外源溶解性有机碳对抚仙湖甲壳类浮游动物碳源的贡献
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摘要
外源溶解性有机碳(DOC)是湖泊碳库的重要组成部分,关于外源DOC对浮游动物的贡献及途径需要深入研究.本研究在抚仙湖受控实验中添加13C标记的葡萄糖,通过分析样品中浮游植物与浮游动物的种类、数量、磷脂脂肪酸生物标志物及其稳定同位素特征,研究外源DOC对湖泊甲壳类浮游动物碳源的贡献比例及其变化.结果表明:细菌、甲壳类浮游动物(象鼻溞)的δ~(13)C值在加入葡萄糖后分别从-16.28‰和-23.88‰快速增加到5408.25‰和1974.7‰,而藻类磷脂脂肪酸(C18∶2ω6)δ~(13)C值从-27.07‰增加到342.44‰,增长的幅度表明添加的葡萄糖首先被细菌和浮游动物快速利用,而藻类只利用了一小部分.同时细菌、颗粒性有机物(POM)和浮游动物的δ~(13)C值在第1 d急剧增加,细菌的δ~(13)C值远大于浮游动物和POM的δ~(13)C值,之后细菌和POM的δ~(13)C值开始下降,但浮游动物的δ~(13)C值却仍在缓慢增加,进一步表明了DOC进入湖泊后首先被细菌吸收利用,而细菌吸收DOC后通过自身代谢作用形成细胞颗粒,浮游甲壳类动物通过摄食细胞颗粒来获得外源DOC.
Allochthonous dissolved organic carbon( DOC) is of importance in lake carbon stock,therefore,the contribution and pathways of allochthonous DOC to planktonic crustaceans deserves further study. A controlled experiment was performed through adding glucose labeled with δ~(13)C in water from Lake Fuxian. By analyzing the samples of phytoplankton and zooplankton species,quantity and phospholipid fatty acid( PLFA) biomarkers and their carbon stable isotope,the contribution of allochthonous DOC to crustacean zooplankton carbon source was investigated. The results showed that after adding glucose,δ~(13)C values of bacteria and planktonic crustaceans( Bosmina) increased rapidly from-16. 28‰ to 5408. 25‰ and-23. 88‰ to 1974. 7‰,respectively,whereas the δ~(13)C value of chlorophyceae increased from-27. 07‰ to 342. 44‰. The growth rate showed that bacteria and zooplankton firstly utilized the adding glucose. The δ~(13)C values of bacteria,particulate organic matter( POM) and zooplankton increased sharply at the first day,moreover,the δ~(13)C value of bacteria is much higher than that of zooplankton and POM. After the first day,δ~(13)C values of bacteria and POM decreased,by contrast,zooplankton δ~(13)C value increased slowly. This suggests that allochthonous DOC is firstly utilized by bacteria and form cell particles via the metabolism of bacteria. Subsequently,planktonic crustaceans can utilize allochthonous DOC by grazing cell particles.
Allochthonous dissolved organic carbon( DOC) is of importance in lake carbon stock,therefore,the contribution and pathways of allochthonous DOC to planktonic crustaceans deserves further study. A controlled experiment was performed through adding glucose labeled with δ~(13)C in water from Lake Fuxian. By analyzing the samples of phytoplankton and zooplankton species,quantity and phospholipid fatty acid( PLFA) biomarkers and their carbon stable isotope,the contribution of allochthonous DOC to crustacean zooplankton carbon source was investigated. The results showed that after adding glucose,δ~(13)C values of bacteria and planktonic crustaceans( Bosmina) increased rapidly from-16. 28‰ to 5408. 25‰ and-23. 88‰ to 1974. 7‰,respectively,whereas the δ~(13)C value of chlorophyceae increased from-27. 07‰ to 342. 44‰. The growth rate showed that bacteria and zooplankton firstly utilized the adding glucose. The δ~(13)C values of bacteria,particulate organic matter( POM) and zooplankton increased sharply at the first day,moreover,the δ~(13)C value of bacteria is much higher than that of zooplankton and POM. After the first day,δ~(13)C values of bacteria and POM decreased,by contrast,zooplankton δ~(13)C value increased slowly. This suggests that allochthonous DOC is firstly utilized by bacteria and form cell particles via the metabolism of bacteria. Subsequently,planktonic crustaceans can utilize allochthonous DOC by grazing cell particles.
