海洋浮游植物胞外多聚物的研究进展
|
摘要
海洋中胞外多聚物(extracellular polymeric substances,EPS)是一类富含碳的黏性物质,由酸性多糖组成,主要由浮游植物产生。EPS主要分为三类:细胞涂层EPS、溶解性EPS和透明胞外聚合颗粒物(transparent exopolymer particles,TEP)。EPS本身呈溶解态,属于溶解有机碳库;但EPS可通过凝结或起泡方式转换形成颗粒态的TEP,属于颗粒有机碳库。EPS能被微生物降解或吸收,直接参与微食物环的碳循环。颗粒态的TEP可聚集生物有机颗粒形成"海洋雪",加快有机碳的沉降。因此EPS成为连接溶解态和颗粒态有机碳的桥梁,有效改变了海洋生态系统中有机碳库的分配,在塑造海洋生态系统结构和功能中具有重要地位。本文从EPS的特点、形成机制、对细胞积聚和海洋碳循环的影响等方面综述了目前EPS的研究进展。
The extracellular polymeric substances( EPS),a class of sticky material with high concentration of carbon,is consists of acidic polysaccharides. EPS is mainly produced by phytoplankton in marine environment. EPS is grouped generally into three categories: EPS cell coating,soluble EPS,and transparent exopolymer particles( TEP). EPS can dissolve in the water column,belonging to the dissolved organic carbon pool.However,EPS can form TEP of particles,belonging to the particulate organic carbon pool,via coagulate or bubbling.EPS can be absorbed or degraded by microorganisms and thus directly involve in carbon cycle within microbial loop. TEP can form marine snow by aggregating bio-organic particles,accelerating the sinking of organic carbon. Therefore,EPS is the bridge of dissolved organic carbon and particulate organic carbon,and plays key roles in shaping of the structure and function of in the marine ecosystems. This paper overviewed the current research progress of EPS,including,characteristics,formation mechanism,impact on the accumulation of cells and the contribution to the marine carbon cycle.
The extracellular polymeric substances( EPS),a class of sticky material with high concentration of carbon,is consists of acidic polysaccharides. EPS is mainly produced by phytoplankton in marine environment. EPS is grouped generally into three categories: EPS cell coating,soluble EPS,and transparent exopolymer particles( TEP). EPS can dissolve in the water column,belonging to the dissolved organic carbon pool.However,EPS can form TEP of particles,belonging to the particulate organic carbon pool,via coagulate or bubbling.EPS can be absorbed or degraded by microorganisms and thus directly involve in carbon cycle within microbial loop. TEP can form marine snow by aggregating bio-organic particles,accelerating the sinking of organic carbon. Therefore,EPS is the bridge of dissolved organic carbon and particulate organic carbon,and plays key roles in shaping of the structure and function of in the marine ecosystems. This paper overviewed the current research progress of EPS,including,characteristics,formation mechanism,impact on the accumulation of cells and the contribution to the marine carbon cycle.
引文
[1] FIELD C B,BEHRENFELD M J,RANDERSON J T,et al.Primary Production of the biosphere:integrating terrestrial and oceanic components[J].Science,1998,281(5374):237-240.
[2] EPPLEY R W,PETERSON B J. Particulate organic matter flux and planktonic new production in the deep ocean[J]. Nature,1979,282(5740):677-680.
[3] PASSOW U,CARLSON C A. The biological pump in a high CO2world[J]. Marine Ecology Progress Series,2012,470:249-271.
[4]LEPPARD G G.The characterization of algal and microbial mucilages and their aggregates in aquatic ecosystems[J]. Science of the Total Environment,1995,165(1/2/3):103-131.
[5]ALLDREDGE A L,SILVER M W.Characteristics,dynamics and significance of marine snow[J].Progress in Oceanography,1988,20(1):41-82.
[6]PASSOW U,DUNNE J,MURRAY J W,et al.Organic carbon to234Th ratios of marine organic matter[J]. Marine Chemistry,2006,100(3/4):323-336.
[7]VERDUGO P,ALLDREDGE A L,AZAM F,et al.The oceanic gel phase:a bridge in the DOM-POM continuum[J].Marine Chemistry,2004,92(1):67-85.
[8] MYKLESTAD SM. Release of extracellular products by phytoplankton with special emphasis on polysaccharides[J].Science of the Total Environment,1995,165(1/2/3):155-164.
[9] THORNTON D C O.Diatom aggregation in the sea:mechanisms and ecological implications[J]. European Journal of Phycology,2002,37(2):149-161.
