自然海区混合营养浮游鞭毛虫和纤毛虫生态学
Ecology of Mixotrophic Planktonic Flagellate and Ciliate in the Sea
DOI: 10.12677/AMS.2016.33011, PDF, HTML, XML,  被引量 下载: 1,942  浏览: 4,931  国家自然科学基金支持
作者: 王超锋, 张武昌, 赵 丽, 赵 苑, 肖 天:中国科学院海洋研究所海洋生态与环境重点实验室,山东 青岛
关键词: 混合营养鞭毛虫纤毛虫微型浮游动物Mixotrophy Ciliate Flagellate Microzooplankton
摘要: 海洋浮游生物的混合营养是指一种生物既能进行光合自养,又能进行摄食营养。混合营养使得这种生物既是生产者,又是消费者,因此对传统的食物链和生态模型研究是个挑战。混合营养的海洋浮游生物在自然海区的分布是理解它们在自然海区生态功能的基础,本文综述了自然海区混合营养浮游生物的重要类群--鞭毛虫和纤毛虫的计数方法和结果。混合营养鞭毛虫在不同海区占含色素鞭毛虫的比例多小于50%。混合营养寡毛类纤毛虫的种类大约十余种,其中球果螺体虫(Laboea strobila)是混合营养纤毛虫的重要类群,多分布在温带和极地海区;不同海区混合营养纤毛虫丰度和生物量所占比例多低于70%。总体来看,混合营养型海洋浮游生物生态特征在自然海区的研究较少,我国在这方面的研究才刚起步。
Abstract: Mixotrophic microplankton are those that can carry out photosynthesis and ingestive behavior simultaneously. Mixotrophy makes mixotrophic microzooplankton be producer and consumer. Thus it’s a challenge for traditional phytoplankton-zooplankton dichotomy and the study of eco-logical model. The distribution of mixotrophic microzooplankton in the sea is the foundation to understand their ecological function. In this study, we summarized the counting method and the result of mixotrophic flagellate and ciliate in the sea in previous studies. For mixotrophic flagellate, its percentage of abundance in flagellate with plastid in different sea regions was mostly less than 50%. Taxonomically, more than ten mixotrophic ciliates species were identified up to now. As the most easily identified mixotrophic ciliates, Laboea strobila mainly appeared in temperate zone and polar sea region. Percentage of mixotrophic ciliates abundance and biomass in different sea regions was mostly less than 70%. Overall, the research about taxonomy and ecology of mixotrophic flagellate and ciliate in natural sea region was scarcely carried out. In China the knowledge gap of mixotrophic microzooplankton has been indentified and is being focused on in some related research.
文章引用:王超锋, 张武昌, 赵丽, 赵苑, 肖天. 自然海区混合营养浮游鞭毛虫和纤毛虫生态学[J]. 海洋科学前沿, 2016, 3(3): 73-83. http://dx.doi.org/10.12677/AMS.2016.33011

参考文献

[1] Lewin, J.C. (1953) Heterotrophy in Diatoms. Microbiology, 9, 305-313.
http://dx.doi.org/10.1099/00221287-9-2-305
[2] Eiler, A. (2006) Evidence for the Ubiquity of Mixotrophic Bacteria in the Upper Ocean: Implications and Consequences. Applied and Environmental Microbiology, 72, 7431-7437.
http://dx.doi.org/10.1128/AEM.01559-06
[3] Flynn, K.J., Stoecker, D.K., Mitra, A., et al. (2012) Misuse of the Phy-toplankton-Zooplankton Dichotomy: The Need to Assign Organisms as Mixotrophs within Plankton Functional Types. Journal of Plankton Research, 35, 3-11.
http://dx.doi.org/10.1093/plankt/fbs062
[4] Stoecker, D.K. (1998) Conceptual Models of Mixotrophy in Planktonic Protists and Some Ecological and Evolutionary Implications. European Journal of Protistology, 34, 281-290.
http://dx.doi.org/10.1016/S0932-4739(98)80055-2
[5] Caron, D.A. (2000) Symbiosis and Mixotrophy among Pelagic Microorganisms. In: Kirchmann, D.L., Ed., Microbial Ecology of the Oceans, Wiley Series in Ecological and Applied Mi-crobiology, Liss/Wiley, New York, 495-523.
