摘要
本论文采用流式细胞仪计数法对我国黄海和东海聚球藻、微微型真核浮游生物、异养细菌在不同海区、不同季节、不同水文情况下的分布特点及影响因素进行了研究,同时采用表面荧光显微镜计数法对胶州湾浮游病毒的分布进行了探讨,不仅可以为我国近海海洋微生物生态学的深入研究提供基础性资料,更重要的是希望能为研究海域生态保护措施的制定提供科学参考。
1.构建了通过流式细胞仪FSC信号转换细胞粒径的关系,为海洋微微型浮游生物调查提供了更可靠的生物量估算方法。
2.对南黄海微微型浮游生物进行了7个月航次调查发现：聚球藻月平均丰度和生物量范围分别为4.0-54.2×103 cell/ml和0.29-5.76μgC/L；微微型真核浮游生物为1.4-8.0×103 cell/ml和0.30-1.64μgC/L；异养细菌为4.1-8.9×105 cell/ml和8.11-17.69μgC/L；对南黄海微微型浮游生物的年变化进行研究,发现聚球藻和异养细菌的丰度和生物量在春末至秋季最高,冬季和初春偏低,微微型真核浮游生物则是在春季最高,其他季节普遍较低；研究发现聚球藻和异养细菌的丰度与温度呈正相关；黄海冷水团区域内微微型浮游生物分布较少。
3.对黄海春季水华过程中微微型浮游生物的变化特点进行研究,结果表明：在所调查的4个水华过程中,异养细菌变化始终与叶绿素a含量保持一致趋势,而聚球藻和微微型真核浮游生物则出现了随水华发生,生物量升高／降低两种相反趋势；导致聚球藻和微微型真核浮游生物在水华期生物量下降的原因可能有：水华浮游植物优势种差异,光照竞争作用,营养盐消耗,浮游病毒的作用等,具体机制有待进一步研究和讨论。
4.通过对胶州湾的聚球藻、异养细菌进行了4年连续观测,发现2006年至2009年胶州湾聚球藻的表层丰度范围为257 cell/ml至263.9×103 cell/ml,季节变化趋势为春季<冬季<秋季<夏季；异养细菌表层丰度范围为1.07×104 cell/ml到86.59×105 cell/ml,季节变化趋势为冬季<秋季<春季<夏季；聚球藻和异养细菌的丰度变化都与温度存在极显著相关性,异养细菌丰度变化还与叶绿素a含量存在极显著正相关关系；对胶州湾浮游病毒的丰度和分布特点进行研究,发现2007年6-8月胶州湾的浮游病毒丰度为0.48×107/ml至22.78×107/ml,并且呈现从湾内到湾外递减的趋势,胶州湾浮游病毒丰度与异养细菌、聚球藻丰度和叶绿素a含量均呈正相关。
5.通过对长江口及邻近海域3个航次的调查,发现夏季至秋季在长江口及邻近海域聚球藻的丰度平均值约为18.73-41.25×103 cell/ml;微微型真核浮游生物为1.07.6.47×10.cell/ml;异养细菌为3.03.20.30×105 cell/ml;聚球藻的丰度主要受到温度和营养盐的影响；异养细菌丰度则主要受到叶绿素a含量和营养盐的影响；在2006年8月,长江口及邻近海区底层可以观察到大面积的水体缺氧现象,未发现聚球藻和微微型真核浮游生物的丰度和分布与低氧有明显相关；对于异养细菌,在低氧区其丰度也未出现明显变化,水体中异养细菌的呼吸作用对低氧的产生和发展贡献可能较为有限。
In this dissertation, the author used flow cytometry to study the spatial and temporal distribution pattern of Cyanobacterium Synechococcus,picoeukaryotes, heterotrophic bacteria in South Yellow Sea and East China Sea. The author also studied the distribution of virioplankton in Jiaozhou Bay with epifluorescence microscopy. With the data got, the author hope to accumulate basic information regarding marine microbiol study for future research as well as to provide scientific reference for ecological protection measurements.
1.Established the relationship between FSC signal of flow cytometry and cell diameter. With this relationship,researchers could get a better understanding of the marine picoplankton biomass in future studies.
2.During the 7 cruises in South Yellow Sea, the Synechococcus average abundance and biomass was 4.0-54.2×103 cell/ml and 0.29-5.76μgC/L. For picoeukatyotes and heterotrophic bacteria, the average abundance and biomass was 1.4-8.0×103 cell/ml and 0.30-1.64μgC/L,4.1-8.9×105cell/ml and 8.11-17.69μgC/L, respectively. Synechococcus and heterotrophic bacteria had a same seasonal variation pattern, with highest value found in late spring to autumn, and the lowest value in winter and early spring. Picoeukaryotes had high abundance and biomass in spring, and during the rest season of the year, the abundance and biomass was relatively low. A positive correlation between water temperature and Synechococcus, heterotrophic bacteria biomass was found. In Yellow Sea Cold Water Mass, picoplankton had low abundance and biomass.
3.The author reported the picoplankton variation during spring bloom in Yellow Sea. In all the 4 bloom studied, heterotrophic bacteria showed a same variation pattern, with a positive correlation with water chlorophyll a concentration.For Synechococcus and picoeukaryotes,the variation had two different trends.During the process of spring bloom, their biomass could either increase or decrease. There were several possible causes for the biomass decrease, which need further study in the future: different preponderant bloom phytoplankton species, light and nutrient competition with large phytoplarikton, and the lysis of virioplankton.