[10] ALLDREDGE A L,PASSOW U,LOGAN B E.The abundance and significance of a class of large,transparent organic particles in the ocean[J].Deep Sea Research Part I:Oceanographic Research Papers,1993,40(6):1131-1140.
[11] PASSOW U. Production of transparent exopolymer particles(TEP)by phyto-and bacterioplankton[J].Marine Ecology Progress Series,2002,236:1-12.
[12]ENGEL A,RIEBESELL U,ZONDERVAN I. The role of transparent exopolymer particles(TEP)for variations of the C:N:Pratios during phytoplankton blooms[C]//EGS XXVII General Assembly.Nice:EGS,2002:27.
[13]PASSOW U.Transparent exopolymer particles(TEP)in aquatic environments[J]. Progress in Oceanography,2002,55(3/4):287-333.
[14]CLAQUIN P,PROBERT I,LEFEBVRE S,et al.Effects of temperature on photosynthetic parameters and TEP production in eight species of marine microalgae[J].Aquatic Microbial Ecology,2010,51(1):1-11.
[15]王大志,黄世玉,程兆第.三种海洋硅藻胞外多聚物形态、微细结构及组成的初步研究[J].海洋与湖沼,2004,35(3):273-278.
[16]BORCHARD C,ENGEL A. Organic matter exudation by Emilianiahuxleyi under simulated future ocean conditions[J]. Biogeosciences,2012,9(8):3405-3423.
[17]ENGEA.Determination of marine gel particles[M]//WURL O.Practical Guidelines for the Analysis of Seawater. Boca Raton:CRC Press,2009:125-142.
[18]UNDERWOOD G J C,BOULCOTT M,RAINES C A,et al.Environmental effects on exopolymer production by marine benthic diatoms:dynamics,changes in composition,and pathways of production[J].Journal of Phycology,2010,40(2),293-304.
[19] ZHOU J,MOPPER K,PASSOW U. The role of surface-active carbohydrates in the formation of transparent exopolymer particles by bubble adsorption of seawater[J].Limnology and Oceanography,1998,43(8):1860-1871.
[20] FUKAO T,KIMOTO K,KOTANI Y. Production of transparent exopolymer particles by four diatom species[J]. Fisheries Science,2010,76(5):755-760.
[21]陈琪,郑纪勇,杨靖亚,等.海洋底栖硅藻胞外多聚物化学成分的定量研究[J].海洋环境科学,2016,35(5):641-646.
[22] MYKLESTAD S. Production of carbohydrates by marine planktonic diatoms. II. Influence of the NP ratio in the growth medium on the assimilation ratio,growth rate,and production of cellular and extracellular carbohydrates by Chaetoceros affinis var. willei(Gran)Hustedt and Skeletonema costatum(Grev.)Cleve[J].Journal of Experimental Marine Biology and Ecology,1977,29(2):161-179.
[23] MAGALETTI E,URBANI R,SIST P,et al. Abundance and chemical characterization of extracellular carbohydrates released by the marine diatom Cylindrotheca fusiformis under N-and P-limitation[J]. European Journal of Phycology,2004,39(2):133-142.
[24]CHEN J,THORNTON D C.The effect of temperature and growth rate on tep production by Thalassiosira weissflogii[J].Journal of Phycology,2011,47:S64-S64.
[25]王大志,黄世玉,程兆第.光暗周期对3种海洋浮游硅藻胞外多糖生产的影响[J].厦门大学学报:自然科学版,2004,43(2):244-248.
[26]陈长平,徐华林,梁君荣,等.镉离子对红树林底栖硅藻新月筒柱藻胞外多糖的影响[J].厦门大学学报:自然科学版,2013,52(1):122-126.
[27]陈琪,武福源,郑纪勇,等.氧含量及附着面积对海洋底栖硅藻生长和胞外多聚物的影响[J].海洋学研究,2017,35(1):66-72.
[28]ENGEL A.The role of transparent exopolymer particles(TEP)in the increase in apparent particle stickiness(α)during the decline of a diatom bloom[J]. Journal of Plankton Research,2000,22(3):485-497.
[29]MARI X,PASSOW U,MIGON C,et al.Transparent exopolymer particles:effects on carbon cycling in the ocean[J].Progress in Oceanography,2017,151:13-37.
[30] CHOW J S,LEE C,ENGEL A. The influence of extracellular polysaccharides,growth rate,and free coccoliths on the coagulation efficiency of Emiliania huxleyi[J]. Marine Chemistry,2015,175:5-17.
