海洋生物泵研究进展
详细信息    查看全文 | 推荐本文 |
  • 英文篇名:Progress in oceanic biological pump
  • 作者:孙军 ; 李晓倩 ; 陈建芳 ; 郭术津
  • 英文作者:Sun Jun;Li Xiaoqian;Chen Jianfang;Guo Shujin;College of Marine and Environmental Sciences,Tianjin University of Science and Technology;Tianjin Key Laboratory of Marine Resources and Chemistry,Tianjin University of Science and Technology;State Oceanic Administration Key Laboratory of Marine Ecosystems and Biogeochemistry,Second Institute of Oceanography,State Oceanic Administration;State Key Laboratory of Satellite Ocean Environment Dynamics,Second Institute of Oceanography,State Oceanic Administration;
  • 关键词:生物泵 ; 南海 ; 碳循环
  • 英文关键词:biological pump;;the South China Sea;;carbon cycle
  • 中文刊名:SEAC
  • 机构:天津科技大学海洋与环境学院;天津科技大学天津市海洋资源与化学重点实验室;国家海洋局第二海洋研究所国家海洋局海洋生态系统与生物地球化学实验室;国家海洋局第二海洋研究所卫星海洋环境动力学国家重点实验室;
  • 出版日期:2016-04-15
  • 出版单位:海洋学报
  • 年:2016
  • 期:v.38
  • 基金:国家重点基础研究发展计划(2015CB954002);; 全球变化与海气相互作用专项(GASI-03-01-03-03);; 教育部新世纪优秀人才计划(NCET-12-1065);; 国家自然科学基金(91128212,41176136,41276124,41306118)
  • 语种:中文;
  • 页:SEAC201604001
  • 页数:21
  • CN:04
  • ISSN:11-2055/P
  • 分类号:3-23
摘要
海洋生物泵是以一系列海洋生物为介质将大气中的碳输运到海洋深层的过程,是海洋碳循环的重要组成部分以及未来的研究重点。本文系统地描述了海洋生物泵碳汇几个主要阶段:浮游植物沉降,浮游动物粪球颗粒沉降,透明胞外聚合颗粒物(TEP)沉降和海雪沉降以及碳酸盐反向泵过程。同时,本文对南海生物泵的研究进展进行简要介绍,服务于中国海碳循环。
        Oceanic biological pump,as an important component of oceanic carbon cycle,transfers the atmospheric carbon into the deep ocean.In this paper,the processes in oceanic carbon sink and biological pump are discussed,including sinking of phytoplankton cells,zooplankton fecal pellet package effects,transparent exopolymer particles(TEP)sinking,marine snow sinking and carbonate counter pump.Meanwhile,the present paper attempts to state the progress and perspectives of the biological pump in the South China Sea,which contributes to the carbon cycle in China Seas.
引文
[1]Falkowski P,Scholes R,Boyle E,et al.The global carbon cycle:a test of our knowledge of earth as a system[J].Science,2000,290(5490):291-296.
    [2]Sarmiento J L,Gruber N.Ocean Biogeochemical Dynamics[M].Cambridge Univ Press,2006,503.
    [3]Hansell D A,Carlson C A.Deep-ocean gradients in the concentration of dissolved organic carbon[J].Nature,1998,395(6699):263-266.
    [4]Eglinton T,Repeta D.Organic matter in the contemporary ocean[M]//Treatise on Geochemistry.Holland H D,Turekian K K.The Oceans and Marine Geochemistry,Elsevier Pergamon,Amsterdam,2004,6,145-180.
    [5]孙军.海洋浮游植物与生物碳汇[J].生态学报,2011,31(18):5372-5378.Sun Jun.Marine phytoplankton and biological carbon sink[J].Acta Ecologica Sinica,2011,31(18):5372-5378.
    [6]Passow U.Transparent exopolymer particles(TEP)in aquatic environments[J].Progress in Oceanography,2002,55(3/4):287-333.
    [7]Reynolds C S,Jaworski G H M,Cmiech H A,et al.On the annual cycle of the blue-green alga Microcystis aeruginosa Kutz.emend.Elenkin[J].Philosophical Transactions of the Royal Society of London B:Biological Sciences,1981,293(1068):419-477.
    [8]Stokes G G.On the effect of the internal friction of fluids on the motion of pendulums[M].Cambridge:Pitt Press,1851.
    [9]Reynolds C S.The ecology of phytoplankton[M].Cambridge:Cambridge University Press,2006.
    [10]Peperzak L,Colijn F,Koeman R,et al.Phytoplankton sinking rates in the Rhine region of freshwater influence[J].Journal of Plankton Research,2003,25(4):365-383.
    [11]Gross F,Zeuthen E.The buoyancy of plankton diatoms:aproblem of cell physiology[J].Proceedings of the Royal Society of London B:Biological Sciences,1948,135(880):382-389.
    [12]Anderson L W J,Sweeney B M.Role of inorganic ions in controlling sedimentation rate of a marine centric diatom ditylum brightwell[J].Journal of Phycology,1978,14(2):204-214.
    [13]Kahn N,Swift E.Positive buoyancy through ionic control in the nonmotile marine dinoflagellate Pyrocystis noctiluca Murray ex Schuett[J].Limnology and Oceanography,1978,23(4):649-658.
    [14]Malins D C,Sargent J R.Biochemical and biophysical perspectives in marine biology[M].New York:Academic Press,1974.
    [15]Smayda T J.The suspension and sinking of phytoplankton in the sea[J].Oceanography and Marine Biology,1970,8:353-414.
    [16]Fogg G E,Thake B.Algal cultures and phytoplankton ecology[M].Wisconsin:University of Wisconsin Press,1987.
    [17]Belcher J H.Notes on the physiology of Botryococcus braunii Kützing[J].Archives of Microbiology,1968,61(4):335-346.
    [18]Reynolds C S.The ecology of freshwater phytoplankton[M].Cambridge:Cambridge University Press,1984.
    [19]Dinsdale M T,Walsby A E.The interrelations of cell turgor pressure,gas-vacuolation,and buoyancy in a blue-green alga[J].Journal of Experimental Botany,1972,23(2):561-570.