4. In Jiaozhou Bay from 2006 to 2009,the Synechococcus surface average biomass ranged from 257 cell/ml to 263.9×103 cell/ml, with a seasonal variation of spring
5.In Changjiang Estuary and adjacent area, Synechococcus, picoeukaryotes and heterotrohic bacteria abundance ranged from 18.73-41.25×103 cell/ml,1.07-6.47×103 cell/ml, and 3.03-20.30×105cell/ml,respectively, from summer to autumn. Result showed that Synechococcus abundance was influenced by water temperature and nutrients, while the heterotrophic bacteria abundance was influenced by chlorophyll a and nutrients.In August 2006, when hypoxia was found outside Changjiang estuary, no apparent influence was found with water DO to Synechococcus,picoeukaryotes abundance. Heterotrophic bacteria abundance didn't change significantly in hypoxia zone, either.
引文
[1]Azam F, Fenchel T, Field J G, Gray J S, Meyer-Reil L A, Thingstad F.The ecological role of water-column microbes in the sea. Mar Ecol Prog Ser,1983,10:257-263
[2]焦念志.海洋微微型生物生态学.北京：科学出版社,2006.
[3]Burkill P H, Leakey R J G,Owens N J P, et al. Synechococcus and its importance to the microbial foodweb of the northwestern Indian Ocean. Deep Sea Research,1993,40(3): 773-782
[4]Gallagar S M, Waterbury J B,Stoecker D K. Efficient grazing and utilization of the marine cyanobacteriun Synechococcus sp. by larvae of the bivalve Mercenara mercenaria. Mar Biol, 1994,119:251-259
[5]Christaki U, Jacque S, Dolan J R, Vaulot D, et al. Growth and grazing on Prochlorococcus and Synechococcus by two marine ciliate. Limnol Oceanogr,1999,44(1):52-61
[6]Fogg G E. Some comments on picoplankton and its importance in the pelagic ecosystem. Aquat Microb Ecol,1995,9:33-39
[7]Li W K W. Primary Production of Prochlorophytes, Cyanobacteria, and Eucaryotic Ultraphytoplankton:Measurements from Flow Cytometric Sorting. Limnol Oceanogr,1994, 39:169-175
[8]Fuhrman J A, Azam F. Bacterioplankton Secondary Production Estimates for Coastal Waters of British Columbia, Antarctica, and California. Appl Envir Microbiol,1980,39:1085-1095.
[9]Wilhelm S W, Suttle C A. Virus and nutrient cycles in the sea. BioScience,1999,49(10): 781-788
[10]Sieburth J M, Smetacek V, Lena J. Pelagic ecosystem structure:compartments of the plankton and their relationship to plankton size fractions. Limnol Oceanogr,1978,23: 1256-1263
[11]Waterbury J B, Watson S W, Guillard R R L, Brand L E.Widespread occurrence of a unicellular, marine, planktonic, cyanobacterium. Nature,1979,277:293-294
[12]Chisholm S W,Olson R J, Zettler E R, et al. A novel free-living prochlorophyte abundant in the oceanic euphotic zone. Nature,1988,334:340-343
[13]Bergh Φ, Bφrsheim K Y, Bratbak G,Heldal M.High abundance of viruses found in aquatic environments. Nature, London,1989,340:467-468
[14]Pomeroy L R. The ocean's food web, a changing paradigm. BioScience,1974,24(9):499-504
[15]Hobbie J E, Daley R J, Jasper S. Use of nuclepore filters for counting bacteria by fluorescence microscopy. Appl Environ Microbiol,1977,33:1225-1228
[16]Scaria D.On the role of bacteria production. Limnol Oceanogr,1988,33:1220-1224
[17]Pomeroy L R, Wiebe W J. Energetics of microbial food webs. Hydrobiologia,1988,159: 7-18
[18]宁修仁.微型生物食物环.东海海洋,1997,15(1)：66-68
[19]Sherr E B,Sherr B F.Role of microbes in pelagic food webs:a revised concept. Limnol Oceanogr,1988,33:1225-1227
[20]Pomeroy L R, Williams P J leB, Azam F, Hobbis J E. The Microbial Loop. Oceanography, 2007,20(2):28-33
[21]Fuhrman J A. Marine viruses and their biogeochemical and ecological effects. Nature, London,1999,399:541-548
[22]Fuhrman, J.A.,Campbell, L.,1998.Microbial microdiversity. Nature,393,410-411
[23]Johnson P W,Sieburth J. Chroococcoid cyanobacteria in the sea:A ubiquitous and diverse phototrophic biomass. Limnol Oceanogr,1979,24:928-935
[24]Stockner J G, Antia N J. Algal picoplankton from marine and freshwater ecosystems:A multidisciplinary perspective. Can J Fish Aquat Sci,1986,43:2472-2503
