海洋硅质化超微型浮游植物迅游藻（Bolidophyceae）的分类和生态学研究
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摘要
超微型浮游植物(大小为0.2～2μm)在海洋生态系统中起着关键作用,也是海洋生态系统能量流动和物质循环的重要组成部分,特别是在寡营养海区初级生产力方面起着主导地位,对硅、碳颗粒物在大洋深层的输出具有重要意义。目前对超微型浮游植物的研究还远远落后于实际需要,主要是因为该类群难以分离培养、鉴定和定量测定。迅游藻(Bolidophyceae)是超微型浮游植物中的一类真核生物,它包括迅游单胞藻目(Bolidomonadales)和帕玛藻目(Parmales)。尽管迅游藻对浮游植物群落的细胞丰度贡献相对较小(<0.1%),但是其分布广泛,是沿海和上升流区域中最丰富的浮游植物类群之一,普遍存在于热带到极地海洋生态系统中。迅游藻是系统发育上最接近硅藻的原生藻群,不同的是迅游藻细胞壁没有全部的硅质化。本文阐述了迅游藻的研究现状,总结了迅游藻的发现,以及近年来通过对迅游藻系统发育(基于SSU rDNA和rbc L基因的序列分析)、生态学和生理学研究获得的最新信息,详细说明了迅游藻作为硅藻姊妹群的分类学地位,对于厘清这些硅化藻类之间的进化联系有重要意义。同时还强调了迅游藻在海洋硅碳循环中的作用,对海洋超微型浮游植物在全球生物地球化学循环中的地位做出了进一步的说明。
Picophytoplankton(0.2～2 μm) is an important part of the energy flow and material cycle of marine ecosystems,thus plays a key role in marine ecosystem. Picophytoplankton contributes greatly to primary productivity in oligotrophic waters, which is of great significance to the export of silicon and carbon particulates in the deep ocean. At present, the research on picophytoplankton can not meet the actual needs, because it is difficult to culture, identify and quantify. Bolidophyceae is a group of picoeukaryotes, including Bolidomonadales and Palmales. Although the contribution of Bolidophyceae to phytoplankton abundance is small(< 0.1%), it is widely distributed and is one of the most abundant phytoplankton in coastal and upwelling regions. Bolidohyceae spreads widely throughout marine ecosystem, from tropical to polar regions. Bolidophyceae is the Stramenopiles group phylogenetically nearest to the diatoms. Unlike diatoms, Bolidophyceae cell walls are not completely silicified. This paper describes the state of art of Bolidophyceae, summarizes its discovery, and the current information on taxonomy(sequence analysis based on SSU r DNA and rbc L genes), ecology and physiology. It explains in detail the taxonomic status of Bolidophyceae as a sister group of diatoms, which is also important to clarify the evolutionary relationship among these silicated alga. At the same time, this paper also emphasizes the role of Bolidophyceae in marine silicon-carbon coupling cycle, and further explains the status of marine picophytoplankton in the global biogeochemical cycle.
Picophytoplankton(0.2～2 μm) is an important part of the energy flow and material cycle of marine ecosystems,thus plays a key role in marine ecosystem. Picophytoplankton contributes greatly to primary productivity in oligotrophic waters, which is of great significance to the export of silicon and carbon particulates in the deep ocean. At present, the research on picophytoplankton can not meet the actual needs, because it is difficult to culture, identify and quantify. Bolidophyceae is a group of picoeukaryotes, including Bolidomonadales and Palmales. Although the contribution of Bolidophyceae to phytoplankton abundance is small(< 0.1%), it is widely distributed and is one of the most abundant phytoplankton in coastal and upwelling regions. Bolidohyceae spreads widely throughout marine ecosystem, from tropical to polar regions. Bolidophyceae is the Stramenopiles group phylogenetically nearest to the diatoms. Unlike diatoms, Bolidophyceae cell walls are not completely silicified. This paper describes the state of art of Bolidophyceae, summarizes its discovery, and the current information on taxonomy(sequence analysis based on SSU r DNA and rbc L genes), ecology and physiology. It explains in detail the taxonomic status of Bolidophyceae as a sister group of diatoms, which is also important to clarify the evolutionary relationship among these silicated alga. At the same time, this paper also emphasizes the role of Bolidophyceae in marine silicon-carbon coupling cycle, and further explains the status of marine picophytoplankton in the global biogeochemical cycle.