    [20]Thomas R H,Walsby A E.Buoyancy regulation in a strain of Microcystis[J].Microbiology,1985,131(4):799-809.
    [21]Utkilen H C,Oliver R L,Walsby A E.Buoyancy regulation in a red Oscillatoriaunable to collapse gas vacuoles by turgor pressure[J].Archiv für Hydrobiologie,1985,102(3):319-329.
    [22]Walsby A E,Kinsman R,Ibelings B W,et al.Highly buoyant colonies of the cyanobacterium Anabaena-Lemmermannii form persistent surface waterblooms[J].Archiv für Hydrobiologie,1991,121(3):261-280.
    [23]Walsby A E.Gas vesicles[J].Microbiological Reviews,1994,58(1):94-144.
    [24]Reynolds C S.Cyanobacterial water-blooms[J].Advances in Botanical Research,1987,13:67-143.
    [25]Kromkamp J C,Mur L R.Buoyant density changes in the cyanobacteriumMicrocystis aeruginosadue to changes in the cellular carbohydrate content[J].FEMS Microbiology Letters,1984,25(1):105-109.
    [26]Walsby A E.The properties and buoyancy-providing role of gas vacuoles in Trichodesmium Ehrenberg[J].British Phycological Journal,1978,13(2):103-116.
    [27]Pitcher G C,Walker D R,Mitchel-Innes B A.Phytoplankton sinking rate dynamics in the southern Benguela upwelling system[J].Marine Ecology Progress Series,1989,55(2/3):261-269.
    [28]Talling J F.Underwater light climate as a controlling factor in the production ecology of freshwater phytoplankton[C]//Proceedings of the International Association of Theoretical and Applied Limnology Symposium,Factors Regul Wax Wane Algal Pop.1971.
    [29]Pollingher U.Freshwater armored dinoflagellates:growth,reproduction strategies,and population dynamics[M]//Sandgren C.Growth and Reproductive Strategies of Freshwater Phytoplankton.Cambridge:Cambridge University Press,1988:134-174.
    [30]Smayda T J.Turbulence,watermass stratification and harmful algal blooms:an alternative view and frontal zones as“pelagic seed banks”[J].Harmful Algae,2002,1(1):95-112.
    [31]Sommer U.The periodicity of phytoplankton in Lake Constance(Bodensee)in comparison to other deep lakes of central Europe[J].Hydrobiologia,1986,138(1):1-7.
    [32]Bienfang P,Laws E,Johnson W.Phytoplankton sinking rate determination:technical and theoretical aspects,an improved methodology[J].Journal of Experimental Marine Biology and Ecology,1977,30(3):283-300.
    [33]Riley G A,Stommel H M,Bumpus D F.Quantitative ecology of the plankton of the western North Atlantic[M].Bingham:Bingham Oceanographic Laboratory,1949.
    [34]Smayda T J,Boleyn B J.Experimental observations on the flotation of marine diatoms.Ⅰ.Thalassiosira nana,Thalassiosira rotula and Nitzschia seriata[J].Limnology and Oceanography,1965,10(4):499-509.
    [35]Smayda T J,Boleyn B J.Experimental observations on the flotation of marine diatoms.Ⅱ.Skeletonema costatum and Rhizosolenia setigera[J].Limnology and Oceanography,1966,11(1):18-34.
    [36]Smayda T J,Boleyn B J.Experimental observations on the flotation of marine diatoms.Ⅲ.Bacteriastrum hyalinum and Chaetoceros lauderi[J].Limnology and Oceanography,1966,11(1):35-43.
    [37]Steele J H,Yentsch C S.The vertical distribution of chlorophyll[J].Journal of the Marine Biological Association of the United Kingdom,1960,39(2):217-226.
    [38]Eppley R W,Holmes R W,Strickland J D H.Sinking rates of marine phytoplankton measured with a fluorometer[J].Journal of Experimental Marine Biology and Ecology,1967,1(2):191-208.
    [39]Titman D.A fluorometric technique for measuring sinking rates of freshwater phytoplankton[J].Limnology and Oceanography,1975,20(5):869-875.
    [40]Bienfang P K.SETCOL-a technologically simple and reliable method for measuring phytoplankton sinking rates[J].Canadian Journal of Fisheries and Aquatic Sciences,1981,38(10):1289-1294.
    [41]Lnnergren C.Buoyancy of natural populations of marine phytoplankton[J].Marine Biology,1979,54(1):1-10.
    [42]Waite A M,Nodder S D.The effect of in situ iron addition on the sinking rates and export flux of Southern Ocean diatoms[J].Deep-Sea Research PartⅡ:Topical Studies in Oceanography,2001,48(11/12):2635-2654.
    [43]Mei Zhiping,Legendre L,Gratton Y,et al.Phytoplankton production in the North Water Polynya:size-fractions and carbon fluxes,April to July1998[J].Marine Ecology Progress Series,2003,256:13-27.
    [44]O'brien K R,Waite A M,Alexander B L,et al.Particle tracking in a salinity gradient:A method for measuring sinking rate of individual phytoplankton in the laboratory[J].Limnology and Oceanography:Methods,2006,4(9):329-335.
    [45]Walsby A E,Holland D P.Sinking velocities of phytoplankton measured on a stable density gradient by laser scanning[J].Journal of the Royal Society Interface,2006,3(8):429-439.
    [46]Bach L T,Riebesell U,Sett S,et al.An approach for particle sinking velocity measurements in the 3-400μm size range and considerations on the effect of temperature on sinking rates[J].Marine Biology,2012,159(8):1853-1864.
    [47]Passow U.Species-specific sedimentation and sinking velocities of diatoms[J].Marine Biology,1991,108(3):449-455.
    [48]Muggli D L,Lecourt M,Harrison P J.Effects of iron and nitrogen source on the sinking rate,physiology and metal composition of an oceanic diatom from the subarctic Pacific[J].Marine Ecology Progress Series,1996,132(1):215-227.
    [49]Anderson L,Sweeney B.Diel changes in sedimentation characteristics of Ditylum brightwelli:Changes in cellular lipid and effects of respiratory inhibitors and ion-transport modifiers[J].Limnol Oceanogr,1977,22(3):539-552.