[25]Stockner J G. Phototrophic picoplankton:An overview from marine and freshwater ecosystems.Limnol Oceanogr,1988,33(4):765-775
[26]Stockner J G Autotrophic picoplankton in freshwater ecosystems:The view from the Summit. Intern Revue Der Gesamten Hydrobiol,1991,76:483-492
[27]Marchant H J, Wright A T, Wright S W. The distribution and abundance if chrococcoid cyanobacteria in the Southern Ocean. Polar Biol,1987,1:1-9
[28]宁修仁,史君贤,刘子琳等.南大洋蓝细菌和微微型光合真核生物的丰度与分布.中国科学C辑.1996,26(2)：164-171
[29]Murphy L S,Haugen E M. The distribution and abundance of phototrophic ultraplankton in the North Atlantic. Limnol Oceanogr,1985,30:47-58
[30]Gallagar S M, Waterbury J B,Stoecker D K. Efficient grazing and utilization of the marine cyanobacteriun Synechococcus sp. by larvae of the bivalve Mercenara mercenaria. Mar Biol, 1994,119:251-259
[31]Uysal Z. Vertical distribution of marine cyanobacteria Synechococcus spp. in the Black, Marmara, Aegean and eastern Mediterranean Seas. Deep Sea Research Ⅱ,2006,53: 1976-1987
[32]Li W K W, Rao D V S, Harrison W G, Smith J C, et al. Autotrophic picoplankton in the tropical ocean. Science,1983,219:292-295
[33]Caron D A, Lim E L, Miceli G, et al.Grazing and utilization of chroococcoid cyanobacteria and heterotrophic bacteria by protozoa in labortary cultures and a coastal plankton community. Mar Ecol Prog Ser,1991,76:205-217
[34]Jacquet S, Lennon J F, Marie D,Vaulot D.Picoplankton population dynamics in costal waters of the northwestern Mediterranean Sea. Limnol Oceanogr,1998,43(8):1916-1931
[35]Worden A Z, Nolan J K, Palenik B.Assessing the dynamics and ecology of marine picophytoplankton:The importance of the eukaryotic component. Limnol Oceanogr,2004, 49(1):168-179
[36]Shalapyonok A, Olson R J, Shalapyonok L S. Arabian Sea phytoplankton during Southwest and Northeast Monsoons 1995:composition, size structure and biomass from individual cell properties measured by flow cytometry. Deep-Sea Research Ⅱ,2001,48:1231-1261
[37]DuRand M D, Olson R J, Chisholm S W. Phytoplankton population dynamics at the Bermuda Atlantic Time-series station in the Sargasso Sea. Deep Sea Research Part Ⅱ:Topical Studies in Oceanography,2001,48(8-9):1983-2003
[38]Calvo-Diaz A, Moran, X A G.Seasonal dynamics of picoplankton in shelf waters of the southern Bay of Biscay. Aquatic Microbial Ecology,2006,42(2):159-174
[39]Fiala M, Kopczynska E E, Oriol L, Machado M C. Phytoplankton variability in the Crozet Basin frontal zone (Southwest Indian Ocean) during austral summer. Journal of Marine System,2004,50:243-261
[40]Fernandez C, Thyssen M, Denis M. Microbial community structure along 18°W (39°N-44.5°N) in the NE Atlantic in late summer 2001 (POMME programme).Journal of Marine System,2008,71:46-62
[41]Tarran G A, Zubkov M V, Sleigh M A, et al. Microbial community structure and standing stocks in the NE Atlantic in June and July of 1996.Deep-Sea Research Ⅱ,2001,48:963-985
[42]Li H B,Xiao T, Lv R H, Ding T, Lin Y A. Impact of tidal front on the distribution of bacterioplankton in the southern Yellow Sea, China. Journal of Marine Systems,2007,67: 263-271
[43]Li H B,Xiao T, Ding T, Lv R H.Effect of the Yellow Sea Cold Water Mass (YSCWM) on distribution of bacterioplankton. Acta Ecologica Sinica,2006,26(4):1012-1020
[44]赵三军,肖天,李洪波,徐剑虹.胶州湾聚球菌(Synechococcus spp.)蓝细菌的分布及其对初级生产力的贡献.海洋与湖沼,2005,36(6)：534-540
[45]宁修仁,沃洛D.长江口及毗连东海水域蓝细菌的分布和细胞特性及其环境调节.海洋学报,1991,13：552-559
[46]Jiao N Z, Yang Y H, Hong N, et al. Dynamics of autotrophic picoplankton and heterotrophic bacteria in the East China Sea. Continental Shelf Research,2005,25:1265-1279
[47]Liu H B, Chang J, Tseng C M, Wen L S, Liu K K. Seasonal varibility of picoplankton in the Northern South China Sea at the SEATS station. Deep Sea Research Ⅱ,2007,54:1602-1616
[48]肖天,王荣.春季与秋季渤海蓝细菌(聚球蓝细菌属)的分布特点.生态学报,2002,22(12)：2071-2078
[49]孙晟.肖天.岳海东.秋季与冬季东、黄海蓝细菌(Synechococcus spp)生态分布特点.海洋与湖沼,2003,34(2)：161-168
[50]赵三军.黄、东海海洋浮游细菌的生态学研究.[博士学位论文].青岛：中国科学院海洋研究所,2007
