六株球形棕囊藻的色素组成特征研究
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摘要
球形棕囊藻(Phaeocystisglobosa)是我国南方沿海近年来主要的赤潮原因种之一,由球形棕囊藻形成的赤潮对海水养殖业发展和海域生态环境构成了严重威胁。棕囊藻通常以囊状群体形式形成赤潮,很难获取其丰度数据,以往研究中多以19′-己酰氧基岩藻黄素(Hex-fuco)或19′-丁酰氧基岩藻黄素(But-fuco)作为其特征色素,利用化学分类软件CHEMTAX计算其生物量。为了解我国近海球形棕囊藻的色素组成特征,本文采用高效液相色谱方法,分析了6株球形棕囊藻的色素组成与含量状况,其中5株分离自我国近海。结果表明, 6株球形棕囊藻均以岩藻黄素和叶绿素a为主要色素,但其特征色素Hex-fuco却存在显著的株系间差异,即便是分离自相同海域的不同球形棕囊藻藻株也存在差别。对比棕囊藻游离细胞和囊状群体的色素组成,可以看出两者在色素组成上基本一致,但囊状群体中捕光色素(Light-harvesting pigment)含量低于游离细胞,而光保护色素(Photoprotective pigment)则高于游离细胞,可能与不同存在形态的棕囊藻对光照的适应特征差异有关。以上研究表明,在以CHEMTAX方法计算球形棕囊藻生物量时,需要充分调查海域棕囊藻的特征色素组成情况,获取其特征色素信息,构建合理色素比例初始矩阵,为球形棕囊藻赤潮监测奠定基础。
Phaeocystis globosa, a major red-tide causative species in the South China Sea, posed big threats to the local aquaculture and ecosystems. Due to the co-existence of both free-swimming cells and big colonies of P. globosa, it is often difficult to obtain the accurate abundance of P. globosa during red tides. Therefore, the CHEMTAX method is often adopted to calculate the biomass of P. globosa using 19′-hexanoyloxyfucoxanthin(Hex-fuco) or 19'-butanoyloxyfucoxanthin(But-fuco)as its representative pigments. To find out the representative pigment of P. globosa, we analyzed the pigment composition and content of six strains of P. globosa with high performance liquid chromatography, of which five strains were isolated from Chinese coastal waters. The results show that the pigment compositions of the six strains are largely the same, the major pigment components are fucoxanthin and chlorophyll a. However, each representative pigment, especially Hex-fuco,features own strain-specific variation pattern, and even the strains isolated from the same sea area had different pigment profiles. In addition, we compared the pigment compositions of the free-swimming cells and colonies of P. globosa in the ratio of each pigment to chlorophyll a, and found that the free-swimming cells contain more light-harvesting pigments such as chl c, fucoxanthin, and Hex-fuco, while the colonie contain more photo-protection pigments such as diatoxanthin and carotenes. This difference in pigment profile reflected their own unique light adaptive strategy. Therefore, the pigment composition of P. globosa in the selected region of red tide monitoring should be examined carefully to determine their most representative pigments to help monitoring P. globosa red tides using the CHEMTAX method.
Phaeocystis globosa, a major red-tide causative species in the South China Sea, posed big threats to the local aquaculture and ecosystems. Due to the co-existence of both free-swimming cells and big colonies of P. globosa, it is often difficult to obtain the accurate abundance of P. globosa during red tides. Therefore, the CHEMTAX method is often adopted to calculate the biomass of P. globosa using 19′-hexanoyloxyfucoxanthin(Hex-fuco) or 19'-butanoyloxyfucoxanthin(But-fuco)as its representative pigments. To find out the representative pigment of P. globosa, we analyzed the pigment composition and content of six strains of P. globosa with high performance liquid chromatography, of which five strains were isolated from Chinese coastal waters. The results show that the pigment compositions of the six strains are largely the same, the major pigment components are fucoxanthin and chlorophyll a. However, each representative pigment, especially Hex-fuco,features own strain-specific variation pattern, and even the strains isolated from the same sea area had different pigment profiles. In addition, we compared the pigment compositions of the free-swimming cells and colonies of P. globosa in the ratio of each pigment to chlorophyll a, and found that the free-swimming cells contain more light-harvesting pigments such as chl c, fucoxanthin, and Hex-fuco, while the colonie contain more photo-protection pigments such as diatoxanthin and carotenes. This difference in pigment profile reflected their own unique light adaptive strategy. Therefore, the pigment composition of P. globosa in the selected region of red tide monitoring should be examined carefully to determine their most representative pigments to help monitoring P. globosa red tides using the CHEMTAX method.
