密闭培养对纤细角毛藻稳定同位素组成的影响
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摘要
为研究碳源对赤潮藻生长的影响,以赤潮藻纤细角毛藻(Chaetoceros gracilis)为研究对象,在密闭条件下进行培养,模拟赤潮发生时碳源缺乏的生长环境;分析碳源不足对纤细角毛藻不同生长阶段的差异影响,采用密闭培养的方法,分别在正常组和缺氮组的营养条件下培养纤细角毛藻,测定生物量、营养盐浓度和碳氮稳定同位素组成等重要监测指标.结果表明:不同营养条件对纤细角毛藻的生物量存在显著影响.在正常组,c(NO3--N)(NO3--N为硝酸盐)随着培养时间呈下降趋势,ρ(NO2--N)(NO2--N为亚硝酸盐)和ρ(NH4+-N)(NH4+-N为铵盐)与藻类生物量呈显著正相关,ρ(PO43--P)(PO43--P为磷酸盐)和c(Si O42--Si)(Si O42--Si为硅酸盐)与藻类生物量呈显著负相关;在缺氮组,c(NO3--N)、ρ(NO2--N)、ρ(NH4+-N)、ρ(PO43--P)和c(Si O42--Si)均与藻类生物量呈显著负相关.整个试验周期中,正常组和缺氮组δ13C和δ15N值越来越正.其中,正常组的δ13C值比缺氮组的更正,平均高2.286‰;缺氮组的δ15N值比正常组的更正,平均高3.307‰.正常组和缺氮组的δ13C、δ15N值与生物量均呈显著正相关.研究显示,碳源和氮源的不足分别会导致更正的δ13C和δ15N值,推测赤潮发生会导致δ13C和δ15N值偏正,赤潮藻的碳氮稳定同位素组成可以作为赤潮监测的指标和方法.
Chaetoceros gracilis was cultured in a closed environment to simulate carbon source limitation during the occurrence of red tides.The microalgae were maintained under both normal nutrient conditions and nitrogen deficient conditions. Biomass,stable carbon,nitrogen isotopes and nutrient concentration were measured to determine how the two different nutrient condition affects C. gracilis growth at different growth phases. The results showed that the two different conditions produced significant differences in biomass during different growth phases. Under the normal nutrient condition,the concentration of nitrate,phosphate,and silicate decreased over time,while nitrite and ammonium increased. Under the nitrogen deficient condition,all five nutrient conditions were negatively correlated with the biomass of the algae. δ13 C and δ15 N values increased throughout the experiment. The δ13 C value of the normal nutrient group was 2. 286‰ higher than that of the nitrogen deficient group. The δ15 C value of the nitrogen deficient group was 3. 307‰ higher than that of the normal nutrient group. Both δ13 C and δ15 N values had a significant positive correlation with biomass in both groups. The occurrence of red tides resulted in higher levels of δ13 C and δ15 N,which could be used as an indicator of red tide events.
Chaetoceros gracilis was cultured in a closed environment to simulate carbon source limitation during the occurrence of red tides.The microalgae were maintained under both normal nutrient conditions and nitrogen deficient conditions. Biomass,stable carbon,nitrogen isotopes and nutrient concentration were measured to determine how the two different nutrient condition affects C. gracilis growth at different growth phases. The results showed that the two different conditions produced significant differences in biomass during different growth phases. Under the normal nutrient condition,the concentration of nitrate,phosphate,and silicate decreased over time,while nitrite and ammonium increased. Under the nitrogen deficient condition,all five nutrient conditions were negatively correlated with the biomass of the algae. δ13 C and δ15 N values increased throughout the experiment. The δ13 C value of the normal nutrient group was 2. 286‰ higher than that of the nitrogen deficient group. The δ15 C value of the nitrogen deficient group was 3. 307‰ higher than that of the normal nutrient group. Both δ13 C and δ15 N values had a significant positive correlation with biomass in both groups. The occurrence of red tides resulted in higher levels of δ13 C and δ15 N,which could be used as an indicator of red tide events.