[51]Pan L A, Zhang J, Zhang L H. Picophytoplankton, nanophytoplankton, heterotrohpic bacteria and viruses in the Changjiang Estuary and adjacent coastal waters. Journal of Plankton Research,2007,29(2):187-197
[52]Chiang K P, Kuo M C, Chang J, Wang R H, Gong G C.Spatial and temporal variation of the Synechococcus population in the East China Sea and its contribution to phytoplankton biomass. Continental Shelf Research,2002,22(1):3-13
[53]郑天凌,默哈默得,陈进才,等.台湾海峡海海域细菌生物量及其环境因子研究,中国海洋文集.北京.海洋出版社,1997：146-152
[54]郑天凌,王斐,徐美珠,洪华生.台湾海峡海域细菌产量、生物量及其在微食物环中的作用.海洋与湖沼,2002,33(4)：415-423
[55]宁修仁,蔡昱明,李国为,史君贤.南海北部微微型光合浮游生物的丰度及环境调控.海洋学报,2003,25(3)：83-97
[56]Ma Y, Jiao N Z, Zeng, Y H. Natural community structure of cyanobacteria in the South China Sea as revealed by rpoC1 gene sequence analysis. Letters in applied microbiology,2004, 39(4):353-358
[57]王晨阳,赵三军,岳海东,肖天.两株海洋聚球蓝细菌(Synechococcus spp.)的16s rDNA系统发生及生理生态特征分析.高技术通讯,2006,16(1)：100-105
[58]Simon N, Barlow RG, Marie D, et al.Characterization of oceanic photosynthetic picoeukaryotes by flow cytometry. Journal of Phycology,1994,30:922-935
[59]Diez B,Pedros-Alio C, Massana R. Study of Genetic Diversity of Eukaryotic Picoplankton in Different Oceanic Regions by Small-Subunit rRNA Gene Cloning and Sequencing. Appl Envir Microbiol,2001,67:2932-2941
[60]Biegala I C, Not F, Vaulot D, Simon N, Quantitative Assessment of Picoeukaryotes in the Natural Environment by Using Taxon-Specific Oligonucleotide Probes in Association with Tyramide Signal Amplification-Fluorescence In Situ Hybridization and Flow Cytometry. Appl Envir Microbiol,2003,69(9):5519-5529
[61]Pan L A, Zhang L H, Zhang J, et al. On-board flow cytometry observation of picoplankton community structure in the East China Seas during the fall of different years.FEMS Microbiol Ecol,2005,52:243-253
[62]黄邦钦,洪华生,林学举,陈纪新.台湾海峡微微型浮游植物的生态研究—Ⅰ.时空分布及其调控机制.海洋学报,2003,25(4)：72-82
[63]袁洁,邵鹏,陈月琴等.南沙群岛微型与超微型真核藻类遗传多样性的初步研究.海洋科学,2003,27(7)：43-47
[64]Yuan J, Chen M Y, Shao P, et al.Genetic diversity of small eukaryotes from the coastal waters of Nansha Islands in China. FEMS Microbiol Lett,2004,240(2):163-170
[65]Partensky F, Blanchot J, Lantoine F, et al. Vertical structure of picophytoplankton at different trophic sites of the tropical northeastern Atlantic Ocean. Deep-Sea Research Ⅰ,1996,43(8): 1191-1213
[66]Liu H, Suzuki K, Minami C, et al. Picoplankton community structure in the subarctic Pacific Ocean and the Bering Sea during summer 1999. Mar Ecol Prog Ser,2002,237:1-14
[67]Iriarte A, Purdie D A. Size distribution of chlorophyll a biomass andprimary production in a temperate estuary (Southampton Water):the contribution ofphotosynthetic picoplankton. Mar Ecol Prog Ser,1994,115:283-297
[68]Ferguson R L, Rublee P. Contribution of bacteria to standing crop of coastal plankton. Limnol Oceanogr,1976,21(1):141-145
[69]Marie D, Simon N, Guillou L, et al. Flow cytometry analysis of marine picoplankton. In:In Living Color:Protocols in Flow Cytometry and Cell Sorting. Berlin:Springer-Verlag,2000: 421-454
[70]Krichman D L,Keil R G,Simon M, et al. Biomass and production of heterotrophic bacterioplankton in the oceanic subarctic pacific. Deep-Sea Research Ⅰ,1993,40:967-988
[71]Williams P J L. Incorporation of microheterotrophic processes into the classical paradigm of the plankton food web. Kiel. Meeresforsch. Sonderh,1981,5:1-28
[72]林凤翱,贺杰.北黄海三类废弃物试验倾倒区海洋细菌的生态学研究：Ⅰ海洋异养细菌的分布.海洋环境科学,1989,8(3)：10-15
[73]刘志刚,韩建新.湛江港南三岛近岸海区异养细菌数量变动与潮汐周期关系研究.湛江水产学院学报,1996,16(1)：19-24
[74]Hewson I, Fuhrman J A. Viriobenthos Production and Virioplankton Sorptive Scavenging by Suspended Sediment Particles in Coastal and Pelagic Waters.Microb Ecol,2003,46: 337-347
[75]Caron D A, Peele, E R, Lim E L, Dennett, M R. Picoplankton and nanoplankton and their trophic coupling in surface waters of the Sargasso Sea south of Bermuda. Limnol Oceanogr, 1999,44(2):259-272
[76]Liu H, Imai K, Suzuki K, Nojiri Y, Tsurushima N, Saino T. Seasonal variability of picophytoplankton and bacteria in the western subarctic Pacific Ocean at station KNOT. Deep Sea Research Ⅱ,2002,49:5409-5420