    [50]Bienfang P K,Harrison P J,Quarmby L M.Sinking rate response to depletion of nitrate,phosphate and silicate in four marine diatoms[J].Marine Biology,1982,67(3):295-302.
    [51]Culver M E,Smith W O.Effects of environmental variation on sinking rates of marine phytoplankton[J].Journal of Phycology,1989,25(2):262-270.
    [52]Bienfang P K.Size structure and sinking rates of various microparticulate constituents in oligotrophic Hawaiian waters[J].Marine Ecology Progress Series,1985,23(2):143-151.
    [53]Johnson T O,Smith W O.Sinking rates of phytoplankton assemblages in the weddell sea marginal ice-zone[J].Marine Ecology Progress Series,1986,33(2):131-137.
    [54]Riebesell U.Comparison of sinking and sedimentation rate measurements in a diatom winter/spring bloom[J].Marine Ecology Progress Series,1989,54(1/2):109-119.
    [55]Waite A,Bienfang P K,Harrison P J.Spring bloom sedimentation in a subarctic ecosystem.Ⅰ.Nutrient sensitivity[J].Marine Biology,1992,114(1):119-129.
    [56]Titman D,Kilham P.Sinking in freshwater phytoplankton:some ecological implications of cell nutrient status and physical mixing processes[J].Limnology and Oceanography,1976,21(3):409-417.
    [57]Mcnown J S,Malaika J.Effects of particle shape on settling velocity at low Reynolds numbers[J].Eos,Transactions American Geophysical Union,1950,31(1):74-82.
    [58]Hutchinson G E.A Treatise on Limnology.Ⅱ.Introduction to lake biology and their limnoplankton[M].New York:Wiley,1967.
    [59]Komar P D.Settling velocities of circular cylinders at low Reynolds numbers[J].The Journal of Geology,1980,88(3):327-336.
    [60]Davey M C,Walsby A E.The form resistance of sinking algal chains[J].British Phycological Journal,1985,20(3):243-248.
    [61]Padisk J,Soróczki-Pintér,Rezner Z.Sinking properties of some phytoplankton shapes and the relation of form resistance to morphological diversity of plankton-an experimental study[J].Hydrobiologia,2003,500(1/3):243-257.
    [62]Holland D P.Sinking rates of phytoplankton filaments orientated at different angles:theory and physical model[J].Journal of Plankton Research,2010,32(9):1327-1336.
    [63]Morris I.The physiological ecology of phytoplankton[M].Oxford:Blackwell,1980.
    [64]Lande R,Wood A M.Suspension times of particles in the upper ocean[J].Deep-Sea Research Part A Oceanographic Research Papers,1987,34(1):61-72.
    [65]Ruiz J,García C M,Rodríguez J.Sedimentation loss of phytoplankton cells from the mixed layer:effects of turbulence levels[J].Journal of Plankton Research,1996,18(9):1727-1734.
    [66]Ruiz J,Macías D,Peters F.Turbulence increases the average settling velocity of phytoplankton cells[J].Proceedings of the National academy of Sciences of the United States of America,2004,101(51):17720-17724.
    [67]Huisman J,Sommeijer B.Maximal sustainable sinking velocity of phytoplankton[J].Marine Ecology Progress Series,2002,244:39-48.
    [68]Maxey M R.The gravitational settling of aerosol particles in homogeneous turbulence and random flow fields[J].Journal of Fluid Mechanics,1987,174:441-465.
    [69]Turner J T,Ferrante J G.Zooplankton fecal pellets in aquatic ecosystems[J].BioScience,1979,29(11):670-677.
    [70]张武昌,张芳,王克.海洋浮游动物粪便通量[J].地球科学进展,2001,16(1):113-119.Zhang Wuchang,Zhang Fang,Wang Ke.Marine zooplankton fecal pellets flux[J].Advances in Earth Science,2001,16(1):113-119.
    [71]Lane P V Z,Smith S L,Urban J L,et al.Carbon flux and recycling associated with zooplanktonic fecal pellets on the shelf of the Middle Atlantic Bight[J].Deep-Sea Research PartⅡ:Topical Studies in Oceanography,1994,41(2/3):437-457.
    [72]Mller E F,Borg C M A,Jónasdóttir S H,et al.Production and fate of copepod fecal pellets across the Southern Indian Ocean[J].Marine Biology,2011,158(3):677-688.
    [73]Riser C W,Wassmann P,Olli K,et al.Production,retention and export of zooplankton faecal pellets on and off the Iberian shelf,north-west Spain[J].Progress in Oceanography,2001,51(2/4):423-441.
    [74]Riser C W,Wassmann P,Olli K,et al.Seasonal variation in production,retention and export of zooplankton faecal pellets in the marginal ice zone and central Barents Sea[J].Journal of Marine Systems,2002,38(1/2):175-188.
    [75]Wassmann P,Hansen L,Andreassen I J,et al.Distribution and sedimentation of faecal on the Nordvestbanken shelf,northern Norway,in 1994[J].Sarsia,1999,84(3/4):239-253.
    [76]Raymont J E G,Gross F.XX.On the feeding and breeding of Calanus finmarchicus under laboratory conditions[J].Proceedings of the Royal Society of Edinburgh Section B Biology,1942,61(3):267-287.
    [77]Marshall S M,Orr A P.On the biology of Calanus finmarchicus VIII.Food uptake,assimilation and excretion in adult and stage VCalanus[J].Journal of the Marine Biological Association of the United Kingdom,1955,34(3):495-529.
    [78]Paffenhfer G A,Knowles S C.Ecological implications of fecal pellet size,production and consumption by copepods[J].J Mar Res,1979,37(1):35-49.
    [79]Butler M,Dam H G.Production rates and characteristics of fecal pellets of the copepod Acartia tonsaunder simulated phytoplankton bloom conditions:implications for vertical fluxes[J].Marine Ecology Progress Series,1994,114(1/2):81-91.
    [80]Corner E D S,Head R N,Kilvington C C.On the nutrition and metabolism of zooplankton.VIII.The grazing of Biddulphiacells by Calanus helgolandicus[J].Journal of the Marine Biological Association of the United Kingdom,1972,52(4):847-861.
