基于~(15)N稳定同位素示踪技术的太湖梅梁湾浮游植物群落氮吸收动力学研究
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摘要
浮游植物对氮的吸收与其生长繁殖密切相关,太湖梅梁湾湖区蓝藻水华频频暴发,对该水域浮游植物氮吸收进行研究具有重要意义.本文分别在冬、春、夏、秋4个季节于梅梁湾采样,对水体常规理化指标和浮游植物群落结构进行分析,并利用~(15)N稳定同位素示踪技术研究了浮游植物对铵态氮(NH_4~+-N)、硝态氮(NO_3~--N)和尿素态氮(Urea-N)吸收的动力学特征.结果表明,太湖梅梁湾浮游植物群落除了秋季对NH_4~+-N的吸收不符合米氏方程外,其余均符合.冬季和春季3种形态氮最大吸收速率(V_(max))的大小依次为:NH_4~+-N> NO_3~--N>Urea-N,而夏季为:NH_4~+-N>Urea-N>NO_3~--N. 3种形态氮Vmax的季节变化规律为夏季>秋季>春季>冬季. V_(max)在不同季节以及不同形态氮之间的差异性可能与浮游植物群落组成以及水体中NH_4~+-N浓度不同有关.浮游植物对NH_4~+-N吸收的KS值在冬、春季高于夏季,对Urea-N吸收的K_s值则在夏、秋季高于冬、春季,而对NO_3~--N吸收的K_s值则在夏季显著高于其他3个季节.冬季和春季梅梁湾浮游植物群落最容易受到NO_3~--N限制,而最不容易受到Urea-N的限制;而夏季,则最容易受到NO_3~--N限制,而最不容易受到NH_4~+-N的限制,且浮游植物群落对NH_4~+-N的亲和力最高.与NO_3~--N相比,秋季浮游植物更容易受到Urea-N的限制.不同季节,容易对浮游植物产生限制作用的氮的形态不同.
Nitrogen is the essential element for phytoplankton. Its concentration and form affect the biomass and community composition of phytoplankton directly or indirectly. In order to discover the restriction of different forms of nitrogen on phytoplankton growth in Lake Taihu,and supply scientific ground for further study of cyanobacteria blooms and eutrophication control,the especial uptake rates of ammonium,nitrate and urea by phytoplankton using ~(15)N-tracer technology as well as physicochemical parameters,urea concentration and phytoplankton community in Mailiang Bay were determined in the middle of January,April,July and October. The result showed that the uptake of ammonium,nitrate and urea,except the uptake of ammonium in August,exhibited the classic hyperbolic Michaelis-Menten kinetic curves. The maximum uptake speed rates(V_(max)) of three forms of nitrogen were:ammonium> nitrate> urea in winter and spring while ammonium>urea> nitrate in summer. The differences of V_(max) between seasons and forms were owing to the different phytoplankton community compositions and ammonium concentration in water. According to the Ks,the growth of phytoplankton community was more likely to be limited by nitrate and not prone to be limited by urea in winter and spring,while more likely to be limited by ammonium in summer. In autumn,it was prone to be limited by urea other than nitrate. In a word,the growth of phytoplankton community was limited by different forms of nitrogen in different seasons.
Nitrogen is the essential element for phytoplankton. Its concentration and form affect the biomass and community composition of phytoplankton directly or indirectly. In order to discover the restriction of different forms of nitrogen on phytoplankton growth in Lake Taihu,and supply scientific ground for further study of cyanobacteria blooms and eutrophication control,the especial uptake rates of ammonium,nitrate and urea by phytoplankton using ~(15)N-tracer technology as well as physicochemical parameters,urea concentration and phytoplankton community in Mailiang Bay were determined in the middle of January,April,July and October. The result showed that the uptake of ammonium,nitrate and urea,except the uptake of ammonium in August,exhibited the classic hyperbolic Michaelis-Menten kinetic curves. The maximum uptake speed rates(V_(max)) of three forms of nitrogen were:ammonium> nitrate> urea in winter and spring while ammonium>urea> nitrate in summer. The differences of V_(max) between seasons and forms were owing to the different phytoplankton community compositions and ammonium concentration in water. According to the Ks,the growth of phytoplankton community was more likely to be limited by nitrate and not prone to be limited by urea in winter and spring,while more likely to be limited by ammonium in summer. In autumn,it was prone to be limited by urea other than nitrate. In a word,the growth of phytoplankton community was limited by different forms of nitrogen in different seasons.