[6] Sanders, R.W. and Porter, K.G. (1987) Phagotrophic Phytoflagellates. Advances in Microbial Ecology, 10, 167-192.
http://dx.doi.org/10.1007/978-1-4684-5409-3_5
[7] Kofoid, C.A. and Swezy, O. (1921) The Free-Living Unarmored Dinoflagellata. Memorial University of California, 5, 1-562.
[8] Porter, K.G. (1988) Phagotrophic Phytoflagellates in Mi-crobial Food Webs. Hydrobiologia, 159, 89-97.
http://dx.doi.org/10.1007/BF00007370
[9] Fenchel, T. (1982) Ecology of Heterotrophic Microflagellates. IV. Quanti-tative Occurrence and Importance as Bacterial Consumers. Marine Ecology Progress Series, 9, 35-42.
http://dx.doi.org/10.3354/meps009035
[10] Sherr, E.B. and Sherr, B.F. (1983) Double-Staining Epifluorescence Tech-nique to Assess Frequency of Dividing Cells and Bacteriovory in Natural-Populations of Heterotrophic Microprotozoa. Applied and Environmental Microbiology, 46, 1388-1393.
[11] Davis, P.G. and Sieburth, J.M. (1984) Differentiation and Characterization of Individual Phototrophic and Heterotrophic Microflagellates by Sequential Epifluorescence and Elec-tron-Microscopy. Transactions of the American Microscopical Society, 103, 221-227.
http://dx.doi.org/10.2307/3226182
[12] Porter, K.G., Sherr, E.B., Sherr, B.F., et al. (1985) Protozoa in Planktonic Food Webs. Journal of Protozoology, 32, 409-415.
http://dx.doi.org/10.1111/j.1550-7408.1985.tb04036.x
[13] Bird, D.F. and Kalff, J. (1986) Bacterial Grazing by Planktonic Lake Algae. Science, 231, 493-495.
http://dx.doi.org/10.1126/science.231.4737.493
[14] Estep, K.W., Davis, P.G., Keller, M.D., et al. (1986) How Important Are Oceanic Algal Nanoflagellates in Bacterivory? Limnology and Oceanography, 31, 646-650.
http://dx.doi.org/10.4319/lo.1986.31.3.0646
[15] Ballantine, D. (1953) Comparison of the Different Methods of Esti-mating Nanoplankton. Journal of the Marine Biological Association of the United Kingdom, 32, 129-147.
http://dx.doi.org/10.1017/S0025315400011462
[16] Pratt, D.M. (1959) The Phytoplankton of Narragansett Bay. Lim-nology and Oceanography, 4, 425-440.
http://dx.doi.org/10.4319/lo.1959.4.4.0425
[17] Booth, B.C. (1975) Growth of Oceanic Phytoplankton in Enrichment Cultures. Limnology and Oceanography, 20, 865- 869.
http://dx.doi.org/10.4319/lo.1975.20.5.0865
[18] Beers, J.R., Reid, F.M.H. and Stewart, G.L. (1975) Microplankton of the North Pacific Central Gyre. Population Structure and Abundance, June 1973. Internationale Revue der gesamten Hydrobiologie, 60, 607-638.
[19] Beers, J.R., Reid, F.M.H. and Stewart, G.L. (1980) Microplankton Population Structure in Southern California Nearshore Waters in Late Spring. Marine Biology, 60, 209-226.
http://dx.doi.org/10.1007/BF00389164
[20] Davis, P.G. and Sieburth, J.M. (1982) Differentiation of Phototrophic and Heterotrophic Nanoplankton Populations in Marine Waters by Epifluorescence Microscopy. Annales de l Institut Océanographique, 58, 249-260.
[21] Arenovski, A.L., Lim, E.L. and Caron, D.A. (1995) Mixotrophic Nanoplankton in Oligotrophic Surface Waters of the Sargasso Sea May Employ Phagotrophy to Obtain Major Nutrients. Journal of Plankton Research, 17, 801-820.
http://dx.doi.org/10.1093/plankt/17.4.801
[22] Chan, Y.F., Tsai, A.Y., Chiang, G.P. and Hsieh, C. (2009) Pigmented Nanoflagellates Grazing on Synechococcus: Seasonal Variations and Effect of Flagellate Size in the Coastal Ecosystem of Subtropical Western Pacific. Microbial Ecology, 58, 548-557.