    [81]Ayukai T,Nishizawa S.Defecation rate as a possible measure of ingestion rate of Calanus pacificus pacificus(Copepoda:Calanoida)[J].Bulletin of the Plankton Society of Japan,1986,33(1):3-10.
    [82]Gamble J C.Copepod grazing during a declining spring phytoplankton bloom in the northern North Sea[J].Marine Biology,1978,49(4):303-315.
    [83]Poulsen L K,Kirboe T.Vertical flux and degradation rates of copepod fecal pellets in a zooplankton community dominated by small copepods[J].Marine Ecology Progress Series,2006,323:195-204.
    [84]Frangoulis C,Belkhiria S,Goffart A,et al.Dynamics of copepod faecal pellets in relation to a Phaeocystis dominated phytoplankton bloom:characteristics,production and flux[J].Journal of Plankton Research,2001,23(1):75-88.
    [85]Juul-Pedersen T,Nielsen T G,Michel C,et al.Sedimentation following the spring bloom in Disko Bay,West Greenland,with special emphasis on the role of copepods[J].Marine Ecology Progress Series,2006,314:239-255.
    [86]Riser C W,Reigstad M,Wassmann P,et al.Export or retention?Copepod abundance,faecal pellet production and vertical flux in the marginal ice zone through snap shots from the northern Barents Sea[J].Polar Biology,2007,30(6):719-730.
    [87]Beaumont K L,Plummer A J,Hosie G W,et al.Production and fate of faecal pellets during summer in an East Antarctic fjord[J].Hydrobiologia,2001,453-454(1):55-65.
    [88]Urban-Rich J L.Latitudinal variations in the contribution by copepod fecal pellets to organic carbon and amino acid flux[M].College Park,Md.:University of Maryland,1997.
    [89]Blaxter J H,Douglas B,Tyler P A,et al.The biology of calanoid copepods:the biology of calanoid copepods[M].New York:Academic Press,1998.
    [90]Smayda T J.Normal and accelerated sinking of phytoplankton in the sea[J].Marine Geology,1971,11(2):105-122.
    [91]Turner J T.Sinking rates of fecal pellets from the marine copepod Pontella meadii[J].Marine Biology,1977,40(3):249-259.
    [92]Yoon W,Kim S,Han K.Morphology and sinking velocities of fecal pellets of copepod,molluscan,euphausiid,and salp taxa in the northeastern tropical Atlantic[J].Marine Biology,2001,139(5):923-928.
    [93]Fowler S W,Small L F.Sinking rates of euphausiid fecal pellets[J].Limnology and Oceanography,1972,17(2):293-296.
    [94]Bruland K W,Silver M W.Sinking rates of fecal pellets from gelatinous zooplankton(salps,pteropods,doliolids)[J].Marine Biology,1981,63(3):295-300.
    [95]Deibel D.Still-water sinking velocity of fecal material from the pelagic tunicate Dolioletta gegenbauri[J].Marine Ecology Progress Series,1990,62:55-60.
    [96]Gorsky G,Fisher N S,Fowler S W.Biogenic debris from the pelagic tunicate,Oikopleura dioica,and its role in the vertical transport of a transuranium element[J].Estuarine,Coastal and Shelf Science,1984,18(1):13-23.
    [97]Dilling L,Alldredge A L.Can chaetognath fecal pellets contribute significantly to carbon flux?[J].Marine Ecology Progress Series,1993,92:51-58.
    [98]Madin L P.Production,composition and sedimentation of salp fecal pellets in oceanic waters[J].Marine Biology,1982,67(1):39-45.
    [99]Small L F,Fowler S W,nlüM Y.Sinking rates of natural copepod fecal pellets[J].Marine Biology,1979,51(3):233-241.
    [100]Bienfang P K.Herbivore diet affects fecal pellet settling[J].Canadian Journal of Fisheries and Aquatic Sciences,1980,37(9):1352-1357.
    [101]Dagg M J,Walser Jr W E.The effect of food concentration on fecal pellet size in marine copepods[J].Limnology and Oceanography,1986,31(5):1066-1071.
    [102]Tsuda A,Nemoto T.The effect of food concentration on the faecal pellet size of the marine copepod Pseudocalanus newmani Frost[J].Bulletin of the Plankton Society of Japan Hiroshima,1990,37(1):83-90.
    [103]Bishop J K B,Edmond J M,Ketten D R,et al.The chemistry,biology,and vertical flux of particulate matter from the upper 400mof the equatorial Atlantic Ocean[J].Deep-Sea Research,1977,24(6):511-548.
    [104]Urrère M A,Knauer G A.Zooplankton fecal pellet fluxes and vertical transport of particulate organic material in the pelagic environment[J].Journal of Plankton Research,1981,3(3):369-387.
    [105]Fowler S W,Small L F,Larosa J L.Seasonal particulate carbon flux in the coastal northwestern mediterranean-sea,and the role of zooplankton fecal matter[J].Oceanologica Acta,1991,14(1):77-85.
    [106]Graf G.Benthic-pelagic coupling in a deep-sea benthic community[J].Nature,1989,341(6241):437-439.
    [107]Pilskaln C H,Honjo S.The fecal pellet fraction of biogeochemical particle fluxes to the deep sea[J].Global Biogeochemical Cycles,1987,1(1):31-48.
    [108]Maita Y,Odate T,Yanada M.Vertical transport of organic carbon by sinking particles and the role of zoo-and phytogenic matters in neritic waters[J].Bulletin of the Faculty of Fisheries Hokkaido University,1988,39(4):265-274.
    [109]Asper V L.Measuring the flux and sinking speed of marine snow aggregates[J].Deep-Sea Research Part A Oceanographic Research Papers,1987,34(1):1-17.
    [110]Taylor G T.Variability in the vertical flux of microorganisms and biogenic material in the epipelagic zone of a North Pacific central gyre station[J].Deep Sea Research Part A Oceanographic Research Papers,1989,36(9):1287-1308.
    [111]Roman M R,Gauzens A L.Copepod grazing in the equatorial Pacific[J].Limnology and Oceanography,1997,42(4):623-634.
