water purification with laser-induced breakdown detectin (LIBD), Acta Hydrochim. Hydrobiol., 29, 7, 10.1002/1521-401X()29:1<7::AID-AHEH7>3.0.CO;2-I

Chin, 1998, Spontaneous assembly of marine dissolved organic matter into polymer gels, Nature, 391, 568, 10.1038/35345

Ciglenecki, 2000, The role of reduced sulfur species in the coalescence of polysaccharides in the Adriatic Sea, Mar. Chem., 71, 233, 10.1016/S0304-4203(00)00051-7

Costerton, 1984, 115

Cottrell, 2000, Natural assemblages of marine proteobacteria and members of the Cytophaga–Flavobacter cluster consuming low- and high- molecular-weight dissolved organic matter, Appl. Environ. Microbiol., 66, 1692, 10.1128/AEM.66.4.1692-1697.2000

Cowen, 1985, Metal deposits associated with bacteria—implications for Fe and Mn marine biogeochemistry, Deep-Sea Res., 32, 253, 10.1016/0198-0149(85)90078-0

Cowie, 1996, Digestion and alteration of the biochemical constituents of a diatom (Thalassiosira weisflogii) ingested by an herbivorous zooplankton (Calanus pacificus), Limnol. Oceanogr., 41, 581, 10.4319/lo.1996.41.4.0581

Crump, 1999, Phylogenetic analysis of particle-attached and free-living bacterial communities in the Columbia River, its estuary, and the adjacent coastal ocean, Appl. Environ. Microbiol., 65, 3192, 10.1128/AEM.65.7.3192-3204.1999

De Gennes, 1992

De Gennes, 1982, Dynamics of entangled polymer chains, Annu. Rev. Chem., 33, 49, 10.1146/annurev.pc.33.100182.000405

De Philippis, 1998, Exocellular polysaccharides from cyanobacteria and their possible applications, FEMS Microbiol. Rev., 22, 151, 10.1016/S0168-6445(98)00012-6

De Ruiter, 1997, Carrageenan biotechnology, Trends Food Sci. Technol., 8, 389, 10.1016/S0924-2244(97)01091-1

Decho, 1990, Microbial exopolymer secretions in the ocean environments their role(s) in food webs and marine processes, Oceanogr. Mar. Biol. Annu. Rev., 28, 73

Degueldre, 2000, Groundwater colloid properties: a global approach, Appl. Geochem., 15, 1043, 10.1016/S0883-2927(99)00102-X

Delong, 1993, Phylogenetic diversity of aggregate-attached versus free-living marine bacterial assemblages, Limnol. Oceanogr., 38, 924, 10.4319/lo.1993.38.5.0924

Doi, 1984

Edwards, 1974, The dynamics of polymer networks, J. Phys. A, 7, 318, 10.1088/0305-4470/7/2/018

Edwards, 1986, The theory of macromolecular networks, Biorheology, 23, 589, 10.3233/BIR-1986-23610

Emery, 1984, Organic films on particulate matter in surface waters off Eastern Asia, Sedimentology, 31, 503, 10.1111/j.1365-3091.1984.tb01816.x

Engel, 2001, Carbon and nitrogen content of transparent exopolymer particles (TEP) in relation to their Alcian Blue adsorption, Mar. Ecol., Prog. Ser., 219, 1, 10.3354/meps219001

Gardner, 2003, Determining true particulate organic carbon: bottles, pumps and methodologies, Deep-Sea Res., Part 2, Top. Stud. Oceanogr., 50, 655, 10.1016/S0967-0645(02)00589-1

Geesey, 1982, Microbial exopolymers: ecological and economic considerations, Am. Soc. Microbiol. News, 48, 9

Grossart, 2001, Bacterial motility in the sea and its ecological implications, Aquat. Microb. Ecol., 25, 247, 10.3354/ame025247

Guo, 1997, Composition and cycling of colloids in marine environments, Rev. Geophys., 35, 17, 10.1029/96RG03195