引文
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[13]Nanjing Institute of Geography and Limnology,Chinese Academy of Sciences ed.Lake Fuxian.Beijing:Ocean Press,1990.[中国科学院南京地理与湖泊研究所.抚仙湖.北京:海洋出版社,1990.]
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[15]Pan Jizheng,Xiong Fei,Li Wenchao et al.Spatial-temporal dynamic changes of the water transparency and their influencing factors in Lake Fuxian,Yunnan Province.J Lake Sci,2008,20(5):681-686.DOI:10.18307/2008.0519.[潘继征,熊飞,李文朝等.云南抚仙湖透明度的时空变化及影响因子分析.湖泊科学,2008,20(5):681-686.]
[16]Lei Ligai,Ma Xiaozhen,Wei Fuxiang et al.Research progress of determination of total nitrogen and total phosphorus in seawater.Hebei Journal of Industrial Science and Technology,2011,28(1):72-76.[雷立改,马晓珍,魏福祥等.水中总氮、总磷测定方法的研究进展.河北工业科技,2011,28(1):72-76.]
[17]Pápista,cs,B9ddi B.Chlorophyll-a determination with ethanol—A critical test.Hydrobiologi,2002,485(123):191-198.
[18]Uterm9hl H.Neue wege in der quantitativan erfassung des planktons(Mit besondere Berü-cksichtigung des Ultraplanktons).Verh Int Verein Theor Angew Limnol,1931,5:567-595.
[19]Zhang Zongshe,Huang Xiangfei eds.Research methods of freshwater plankton.Beijing:Science Press,1991.[章宗涉,黄祥飞.淡水浮游生物研究方法.北京:科学出版社,1991.]
[20]Hu Hongjun,Wei Yinxin eds.The freshwater algae of China—Systematics,taxonomy and ecology.Beijing:Science Press,2006.[胡鸿钧,魏印心.中国淡水藻类——系统、分类及生态.北京:科学出版社,2006.]
[21]Research Group of Carcinology,Institute of Zoology,Academia Sinica ed.Fauna sinica crustacea·freshwater copepods Beijing:Science Press,1979.[中国科学院动物研究所甲壳动物研究组.中国动物志甲壳纲:淡水桡足类.北京:科学出版社,1979.]
[22]Jiang Xiezhi,Du Nanshan eds.Fauna Sinica Crustacea·Freshwater Cladocera.Beijing:Science Press,1979.[蒋燮治,堵南山.中国动物志甲壳纲:淡水枝角类.北京:科学出版社,1979.]
[23]Bligh EG,Dyer WJ.A rapid method of total lipid extraction and purification.Canadian Journal of Biochemistry and Physiology,1959,37(8):911-917.
[24]Van den Meersche KK,Van Rijswijk P,Karline S et al.Autochthonous and allochthonous contributions to mesozooplankton diet in a tidal river and estuary:Integrating carbon isotope and fatty acid constraints.Limnology and Oceanography,2009,54(1):62-74.
[25]Zhao Dayong,Yan Wenming,Feng Jingwei et al.Applications of phospholipid fatty acid analysis in the microbial ecology studies of lake sediment.Chemistry&Bioengineering,2009,26(12):17-20.[赵大勇,燕文明,冯景伟等.磷脂脂肪酸分析在湖泊沉积物微生物生态学研究中的应用.化学与生物工程,2009,26(12):17-20.]
[26]Rajendran N,Matsuda O,Imamura N et al.Variation in microbial biomass and community structure in sediments of eutrophic bays as determined by phospholipid ester-linked fatty acids.Applied and Environmental Microbiology,1992,58(2):562-571.
[27]Ge Yuan,He Jizheng,Zheng Yuanming et al.Stable isotope probing and its applications in microbicalecology.Acta Ecologica Sinica,2006,26(5):1574-1582.[葛源,贺纪正,郑袁明等.稳定性同位素探测技术在微生物生态学研究中的应用.生态学报,2006,26(5):1574-1582.]