[77]白洁,李岿然,李正炎等.渤海春季浮游细菌分布与生态环境因子的关系.中国海洋大学学报,2003,33(6)：841-846
[78]Hyun J H, Kim K H. Bacterial abundance and production during the unique spring phytoplankton bloom in the central Yellow Sea. Marine Ecology Progress Series,2003,252: 77-88
[79]赵三军.肖天.岳海东.秋季东、黄海异养细菌(Heterotrophic Bacteria)的分布特点.海洋与湖沼,2003,34(3)：295-305
[80]刘诚刚,宁修仁,蔡昱明,郝锵,乐凤凤.南海北部及珠江口细菌生产力研究.海洋学报,2007,29(2)：112-122
[81]赵三军,肖天,李洪波,徐剑虹.胶州湾异养细菌及大肠菌群的分布及对陆源污染的指示.海洋与湖沼,2005,36(6)：541-547
[82]Wommack K E, Colwell R R. Virioplankton:viruses in aquatic ecosystems. Microbiology and Molecular Biology Reviews,2000,64:69-114
[83]王斐,郑天凌,洪华生.海洋病毒在微生物食物环中的重要作用.海洋科学,1998,4：41-43
[84]Hara S, Koike I, Terauchi K, et al. Abundance of viruses in deep oceanic waters. Mar Ecol Prog Ser,1996,145:269-277
[85]Marie D, Brussard C P D, Thyrhaug, R, et al. Enumeration of marine viruses in culture and natural samples by flow cytometry. Appl Envir Microbiol,1999,65:45-52
[86]Hara S, Terauchi K, Koike I, et al. Abundance of viruses in marine waters:assessment by Epifluorescence and Transmission Electron Microscopy. Appl Envir Microbiol,1991,57(9): 2731-2734
[87]Steward G F, Fandino L B,Hollibaugh J T, et al. Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of thecentral Arctic Ocean. Deep-Sea Research I,2007,54:1744-1757
[88]Marchant H, Davidson A, Wright S, et al. The distribution and abundance of viruses in the Southern Ocean during spring. Antarctic Science,2000,12(4):414-417
[89]时瑶.黄海西北部细菌和浮游病毒的生态分布特征及其影响因素研究.[硕士学位论文].青岛：中国海洋大学,2009
[90]白晓歌,汪岷,马晶晶,等.冬季和春季长江口及其近海水域浮游病毒丰度的分析.海洋与湖沼,2007,38(4)：367-372
[91]Jiao N Z, Zhao Y L, Luo T W, et al. Natural and anthropogenic forcing on the dynamics of virioplankton in the Yangtze river estuary. J Mar Biol Assoc UK,2006,86,543-550
[92]季倩.长江口邻近海域水体及底栖微微型浮游生物研究.[硕士学位论文].上海：华东师范大学,2008
[93]Shiah F K, Ducklow H W. Temperature and substrate regulation of bacterial abundance, production and specific growth rate in Chesapeake Bay, Mar Ecol Prog Ser,1994,103: 297-308
[94]Shiah F K, Ducklow H W. Temperature regulation of heterotrophic bacterioplankton abundance, production and specific growth rate in Chesapeake Bay. Limnol.Oceanogr.,1994, 39(6):1243-1258
[95]李清雪,赵海萍,陶建华.渤海湾海域浮游细菌的生态研究.海洋技术,2005,24(4)：50-53
[96]Li W K W. Annual average abundance of heterotrophic bacteria and Synechococcus on surface ocean waters. Limnol Oceanogr,1998,43(7):1746-1753
[97]刘子琳,越川海,宁修仁等.长江冲淡水区细菌生产力研究.海洋学报,2001,23(4)：93-99
[98]Jiao N Z, Yang Y H, Koshikawa H, et al. Influence of hydrographic conditions on picoplankton and heterotrophic bacteria in the East China Sea, a marginal sea of the Northwest Pacific. Aquat Micro Ecol,2002,30(1):37-48
[99]Amon R M W, Benner R. Bacterial utilization of different size classes of dissolved organic matter. Limnol. Oceanogr,1996,41(1):41-51
[100]Bernard L, Curties C, Servais P, et al. Relationship among bacterial cell size, productivity and genetic diversity in aquatic environments using cell sorting and flow cytometry. Micro Ecol,2000,40:148-158
[101]Wheeler P A, Krichman D L. Utilization of inorganic and organic nitrogen by bacteria in marine systems. Limnol. Oceanogr,1986,31(5):998-1009
[102]Rivkin R B, Anderson M R. Inorganic nutrient limitation of oceanic bacterioplankton. Limnol. Oceanogr,1997,42(4):730-740
[103]Scanlan D J, West N J. Molecular ecology of the marine cyanobacterial genera Prochlorococcus and Synechococcus. FEMS Microbiol Ecol,2002,40:1-12
[104]Ducklow H W, Hill S M. The growth of heterotrophic bacteria in the surface waters of warm core rings. Limnol.Oceanogr,1985,30:239-259
[105]Rassoulzadegan F, Laval-Peuto M,.Sheldon R W. Partitioning of the food ration of marine ciliates between pico-and nanoplankton. Hydrobiologia,1988,159:75-88.
[106]Rassoulzadegan F, Sheldon R W. Predator-prey interactions of nanozooplankton and bacteria in an oligotrophic marine environment. Limnol.Oceanogr,1986,31:1010-1021.