    [112]Roy S,Silverberg N,Romero N,et al.Importance of mesozooplankton feeding for the downward flux of biogenic carbon in the Gulf of St.Lawrence(Canada)[J].Deep-Sea Research PartⅡ:Topical Studies in Oceanography,2000,47(3/4):519-544.
    [113]Small L F,Fowler S W,Moore S A,et al.Dissolved and fecal pellet carbon and nitrogen release by zooplankton in tropical waters[J].Deep Sea Research Part A Oceanographic Research Papers,1983,30(12):1199-1220.
    [114]Wassmann P,Ypma J E,Tselepides A.Vertical flux of faecal pellets and microplankton on the shelf of the oligotrophic Cretan Sea(NE Mediterranean Sea)[J].Progress in Oceanography,2000,46(2/4):241-258.
    [115]Lapoussière A,Michel C,Gosselin M,et al.Spatial variability in organic material sinking export in the Hudson Bay system,Canada,during fall[J].Continental Shelf Research,2009,29(9):1276-1288.
    [116]Juul-Pedersen T,Michel C,Gosselin M.Sinking export of particulate organic material from the euphotic zone in the eastern Beaufort Sea[J].Marine Ecology Progress Series,2010,410:55-70.
    [117]Gleiber M R.Time series of vertical flux of zooplankton fecal pellets on the continental shelf of the western Antarctic Peninsula[D].Williamsburg:The College of William and Mary,2010.
    [118]Ayukai T,Hattori H.Production and downward flux of zooplankton fecal pellets in the anticyclonic gyre off Shikoku,Japan[J].Oceanologica Acta,1992,15(2):163-172.
    [119]Passow U,Shipe R F,Murray A,et al.The origin of transparent exopolymer particles(TEP)and their role in the sedimentation of particulate matter[J].Continental Shelf Research,2001,21(4):327-346.
    [120]Olli K,Wassmann P,Reigstad M,et al.The fate of production in the central Arctic Ocean-top-down regulation by zooplankton expatriates?[J].Progress in Oceanography,2007,72(1):84-113.
    [121]Goldthwait S A,Steinberg D K.Elevated biomass of mesozooplankton and enhanced fecal pellet flux in cyclonic and mode-water eddies in the Sargasso Sea[J].Deep-Sea Research PartⅡ:Topical Studies in Oceanography,2008,55(10/13):1360-1377.
    [122]Tamelander T,Aubert A,Wexels Riser C.Export stoichiometry and contribution of copepod faecal pellets to vertical flux of particulate organic carbon,nitrogen and phosphorus[J].Marine Ecology Progress Series,2012,459:17-28.
    [123]Carroll M L,Miquel J C,Fowler S W.Seasonal patterns and depth-specific trends of zooplankton fecal pellet fluxes in the Northwestern Mediterranean Sea[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,1998,45(8):1303-1318.
    [124]Miquel J C,Fowler S W,La Rosa J,et al.Dynamics of the downward flux of particles and carbon in the open northwestern Mediterranean Sea[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,1994,41(2):243-261.
    [125]Gowing M M,Garrison D L,Kunze H B,et al.Biological components of Ross Sea short-term particle fluxes in the austral summer of 1995-1996[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,2001,48(12):2645-2671.
    [126]Manno C,Tirelli V,Accornero A,et al.Importance of the contribution of Limacina helicinafaecal pellets to the carbon pump in Terra Nova Bay(Antarctica)[J].Journal of Plankton Research,2010,32(2):145-152.
    [127]Lalande C,Bauerfeind E,Nthig E M,et al.Impact of a warm anomaly on export fluxes of biogenic matter in the eastern Fram Strait[J].Progress in Oceanography,2013,109:70-77.
    [128]Turner J T.Zooplankton fecal pellets,marine snow,phytodetritus and the ocean's biological pump[J].Progress in Oceanography,2015,130:205-248.
    [129]Turner J T.Zooplankton fecal pellets,marine snow and sinking phytoplankton blooms[J].Aquatic Microbial Ecology,2002,27(1):57-102.
    [130]Svensen C,Riser C W,Reigstad M,et al.Degradation of copepod faecal pellets in the upper layer:Role of microbial community and Calanus finmarchicus[J].Marine Ecology Progress Series,2012,462:39-49.
    [131]Gowing M M,Wishner K F.Trophic relationships of deep-sea calanoid copepods from the benthic boundary layer of the Santa Catalina Basin,California[J].Deep-Sea Research Part A Oceanographic Research Papers,1986,33(7):939-961.
    [132]Green E P,Harris R P,Duncan A.The production and ingestion of faecal pellets by nauplii of marine calanoid copepods[J].Journal of Plankton Research,1992,14(12):1631-1643.
    [133]Lampitt R S,Noji T,Von Bodungen B.What happens to zooplankton faecal pellets?Implications for material flux[J].Marine Biology,1990,104(1):15-23.
    [134]Gonzlez H E,Smetacek V.The possible role of the cyclopoid copepod Oithona in retarding vertical flux of zooplankton faecal material[J].Marine Ecology Progress Series,1994,113(3):233-246.
    [135]Svensen C,Nejstgaard J C.Is sedimentation of copepod faecal pellets determined by cyclopoids?Evidence from enclosed ecosystems[J].Journal of Plankton Research,2003,25(8):917-926.
    [136]Noji T T,Estep K W,Macintyre F,et al.Image analysis of faecal material grazed upon by three species of copepods:evidence for coprorhexy,coprophagy and coprochaly[J].Journal of the Marine Biological Association of the United Kingdom,1991,71(2):465-480.
    [137]Alldredge A L,Passow U,Logan B E.The abundance and significance of a class of large,transparent organic particles in the ocean[J].DeepSea Research PartⅠ:Oceanographic Research Papers,1993,40(6):1131-1140.
    [138]孙军.海洋中的凝集网与透明胞外聚合颗粒物[J].生态学报,2005,25(5):1191-1198.Sun Jun.Transparent Exopolymer Particles(TEP)and aggregation web in marine environments[J].Acta Ecologica Sinica,2005,25(5):1191-1198.
    [139]Engel A.Carbon and nitrogen content of transparent exopolymer particles(TEP)in relation to their Alcian Blue adsorption[J].Mar Ecol Prog Ser,2001,219(8):1-10.