http://dx.doi.org/10.1007/s00248-009-9569-x
[23] Christaki, U., Van Wambeke, F. and Dolan, J.R. (1999) Nanoflagel-lates (Mixotrophs, Heterotrophs and Autotrophs) in the Oligotrophic Eastern Mediterranean: Standing Stocks, Bacterivory and Relationships with Bacterial Production. Marine Ecology Progress Series, 181, 297-307.
http://dx.doi.org/10.3354/meps181297
[24] Czypionka, T., Vargas, C.A. and Silva, N. (2011) Importance of Mixotrophic Nanoplankton in Aysen Fjord (Southern Chile) during Austral Winter. Continental Shelf Research, 31, 216-224.
http://dx.doi.org/10.1016/j.csr.2010.06.014
[25] Havskum, H. and Riemann, B. (1996) Ecological Importance of Bac-terivorous, Pigmented Flagellates (Mixotrophs) in the Bay of Aarhus, Denmark. Marine Ecology Progress Series, 137, 251-263.
http://dx.doi.org/10.3354/meps137251
[26] Moorthi, S.D., Caron, D.A., Gast, R.J. and Sanders, R.W. (2009) Mixotrophy: A Widespread and Important Ecological Strategy for Planktonic and Sea-Ice Nanoflagellates in the Ross Sea, Antarctica. Aquatic Microbial Ecology, 54, 269- 277.
http://dx.doi.org/10.3354/ame01276
[27] Sanders, R.W., Berginger, U.G., Lim, E.L., Kemp, P.F. and Caron, D.A. (2000) Heterotrophic and Mixotrophic Nanoplankton Predation on Picoplankton in the Sargasso Sea and on Georges Bank. Marine Ecology Progress Series, 192, 103-118.
http://dx.doi.org/10.3354/meps192103
[28] Unrein, F., Massana, R., Alonso-Saez, L. and Gasol, J.M. (2007) Significant Year-Round Effect of Small Mixotrophic Flagellates on Bacterioplankton in an Oligotrophic Coastal System. Limnology and Oceanography, 52, 456-469.
http://dx.doi.org/10.4319/lo.2007.52.1.0456
[29] Vargas, C.A., Contreras, P.Y. and Iriarte, J.L. (2011) Relative Im-portance of Phototrophic, Heterotrophic, and Mixotrophic Nanoflagellates in the Microbial Food Web of a River-Influenced Coastal Upwelling Area. Aquatic Microbial Ecology, 65, 233-248.
http://dx.doi.org/10.3354/ame01551
[30] Norris, R.E. (1967) Algal Consortisms in Marine Plankton. In: Krishnamurthy, V., Ed., Pro Seminar on Sea, Salt and Plants, Central Salt and Marine Chemicals Research Institute, Bhavnagar, 178-189.
[31] Blackbourn, D.J., Taylor, F.J.R. and Blackbourn, J. (1973) Foreign Organelle Retention by Ciliates. Journal of Protozoology, 20, 286-288.
http://dx.doi.org/10.1111/j.1550-7408.1973.tb00877.x
[32] Laval-Peuto, M. and Febvre, M. (1986) On Plastid Symbi-osis in Tontonia appendiculariformis (Ciliophora, Oligotrichina). Biosystems, 19, 137-158.
http://dx.doi.org/10.1016/0303-2647(86)90026-2
[33] McManus, G.B. and Fuhrman, J.A. (1986) Bacterivory in Sea-water Studied with the Use of Inert Fluorescent Particles. Limnology and Oceanography, 31, 420-426.
http://dx.doi.org/10.4319/lo.1986.31.2.0420
[34] Jonsson, P.R. (1987) Photosynthetic Assimilation of Inorganic Carbon in Marine Oligotrich Ciliates (Ciliophora, Oligotrichina). Marine Microbial Food Webs, 2, 55-67.