引文
王朝晖,吕颂辉,陈菊芳等,1998.广东沿海几种赤潮生物的分类学研究.武汉植物学研究,16(4):310-314
杨和福,2004.棕囊藻的生物学概述I.形态分类和生理生态学.东海海洋,22(1):49-63
何家菀,施之新,张银华等,1999.一种棕囊藻的形态特征与毒素分析.海洋与湖沼,30(2):172-179
沈萍萍,王艳,齐雨藻等,2000.球形棕囊藻的生长特性及生活史研究.水生生物学报,24(6):635-643
胡晓坤,张清春,陈振帆等,2019.北部湾海域球形棕囊藻遗传多样性分析.海洋与湖沼,50(3):601-610
梁玉波,2012.中国赤潮灾害调查与评价(1933-2009).北京:海洋出版社,112-115
Antajan E,Chrétiennot-Dinet M J,Leblanc C et al,2004.19′-Hexanoyloxy fucoxanthin may not be the appropriate pigment to trace occurrence and fate of Phaeocystis:the case of P.globosa in Belgian coastal waters.Journal of Sea Research,52(3):165-177
Baumann M E M,Lancelot C,Brandini F P et al,1994.The taxonomic identity of the cosmopolitan prymnesiophyte Phaeocystis:a morphological and ecophysiological approach.Journal of Marine Systems,5(1):5-22
Breton E,Brunet C,Sautour B et al,2000.Annual variations of phytoplankton biomass in the Eastern English Channel:comparison by pigment signatures and microscopic counts.Journal of Plankton Research,22(8):1423-1440
Buma A G J,Bano N,Veldhuis M J W et al,1991.Comparison of the pigmentation of 2 strains of the Prymnesiophyte Phaeocystis sp.Netherlands Journal of Sea Research,27(2):173-182
Charlson R J,Lovelock J E,Andreae M O et al,1987.Oceanic phytoplankton,atmospheric sulphur,cloud albedo and climate.Nature,326(6114):655-661
Fragoso G M,Poulton A J,Yashayaev I M et al,2017.Spring phytoplankton communities of the Labrador Sea(2005-2014):pigment signatures,photophysiology and elemental ratios.Biogeosciences,14(5):1235-1259
Gibb S W,Barlow R G,Cummings D G et al,2000.Surface phytoplankton pigment distributions in the Atlantic Ocean:an assessment of basin scale variability between 50°N and50°S.Progress in Oceanography,45(3-4):339-368
Gibberd M J,Kean E,Barlow R et al,2013.Phytoplankton chemotaxonomy in the Atlantic sector of the Southern Ocean during late summer 2009.Deep Sea Research Part I:Oceanographic Research Papers,78:70-78
Jeffrey S W,Wright S W,1994.Photosynthetic pigments in the Haptophyta.In:Green J C,Leadbeater B S C eds.The Haptophyte Algae.Oxford,UK:Clarendon Press,111-132
Liu S X,Yu Z G,Yao P et al,2011.Effects of irradiance on pigment signatures of harmful algae during growth process.Acta Oceanologica Sinica,30(6):46-57
Mackey M D,Mackey D J,Higgins H W et al,1996.CHEMTAX-a program for estimating class abundances from chemical markers:application to HPLC measurements of phytoplankton.Marine Ecology Progress Series,144:265-283
Muylaert K,Gonzales R,Franck M et al,2006.Spatial variation in phytoplankton dynamics in the Belgian coastal zone of the North Sea studied by microscopy,HPLC-CHEMTAXand underway fluorescence recordings.Journal of Sea Research,55(4):253-265
Schoemann V,Wollast R,Chou L et al,2001.Effects of photosynthesis on the accumulation of Mn and Fe by Phaeocystis colonies.Limnology and Oceanography,46(5):1065-1076
Schoemann V,Becquevort S,Stefels J et al,2005.Phaeocystis blooms in the global ocean and their controlling mechanisms:a review.Journal of Sea Research,53(1-2):43-66
Seoane S,Zapata M,Orive E,2009.Growth rates and pigment patterns of haptophytes isolated from estuarine waters.Journal of Sea Research,62(4):286-294
Smith W O Jr,Codispoti L A,Nelson D M et al,1991.Importance of Phaeocystis blooms in the high-latitude ocean carbon cycle.Nature,352(6335):514-516
Stefels J,Van Leeuwe M A,1998.Effects of iron and light stress on the biochemical composition of Antarctic Phaeocystis sp.(Prymnesiophyceae).I.Intracellular DMSP concentrations.Journal of Phycology,34(3):486-495