引文
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[16]王海霞,刘瑀,关春江,等.营养条件对微藻碳、氮稳定同位素组成的影响[J].中国环境科学,2014,34(3):727-733.WANG Haixia,LIU Yu,GUAN Chunjiang,et al.Effects of nutritional conditions on the stable carbon and nitrogen isotope of microalgae[J].China Environmental Science,2014,34(3):727-733.
[17]金海燕,陈建芳,翁焕新,等.长江口外赤潮多发区近几十年来古生产力记录及环境意义[J].海洋学报,2009,31(2):113-119.JIN Haiyan,CHEN Jianfang,WENG Huanxin,et al.Variations of paleoproductivity in the past decades and the environmental implications in the Changjiang Estuary in China[J].Acta Oceanologica Sinica,2009,31(2):113-119.
[18]STREET G T,MONTAGNA P A,PARKER P L.Incorporation of brown tide into an estuarine food web[R].Twenty-Third Bembre Ecology Meeting.NJUSA:Institute of Marine Coastal Science,1997.
[19]俞志明.不同氮源对海洋微藻氮稳定同位素分馏作用的影响[J].海洋与湖沼,2004,35(6):524-529.YU Zhiming.Effect of different nitrogen pools on nitrogen isotopic fractionation during the uptake by marine microalgae[J].Oceanologia et Limnologia Sinica,2004,35(6):524-529.
[20]刘苓.不同营养条件对赤潮藻碳和氮稳定同位素组成的影响[D].大连:大连海事大学,2012.
[21]国家海洋局908专项办公室.海洋化学调查技术规程[M].北京:海洋出版社,2006.
[22]WAITE A M,MUHLING B A,HOLL C M,et al.Food web structure in two counter-rotating eddies based onδ15N andδ13C isotopic analyses[J].Deep-Sea Research Part II,2007,54(8):1055-1075.
[23]赵佩佩,顾文辉,伍松翠,等.氮限制有利于三角褐指藻脂质积累[J].科学通报,2015,60(23):2196-2208.ZHAO Peipei,GU Wenhui,WU Songcui,et al.Changes in central carbon metabolism of Phaeodactylum Tricornutum are beneficial for lipid accumulation under nitrogen starvation conditions[J].Chinese Science Bulletin,2015,60(23):2196-2208.
[24]吕嘉扬,王大志,洪华生,等.两种硝酸盐浓度下威氏海链藻和盐生杜氏藻硝酸还原酶活力研究[J].集美大学学报(自然科学版),2004,9(1):6-9.LU Jiayang,WANG Dazhi,HONG Huasheng,et al.Comparative studies on nitrate reductase activity of Thalassiosira weissflogii and Dunaliella salina cultured in two nitrate concentrations[J].Journal of Jimei University(Natural Science),2004,9(1):6-9.
[25]刘瑀,王海霞,李晓琳,等.赤潮过程中塔玛亚历山大藻(Alexandrium tamareme)C、N稳定同位素组成的变化规律[J].海洋环境科学,2015,34(5):783-787.LIU Yu,WANG Haixia,LI Xiaolin,et al.Variation of carbon and nitrogen stable isotope composition of Alexandrium tamareme in the process of red tide[J].Marine Environmental Science,2015,34(5):783-787.
[26]GU Xingyan,LI Keqiang,PANG Kai,et al.Effects of p H on the growth and NH4-N uptake of Skeletonema costatum and Nitzschia closterium[J].Marine Pollution Bulletin,2017,124(2):946-952.
[27]LUDWIG C A.The availability of different forms of nitrogen to a green alga[J].American Journal of Botany,1938,25(6):448-458.