引文
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[27]Chen M,Huang YP,Qiu YS.Nitrogen uptake rate of the Bering Sea basin using isotope tracer technology.Progress in Natural Science Materials International,1997,17:1672-1684.[陈敏,黄奕普,邱雨生.白令海盆氮吸收速率的同位素示踪.自然科学进展,1997,17:1672-1684.]
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[29]Bronk DA,Sanderson MP,Mulholland MR et al.Organic and inorganic nitrogen uptake kinetics ineld populations dominated by Karenia brevis.In:Steidinger K,Vargo GA,Heil CA eds.Harmful Algae 2002.Florida Fish and Wildlife Conservation Commission,Florida Institute of Oceanography and Intergovernmental Oceanographic Commission of UNESCO,St.Petersburg,FL,2004:80-82.
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[32]Flynn KJ.Algal carbon-nitrogen metabolism:a biochemical basis for modelling the interactions between nitrate and ammonium uptake.Journal of Plankton Research,1991,13:373-387.
[33]Veuger B,Middelburg JJ,Boschker H et al.Microbial uptake of dissolved organic and inorganic nitrogen in Randers Fjord.Estuarine Coastal and Shelf Science,2004,61:507-515.
[34]Dugdale RC,Wilkerson FP,Hogue VE et al.The role of ammonium and nitrate in spring bloom development in San Francisco Bay.Estuarine Coastal and Shelf Science,2007,73:17-29.
[35]Herndon J,Cochlan WP.Nitrogen utilization by the raphidophyte Heterosigma akashiwo:Growth and uptake kinetics in laboratory cultures.Harmful Algae,2007,6:260-270.
[36]Berg GM,Balode M,Purina I et al.Plankton community composition in relation to availability and uptake of oxidized and reduced nitrogen.Aquatic Microbial Ecology,2003,30:263-274.
[37]Lomas MW,Glibert PM.Temperature regulation of nitrate uptake:A novel hypothesis about nitrate uptake and reduction in cool-water diatoms.Limnology and Oceanography,1999,44:556-572.
[38]Goldman JC.Potential role of large oceanic diatoms in new primary production.Deep Sea Research Part I:Oceanographic Research,1993,40:159-168.
[39]Domingues RB,Barbosa AB,Sommer U et al.Ammonium,nitrate and phytoplankton interactions in a freshwater tidal estuarine zone:potential effects of cultural eutrophication.Aquatic Sciences,2011,73(3):331-343.
[40]L'helguen S,Maguer JF,Caradec J.Inhibition kinetics of nitrate uptake by ammonium in size-fractionated oceanic phytoplankton communities:implications for new production and f-ratio estimates.Journal of Plankton Research,2008,30(10):1179-1188.
[41]Glibert PM,Wilkerson FP,Dugdale RC et al.Phytoplankton communities from San Francisco Bay Delta respond differently to oxidized and reduced nitrogen substrates-even under conditions that would otherwise suggest nitrogen sufficiency.Frontiers in Marine Science,2014,(1):1-16.
[42]Shen GY,Shi BZ eds.Marine ecology.Beijing:Science Press,2002:197-198.[沈国英,施并章.海洋生态学.北京:科学出版社,2002:197-198.]
[43]Reynolds C ed.Ecology of phytoplankton.Cambridge:Cambridge University Press,2006:145-175.
[44]Zhang X,Yi QT,Xie K et al.Nutrient kinetics of dominant species of phytoplankton in the artificial lakes formed from land subsidence by mining activities in the Huainan coalmine subsidence areas.J Lake Sci,2018,30(3):713-722.DOI:10.18307/2018.0313.[张鑫,易齐涛,谢凯等.淮南采煤塌陷湖泊浮游植物优势种的营养动力学.湖泊科学,2018,30(3):713-722.]
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[2]Berg GM,Glibert PM,Lomas MW et al.Organic nitrogen uptake and growth by the chrysophyte Aureococcus anophagefferens during a brown tide event.Marine Biology,1997,129:377-387.
[3]Lomas MW,Glibert PM,Clougherty D.Elevated organic nutrient ratio sassociated with brown tide blooms of Aureococcus anophagefferens(Pelagophyceae).Journal of Plankton Research,2001,23:1339-1344.
[4]Gao XM,Li YP,Du W et al.Suspended solid settlement characteristics in Meiliang Bay of Lake Taihu:in situ observation.J Lake Sci,2017,29(1):52-58.DOI:10.18307/2017.0106.[高小孟,李一平,杜薇等.太湖梅梁湾湖区悬浮物动态沉降特征的野外观测.湖泊科学,2017,29(1):52-58.]