[31]THORNTON D C O,CHEN J.Exopolymer production as a function of cell permeability and death in a diatom(Thalassiosira weissflogii)and a cyanobacterium(Synechococcus elongatus)[J].Journal of Phycology,2016,53(2):245-260.
[32] WINDECKER L,BRZEZINSKI M A,WEAR E,et al.Revising the release of fixed carbon in coastal phytoplankton:the role of transparent exopolymer particles(TEP)[C]//Proceedings of2016 Ocean Sciences Meeting.New Orleans:American Geophysical Union,2016:EC43A-04.
[33]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.
[34]REDFIELD A C.The influence of organisms on the composition of sea-water[M]//HILL MN. The Composition of Seawater:Comparative and Descriptive oce Anography. The Sea:Ideas and Observations on Progress in the Study of the Seas.New York:Interscience Publishers,1963,40(6):640-644.
[35]CHEN J,THORNTON D C O. Transparent exopolymer particle production and aggregation by a marine planktonic diatom(Thalassiosira weissflogii)at different growth rates[J].Journal of Phycology,2015,51(2):381-393.
[36] AZETSU-SCOTT K,PASSOW U. Ascending marine particles:significance of transparent exopolymer particles(TEP)in the upper ocean[J]. Limnology and Oceanography,2004,49(3):741-748.
[37]LEWIS J,HARRIS A S,JONES K J,et al.Long-term survival of marine planktonic diatoms and dinoflagellates in stored sediment samples[J]. Journal of Plankton Research,1999,21(2):343-354.
[2] EPPLEY R W,PETERSON B J. Particulate organic matter flux and planktonic new production in the deep ocean[J]. Nature,1979,282(5740):677-680.
[3] PASSOW U,CARLSON C A. The biological pump in a high CO2world[J]. Marine Ecology Progress Series,2012,470:249-271.
[4]LEPPARD G G.The characterization of algal and microbial mucilages and their aggregates in aquatic ecosystems[J]. Science of the Total Environment,1995,165(1/2/3):103-131.
[5]ALLDREDGE A L,SILVER M W.Characteristics,dynamics and significance of marine snow[J].Progress in Oceanography,1988,20(1):41-82.
[6]PASSOW U,DUNNE J,MURRAY J W,et al.Organic carbon to234Th ratios of marine organic matter[J]. Marine Chemistry,2006,100(3/4):323-336.
[7]VERDUGO P,ALLDREDGE A L,AZAM F,et al.The oceanic gel phase:a bridge in the DOM-POM continuum[J].Marine Chemistry,2004,92(1):67-85.
[8] MYKLESTAD SM. Release of extracellular products by phytoplankton with special emphasis on polysaccharides[J].Science of the Total Environment,1995,165(1/2/3):155-164.
[9] THORNTON D C O.Diatom aggregation in the sea:mechanisms and ecological implications[J]. European Journal of Phycology,2002,37(2):149-161.
[10] ALLDREDGE A L,PASSOW U,LOGAN B E.The abundance and significance of a class of large,transparent organic particles in the ocean[J].Deep Sea Research Part I:Oceanographic Research Papers,1993,40(6):1131-1140.
[11] PASSOW U. Production of transparent exopolymer particles(TEP)by phyto-and bacterioplankton[J].Marine Ecology Progress Series,2002,236:1-12.
[12]ENGEL A,RIEBESELL U,ZONDERVAN I. The role of transparent exopolymer particles(TEP)for variations of the C:N:Pratios during phytoplankton blooms[C]//EGS XXVII General Assembly.Nice:EGS,2002:27.
[13]PASSOW U.Transparent exopolymer particles(TEP)in aquatic environments[J]. Progress in Oceanography,2002,55(3/4):287-333.
[14]CLAQUIN P,PROBERT I,LEFEBVRE S,et al.Effects of temperature on photosynthetic parameters and TEP production in eight species of marine microalgae[J].Aquatic Microbial Ecology,2010,51(1):1-11.
[15]王大志,黄世玉,程兆第.三种海洋硅藻胞外多聚物形态、微细结构及组成的初步研究[J].海洋与湖沼,2004,35(3):273-278.
[16]BORCHARD C,ENGEL A. Organic matter exudation by Emilianiahuxleyi under simulated future ocean conditions[J]. Biogeosciences,2012,9(8):3405-3423.
[17]ENGEA.Determination of marine gel particles[M]//WURL O.Practical Guidelines for the Analysis of Seawater. Boca Raton:CRC Press,2009:125-142.