[28]GARDNER W S,LAVRENTYEV P J,CAVALETTO J F,et al.Distribution and dynamics of nitrogen and microbial plankton in southern Lake Michigan during spring transition 1999-2000[J].Journal of Geophysical Research Oceans,2004,109:325-347.
[29]PAERL H W,XU Hai,MCCARTHY M J,et al.Controlling harmful cyanobacterial blooms in a hyper-eutrophic lake(Lake Taihu,China):the need for a dual nutrient(N&P)management strategy[J].Water Research,2011,45(5):1973-1983.
[30]杨东方,高振会,陈豫,等.硅的生物地球化学过程的研究动态[J].海洋科学,2002,26(3):39-42.YANG Dongfang,GAO Zhenhui,CHEN Yu,et al.The biogeochemical process of silicon[J].Marine Science,2002,26(3):39-42.
[31]高春梅,郑伊汝,张硕.海州湾海洋牧场沉积物-水界面营养盐交换通量的研究[J].大连海洋大学学报,2016,31(1):95-102.GAO Chunmei,ZHENG Yiru,ZHANG Shuo.Exchange fluxes of nutrients at sediment-seawater interface in marine ranching area of Haizhou Bay[J].Journal of Dalian Ocean University,2016,31(1):95-102.
[32]BRUTEMARK A,LINDEHOFF E,GRANLI E,et al.Carbon isotope signature variability among cultured microalgae:influence of species,nutrients and growth[J].Journal of Experimental Marine Biology and Ecology,2009,372(1/2):98-105.
[33]GLEITZ M,KUKERT H,RIEBESELL U,et al.Carbon acquisition and growth of Antarctic sea ice diatoms in closed bottle incubations[J].Marine Ecology Progress,1996,135(1/2/3):169-177.
[34]BOSCHKER H T S,MIDDELBURG J J.Stable isotopes and biomarkers in microbial ecology[J].FEMS Microbiology Ecology,2002,40:85-95.
[35]FINLAY J C.Patterns and controls of lotic algal stable carbon isotope ratios[J].Limnology and Oceanography,2004,49(3):850-861.
[2]宋秀贤,俞志明,殷克东,等.香港近岸海域营养盐结构特征及其对浮游植物生长的影响[J].海洋与湖沼,2013,44(4):846-852.SONG Xiuxian,YU Zhiming,YIN Kedong,et al.Temporal and spatial distribution of nutrients and chl a in the coastal area of Hong Kong[J].Oceanologia et Limnologia Sinica,2013,44(4):846-852.
[3]马佳莹,戴家伟,胡芳露,等.大型海藻对富营养化海水养殖区的修复[J].农村经济与科技,2016,27(11):6-7.
[4]NRC(National Research Council).Clean coastal waters:understanding and reducing the effects of nutrient pollution[R].Washington DC:National Academies Press,2000.
[5]GLIBERT P M,ANDERSON D M,GENTIEN P,et al.The global complex phenomena of harmful algal blooms[J].Oceanography,2005,18(2):136-147.
[6]ANDERSON D M,GLIBERT P M,BURKHOLDER J M.Harmful algal blooms and eutrophication:nutrient sources,composition and consequences[J].Estuaries,2002,25(4):704-726.
[7]李鸿妹,石晓勇,陈鹏,等.春季东海赤潮发生前后营养盐及溶解氧的平面分布特征[J].环境科学,2013,34(6):2159-2165.LI Hongmei,SHI Xiaoyong,CHEN Peng,et al.Distribution of dissolved inorganic nutrients and dissolved oxygen in the high frequency area of harmful algal blooms in the East China Sea in spring[J].Environmental Science,2013,34(6):2159-2165.
[8]SMAYDA T J.Harmful algal blooms:their ecophysiology and general relevance to phytoplankton blooms in the sea[J].Limnology and Oceanography,1997,42(5):1137-1153.