    [140]Mari X,Kirboe T.Abundance,size distribution and bacterial colonization of transparent exopolymeric particles(TEP)during spring in the Kattegat[J].Journal of Plankton Research,1996,18(6):969-986.
    [141]Hong Y,Smith W O,White A M.Studies on transparent exopolymer particles(TEP)produced in the ross sea(Antarctica)and by Phaeocystis Antarctica(Prymnesiophyceae)[J].Journal of Phycology,1997,33(3):368-376.
    [142]Alldredge A L,Passow U,Haddock H D.The characteristics and transparent exopolymer particle(TEP)content of marine snow formed from thecate dinoflagellates[J].Journal of Plankton Research,1998,20(3):393-406.
    [143]Berman T,Viner-Mozzini Y.Abundance and characteristics of polysaccharide and proteinaceous particles in Lake Kinneret[J].Aquatic Microbial Ecology,2001,24(3):255-264.
    [144]Grossart H P,Simon M,Logan B E.Formation of macroscopic organic aggregates(lake snow)in a large lake:the significance of transparent exopolymer particles,plankton,and zooplankton[J].Limnology and Oceanography,1997,42(8):1651-1659.
    [145]Riley G A.Organic aggregates in seawater and the dynamics of their formation and utilization[J].Limnology and Oceanography,1963,8(4):372-381.
    [146]Johnson B D,Cooke R C.Organic particle and aggregate formation resulting from the dissolution of bubbles in seawater[J].Limnology and Oceanography,1980,25(4):653-661.
    [147]Leppard G G,West M M,Flannigan D T,et al.A classification scheme for marine organic colloids in the Adriatic Sea:colloid speciation by transmission electron microscopy[J].Canadian Journal of Fisheries and Aquatic Sciences,1997,54(10):2334-2349.
    [148]Leppard G G.The characterization of algal and microbial mucilages and their aggregates in aquatic ecosystems[J].Science of the Total Environment,1995,165(1/3):103-131.
    [149]Leppard G G,Heissenberger A,Herndl G J.Ultrastructure of marine snow.I.Transmission electron microscopy methodology[J].Marine Ecology Progress Series,1996,135:289-298.
    [150]Leppard G G,Massalski A,Lean D R S.Electron-opaque microscopic fibrils in lakes:their demonstration,their biological derivation and their potential significance in the redistribution of cations[J].Protoplasma,1977,92(3/4):289-309.
    [151]Stoderegger K,Herndl G J.Production and release of bacterial capsular material and its subsequent utilization by marine bacterioplankton[J].Limnology and Oceanography,1998,43(5):877-884.
    [152]Baldi F,Minacci A,Saliot A,et al.Cell lysis and release of particulate polysaccharides in extensive marine mucilage assessed by lipid biomarkers and molecular probes[J].Mar Ecol Prog Ser,1997,153:45-57.
    [153]Shibata A,Kogure K,Koike I,et al.Formation of submicron colloidal particles from marine bacteria by viral infection[J].Marine Ecology Progress Series,1997,155:303-307.
    [154]Wells M L,Goldberg E D.Colloid aggregation in seawater[J].Marine Chemistry,1993,41(4):353-358.
    [155]Kepkay P E.Particle aggregation and the biological reactivity of colloids[J].Marine Ecology Progress Series,1994,109:293-304.
    [156]Chin W C,Orellana M V,Verdugo P.Spontaneous assembly of marine dissolved organic matter into polymer gels[J].Nature,1998,391(6667):568-572.
    [157]Passow U.Distribution,size,and bacterial colonization of transparent exopolymer particles(TEP)in the ocean[J].Mar Ecol Prog Ser,1994,113:185-198.
    [158]Passow U,Alldredge A L.Aggregation of a diatom bloom in a mesocosm:The role of transparent exopolymer particles(TEP)[J].Deep-Sea Research Part II:Topical Studies in Oceanography,1995,42(1):99-109.
    [159]Grossart H P,Simon M.Bacterial colonization and microbial decomposition of limnetic organic aggregates(lake snow)[J].Aquatic Microbial Ecology,1998,15(2):127-140.
    [160]Kozlowski W,Vernet M,Lamerdin S.Predominance of cryptomonads and diatoms in Antarctic coastal waters[J].Antarctic Journal of the United States,1995,30:267-268.
    [161]Passow U,Kozlowski W,Vernet M.Distribution of Transparent Exopolymer Particles(TEP)during summer at a permanent station in Antarctica[J].Antarctic Journal of the United States,1995,30:265-266.
    [162]Schuster S,Herndl G J.Formation and significance of transparent exopolymer particles in the Northern Adriatic Sea[J].Marine Ecology Progress Series,1995,124(1/3):227-236.
    [163]Ramaiah N,Yoshikawa T,Furuya K.Temporal variations in transparent exopolymer particles(TEP)associated with a diatom spring bloom in a subarctic ria in Japan[J].Marine Ecology Progress Series,2001,212(1):79-88.
    [164]Wild C.Effekte von“marine snow”-Sedimentation auf Steinkorallen(Hexacorallia,Scleractinia)des Great Barrier Reef,Australia[D].Bremen:University of Bremen,Dept of Biology and Chemistry,2000.
    [165]Passow U,Alldredge A L.A dye-binding assay for the spectrophotometric measurement of transparent exopolymer particles(TEP)[J].Limnology and Oceanography,1995,40(7):1326-1335.
    [166]Mari X,Dam H G.Production,concentration,and isolation of transparent exopolymeric particles using paramagnetic functionalized microspheres[J].Limnology and Oceanography,2004,2(1):13-24.
    [167]Mari X,Burd A.Seasonal size spectra of transparent exopolymeric particles(TEP)in a coastal sea and comparison with those predicted using coagulation theory[J].Marine Ecology Progress Series,1998,163:63-76.
    [168]Krembs C E,Eicken H,Junge K,et al.High concentrations of exopolymeric substances in Arctic winter sea ice:implications for the polar ocean carbon cycle and cryoprotection of diatoms[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,2002,49(12):2163-2181.
    [169]Engel A.Direct relationship between CO2uptake and transparent exopolymer particles production in natural phytoplankton[J].Journal of Plankton Research,2002,24(1):49-53.