[28]Kritzberg ES,Cole J,Pace ML et al.Bacteria growth on allochthonous carbon in humic and nutrient-riched lakes:Results from whole-lake13C addition experiments.Ecosystems,2006,9(3):489-499.
[29]Azam F,Fenchel T,Field J et al.The ecological role of water-column microbes in the sea.Marine Ecology Progress Series,1983,10(3):257-263.
[30]Jansson M,Persson L,De Roos AM et al.Terrestrial carbon and intraspecific size-variation shape lake ecosystems.Trends in Ecology and Evolution,2007,22(6):316-322.
[31]Carignan R,Planas D,Vis C.Planktonic production and respiration in oligotrophic shield lakes.Limnology and Oceanography,2000,45(1):189-199.
[32]Tang Yali,Cheng Dongmei,Liu Zhengwen et al.Differntial support of lake food webs by allochthonous organic carbon.Ecological Science,2014,33(1):161-165.[唐雅丽,程冬梅,刘正文等.外源性有机碳对淡水生态系统食物网的贡献.生态学报,2014,33(1):161-165.]
[33]Institute of Animal,Chinese Academy of Science,crustaceans research group ed.Chinese animal.Beijing:Science Press,1979.[中国科学院中国动物志委员会.中国动物志.北京:科学出版社,1979.]
[34]Banta AM.Population density as related to sexand to evolution in Cladocera.The American Naturalist,1937,71(732):34-49.
[35]Zhang Guangtao,Li Chaolun,Sun Song et al.Feeding habit of Calanussinicus(Crustacea:Copepoda)during spring and autumn in the Bohai Sea studied with herbivore index.Scientia Marina,2006,70(3):381-388.
[36]Hama T,Yanagi K.Production and neutral aldose composition of dissolved carbohydrates excreted by natural marine phytoplankton populations.Limnology and Oceanography,2001,46(8):1945-1955.
[37]Biddanda B,Benner R.Carbon,nitrogen,and carbohydrate fluxes during the production of particulate and dissolved organic matter by marine phytoplankton.Limnology and Oceanography,1997,42(3):506-518.
[38]Pan Jizheng,Xiong Fei,Li Wenchao et al.Structure distribution and it’s impact factors of phytoplankton community in Fuxian Lake.Acta Ecological Sinica,2009,29(10):5376-5385.[潘继征,熊飞,李文朝等.抚仙湖浮游植物群落结构、分布及其影响因子.生态学报,2009,29(10):5376-5385.]
[39]Berggren M,Strom L,Laudon H et al.Lake secondary production fueled by rapid transfer of low molecular weight organic carbon from terrestrial sources to aquatic consumers.Ecology Letters,2010,13(7):870-880.
[40]Yokokawa T,Nagata T.Linking bacterial community structure to carbon fluxes in marine environments.Journal of Oceanography,2010,66(1):1-12.
[41]Algesten G,Sobek S,Bergstro MAK et al.Role of lakes for organic carbon cycling in the boreal zone.Global Change Biology,2005,10(1):141-147.
[2]Wetzel RG.Limnology:Lake and river ecosystems.Eos Transactions American Geophysical Union,2001,21(2):1-9.
[3]Grey J,Jones RI,Sleep D.Stable isotope analysis of the origins of zooplankton carbon in lakes of differing trophic state.Oecologia,2000,123(2):232-240.
[4]Pace ML,Cole JJ,Carpenter SR et al.Whole-lake carbon-13 additions reveal terrestrial support of aquatic food webs.Nature,2004,427(6971):240-243.
[5]Cole JJ,Carpenter SR,Pace ML et al.Differential support of lake food webs by three types of terrestrial organic carbon.Ecology Letters,2006,9(5):558-568.
[6]Fry B.Stable isotope ecology.New York:Springer,2006:204-212.
[7]Boschker HTS,Middelburg JJ.Stable isotopes and biomarkers in microbial ecology.FEMS Microbiology Ecology,2002,40(2):85-95.
[8]Boschker HTS,Kromkamp JC,Middelburg JJ.Biomarker and carbon isotopic constraints on bacterial and algal community structure and functioning in a turbid,tidal estuary.Limnology and Oceanography,2005,50(1):70-80.