[9]李涵,许秋瑾,储昭升,等.不同形态氮对洋河水库螺旋鱼腥藻和惠氏微囊藻生长的影响[J].环境科学研究,2010,23(12):1494-1498.LI Han,XU Qiujin,CHU Zhaosheng,et al.Effects of different forms of nitrogen on the growth of Anabaena sp.and Microcystis sp.from Yanghe Reservoir[J].Research of Environmental Sciences,2010,23(12):1494-1498.
[10]HOWARTH R,CHAN F,CONLEY D J,et al.Coupled biogeochemical cycles:eutrophication and hypoxia in temperate estuaries and coastal marine ecosystems[J].Frontiers in Ecology&the Environment,2011,9(1):18-26.
[11]FROELICH P N.Kinetic control of dissolved phosphate in natural rivers and estuaries:a primer on the phosphate buffer mechanism[J].Limnology&Oceanography,1988,33(4):649-668.
[12]NIXON S W,AMMERMAN J W,ATKINSON L P,et al.The fate of nitrogen and phosphorus at the land-sea margin of the North Atlantic Ocean[J].Biogeochemistry,1996,35(1):141-180.
[13]NMERY J,GARNIER J.Typical features of particulate phosphorus in the Seine estuary(France)[J].Hydrobiologia,2007,588(1):271-290.
[14]王睿喆,王沛芳,任凌霄,等.营养盐输入对太湖水体中磷形态转化及藻类生长的影响[J].环境科学,2015,36(4):1301-1308.WANG Ruizhe,WANG Peifang,REN Lingxiao,et al.Effects of nutrient inputs on changes of phosphorus forms and phytoplankton growth in Taihu Lake[J].Environmental Science,2015,36(4):1301-1308.
[15]岳冬梅,李洁,肖琳.营养盐恢复对氮磷饥饿铜绿微囊藻生长的影响[J].环境科学,2016,37(11):4220-4227.YUE Dongmei,LI Jie,XIAO Lin.Nutrients recovery on the growth of nitrogen and phosphorus starved Microcystis aeruginosa[J].Environmental Science,2016,37(11):4220-4227.
[16]王海霞,刘瑀,关春江,等.营养条件对微藻碳、氮稳定同位素组成的影响[J].中国环境科学,2014,34(3):727-733.WANG Haixia,LIU Yu,GUAN Chunjiang,et al.Effects of nutritional conditions on the stable carbon and nitrogen isotope of microalgae[J].China Environmental Science,2014,34(3):727-733.
[17]金海燕,陈建芳,翁焕新,等.长江口外赤潮多发区近几十年来古生产力记录及环境意义[J].海洋学报,2009,31(2):113-119.JIN Haiyan,CHEN Jianfang,WENG Huanxin,et al.Variations of paleoproductivity in the past decades and the environmental implications in the Changjiang Estuary in China[J].Acta Oceanologica Sinica,2009,31(2):113-119.
[18]STREET G T,MONTAGNA P A,PARKER P L.Incorporation of brown tide into an estuarine food web[R].Twenty-Third Bembre Ecology Meeting.NJUSA:Institute of Marine Coastal Science,1997.
[19]俞志明.不同氮源对海洋微藻氮稳定同位素分馏作用的影响[J].海洋与湖沼,2004,35(6):524-529.YU Zhiming.Effect of different nitrogen pools on nitrogen isotopic fractionation during the uptake by marine microalgae[J].Oceanologia et Limnologia Sinica,2004,35(6):524-529.
[20]刘苓.不同营养条件对赤潮藻碳和氮稳定同位素组成的影响[D].大连:大连海事大学,2012.
[21]国家海洋局908专项办公室.海洋化学调查技术规程[M].北京:海洋出版社,2006.
[22]WAITE A M,MUHLING B A,HOLL C M,et al.Food web structure in two counter-rotating eddies based onδ15N andδ13C isotopic analyses[J].Deep-Sea Research Part II,2007,54(8):1055-1075.