    [170]García C M,Prieto L,Vargas M,et al.Hydrodynamics and the spatial distribution of plankton and TEP in the Gulf of Cdiz(SW Iberian Peninsula)[J].Journal of Plankton Research,2002,24(8):817-833.
    [171]Fabricius K E,Wild C,Wolanski E,et al.Effects of transparent exopolymer particles and muddy terrigenous sediments on the survival of hard coral recruits[J].Estuarine,Coastal and Shelf Science,2003,57(4):613-621.
    [172]Engel A.Distribution of transparent exopolymer particles(TEP)in the northeast Atlantic Ocean and their potential significance for aggregation processes[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,2004,51(1):83-92.
    [173]Radifl T,Kraus R,Fuks D,et al.Transparent exopolymeric particles'distribution in the northern Adriatic and their relation to microphytoplankton biomass and composition[J].Science of the Total Environment,2005,353(1/3):151-161.
    [174]Corzo A,Rodríguez-Glvez S,Lubian L,et al.Spatial distribution of transparent exopolymer particles in the Bransfield Strait,Antarctica[J].Journal of Plankton Research,2005,27(7):635-646.
    [175]Prieto L,Navarro G,Cózar A,et al.Distribution of TEP in the euphotic and upper mesopelagic zones of the southern Iberian coasts[J].DeepSea Research PartⅡ:Topical Studies in Oceanography,2006,53(11/13):1314-1328.
    [176]Bhaskar P V,Bhosle N B.Dynamics of transparent exopolymeric particles(TEP)and particle-associated carbohydrates in the Dona Paula bay,west coast of India[J].Journal of Earth System Science,2006,115(4):403-413.
    [177]彭安国,黄奕普.九龙江河口区TEP及其与铀、钍、钋同位素相关性的研究[J].厦门大学学报:自然科学版,2007,46(增刊1):38-42.Peng Anguo,Huang Yipu.Study on TEP and its relationships with uranium,thorium,polonium isotopes in Jiulong estuary[J].Journal of Xiamen University(Natural Science),2007,46(S1):38-42.
    [178]Ortega-Retuerta E,Reche I,Pulido-Villena E,et al.Uncoupled distributions of transparent exopolymer particles(TEP)and dissolved carbohydrates in the Southern Ocean[J].Marine Chemistry,2009,115(1/2):59-65.
    [179]Wetz M S,Robbins M C,Paerl H W.Transparent Exopolymer Particles(TEP)in a river-dominated estuary:spatial-temporal distributions and an assessment of controls upon TEP formation[J].Estuaries&Coasts,2009,32(3):447-455.
    [180]孙翠慈,王友绍,吴梅林,等.夏季珠江口透明胞外聚合颗粒物分布特征[J].热带海洋学报,2010,29(5):81-87.Sun Cuici,Wang Youshao,Wu Meilin,et al.Distribution of transparent exopolymer particles in the Pearl River Estuary in summer[J].Jouenal of Tropical Oceanography,2010,29(5):81-87.
    [181]Mari X.Carbon content and C:N ratio of transparent exopolymeric particles(TEP)produced by bubbling exudates of diatoms[J].Marine Ecology Progress Series,1999,183(3):59-71.
    [182]Sun C C,Wang Y S,Li Q P,et al.Distribution characteristics of transparent exopolymer particles in the Pearl River estuary,China[J].Journal of Geophysical Research Biogeosciences,2012,117(G4):200-209.
    [183]Alldredge A L,Silver M W.Characteristics,dynamics and significance of marine snow[J].Progress in Oceanography,1988,20(1):41-82.
    [184]Simon M,Grossart H P,Schweitzer B,et al.Microbial ecology of organic aggregates in aquatic ecosystems[J].Aquatic Microbial Ecology,2002,28(2):175-211.
    [185]Pilskaln C H,Villareal T A,Dennett M,et al.High concentrations of marine snow and diatom algal mats in the North Pacific Subtropical Gyre:Implications for carbon and nitrogen cycles in the oligotrophic ocean[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,2005,52(12):2315-2332.
    [186]Shanks A L.The abundance,vertical flux,and still-water and apparent sinking rates of marine snow in a shallow coastal water column[J].Continental Shelf Research,2002,22(14):2045-2064.
    [187]Stemmann L,Prieur L,Legendre L,et al.Effects of frontal processes on marine aggregate dynamics and fluxes:An interannual study in a permanent geostrophic front(NW Mediterranean)[J].Journal of Marine Systems,2008,70(1/2):1-20.
    [188]Alldredge A L.The contribution of discarded appendicularian houses to the flux of particulate organic carbon from oceanic surface waters[M]//Gorsky G,Youngbluth M J,Deibel D.Response of Marine Ecosystems to Global Change:Ecological Impact of Appendicularians.Paris:GB Scientific Publisher,2005:309-326.
    [189]Prairie J C,Ziervogel K,Arnosti C,et al.Delayed settling of marine snow at sharp density transitions driven by fluid entrainment and diffusionlimited retention[J].Mar Ecol Prog Ser,2013,487:185-200.
    [190]Goldthwait S,Yen J,Brown J,et al.Quantification of marine snow fragmentation by swimming euphausiids[J].Limnology and Oceanography,2004,49(4):940-952.
    [191]Lombard F,Kirboe T.Marine snow originating from appendicularian houses:age-dependent settling characteristics[J].Deep-Sea Research Part I:Oceanographic Research Papers,2010,57(10):1304-1313.
    [192]Lyons M M,Dobbs F C.Differential utilization of carbon substrates by aggregate-associated and water-associated heterotrophic bacterial communities[J].Hydrobiologia,2012,686(1):181-193.
    [193]Lyons M M,Lau Y T,Carden W E,et al.Characteristics of marine aggregates in shallow-water ecosystems:implications for disease ecology[J].EcoHealth,2007,4(4):406-420.
    [194]Kirboe T.Colonization of marine snow aggregates by invertebrate zooplankton:abundance,scaling,and possible role[J].Limnology and Oceanography,2000,45(2):479-484.
    [195]Green E P,Dagg M J.Mesozooplankton associations with medium to large marine snow aggregates in the northern Gulf of Mexico[J].Journal of Plankton Research,1997,19(4):435-447.