[9]Liu Zhanfeng,Li Liangjun,Ren Zheng et al.The advance in isotope labeled glucose.Nuclear Techniques,2013,36(1):103021-103027.[刘占峰,李良君,任征等.同位素标记葡萄糖的研究进展.核技术,2013,36(1):103021-103027.]
[10]Ziegler SE,White PM,Wolf D et al.Tracing the fate and recycling of 13C-labeled glucose in soil.Soil Science,2005,170(10):767-778.
[11]Shi Ningning,Xu Hong,Yao Jun et al.Investigation of metabolic routes toγ-Poly(glutamic acid)by13C-labeled glucose as medium carbon source.Chinese Journal of Process Engineering,2007,7(1):145-148.[石宁宁,徐虹,姚俊等.利用13C标记葡萄糖分析γ-聚谷氨酸的代谢途径.过程工程学报,2007,7(1):145-148.]
[12]Wang Lin,Tang Jinyan,Liu Yu et al.Study on the vulnerability of environment in Fuxian Lake.Hubei Agricultural Sciences,2012,51(14):2965-2970.[王林,唐金焰,刘宇等.抚仙湖生态环境脆弱性分析研究.湖北农业科学,2012,51(14):2965-2970.]
[13]Nanjing Institute of Geography and Limnology,Chinese Academy of Sciences ed.Lake Fuxian.Beijing:Ocean Press,1990.[中国科学院南京地理与湖泊研究所.抚仙湖.北京:海洋出版社,1990.]
[14]Gao Wei,Chen Yan,Xu Min et al.Trend and driving factors of water quality change in Lake Fuxian(1980-2011).J Lake Sci,2013,25(5):635-642.DOI:10.18307/2013.0503.[高伟,陈岩,徐敏等.抚仙湖水质变化(19802011)趋势和驱动力分析.湖泊科学,2013,25(5):635-642.]
[15]Pan Jizheng,Xiong Fei,Li Wenchao et al.Spatial-temporal dynamic changes of the water transparency and their influencing factors in Lake Fuxian,Yunnan Province.J Lake Sci,2008,20(5):681-686.DOI:10.18307/2008.0519.[潘继征,熊飞,李文朝等.云南抚仙湖透明度的时空变化及影响因子分析.湖泊科学,2008,20(5):681-686.]
[16]Lei Ligai,Ma Xiaozhen,Wei Fuxiang et al.Research progress of determination of total nitrogen and total phosphorus in seawater.Hebei Journal of Industrial Science and Technology,2011,28(1):72-76.[雷立改,马晓珍,魏福祥等.水中总氮、总磷测定方法的研究进展.河北工业科技,2011,28(1):72-76.]
[17]Pápista,cs,B9ddi B.Chlorophyll-a determination with ethanol—A critical test.Hydrobiologi,2002,485(123):191-198.
[18]Uterm9hl H.Neue wege in der quantitativan erfassung des planktons(Mit besondere Berü-cksichtigung des Ultraplanktons).Verh Int Verein Theor Angew Limnol,1931,5:567-595.
[19]Zhang Zongshe,Huang Xiangfei eds.Research methods of freshwater plankton.Beijing:Science Press,1991.[章宗涉,黄祥飞.淡水浮游生物研究方法.北京:科学出版社,1991.]
[20]Hu Hongjun,Wei Yinxin eds.The freshwater algae of China—Systematics,taxonomy and ecology.Beijing:Science Press,2006.[胡鸿钧,魏印心.中国淡水藻类——系统、分类及生态.北京:科学出版社,2006.]
[21]Research Group of Carcinology,Institute of Zoology,Academia Sinica ed.Fauna sinica crustacea·freshwater copepods Beijing:Science Press,1979.[中国科学院动物研究所甲壳动物研究组.中国动物志甲壳纲:淡水桡足类.北京:科学出版社,1979.]
[22]Jiang Xiezhi,Du Nanshan eds.Fauna Sinica Crustacea·Freshwater Cladocera.Beijing:Science Press,1979.[蒋燮治,堵南山.中国动物志甲壳纲:淡水枝角类.北京:科学出版社,1979.]
[23]Bligh EG,Dyer WJ.A rapid method of total lipid extraction and purification.Canadian Journal of Biochemistry and Physiology,1959,37(8):911-917.