[5]Song XL,Liu ZW,Pan HK et al.Phytoplankton community structure in Meiliang Bay and Lake Wutli of Lake Taihu.JLake Sci,2007,19(6):643-651.DOI:10.18307/2007.0604.[宋晓兰,刘正文,潘宏凯等.太湖梅梁湾与五里湖浮游植物群落的比较.湖泊科学,2007,19(6):643-651.]
[6]Yang L,Zhang M,Liu ZW.Uptake of various forms of nitrogen by phytoplankton community in spring in Lake Taihu.JLake Sci,2011,23(4):605-611.DOI:10.18307/2011.0417.[杨柳,章铭,刘正文.太湖春季浮游植物群落对不同形态氮的吸收.湖泊科学,2011,23(4):605-611.]
[7]Yang J,Gao H,Glibert PM et al.Rates of nitrogen uptake by cyanobacterially-dominated assemblages in Lake Taihu,China,during late summer.Harmful Algae,2017,65:71-84.
[8]Xu H,Paerl HW,Qin BQ et al.Determining critical nutrient thresholds needed to control harmful cyanobacterial blooms in eutrophic Lake Taihu,China.Environ Sci Technol,2015,49(2):1051-1059.
[9]Zhou J,Qin BQ,Han XX et al.Turbulence increases the risk of microcystin exposure in a eutrophic lake(Lake Taihu)during cyanobacterial bloom periods.Harmful Algae,2016,55:213-220.
[10]Glibert PM,Maranger R,Sobota DJ et al.The haber-Bosch-harmful algal bloom(HB-HAB)link.Environmental Research Letters,2014,9(10):13.
[11]Dugdale RC,Wilkerson FP.The use of15N to measure nitrogen uptake in eutrophic oceans;experimental considerations.Limnology and Oceanography,1986,31:673-689.
[12]Gu BH,Havens KE,Schelske CL et al.Uptake of dissoved nitrogen by phytoplankton in a eutrophic subtropical lake.Journal of Plankton Research,1997,19:759-770.
[13]Presing M,Herodek S,Preston T et al.Nitrogen uptake and the importance of internal nitrogen loading in Lake Balaton.Freshwater Biology,2001,46:125-139.
[14]Jin XC,Tu QY eds.Specifications for lake eutrophication survey:2nd edition.Beijing:China Environmental Science Press,1990.[金相灿,屠清瑛.湖泊富营养化调查规范:第2版.北京:中国环境科学出版社,1990.]
[15]Koroleff F ed.Determination of urea.In:Grasshoff KEMA eds.Methods of seawater analysis:2nd.New York:VCHPublshers,1983.
[16]Huang ZS,Huang XF eds.Research method 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.]
[17]Hu HJ,Wei YX eds.The freshwater algae of China:Systematics,taxonomy and ecology.Beijing:Science Press,2006.[胡鸿均,魏印心.中国淡水藻类---系统、分类及生态.北京:科学出版社,2006.]
[18]Zhang GW.Studies ondissovled nitrogen,phytoplankton and nitrogen uptake by phytoplankton in Huguangyan Maar Lake[Dissertation].Zhanjiang:Guangdong Ocean University,2014.[张国维.湖光岩玛珥湖溶解态氮与浮游植物及其氮吸收的研究[学位论文].湛江:广东海洋大学,2014.]
[19]Takamura N,Iwakuma T,Yasuno M.Uptake of13C and15N(ammonium,nitrate and urea)by Microcystis in Lake Kasumigaura.Journal of Plankton Research,1987,9:151-165.
[20]Gu BH,Alexander V.Dissolved nitrogen uptake by a cyanobacterial bloom(Anabaena flos-aquae)in a subarctic lake.Applied and Environmental Microbiology,1993,59:422-430.
[21]Kudela RM,Cochlan WP.Nitrogen and carbon uptake kinetics and the influence of irradiance for a red tide bloom off southern California.Aquatic Microbial Ecology,2000,21:31-47.
[22]Cochlan WP,Bronk DA.Nitrogen uptake kinetics in the Ross Sea,Antarctica.Deep-Sea Research Part I-Topical Studies in Oceanography,2001,48:4127-4153.
[23]Herndon J,Cochlan WP.Nitrogen utilization by the raphidophyte Heterosigma akashiwo:Growth and uptake kinetics in laboratory cultures.Harmful Algae,2007,6:260-270.