    [196]Dilling L,Brzezinski M A.Quantifying marine snow as a food choice for zooplankton using stable silicon isotope tracers[J].Journal of Plankton Research,2004,26(9):1105-1114.
    [197]Sano M,Maki K,Nishibe Y,et al.Feeding habits of mesopelagic copepods in Sagami Bay:Insights from integrative analysis[J].Progress in Oceanography,2013,110:11-26.
    [198]Goldthwait S A,Carlson C A,Henderson G K,et al.Effects of physical fragmentation on remineralization of marine snow[J].Marine Ecology Progress Series,2005,305:59-65.
    [199]Larson E T,Shanks A L.Consumption of marine snow by two species of juvenile mullet and its contribution to their growth[J].Marine Ecology Progress Series,1996,130(1):19-28.
    [200]Kamjunke N,Mehner T.Coupling the microbial food web with fish:are bacteria attached to cyanobacteria an important food source for underyearling roach?[J].Freshwater Biology,2001,46(5):633-639.
    [201]Newell C R,Pilskaln C H,Robinson S M,et al.The contribution of marine snow to the particle food supply of the benthic suspension feeder,Mytilus edulis[J].Journal of Experimental Marine Biology and Ecology,2005,321(2):109-124.
    [202]Kach D J,Ward J E.The role of marine aggregates in the ingestion of picoplankton-size particles by suspension-feeding molluscs[J].Marine Biology,2008,153(5):797-805.
    [203]Ward J E,Kach D J.Marine aggregates facilitate ingestion of nanoparticles by suspension-feeding bivalves[J].Marine Environmental Research,2009,68(3):137-142.
    [204]Jordan R W,Kleijne A.A classification system for living coccolithophores[M]//Winter A,Siesser W G.Coccolithophores.Cambridge:Cambridge University Press,1994:83-105.
    [205]孙军.今生颗石藻的有机碳泵和碳酸盐反向泵[J].地球科学进展,2007,22(12):1231-1239.Sun Jun.Organic carbon pump and carbonate counter pump of living coccolithophorid[J].Advances in Earth Science,2007,22(12):1231-1239.
    [206]Baumann K H,Bckel B,Frenz M.Coccolith contribution to South Atlantic carbonate sedimentation[M].Coccolithophores.Springer.2004:367-402.
    [207]Balch W M,Kilpatrick K.Calcification rates in the equatorial Pacific along 140W[J].Deep-Sea Research PartⅡ:Topical Studies in Oceanography,1996,43(4):971-993.
    [208]De Vargas C,Aubry P,Probert I,et al.Origin and Evolution of Coccolithophores:From Coastal Hunters to Oceanic Farmers[J].Evolution of primary producers in the sea,2007,12:251-285.
    [209]Passow U,Carlson C A.The biological pump in a high CO2world[J].Mar Ecol Prog Ser,2012,470(2):249-271.
    [210]Schneider B,Bopp L,Gehlen M.Assessing the sensitivity of modeled air-sea CO2exchange to the remineralization depth of particulate organic and inorganic carbon[J].Global Biogeochemical Cycles,2008,22(3):1-13.
    [211]Rost B,Riebesell U.Coccolithophores and the biological pump:responses to environmental changes[M].Coccolithophores.Springer.2004:99-125.
    [212]Salter I,Schiebel R,Ziveri P,et al.Carbonate counter pump stimulated by natural iron fertilization in the Polar Frontal Zone[J].Nature Geoscience,2014,7(12):1-5.
    [213]王汝建,林隽,郑连福,等.1993~1995年南海中部的硅质生物通量及其季节性变化:季风气候和El Nio的响应[J].科学通报,2000,45(9):974-978.Wang Rujian,Lin Jun,Zhang Lianfu,et al.Siliceous microplankton fluxes and seasonal variations in the central South China Sea during 1993—1995:monsoon climate and El Nio responses[J].Chinese Science Bulletin,2000,45(23):2168-2172.
    [214]陈建芳,郑连福,Wiesner M G,等.基于沉积物捕获器的南海表层初级生产力及输出生产力估算[J].科学通报,1998,43(6):639-642.Chen Jianfang,Zheng Lianfu,Wiesner M G,et al.Estimations of primary production and export production in the South China Sea based on sediment trap experiments[J].Chinese Science Bulletin,1998,43(7):583-586.
    [215]陈蔚芳.南海北部颗粒有机碳输出通量、季节变化及其调控过程[D].厦门:厦门大学,2008.Chen Weifang.On the export flux,seasonality and controls of particulate organic carbon in the northern South China Sea[D].Xiamen:Xiamen University,2008.
    [216]Hung C C,Gong G C.POC/234 Th ratios in particles collected in sediment traps in the northern South China Sea[J].Estuarine,Coastal and Shelf Science,2010,88(3):303-310.
    [217]Wei C L,Lin S Y,Sheu D D,et al.Particle-reactive radionuclides(234 Th,210Pb,210Po)as tracers for the estimation of export production in the South China Sea[J].Biogeosciences,2011,8(12):3793-3808.
    [218]Liu Guimei,Chai Fei.Seasonal and interannual variability of primary and export production in the South China Sea:a three-dimensional physicalbiogeochemical model study[J].ICES Journal of Marine Science,2009,66(2):420-431.
    [219]Liu K K,Chao S Y,Shaw P T,et al.Monsoon-forced chlorophyll distribution and primary production in the South China Sea:observations and a numerical study[J].Deep-Sea Research PartⅠ:Oceanographic Research Papers,2002,49(8):1387-1412.
    [220]Chai F,Liu G,Xue H,et al.Seasonal and interannual variability of carbon cycle in South China Sea:A three-dimensional physical-biogeochemical modeling study[J].Journal of Oceanography,2009,65(5):703-720.
    [221]Ma W,Chai F,Xiu P,et al.Modeling the long-term variability of phytoplankton functional groups and primary productivity in the South China Sea[J].Journal of oceanography,2013,69(5):527-544.

© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号

地址:北京市海淀区学院路29号 邮编:100083

电话:办公室:(+86 10)66554848;文献借阅、咨询服务、科技查新:66554700