[107]白洁,张昊飞,李岿然,等.海洋异养浮游细菌生物量及生产力的制约因素.中国海洋大学学报,2004,34(3)：594-602
[108]Wilhelm S W, Weinbauer M G, Suttle C A, et al. The role of sunlight in the removal and repair of viruses in the sea. Limnol. Oceanogr,1998,43:586-592
[109]Suttle C A, Chan A M, Cottrell M T. Infection of phytoplankton by viruses and reduction of primary productivity. Nature,1990,387:467-469
[110]Suttle C A, Chan A M. Dynamics and distribution of cyanophages and their effect on marine Synechococcus spp. Appl Environ Microbiol,1994,60:3167-3174
[111]赵三军,岳海东,肖天.海洋异养细菌生物量与生产力的研究方法.海洋科学,2002,26(1)：21-23
[112]梁智辉,朱慧芬,陈九武.流式细胞术基本原理与实用技术.武汉：华中科技大学出版社，2008.
[113]Olson R J, Vaulot D, Chisholm S W. Marine phytoplankton distribution measured using shipboard flow cytometry. Deep-Sea Research Part A. Oceanographic Research Papers,1985, 32(10):1273-1280
[114]Duda A M, Sherman K. A new imperative for improving management of large marine ecosystems. Ocean& Coastal Management,2002,45:797-833
[115]曾成奎.中国海洋志.郑州：大象出版社,2003
[116]Wei H, Su J, Wan R J, Wang L, Lin Y A. Tidal front and the convergence of anchovy (Engraulis japonicus) eggs in the Yellow Sea. Fisheries Oceanography,2003,12(4/5): 434-442
[117]袁业立,李惠卿.黄海冷水团环流结构及其生成机制研究I.O.阶解及冷水团的环流结构.中国科学B辑,1993,23(1)：90-103。
[118]苏纪兰,黄大吉.黄海冷水团的环流结构.海洋与湖沼,1995,26(5)：1-7
[119]张启龙,翁学传,杨玉玲.南黄海春季水团分析.海洋与湖沼,1996,27(4)：421-428
[120]梁玉波.海洋生物生态调查技术规程.北京：海洋出版社,2006
[121]Krempin D W,Sullivan C W. The seasonal abundance, vertical distribution, and relative microbial biomass of chroococcoid cyanobacteria at a station in southern California coastal waters,1981,27:1341-1344
[122]Harris G P. Picoplankton Ecology, Structure Function and Fluctuations. London:Chapman and Hall,1986,221-235
[123]Verity P, Robertson C Y, Tronzo C R, Andrews M G, Nelson J R, Sieracki M E. Relationships between cell volume and the carbon and nitrogen content of marine photosynthetic nanoplankton. Limnol Oceanogr,1992,37(7):1434-1446
[124]Mackey K R M, RivlinT, Grossman A R, et al. Picophytoplankton responses to changing nutrient and light regimes during a bloom. Marine Biology,2009,156:1531-1546
[125]Lee S,Fuhrman J A. Relationships between biovolume and biomass of naturally derived marine bactcrioplankton. Applied and Environmental Microbiology,1987,53(6):1298-1303
[126]Blanchot J, Andre J M, Navarette C, Neveuxd J, Radenac M H.Picophytoplankton in the equatorial Pacific:vertical distribution in the warm pool and in the high nutrient low chlorophyll conditions. Deep-Sea Research I,2001,48:297-314
[127]Olson R J, Zettler E R, Anderson O K. Discrimination of eukaryotic phytoplankton cell types from light scatter and autofluorescence properties measured by flow cytometry. Cytometry,1989,10:636-643
[128]Grob C, Ulloa O, Li W K M, Alarcon G, Fukasawa M, Watanabe Shuichi.2007. Picoplankton abundance and biomass across the eastern South Pacific Ocean along latitude 32.5 S.Mar. Ecol. Prog. Ser.,332:53-62.
[129]Campbell L, Nolla H A, Vaulot D.The importance of Prochlorococcus to community structure in the central North Pacific Ocean. Limnol Oceanogr,1994,39:954-961
[130]Fuhrman J A, Sleeter T D, Carlson C A, Proctor L M. Dominance of bacterial biomass in the Sargasso Sea and its ecological implications. Mar Ecol Prog Ser,1989,57:207-217
[131]Zubkov M V, Sleigh M A, Burkill P H, Leakey R J G. Picoplankton community structure on the Atlantic Meridional Transect:a comparison between seasons. Prog Oceanogr,2000,45: 369-386
[132]Fukuda R, Ogawa H, Nagata T, Koike I. Direct determination of carbon and nitrogen contents of natural bacterial assemblages in marine environments. Appl Environ Microb, 1998,64:3352-3358
[133]Agawin N S R, Duarte C M, Agusti S. Growth and abundance of Synechococcus sp. in a Mediterranean Bay:seasonality and relationship with temperature. Mar Ecol Prog Ser.,1998, 170:45-53
[134]Smith R, Hall J. Bacterial abundance and production in different water masses around South Island, New Zealand. New Zealand Journal of Marine and Freshwater Research,1997,31: 515-524
[135]Lemee R, Rochelle-Newall E, Van Wambeke F, et al. Seasonal variation of bacterial production, respiration and growth efficiency in the open NW Mediterranean Sea. Aquat Microb Ecol,2002,29:227-237
[136]冯士笮,李凤岐,李少菁.海洋科学导论.北京：高等教育出版社,1999,272～350
[137]王真良,刘晓丹.北黄海浮游动物昼夜垂直移动的初步研究.黄渤海海洋,1989,7(4)：50～54
[138]左涛,王荣,王克等.夏季南黄海浮游动物的垂直分布与昼夜垂直移动.生态学报,2004,24(3)：524～530
[139]Jochem F.On the distribution and importance of picocyanobacteria in a boreal inshore area (Kiel Bight, Western Baltic). Journal of plankton research,1988,10(55):1009-1022
[140]肖天,岳海东,张武昌,王荣.东海聚球蓝细菌(Synechococcus)的分布特点及在微食物环中的作用.海洋与湖沼,2003,34(1)：33～43
[141]Ducklow H W,Kirchman D L, Quinby H L, et al. Stocks and dynamics of bacteriaplankton carbon during the spring bloom in the eastern North Atlantic Ocean. Deep Sea Research II, 1993,40:245-263
[142]Lancelot C, Billen G.Activity of heterotrophic bacteria and its coupling to primary production during the spring phytoplankton bloom in the southern bight of the North Sea. Limnology and Oceanography,1984,29:721-730.