Guo, 1995, Dynamics of dissolved organic carbon (DOC) in oceanic environments, Limnol. Oceanogr., 40, 1392, 10.4319/lo.1995.40.8.1392

Guo, 2002, 234Th scavenging and its relationship to acid polysaccharide abundance in the Gulf of Mexico, Mar. Chem., 78, 103, 10.1016/S0304-4203(02)00012-9

Gustafsson, 1997, Aquatic colloids: concepts, definitions, and current challenges, Limnol. Oceanogr., 42, 519, 10.4319/lo.1997.42.3.0519

Hamanaka, 2002, Production and export of particulate fatty acids, carbohydrates and combined amino acids in the euphotic zone, Mar. Chem., 77, 55, 10.1016/S0304-4203(01)00075-5

Hedges, 1997, Comparative organic geochemistries of soils and marine sediments, Org. Geochem., 27, 319, 10.1016/S0146-6380(97)00056-9

Herndl, 1988, Ecology of amorphous aggregations (marine snow) in the Northern Adriatic Sea: I. General considerations, Mar. Ecol., 9, 79, 10.1111/j.1439-0485.1988.tb00199.x

Hernes, 1996, Neutral carbohydrate geochemistry of particulate material in the central equatorial Pacific, Deep-Sea Res., Part 2, Top. Stud. Oceanogr., 43, 1181, 10.1016/0967-0645(96)00012-4

Honeyman, 1989, A Brownian-pumping model for trace metal scavenging: evidence from Th isotopes, J. Mar. Res., 47/4, 950

Hung, 2001, Distributions of carbohydrates, including uronic acids, in estuarine waters of Galveston Bay, Mar. Chem., 73, 305, 10.1016/S0304-4203(00)00114-6

Hung, 2003, Distributions of carbohydrate species in the Gulf of Mexico, Mar. Chem., 81, 119, 10.1016/S0304-4203(03)00012-4

Hung, 2003, Production and fluxes of carbohydrate species in the Gulf of Mexico, Glob. Biogeochem. Cycles, 17, 1055, 10.1029/2002GB001988

Jackson, 1990, A model of the formation of marine algal flocs by physical coagulation processes, Deep-Sea Res., 37, 1197, 10.1016/0198-0149(90)90038-W

Jackson, 1998, Aggregation in the marine environment, Environ. Sci. Technol., 32, 2805, 10.1021/es980251w

Jensen, 1982, Abiotic factors of particles from extracellular organic carbon released by phytoplankton, Microb. Ecol., 8, 47, 10.1007/BF02011460

Johnson, 1980, Organic particle and aggregate formation resulting from the dissolution of bubbles in seawater, Limnol. Oceanogr., 25, 653, 10.4319/lo.1980.25.4.0653

Jumars, 1993, Physical constraints on marine osmotrophy in an optimal foraging context, Mar. Microb. Food Webs, 7, 121

Kepkay, 1994, Particle aggregation and the biological reactivity of colloids, Mar. Ecol., Prog. Ser., 109, 293, 10.3354/meps109293

Kepkay, 1993, Low molecular weight and colloidal DOC production during a phytoplankton bloom, Mar. Ecol., Prog. Ser., 100, 233, 10.3354/meps100233

Kerner, 2003, Self-organization of dissolved organic matter to micelle-like microparticles in river water, Nature, 422, 150, 10.1038/nature01469

Kinrade, 2002, Silicon-29 NMR evidence of a transient hexavalent silicon complex in the diatom Navicula pelliculosa, J. Chem. Soc. Dalton Trans., 3, 307, 10.1039/b105379p

Kirchman, 2002, The ecology of Cytophaga–Flavobacteria in aquatic environments, FEMS Microbiol. Ecol., 39, 91

Kirchman, 2003, Fluxes of dissolved combined neutral sugars (polysaccharides) in the Delaware estuary, Estuaries, 26, 894, 10.1007/BF02803348