[35] Stoecker, D.K., Michaels, A.E. and Davis, L.H. (1987) Large Proportion of Marine Planktonic Ciliates Found to Contain Functional Chloro-plasts. Nature, 326, 790-792.
http://dx.doi.org/10.1038/326790a0
[36] Laval-Peuto, M. and Rassoulzadegan, F. (1988) Autofluorescence of Marine Planktonic Oligotrichina and Other Ciliates. Hydrobiologia, 159, 99-110.
http://dx.doi.org/10.1007/BF00007371
[37] Verity, P.G. and Vernet, M. (1992) Microzooplankton Grazing, Pigments, and Composition of Plankton Communities during Late Spring in Two Norwegian Fjords. Sarsia, 77, 263-274.
http://dx.doi.org/10.1080/00364827.1992.10413511
[38] Stoecker, D.K., Taniguchi, A. and Michaels, A.E. (1989) Abundance of Autotrophic, Mixotrophic and Heterotrophic Planktonic Ciliates in Shelf and Slope Waters. Marine Ecology Progress Series, 50, 241-254.
http://dx.doi.org/10.3354/meps050241
[39] Stoecker, D.K. (1991) Mixotrophy in Marine Planktonic Ciliates: Physio-logical and Ecological Aspects of Plastid-Re- tention by Oligotrichs. In: Reid, P.C., Turley, C. and Burkill, P.H., Eds., Pro-tozoa and Their Role in Marine Processes, Springer, Berlin, 161-179.
http://dx.doi.org/10.1007/978-3-642-73181-5_12
[40] Stoecker, D.K., Sieracki, M.E., Verity, P.G., et al. (1994) Nano-plankton and Protozoan Microzooplankton during the JGOFS North Atlantic Bloom Experiment: 1989 and 1990. Journal of the Marine Biological Association of the United Kingdom, 74, 427-443.
http://dx.doi.org/10.1017/S0025315400039448
[41] Stoecker, D.K., Gustafson, D.E. and Verity, P.G. (1996) Micro- and Mesoprotozooplankton at 140 Degree W in the Equatorial Pacific: Heterotrophs and Mixotrophs. Aquatic Microbial Ecology, 10, 273-282.
http://dx.doi.org/10.3354/ame010273
[42] Modigh. M. (2001) Seasonal Variations of Photosynthetic Ciliates at a Med-iterranean Coastal Site. Aquatic Microbial Ecology, 23, 163-175.
http://dx.doi.org/10.3354/ame023163
[43] Stoecker, D.K., Silver, M.W., Michaels, A.E. and Davis, L.H. (1988) Obligate Mixotrophy in Laboea strobila, a Ciliate Which Retains Chloroplasts. Marine Biology, 99, 415-423.
http://dx.doi.org/10.1007/BF02112135
[44] Pitta, P. and Giannakourou, A. (2000) Planktonic Ciliates in the Oligotrophic Eastern Mediterranean: Vertical, Spatial Distribution and Mixotrophy. Marine Ecology Progress Series, 194, 269-282.
http://dx.doi.org/10.3354/meps194269
[45] McManus, G.B., Zhang, H. and Lin, S. (2004) Marine Planktonic Ciliates That Prey on Macroalgae and Enslave Their Chloroplasts. Limnology and Oceanography, 49, 308-313.
http://dx.doi.org/10.4319/lo.2004.49.1.0308
[46] Dolan, J.R. (1991) Guilds of Ciliate Microzooplankton in the Chesapeake Bay. Estuarine, Coastal and Shelf Science, 33, 137-152.
http://dx.doi.org/10.1016/0272-7714(91)90003-T
[47] Dolan, J.R. and Maarasse, C. (1995) Planktonic Ciliate Distribu-tion Relative to a Deep Chlorophyll Maximum: Catalan Sea, NW Mediterranean, June 1993. Deep Sea Research Part I, 42, 1965-1987.
http://dx.doi.org/10.1016/0967-0637(95)00092-5
[48] Dolan, J.R., Vidussi, F. and Claustre, H. (1999) Planktonic Cili-ates in the Mediterranean Sea: Longitudinal Trends. Deep Sea Research Part I, 46, 2025-2039.
http://dx.doi.org/10.1016/S0967-0637(99)00043-6
[49] Rassoulzadegan, F., Laval-Peuto, M. and Sheldon, R.W. (1988) Partitioning of the Food Ration of Marine Ciliates between Pico- and Nanoplankton. Hydrobiologia, 159, 75-88.
http://dx.doi.org/10.1007/BF00007369
[50] Bernard, C. and Rassoulzadegan, F. (1990) Bacteria or Microflagellates as a Major Food Source for Marine Ciliates: Possible Implications for the Microzooplankton. Marine Ecology Progress Series, 64, 147-155.
http://dx.doi.org/10.3354/meps064147
[51] Bernard, C. and Rassoulazdegan, F. (1994) Seasonal Variations of Mixo-trophic Ciliates in the Northwest Mediterranean Sea. Marine Ecology Progress Series, 108, 295-301.