[18]UNDERWOOD G J C,BOULCOTT M,RAINES C A,et al.Environmental effects on exopolymer production by marine benthic diatoms:dynamics,changes in composition,and pathways of production[J].Journal of Phycology,2010,40(2),293-304.
[19] ZHOU J,MOPPER K,PASSOW U. The role of surface-active carbohydrates in the formation of transparent exopolymer particles by bubble adsorption of seawater[J].Limnology and Oceanography,1998,43(8):1860-1871.
[20] FUKAO T,KIMOTO K,KOTANI Y. Production of transparent exopolymer particles by four diatom species[J]. Fisheries Science,2010,76(5):755-760.
[21]陈琪,郑纪勇,杨靖亚,等.海洋底栖硅藻胞外多聚物化学成分的定量研究[J].海洋环境科学,2016,35(5):641-646.
[22] MYKLESTAD S. Production of carbohydrates by marine planktonic diatoms. II. Influence of the NP ratio in the growth medium on the assimilation ratio,growth rate,and production of cellular and extracellular carbohydrates by Chaetoceros affinis var. willei(Gran)Hustedt and Skeletonema costatum(Grev.)Cleve[J].Journal of Experimental Marine Biology and Ecology,1977,29(2):161-179.
[23] MAGALETTI E,URBANI R,SIST P,et al. Abundance and chemical characterization of extracellular carbohydrates released by the marine diatom Cylindrotheca fusiformis under N-and P-limitation[J]. European Journal of Phycology,2004,39(2):133-142.
[24]CHEN J,THORNTON D C.The effect of temperature and growth rate on tep production by Thalassiosira weissflogii[J].Journal of Phycology,2011,47:S64-S64.
[25]王大志,黄世玉,程兆第.光暗周期对3种海洋浮游硅藻胞外多糖生产的影响[J].厦门大学学报:自然科学版,2004,43(2):244-248.
[26]陈长平,徐华林,梁君荣,等.镉离子对红树林底栖硅藻新月筒柱藻胞外多糖的影响[J].厦门大学学报:自然科学版,2013,52(1):122-126.
[27]陈琪,武福源,郑纪勇,等.氧含量及附着面积对海洋底栖硅藻生长和胞外多聚物的影响[J].海洋学研究,2017,35(1):66-72.
[28]ENGEL A.The role of transparent exopolymer particles(TEP)in the increase in apparent particle stickiness(α)during the decline of a diatom bloom[J]. Journal of Plankton Research,2000,22(3):485-497.
[29]MARI X,PASSOW U,MIGON C,et al.Transparent exopolymer particles:effects on carbon cycling in the ocean[J].Progress in Oceanography,2017,151:13-37.
[30] CHOW J S,LEE C,ENGEL A. The influence of extracellular polysaccharides,growth rate,and free coccoliths on the coagulation efficiency of Emiliania huxleyi[J]. Marine Chemistry,2015,175:5-17.
[31]THORNTON D C O,CHEN J.Exopolymer production as a function of cell permeability and death in a diatom(Thalassiosira weissflogii)and a cyanobacterium(Synechococcus elongatus)[J].Journal of Phycology,2016,53(2):245-260.
[32] WINDECKER L,BRZEZINSKI M A,WEAR E,et al.Revising the release of fixed carbon in coastal phytoplankton:the role of transparent exopolymer particles(TEP)[C]//Proceedings of2016 Ocean Sciences Meeting.New Orleans:American Geophysical Union,2016:EC43A-04.
[33]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.
[34]REDFIELD A C.The influence of organisms on the composition of sea-water[M]//HILL MN. The Composition of Seawater:Comparative and Descriptive oce Anography. The Sea:Ideas and Observations on Progress in the Study of the Seas.New York:Interscience Publishers,1963,40(6):640-644.
[35]CHEN J,THORNTON D C O. Transparent exopolymer particle production and aggregation by a marine planktonic diatom(Thalassiosira weissflogii)at different growth rates[J].Journal of Phycology,2015,51(2):381-393.
[36] AZETSU-SCOTT K,PASSOW U. Ascending marine particles:significance of transparent exopolymer particles(TEP)in the upper ocean[J]. Limnology and Oceanography,2004,49(3):741-748.
[37]LEWIS J,HARRIS A S,JONES K J,et al.Long-term survival of marine planktonic diatoms and dinoflagellates in stored sediment samples[J]. Journal of Plankton Research,1999,21(2):343-354.