Kirchman, 1991, High turnover rates of dissolved organic carbon during a spring phytoplankton bloom, Nature, 352, 612, 10.1038/352612a0

Kohn, 1972, Ion binding on polyuronides—alginate and pectin, Pure Appl. Chem., 42, 371, 10.1351/pac197542030371

Koike, 1990, Role of sub-micrometre particles in the ocean, Nature, 345, 242, 10.1038/345242a0

Leppard, 1992, Evaluation of electron microscopic techniques for the description of aquatic colloids, vol. 1, 231

Leppard, 1993, 169

Leppard, 1995, The characterization of algal and microbial mucilages and their aggregates in aquatic ecosystems, Sci. Total Environ., 165, 103, 10.1016/0048-9697(95)04546-D

Leppard, 1997, Colloidal organic fibrils of acid polysaccharides in surface water: electron-optical characteristics, activities and chemical estimates of abundance, Colloids Surf., A Physicochem. Eng. Asp., 120, 1, 10.1016/S0927-7757(96)03676-X

Leppard, 1977, Electron-opaque microscopic fibrils in lakes: their demonstration, their biological derivation and their potential significance in the redistribution of cations, Protoplasma, 92, 289, 10.1007/BF01279466

Leppard, 1996, Ultrastructure of marine snow: I. Transmission electron microscopy methodology, Mar. Ecol., Prog. Ser., 135, 289, 10.3354/meps135289

Leppard, 1996, Ultrastructure of marine snow: I. Transmission electron microscopy methodology, Mar. Ecol., Prog. Ser., 135, 289, 10.3354/meps135289

Leppard, 1997, A classification scheme for marine organic colloids in the Adriatic Sea: colloid speciation by transmission electron microscopy, Can. J. Fish. Aquat. Sci., 54, 2334, 10.1139/cjfas-54-10-2334

Leveille, 2000, Geomicrobiology of carbonate–silicate microbioalites from Hawaiian basaltic sea caves, Chem. Geol., 169, 339, 10.1016/S0009-2541(00)00213-8

Libes, 1992, 397

Ling, 2003, Does the marine copepod Calanus pacificus consume transparent exopolymer particles (TEP), J. Plankton Res., 25, 507, 10.1093/plankt/25.5.507

Logan, 1995, Rapid formation and sedimentation of large aggregates is predictable from coagulation rates (half lives) of transparent exopolymer particles (TEP), Deep-Sea Res., 42, 203, 10.1016/0967-0645(95)00012-F

Long, 1996, Abundant protein-containing particles in the sea, Aquat. Microb. Ecol., 10, 213, 10.3354/ame010213

Long, 2001, Microscale patchiness of bacterioplankton assemblage richness in seawater, Aquat. Microb. Ecol., 26, 103, 10.3354/ame026103

Longhurst, 1992, Submicron particles in Northwest Atlantic shelf water, Deep-Sea Res., Part 1, Oceanogr. Res. Pap., 39, 1, 10.1016/0198-0149(92)90016-M

Mari, 1999, Carbon content and C:N ratio of transparent exopolymeric particles (TEP) produced by bubbling exudates of diatoms, Mar. Ecol., Prog. Ser., 183, 59, 10.3354/meps183059

Mari, 1998, Seasonal size spectra of transparent exopolymeric particles (TEP) in a coastal sea and comparison with those predicted using coagulation theory, Mar. Ecol., Prog. Ser., 163, 63, 10.3354/meps163063

Mari, 2001, Non-Redfield C:N ratio of transparent exopolymeric particles in the northwestern Mediterranean Sea, Limnol. Oceanogr., 46, 1831, 10.4319/lo.2001.46.7.1831

McCarthy, 1996, Major biochemical composition of dissolved high molecular weight organic matter in seawater, Mar. Chem., 55, 281, 10.1016/S0304-4203(96)00041-2

McCave, 1984, Size spectra and aggregation of suspended particles in the deep ocean, Deep-Sea Res., 31, 329, 10.1016/0198-0149(84)90088-8