引文
Andersen R A,Bidigare R R,Keller M D,et al,1996.A comparison of HPLC pigment signatures and electron microscopic observations for oligotrophic waters of the North Atlantic and Pacific Oceans.DeepSea Research.Part II Top.Stud.Oceanogr,43:517-537.
Bailey J C,Bldigare R R,Christensen S J,et al,1995.Phaeothamniophyceae classis nova:a new lineage of chromophytes based upon photosynthetic pigments,rbc L sequence analysis and ultrastructure.Protist,149:245-263.
Booth B C,Marchant H J,1987.Parmales,a new order of marine Chrysophytes,with descriptions of three new genera and seven new species.Joural of Phycology,23:245-260.
Booth B C,Marchant H J,1988.Triparmaceae,a substitute name for a family in the order Parmales(Chrysophyceae).Joural of Phycology,24:124.
Bravo-Sierra E,Hernández-Becerril D U,2003.Parmales(Chrysophyceae)from the Gulf of Tehuantepec,Mexico,including the description of a new species,Tetraparma insecta sp Nov.,and a proposal to the taxonomy of the group.Joural of Phycology,39:577-583.
Daugbjerg N,Guillou L,2001.Phylogenetic analyses of Bolidophyceae(Heterokontophyta)using rbc L gene sequences support their sister group relationship to diatoms.Phycologia,40(2):153-161.
Derelle R,López-García P,Timpano H,et al,2016.A phylogenomic framework to study the diversity and evolution of stramenopiles(=Heterokonts).Molecular.Biology.Evolution,33:2890-2898.
Drebes G,1997.“Sexuality,”in The Biology of Diatoms,Oxford:Blackwell Scientific Publications,250-283.
Edgar L A,Pickett-Heaps J D,1984.Valve morphogenesis in the pennate diatom Navicula cuspidata.Joural of Phycology,,20:47-61.
Guillou L,2011.Characterization of the Parmales:much more than the resolution of a taxonomic enigma.Joural of Phycology,,47:2-4.
Guillou L,Chrétiennot-Dinet M J,Medlin L K,et al,1999a.Bolidomonas:a new genus with two species belonging to a new algal class,the Bolidophyceae(Heterokonta).Joural of Phycology,,35:368-381.
Guillou L,Claustre H,Vaulot D,et al,1999b.Diversity and Abundance of Bolidophyceae(Heterokonta)in Two Oceanic Regions.Environmental.Microbiology,4528-4536.
Ichinomiya M,Dos Santos A L,Gourvil P,et al,2016.Diversity and oceanic distribution of the Parmales(Bolidophyceae),a picoplanktonic group closely related to diatoms.Microbial.Ecology,10:2419-2434.
Ichinomiya M,Yoshikawa S,Kamiya M,et al,2011.Isolation and characterization of Parmales(Heterokonta/Heterokontophyta/Stramenopiles)from the Oyashio region,Western North Pacific.Joural of Phycology,47:144-151.
Idei M,Osada K,Sato S,et al,2012.Gametogenesis and auxospore development in Actinocyclus(Bacillariophyta).Plos One,7:e41890.
Kawachi M,Inouye I,Honda D,et al,2000.The Pinguiophyceae classis nova,chromophytes algae producing large amounts of omega-3 fatty acids.Joural of Phycology,36:35.
Kessenich C R,Ruck E C,SchurkoA M,et al,2014.Transcriptomic Insights into the Life History of Bolidophytes,the Sister Lineage to Diatoms.Joural of Phycology,50:977-983.