[143]Cho B C,Choi J K, Chung C S, et al. Uncoupling of bacteria and phytoplankton during a spring diatom bloom in the mouth of the Yellow Sea.Marine Ecology Progress Series,1994, 115:181-190.
[144]Campbell L, Carpenter E J,Montoya J P, et al.Picoplankton community structure within and outside a Trichodesmium bloom in the southwestern Pacific Ocean. Vie et Milieu,2005, 55(3-4):185-195.
[145]Iriarte A, Purdie D A. Size distribution of chlorophyll a biomass and primary production in a temperate estuary (Southampton Water)-the contribution of photosynthetic picoplankton. Marine Ecology Progress Series,1994,115:283-297
[146]张传松,王修林,石晓勇等.东海赤潮高发区营养盐时空分布特征及其与赤潮的关系环境科学,2007,28(11)：2416-2424
[147]Mackey K R M, RivlinT, Grossman A R, et al. Picophytoplankton responses to changing nutrient and light regimes during a bloom. Marine Biology,2009,156:1531-1546
[148]黄凌风,郭丰,黄邦钦等.初夏黄海中部和北部海洋鞭毛虫的分布特征及其影响因素,海洋学报,2003,25(Supp.)2：82-87
[149]Chiang K P, Lin C Y, Lee C H, et al. The coupling of oligotrich populations and hydrography in the East China Sea:spatial and temporal variation. Deep Sea Research,2003, 50:1279-1293
[150]刘瑞玉.胶州湾生态学和生物资源.北京：科学出版社,1992.2-3
[151]Noble R T, Fuhrman J A. Virus decay and its causes in coastal waters. Appl Environ Microbiol,1997,63:77-83
[152]肖天,焦念志,王荣.胶州湾蓝细菌、细菌的数量分布特点.见：董金海,焦念志,胶州湾生态学研究.北京：科学出版社,1995.118—124
[153]白晓歌,汪岷,梁彦韬,等.运用荧光显微技术分析北黄海夏季浮游病毒的分布.中国海洋大学学报,2008,38(4)：609-613
[154]Boehme J, Frischer M E, Jiang S C, et al. Viruses, bacterioplankton, and phytoplankton in the southeastern Gulf of Mexico:Distribution and contribution to oceanic DNA pools. Marine ecology progress series, Oldendorf,1993,97(1):1-10
[155]Weinbauer M G, Fuks D, Puskaric S, et al.Diel, seasonal, and depth-related variability of viruses and dissolved DNA in the northern Adriatic Sea. Microbial Ecology,1995,30(1): 25-41
[156]孙优善,孙鹤鲲,王学昌,等.胶州湾近岸海域水质状况调查与评价.海洋湖沼通报,2007，4：93-97
[157]Bettarel Y, Arfi R, Bouvier T, et al. Virioplankton distribution and activity in a tropical eutrophicated bay. Estuarine, Coastal and Shelf Science,2008,80(3):425-429
[158]翁学传,朱兰部,王一飞.胶州湾的自然环境：水文要素的结构和变化[A].刘瑞玉.胶州湾生态学和生物资源.北京：科学出版社,1992.20-38
[159]Wommack K E, Hill R T, Kessel M, et al.Distribution of viruses in the Chesapeake Bay. Applied and Environmental Microbiology,1992,58(9):2965-2970
[160]Auguet J C, Montanie H, Delmas D, et al.Dynamic of virioplankton abundance and its environmental control in the Charente estuary (France). Microbial ecology,2005,50(3): 337-349
[161]Corinaldesi C, Crevatin E, Del Negro P, et al. Large-scale spatial distribution of virioplankton in the Adriatic Sea:testing the trophic state control hypothesis. Applied and Environmental Microbiology,2003.69(5):2664-2673
[162]Jiang S C, Paul J H. Seasonal and diel abundance of viruses and occurrence of lysogeny/ bacteriocinogeny in the marine environment. Marine Ecology Progress Series. Oldendorf, 1994,104:163-172
[163]Brussaard C P D, Kempers R S,Kop A J, et al.Virus-like particles in a summer bloom of Emiliania huxleyi in the North Sea. Aquatic Microbial Ecology,1996,10:105-113
[164]Tomaru Y, Tarutani K, Yamaguchi M, et al. Quantitative and qualitative impacts of viral infection on a Heterosigma akashiwo (Raphidophyceae) bloom in Hiroshima Bay, Japan. Aquatic Microbial Ecology,2004,34(3):227-238
[165]Bratbak G, Egge J K, Heldal M.1993.Viral mortality of the marine alga Emiliania huxleyi (Haptophyceae) and termination of algal blooms. Marine Ecology Progress Series. Oldendorf 1993,93:39-48
[166]Jacquet S, Heldal M, Iglesias-Rodriguez D, et al.2002.Flow cytometric analysis of an Emiliania huxleyi bloom terminated by viral infection. Aquatic Microbial Ecology, 27:111-124