http://dx.doi.org/10.3354/meps108295
[52] Martin, A.J. and Montagnes, D.J. (1993) Winter Ciliates in a British Co-lumbian Fjord: Six New Species and an Analysis of Ciliate Putative Prey. Journal of Eukaryotic Microbiology, 40, 535-549.
http://dx.doi.org/10.1111/j.1550-7408.1993.tb06105.x
[53] Putt, M. (1991) Abundance, Chlorophyll Content and Pho-tosynthetic Rates of Ciliates in the Nordic Seas during Summer. Deep Sea Research Part A, 37, 1713-1731.
http://dx.doi.org/10.1016/0198-0149(90)90073-5
[54] Lynn, D.H., Montagnes, D.J.S., Dale, T., Gilrona, G.L. and Strom, S.L. (1991) A Reassessment of the Genus Strombidinopsis (Ciliophora, Choreotrichida) with Descriptions of Four New Planktonic Species and Remarks in Its Taxonomy and Phylogeny. Journal of the Marine Biological Association of the United Kingdom, 71, 597-612.
http://dx.doi.org/10.1017/S0025315400053170
[55] Putt, M. (1990) Metabolism of Photosynthate in the Chloro-plast-Retaining Ciliate Laboea strobila. Marine Ecology Progress Series, 60, 271-282.
http://dx.doi.org/10.3354/meps060271
[56] Booth, B.C., Lewin, J. and Postel, J.R. (1993) Temporal Variation in the Structure of Autotrophic and Heterotrophic Communities in the Subarctic Pacific. Progress in Oceanography, 32, 57-99.
http://dx.doi.org/10.1016/0079-6611(93)90009-3
[57] Gowing, M.M. and Garrison, D.L. (1992) Abundance and Feed-ing Ecology of Larger Protozooplankton in the Ice Edge Zone of the Weddell and Scotia Seas during the Austral Winter. Deep Sea Research Part A, 39, 893-919.
http://dx.doi.org/10.1016/0198-0149(92)90128-G
[58] Perez, M.T., Dolan, J.R., Vidussi, F. and Fukai, E. (2000) Diel Vertical Distribution of Planktonic Ciliates within the Surface Layer of the NW Mediterranean (May 1995). Deep Sea Research Part I, 47, 479-503.
http://dx.doi.org/10.1016/S0967-0637(99)00099-0
[59] Paranjape, M.A. (1987) The Seasonal Cycles and Vertical Dis-tribution of Tintinnines in Bedford Basin, Nova Scotia, Canada. Canadian Journal of Zoology, 65, 41-48.
http://dx.doi.org/10.1139/z87-007
[60] James, M.R., Hall, J.A. and Barrett D.P. (1996) Grazing by Protozoa in Marine Coastal and Oceanic Ecosystems off New Zealand. New Zealand Journal of Marine and Freshwater Research, 30, 313-324.
http://dx.doi.org/10.1080/00288330.1996.9516718
[61] Dale, T. and Dahl, E. (1987) Mass Occurrence of Planktonic Oligotrichous Ciliates in a Bay in Southern Norway. Journal of Plankton Research, 9, 871-879.
http://dx.doi.org/10.1093/plankt/9.5.871
[62] Flynn, J., Stoecker, K., Mitra, A., et al. (2013) Misuse of the Phyto-plankton-Zooplankton Dichotomy: The Need to Assign Organisms as Mixotrophs within Plankton Functional Types. Journal of Plankton Research, 35, 3-11.
http://dx.doi.org/10.1093/plankt/fbs062
[63] 孙松, 唐启升. 海洋生态学研究现状与发展趋势[J]. 海洋与湖沼, 2002(5): 1-9.
[64] 戴聪杰, 何剑锋, 王桂忠, 李少菁. 混合营养型浮游生物生态学研究进展[J]. 生态学报, 2005, 25(9): 2399-2405.
[65] Zhang, W.C., Xu, K.D., Wan, R., et al. (2002) Spatial Distribution of Ciliates, Copepod Nauplii and Eggs, Engraulis japonicus Post-Larvae and Microzooplankton Herbivorous Activity in the Yellow Sea, China. Aquatic Microbial Ecology, 27, 249-259.
http://dx.doi.org/10.3354/ame027249