Mecozzi, 2001, Characterization of mucilage aggregates in Adriatic and Tyrrhenian Sea: structure similarities between mucilage samples and the insoluble fractions of marine humic substance, Chemosphere, 44, 709, 10.1016/S0045-6535(00)00375-1

Mopper, 1995, Role of surface-active carbohydrates in the flocculation of a diatom bloom in a mesocosm, Deep-Sea Res., Part 2, Top. Stud. Oceanogr., 42, 47, 10.1016/0967-0645(95)00004-A

Nagata, 1999, Bacterial mortality: a pathway for the formation of refractory DOM

Nissen, 1984, Multiphasic uptake of d-glucose by an oligotrophic marine bacterium, Mar. Ecol., Prog. Ser., 16, 155, 10.3354/meps016155

O'Melia, 1972, An approach to modeling lakes, Schweiz. Z. Hydrol., 34, 1, 10.1007/BF02502798

Orellana, 2000, The assembly of DOM polymers into POM microgels enhances their susceptibility to bacterial degradation, EOS, Trans. AGU., 49, 128

Orellana, 2003, Ultraviolet radiation hinders the exchange between dissolved and particulate organic carbon in the ocean, Limnol. Oceanogr., 48, 1618, 10.4319/lo.2003.48.4.1618

Orellana, 2003, Tracing the source and fate of biopolymers in seawater: application of an immunological technique, Mar. Chem., 83, 89, 10.1016/S0304-4203(03)00098-7

Pakulski, 1994, Abundance and distribution of carbohydrates in the ocean, Limnol. Oceanogr., 39, 930, 10.4319/lo.1994.39.4.0930

Passow, 2000, Formation of transparent exopolymer particles, TEP, from dissolved precursor material, Mar. Ecol., Prog. Ser., 192, 1, 10.3354/meps192001

Passow, 2002, Production of transparent exopolymer particles (TEP) by phyto- and bacterioplankton, Mar. Ecol., Prog. Ser., 236, 1, 10.3354/meps236001

Passow, 2002, Transparent exopolymer particles (TEP) in aquatic environments, Prog. Oceanogr., 55, 287, 10.1016/S0079-6611(02)00138-6

Passow, 1994, Distribution, size and bacterial colonization of transparent exopolymer particles (TEP) in the ocean, Mar. Ecol., Prog. Ser., 113, 185, 10.3354/meps113185

Passow, 1995, A dye-binding assay for the spectrophotometric measurement of transparent exopolymer particles (TEP), Limnol. Oceanogr., 40, 1326, 10.4319/lo.1995.40.7.1326

Passow, 1999, Grazing of euphausids on cells in the presence of TEP, J. Plankton Res., 21, 2203, 10.1093/plankt/21.11.2203

Quigley, 2001, Sorption irreversibility and coagulation behavior of 234Th with surface-active marine organic matter, Mar. Chem., 76, 27, 10.1016/S0304-4203(01)00045-7

Quigley, 2002, Importance of polysaccharides for 234Th complexation to marine organic matter, Limnol. Oceanogr., 47, 367, 10.4319/lo.2002.47.2.0367

Rees, 1972, Polysaccharide gels, a molecular view. Chemistry and industry, London, Soc. Chem. Ind., 19, 630

Rees, 1977

Repeta, 2002, Chemical characterization of high molecular weight dissolved organic matter in fresh and marine waters, Geochim. Cosmochim. Acta, 66, 955, 10.1016/S0016-7037(01)00830-4

Riley, 1963, Organic aggregates in seawater and the dynamics of their formation and utilization, Limnol. Oceanogr., 8, 372, 10.4319/lo.1963.8.4.0372

Sakugawa, 1985, Chemical studies on dissolved carbohydrates in the water wimples collected from the North Pacific and Bering Sea, Oceanol. Acta, 8, 185

Santschi, 1995, Isotopic evidence for contemporary origin of high-molecular weight organic matter in oceanic environments, Geochim. Cosmochim. Acta, 59, 625, 10.1016/0016-7037(94)00378-Y