[24]Van den Meersche KK,Van Rijswijk P,Karline S et al.Autochthonous and allochthonous contributions to mesozooplankton diet in a tidal river and estuary:Integrating carbon isotope and fatty acid constraints.Limnology and Oceanography,2009,54(1):62-74.
[25]Zhao Dayong,Yan Wenming,Feng Jingwei et al.Applications of phospholipid fatty acid analysis in the microbial ecology studies of lake sediment.Chemistry&Bioengineering,2009,26(12):17-20.[赵大勇,燕文明,冯景伟等.磷脂脂肪酸分析在湖泊沉积物微生物生态学研究中的应用.化学与生物工程,2009,26(12):17-20.]
[26]Rajendran N,Matsuda O,Imamura N et al.Variation in microbial biomass and community structure in sediments of eutrophic bays as determined by phospholipid ester-linked fatty acids.Applied and Environmental Microbiology,1992,58(2):562-571.
[27]Ge Yuan,He Jizheng,Zheng Yuanming et al.Stable isotope probing and its applications in microbicalecology.Acta Ecologica Sinica,2006,26(5):1574-1582.[葛源,贺纪正,郑袁明等.稳定性同位素探测技术在微生物生态学研究中的应用.生态学报,2006,26(5):1574-1582.]
[28]Kritzberg ES,Cole J,Pace ML et al.Bacteria growth on allochthonous carbon in humic and nutrient-riched lakes:Results from whole-lake13C addition experiments.Ecosystems,2006,9(3):489-499.
[29]Azam F,Fenchel T,Field J et al.The ecological role of water-column microbes in the sea.Marine Ecology Progress Series,1983,10(3):257-263.
[30]Jansson M,Persson L,De Roos AM et al.Terrestrial carbon and intraspecific size-variation shape lake ecosystems.Trends in Ecology and Evolution,2007,22(6):316-322.
[31]Carignan R,Planas D,Vis C.Planktonic production and respiration in oligotrophic shield lakes.Limnology and Oceanography,2000,45(1):189-199.
[32]Tang Yali,Cheng Dongmei,Liu Zhengwen et al.Differntial support of lake food webs by allochthonous organic carbon.Ecological Science,2014,33(1):161-165.[唐雅丽,程冬梅,刘正文等.外源性有机碳对淡水生态系统食物网的贡献.生态学报,2014,33(1):161-165.]
[33]Institute of Animal,Chinese Academy of Science,crustaceans research group ed.Chinese animal.Beijing:Science Press,1979.[中国科学院中国动物志委员会.中国动物志.北京:科学出版社,1979.]
[34]Banta AM.Population density as related to sexand to evolution in Cladocera.The American Naturalist,1937,71(732):34-49.
[35]Zhang Guangtao,Li Chaolun,Sun Song et al.Feeding habit of Calanussinicus(Crustacea:Copepoda)during spring and autumn in the Bohai Sea studied with herbivore index.Scientia Marina,2006,70(3):381-388.
[36]Hama T,Yanagi K.Production and neutral aldose composition of dissolved carbohydrates excreted by natural marine phytoplankton populations.Limnology and Oceanography,2001,46(8):1945-1955.
[37]Biddanda B,Benner R.Carbon,nitrogen,and carbohydrate fluxes during the production of particulate and dissolved organic matter by marine phytoplankton.Limnology and Oceanography,1997,42(3):506-518.
[38]Pan Jizheng,Xiong Fei,Li Wenchao et al.Structure distribution and it’s impact factors of phytoplankton community in Fuxian Lake.Acta Ecological Sinica,2009,29(10):5376-5385.[潘继征,熊飞,李文朝等.抚仙湖浮游植物群落结构、分布及其影响因子.生态学报,2009,29(10):5376-5385.]
[39]Berggren M,Strom L,Laudon H et al.Lake secondary production fueled by rapid transfer of low molecular weight organic carbon from terrestrial sources to aquatic consumers.Ecology Letters,2010,13(7):870-880.
[40]Yokokawa T,Nagata T.Linking bacterial community structure to carbon fluxes in marine environments.Journal of Oceanography,2010,66(1):1-12.
[41]Algesten G,Sobek S,Bergstro MAK et al.Role of lakes for organic carbon cycling in the boreal zone.Global Change Biology,2005,10(1):141-147.