Stuart V,Sathyendranath S,Head E J H et al,2000.Bio-optical characteristics of diatom and prymnesiophyte populations in the Labrador Sea.Marine Ecology Progress Series,201:91-106
Van Leeuwe M A,Visser R J W,Stefels J,2014.The pigment composition of Phaeocystis antarctica(Haptophyceae)under various conditions of light,temperature,salinity,and iron.Journal of Phycology,50(6):1070-1080
Vaulot D,Birrien J L,Marie D et al,1994.Morphology,ploidy,pigment composition,and genome size of cultured strains of Phaeocystis(Prymnesiophyceae).Journal of Phycology,30(6):1022-1035
Wright S W,Jeffrey S W,1987.Fucoxanthin pigment markers of marine phytoplankton analysed by HPLC and HPTLC.Marine Ecology Progress Series,38:259-266
Wright S W,Thomas D P,Marchant H J et al,1996.Analysis of phytoplankton of the Australian sector of the Southern Ocean:Comparisons of microscopy and size-frequency data with interpretations of pigment HPLC data using the CHEMTAX matrix factorisation program.Marine Ecology Progress Series,144:285-298
Wright S W,Van Den Enden R L,Pearce I et al,2010.Phytoplankton community structure and stocks in the Southern Ocean(30°-80°E)determined by CHEMTAXanalysis of HPLC pigment signatures.Deep Sea Research Part II:Topical Studies in Oceanography,57(9-10):758-778
Zapata M,Rodríguez F,Garrido J L,2000.Separation of chlorophylls and carotenoids from marine phytoplankton:a new HPLC method using a reversed phase C8 column and pyridine-containing mobile phases.Marine Ecology Progress Series,195:29-45
Zapata M,Jeffrey S W,Wright S W et al,2004.Photosynthetic pigments in 37 species(65 strains)of Haptophyta:implications for oceanography and chemotaxonomy.Marine Ecology Progress Series,270:83-102
Zapata M,2005.Recent advances in pigment analysis as applied to picophytoplankton.Vie et Milieu,55(3-4):233-248
杨和福,2004.棕囊藻的生物学概述I.形态分类和生理生态学.东海海洋,22(1):49-63
何家菀,施之新,张银华等,1999.一种棕囊藻的形态特征与毒素分析.海洋与湖沼,30(2):172-179
沈萍萍,王艳,齐雨藻等,2000.球形棕囊藻的生长特性及生活史研究.水生生物学报,24(6):635-643
胡晓坤,张清春,陈振帆等,2019.北部湾海域球形棕囊藻遗传多样性分析.海洋与湖沼,50(3):601-610
梁玉波,2012.中国赤潮灾害调查与评价(1933-2009).北京:海洋出版社,112-115
Antajan E,Chrétiennot-Dinet M J,Leblanc C et al,2004.19′-Hexanoyloxy fucoxanthin may not be the appropriate pigment to trace occurrence and fate of Phaeocystis:the case of P.globosa in Belgian coastal waters.Journal of Sea Research,52(3):165-177
Baumann M E M,Lancelot C,Brandini F P et al,1994.The taxonomic identity of the cosmopolitan prymnesiophyte Phaeocystis:a morphological and ecophysiological approach.Journal of Marine Systems,5(1):5-22
Breton E,Brunet C,Sautour B et al,2000.Annual variations of phytoplankton biomass in the Eastern English Channel:comparison by pigment signatures and microscopic counts.Journal of Plankton Research,22(8):1423-1440
Buma A G J,Bano N,Veldhuis M J W et al,1991.Comparison of the pigmentation of 2 strains of the Prymnesiophyte Phaeocystis sp.Netherlands Journal of Sea Research,27(2):173-182
Charlson R J,Lovelock J E,Andreae M O et al,1987.Oceanic phytoplankton,atmospheric sulphur,cloud albedo and climate.Nature,326(6114):655-661
Fragoso G M,Poulton A J,Yashayaev I M et al,2017.Spring phytoplankton communities of the Labrador Sea(2005-2014):pigment signatures,photophysiology and elemental ratios.Biogeosciences,14(5):1235-1259
Gibb S W,Barlow R G,Cummings D G et al,2000.Surface phytoplankton pigment distributions in the Atlantic Ocean:an assessment of basin scale variability between 50°N and50°S.Progress in Oceanography,45(3-4):339-368
Gibberd M J,Kean E,Barlow R et al,2013.Phytoplankton chemotaxonomy in the Atlantic sector of the Southern Ocean during late summer 2009.Deep Sea Research Part I:Oceanographic Research Papers,78:70-78