Santschi, 1998, Fibrillar polysaccharides in marine macromolecular organic matter, as imaged by atomic force microscopy and transmission electron microscopy, Limnol. Oceanogr., 43, 896, 10.4319/lo.1998.43.5.0896

Santschi, 2003, Control of acid polysaccharide production, and 234Th and POC export fluxes by marine organisms, Geophys. Res. Lett., 30, 10.1029/2002GL016046

Sherr, 1988, Direct use of high molecular weight polysaccharide by heterotrophic flagellates, Nature, 335, 348, 10.1038/335348a0

Simpson, 2002, Molecular structures and associations of humic substances in the terrestrial environment, Naturwissenschaftern, 89, 84, 10.1007/s00114-001-0293-8

Simpson, 2003, New insights on the structure of algaenan from Botryoccocus braunii race A and its hexane insoluble botryals based on multidimensional NMR spectroscopy and electrospray-mass spectrometry techniques, Phytochemistry, 62, 783, 10.1016/S0031-9422(02)00628-3

Simpson, 2003, The identification of plant derived structures in humic materials using three-dimensional NMR spectroscopy, Environ. Sci. Technol., 37, 337, 10.1021/es025956j

Skoog, 1997, Aldoses in various size fractions of marine organic matter: implications for carbon cycling, Limnol. Oceanogr., 42, 1803, 10.4319/lo.1997.42.8.1803

Smetacek, 1986, Nutrient cycling in pelagic systems a reappraisal of the conceptual framework, Ophelia, 26, 401, 10.1080/00785326.1986.10422003

Stordal, 1996, Colloidal pumping: evidence for the coagulation process using natural colloids tagged with 203Hg, Environ. Sci. Technol., 30, 3335, 10.1021/es9601806

Sutcliffe, 1963, Sea surface chemistry and Langmuir circulation, Deep-Sea Res., 10, 233

Tanaka, 1992, Phase transitions of gels, A.C.S. Symp. Ser., 480, 1, 10.1021/bk-1992-0480.ch001

Twomey, 1977

van der Weerd, 1999, Modeling the dynamic adsorption/desorption of a NOM mixture: effects of physical and chemical heterogeneity, Environ. Sci. Technol., 33, 1675, 10.1021/es980815w

Viney, 1993, Processing biological polymers in the liquid crystaline state, 209

Walters, 1988, Simultaneous determination of uronic acids and aldoses in Pleankton, plant tissues, and sediment by capillary gas chromatography of n-hexylaldonamide and alditol acetates, Anal. Chem., 60, 988, 10.1021/ac00161a010

Wells, 1998, Marine colloids: a neglected dimension. News and Views, Nature, 391, 530, 10.1038/35248

Wells, 1992, Occurrence of small colloids in seawater, Nature, 353, 342, 10.1038/353342a0

Wells, 1994, The distribution of colloids in the North Atlantic and Southern Oceans, Limnol. Oceanogr., 39, 286, 10.4319/lo.1994.39.2.0286

Wen, 1997, Interactions between radioactively labeled colloids and natural particles: evidence for colloidal pumping, Geochim. Cosmochim. Acta, 61, 2867, 10.1016/S0016-7037(97)00139-7

Wilkinson, 1999, Characteristic features of the major components of freshwater colloidal organic matter revealed by transmission electron and atomic force microscopy, Colloids Surf., A Physicochem. Eng. Asp., 155, 287, 10.1016/S0927-7757(98)00874-7

Witter, 2002, Spectrophotometric measurement of seawater carbohydrate concentrations in neritic and oceanic waters from the U.S. Middle Atlantic Bight and the Delaware estuary, Mar. Chem., 77, 143, 10.1016/S0304-4203(01)00083-4

Zhou, 1998, The role of surface-active carbohydrates in the formation of transparent exopolymer particles by bubble adsorption of seawater, Limnol. Oceanogr., 43, 1860, 10.4319/lo.1998.43.8.1860