[167]Evans C G T, Archer S D, Jacquet S, et al.Direct measurements of the contribution of viral lysis and microzooplankton grazing to the decline of a Micromonas spp. population. Aquatic Microbial Ecology,2003,30:207-219
[168]杨小茹,郑天凌,苏建强,等.海洋病毒——一种新的,潜力巨大的赤潮防治工具.应用与环境生物学报,2005,11(5)：651-656
[169]Maranger R, Bird D F. Viral abundance in aquatic systems:a comparison between marine and fresh waters. Marine Ecology Progress Series. Oldendorf,1995,121:217-226
[170]Liu Y M, Zhang Q Y, Yuan X P, et al.Seasonal Variation of Virioplankton in a Eutrophic Shallow Lake. Hydrobiologia,2006,560(1):323-334
[171]刘艳鸣,张奇亚,袁秀平.武汉东湖浮游病毒的丰度及多样性.水生生物学报,2005,29(1)：1-6
[172]Maranger R, Bird D F, Juniper S K. Viral and bacterial dynamics in Arctic sea ice during the spring algal bloom near Resolute, N.W.T.,Canada. Marine Ecology Progress Series. Oldendorf 1994,111:121-127
[173]Tuomi P, Fagerbakke K M, Bratbak G, et al.Nutritional enrichment of a microbial community:The effects on activity, elemental composition, community structure and virus production. FEMS Microbiology Ecology,1995,16(2):123-134
[174]沈志良.长江干流营养盐通量的初步研究.海洋与湖沼,1997,28(5)：522～528
[175]全为民,沈新强,韩金娣,陈亚瞿.长江口及邻近水域富营养化现状及变化趋势的评价与分析.海洋环境科学,2005,24(3)：13～16
[176]Pavela J S, Ross J L, Chittended,M E Jr. Sharp reductions in abundances of fishes and benthic macroinvertebrates in the Gulf of Mexico off Texas associated with hypoxia. Northeast Gulf Science,1983,6:167-173
[177]Parker C A, O'Reilly J E. Oxygen depletion in Long Island Sound:A historical perspective. Estuaries,1991,14:248-264
[178]Limeburner R, Beardsley R C, Zhao J. Water Masses and Circulation in the East China Sea. Proceedings of International symposium on sedimentation on the continental shelf, with special reference to the East China Sea, April 12-16,1983,Hangzhou, China, Vol.1.Beijing: China Ocean Press,1983.285-294
[179]陈吉余,陈祥禄,杨启伦.上海海岸带和海涂资源综合调查报告.上海：上海科技出版社,1988.114～116
[180]李道季,张经,黄大吉等.长江口外氧的亏损.中国科学,2002,32(8)：686～694
[181]石晓勇,陆茸,张传松,王修林.长江口邻近海域溶解氧分布特征及主要影响因素.中国海洋大学学报(自然科学版),2006,36(02)：287～290
[182]张莹莹,张经,吴莹,朱卓毅.长江口溶解氧的分布特征及影响因素研究.环境科学,2007,28(8)：1649-1654
[183]刘才才,项凌云,张昊飞,王金辉.长江口异养细菌生态分布特征及其与环境因子的关系.海洋环境科学,2009,28(A01)：1-4
[184]Wheeler P A, Kirchman D L. Utilization of inorganic and organic nitrogen by bacteria inmarine systems. Limnol Oceanogr,1986,31:998-1009
[185]Richard B R. Inorganic nutrient limitation of oceanic baeterioplankton. Liminol Oceanogr, 1997,42(4):730-740
[186]Kirchman D L, Wheeler P A. Uptake of ammonium and nitrate by heteotrophic bacteria and phytoplankton in the sub-Arctic Pacific. Deep Sea Research(Ⅰ),1998,45:347-365
[187]陈水土,阮五崎.台湾海峡海域表现耗氧量之研究.海洋与湖沼,1996,27(3)：287-295
[188]Rowe G T. Seasonal hypoxia in the bottom water off the Mississippi River delta. Journal of Environmental Quality,2001,30:281-290
[189]Paoli A, Celussi M, Valeri A, et al.Picocyanobacteria in Adriatic transitional environments. Estuarine,Coastal and Shelf Science,2005,75:13-20
[190]Phlips E J, Zeman C, Hansen P. Growth photosynthesis, nitrogen fixation and carbohydrate production by a unicellular cyanobacterium, Synechococcus sp. (Cyanophyta).Journal of Applied Phycology,1989,1,137-145
[191]Dagg M J,Ammerman J W, Amon R M W, et al.A Review of Water Column Processes Influencing Hypoxia in the Northern Gulf of Mexico.Estuaries and Coasts,2007,30(5): 735-752
[192]Fontes M L S,Abreu P C.Spatiotemporal Variation of Bacterial Assemblages in a Shallow Subtropical Coastal Lagoon in Southern Brazil. Microb Ecol,2009,58:140-152
[193]刘敏,王子峰,朱开玲等.应用PCR-DGGE技术分析长江口低氧区的细菌群落组成.高技术通讯,2008,18(6)：649-656