Jeffrey S W,Wright S W,1994.Photosynthetic pigments in the Haptophyta.In:Green J C,Leadbeater B S C eds.The Haptophyte Algae.Oxford,UK:Clarendon Press,111-132
Liu S X,Yu Z G,Yao P et al,2011.Effects of irradiance on pigment signatures of harmful algae during growth process.Acta Oceanologica Sinica,30(6):46-57
Mackey M D,Mackey D J,Higgins H W et al,1996.CHEMTAX-a program for estimating class abundances from chemical markers:application to HPLC measurements of phytoplankton.Marine Ecology Progress Series,144:265-283
Muylaert K,Gonzales R,Franck M et al,2006.Spatial variation in phytoplankton dynamics in the Belgian coastal zone of the North Sea studied by microscopy,HPLC-CHEMTAXand underway fluorescence recordings.Journal of Sea Research,55(4):253-265
Schoemann V,Wollast R,Chou L et al,2001.Effects of photosynthesis on the accumulation of Mn and Fe by Phaeocystis colonies.Limnology and Oceanography,46(5):1065-1076
Schoemann V,Becquevort S,Stefels J et al,2005.Phaeocystis blooms in the global ocean and their controlling mechanisms:a review.Journal of Sea Research,53(1-2):43-66
Seoane S,Zapata M,Orive E,2009.Growth rates and pigment patterns of haptophytes isolated from estuarine waters.Journal of Sea Research,62(4):286-294
Smith W O Jr,Codispoti L A,Nelson D M et al,1991.Importance of Phaeocystis blooms in the high-latitude ocean carbon cycle.Nature,352(6335):514-516
Stefels J,Van Leeuwe M A,1998.Effects of iron and light stress on the biochemical composition of Antarctic Phaeocystis sp.(Prymnesiophyceae).I.Intracellular DMSP concentrations.Journal of Phycology,34(3):486-495
Stuart V,Sathyendranath S,Head E J H et al,2000.Bio-optical characteristics of diatom and prymnesiophyte populations in the Labrador Sea.Marine Ecology Progress Series,201:91-106
Van Leeuwe M A,Visser R J W,Stefels J,2014.The pigment composition of Phaeocystis antarctica(Haptophyceae)under various conditions of light,temperature,salinity,and iron.Journal of Phycology,50(6):1070-1080
Vaulot D,Birrien J L,Marie D et al,1994.Morphology,ploidy,pigment composition,and genome size of cultured strains of Phaeocystis(Prymnesiophyceae).Journal of Phycology,30(6):1022-1035
Wright S W,Jeffrey S W,1987.Fucoxanthin pigment markers of marine phytoplankton analysed by HPLC and HPTLC.Marine Ecology Progress Series,38:259-266
Wright S W,Thomas D P,Marchant H J et al,1996.Analysis of phytoplankton of the Australian sector of the Southern Ocean:Comparisons of microscopy and size-frequency data with interpretations of pigment HPLC data using the CHEMTAX matrix factorisation program.Marine Ecology Progress Series,144:285-298
Wright S W,Van Den Enden R L,Pearce I et al,2010.Phytoplankton community structure and stocks in the Southern Ocean(30°-80°E)determined by CHEMTAXanalysis of HPLC pigment signatures.Deep Sea Research Part II:Topical Studies in Oceanography,57(9-10):758-778
Zapata M,Rodríguez F,Garrido J L,2000.Separation of chlorophylls and carotenoids from marine phytoplankton:a new HPLC method using a reversed phase C8 column and pyridine-containing mobile phases.Marine Ecology Progress Series,195:29-45
Zapata M,Jeffrey S W,Wright S W et al,2004.Photosynthetic pigments in 37 species(65 strains)of Haptophyta:implications for oceanography and chemotaxonomy.Marine Ecology Progress Series,270:83-102
Zapata M,2005.Recent advances in pigment analysis as applied to picophytoplankton.Vie et Milieu,55(3-4):233-248
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