[23]赵佩佩,顾文辉,伍松翠,等.氮限制有利于三角褐指藻脂质积累[J].科学通报,2015,60(23):2196-2208.ZHAO Peipei,GU Wenhui,WU Songcui,et al.Changes in central carbon metabolism of Phaeodactylum Tricornutum are beneficial for lipid accumulation under nitrogen starvation conditions[J].Chinese Science Bulletin,2015,60(23):2196-2208.
[24]吕嘉扬,王大志,洪华生,等.两种硝酸盐浓度下威氏海链藻和盐生杜氏藻硝酸还原酶活力研究[J].集美大学学报(自然科学版),2004,9(1):6-9.LU Jiayang,WANG Dazhi,HONG Huasheng,et al.Comparative studies on nitrate reductase activity of Thalassiosira weissflogii and Dunaliella salina cultured in two nitrate concentrations[J].Journal of Jimei University(Natural Science),2004,9(1):6-9.
[25]刘瑀,王海霞,李晓琳,等.赤潮过程中塔玛亚历山大藻(Alexandrium tamareme)C、N稳定同位素组成的变化规律[J].海洋环境科学,2015,34(5):783-787.LIU Yu,WANG Haixia,LI Xiaolin,et al.Variation of carbon and nitrogen stable isotope composition of Alexandrium tamareme in the process of red tide[J].Marine Environmental Science,2015,34(5):783-787.
[26]GU Xingyan,LI Keqiang,PANG Kai,et al.Effects of p H on the growth and NH4-N uptake of Skeletonema costatum and Nitzschia closterium[J].Marine Pollution Bulletin,2017,124(2):946-952.
[27]LUDWIG C A.The availability of different forms of nitrogen to a green alga[J].American Journal of Botany,1938,25(6):448-458.
[28]GARDNER W S,LAVRENTYEV P J,CAVALETTO J F,et al.Distribution and dynamics of nitrogen and microbial plankton in southern Lake Michigan during spring transition 1999-2000[J].Journal of Geophysical Research Oceans,2004,109:325-347.
[29]PAERL H W,XU Hai,MCCARTHY M J,et al.Controlling harmful cyanobacterial blooms in a hyper-eutrophic lake(Lake Taihu,China):the need for a dual nutrient(N&P)management strategy[J].Water Research,2011,45(5):1973-1983.
[30]杨东方,高振会,陈豫,等.硅的生物地球化学过程的研究动态[J].海洋科学,2002,26(3):39-42.YANG Dongfang,GAO Zhenhui,CHEN Yu,et al.The biogeochemical process of silicon[J].Marine Science,2002,26(3):39-42.
[31]高春梅,郑伊汝,张硕.海州湾海洋牧场沉积物-水界面营养盐交换通量的研究[J].大连海洋大学学报,2016,31(1):95-102.GAO Chunmei,ZHENG Yiru,ZHANG Shuo.Exchange fluxes of nutrients at sediment-seawater interface in marine ranching area of Haizhou Bay[J].Journal of Dalian Ocean University,2016,31(1):95-102.
[32]BRUTEMARK A,LINDEHOFF E,GRANLI E,et al.Carbon isotope signature variability among cultured microalgae:influence of species,nutrients and growth[J].Journal of Experimental Marine Biology and Ecology,2009,372(1/2):98-105.
[33]GLEITZ M,KUKERT H,RIEBESELL U,et al.Carbon acquisition and growth of Antarctic sea ice diatoms in closed bottle incubations[J].Marine Ecology Progress,1996,135(1/2/3):169-177.
[34]BOSCHKER H T S,MIDDELBURG J J.Stable isotopes and biomarkers in microbial ecology[J].FEMS Microbiology Ecology,2002,40:85-95.
[35]FINLAY J C.Patterns and controls of lotic algal stable carbon isotope ratios[J].Limnology and Oceanography,2004,49(3):850-861.