[24]Zhang GW,Li CL,Huang XH et al.Uptake of dissolved nitrogen by phytoplankton in spring in Huguangyan Maar Lake.JLake Sci,2015,27(3):527-534.DOI:10.18307/2015.0321.[张国维,李长玲,黄翔鹄等.湖光岩玛珥湖春季浮游植物对溶解态氮的吸收.湖泊科学,2015,27(3):527-534.]
[25]Raphael M,Kudela A,William JQL.The potential roal of anthropogenically derived nitrogen in the growth of harmful algae in California,USA.Harmful Algae,2008,8:103-110.
[26]Gu BH,Alexander V.Seasonal variations in dissolved inorganic nitrogen utilization in a subarctic Alaskan lake.Archiv fur Hydrobiologie,1993,126:273-288.
[27]Chen M,Huang YP,Qiu YS.Nitrogen uptake rate of the Bering Sea basin using isotope tracer technology.Progress in Natural Science Materials International,1997,17:1672-1684.[陈敏,黄奕普,邱雨生.白令海盆氮吸收速率的同位素示踪.自然科学进展,1997,17:1672-1684.]
[28]Mukherjee R,Kumar S,Muduli PR.Spatial variation of nitrogen uptake rates in the largest brackish water lagoon of Asia(Chilika,India).Estuarine,Coastal and Shelf Science,2018:1-11.DOI:10.1016/j.ecss.2018.01.012.
[29]Bronk DA,Sanderson MP,Mulholland MR et al.Organic and inorganic nitrogen uptake kinetics ineld populations dominated by Karenia brevis.In:Steidinger K,Vargo GA,Heil CA eds.Harmful Algae 2002.Florida Fish and Wildlife Conservation Commission,Florida Institute of Oceanography and Intergovernmental Oceanographic Commission of UNESCO,St.Petersburg,FL,2004:80-82.
[30]Bronk DA,See JH,Bradley P et al.DON as a source of bioavailable nitrogen for phytoplankton.Biogeosciences,2007,(4):283-296.
[31]Dortch Q.The interaction between ammonium and nitrate uptake in phytoplankton.Marine Ecology Progress Series,1990,61:183-201.
[32]Flynn KJ.Algal carbon-nitrogen metabolism:a biochemical basis for modelling the interactions between nitrate and ammonium uptake.Journal of Plankton Research,1991,13:373-387.
[33]Veuger B,Middelburg JJ,Boschker H et al.Microbial uptake of dissolved organic and inorganic nitrogen in Randers Fjord.Estuarine Coastal and Shelf Science,2004,61:507-515.
[34]Dugdale RC,Wilkerson FP,Hogue VE et al.The role of ammonium and nitrate in spring bloom development in San Francisco Bay.Estuarine Coastal and Shelf Science,2007,73:17-29.
[35]Herndon J,Cochlan WP.Nitrogen utilization by the raphidophyte Heterosigma akashiwo:Growth and uptake kinetics in laboratory cultures.Harmful Algae,2007,6:260-270.
[36]Berg GM,Balode M,Purina I et al.Plankton community composition in relation to availability and uptake of oxidized and reduced nitrogen.Aquatic Microbial Ecology,2003,30:263-274.
[37]Lomas MW,Glibert PM.Temperature regulation of nitrate uptake:A novel hypothesis about nitrate uptake and reduction in cool-water diatoms.Limnology and Oceanography,1999,44:556-572.
[38]Goldman JC.Potential role of large oceanic diatoms in new primary production.Deep Sea Research Part I:Oceanographic Research,1993,40:159-168.
[39]Domingues RB,Barbosa AB,Sommer U et al.Ammonium,nitrate and phytoplankton interactions in a freshwater tidal estuarine zone:potential effects of cultural eutrophication.Aquatic Sciences,2011,73(3):331-343.
[40]L'helguen S,Maguer JF,Caradec J.Inhibition kinetics of nitrate uptake by ammonium in size-fractionated oceanic phytoplankton communities:implications for new production and f-ratio estimates.Journal of Plankton Research,2008,30(10):1179-1188.
[41]Glibert PM,Wilkerson FP,Dugdale RC et al.Phytoplankton communities from San Francisco Bay Delta respond differently to oxidized and reduced nitrogen substrates-even under conditions that would otherwise suggest nitrogen sufficiency.Frontiers in Marine Science,2014,(1):1-16.
[42]Shen GY,Shi BZ eds.Marine ecology.Beijing:Science Press,2002:197-198.[沈国英,施并章.海洋生态学.北京:科学出版社,2002:197-198.]
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