Konno S,Jordan R W,2007a.An amended terminology for the Parmales(Chrysophyceae).Phycologia,46:612-616.
Konno S,Ohira R,Harada N,et al,2007b.Six new taxa of subarctic Parmales(Chrysophyceae).Joural of Nannoplankton Research,29:108-128.
Kosman C A,Thomsen H A,and覫stergaard J B,1993.Parmales(Chrysophyceae)from Mexican,Californian,Baltic,Arctic and Antarctic waters with the description of a new subspecies and several new forms.Phycologia,32:116-128.
Kuwata A,Yamada K,Ichinomiya M,2018.Bolidophyceae,a sister picoplanktonic group of diatoms-a review.Marine.Biogeochemistry,5.
Lee M,Li C W,1992.The origin of the silica deposition vesicle of diatoms.Bot.Bull.Acad.Sin,33:317-325.
Mann D,Marchant H,1989.The origins of the diatom and its life cycle.The Chromophyte Algae:Problems and Perspectives.Oxford,Clarendon Press,307-323.
Marchant H J,McEldowney A,1986.Nanoplanktonic siliceous cysts from Antarctica are algae.Marine Biology,92:53-57.
Moestrup覫,1982.Flagellar structure in algae:a review,with new observations particularly on the Chrysophyceae,Phaeophyceae(Fucophyceae),Euglenophyceae,and Reckertia.Phycologia,21:427-528.
Preisig H R,1989.The flagellar base ultrastructure and phylogeny of chromophytes.Tn:The chromophyte algae:problems and perspectives.167-187.
Preisig H R,1994.Siliceous structures and silicification in flagellated protists.Protoplasma,181:29-42.
Riisberg I,Orr R J S,Kluge R,et al,2009.Seven gene phylogeny of Heterokonts.Protist,160:191-204.
Sara L.Thomas,Douglas A.Campbell,2013.Photophysiology of Bolidomonas pacifica.J.Plankton Res,35(2):260-269.
Saunders G W,Poter D,Paskind M P,et al,1995.Cladistic analyses of combined traditional and molecular data sets reveal an algal lineage.Proceedings of the National Academy of Sciences of the United States of America,92:244-248.
Secq M P O Le,Green B R,2011.Complex repeat structures and novel features in the mitochondrial genomes of the diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana.Gene,476:20-26.
譒evèíkováT,Horák A,Klime觢V,et al,2015.Updating algal evolutionary relationships through plastid genome sequencing:did alveolate plastids emerge through endosymbiosis of an ochrophyte?Scientific Reports,5:10134.
Silver M W,Mitchell J G,Ringo D L,1980.Siliceous nanoplankton.II.Newly discovered cysts and abundant choanoflagellates from the Weddell Sea,Antarctica.Marine Biology,58:211-217.
Simpson T L,Volcani B E,1981.Silicon and Siliceous Structures in Biological Systems,Hamburg:Springer Verlag Gmbh.
Stoermer E F,Pankratz H S,and Bowen C C,1965.Fine structure of the diatom Amphipleura pellucida.II.Cytoplasmic fine structure and frustule formation.American Joural of Botany,52,1067-1078.
Tajima N,Saitoh K,Sato S,et al,2016.Sequencing and analysis of the complete organellar genomes of Parmales,a closely related group to Bacillariophyta(diatoms).Current.Genetics,62:887-896.
Vaulot D,Eikrem W,Viprey M,et al,2008.The diversity of small eukaryotic phytoplankton(≤3μm)in marine ecosystems.FEMS Microbiology.Reviews,32:795-820.
Véronique Martin-Jézéquel,2000.Silicon metabolism in diatoms:implications for growth.Journal of Phycology,36:821-840.
Waterbury J B,Watson S W,Guillard R R L,et al,1979.Widespread occurrence of a unicellular,marine,planktonic,cyanobacterium.Nature,277:293-294.
Yamada K,Yoshikawa S,Ichinomiya M,et al,2014.Effects of siliconlimitation on growth and morphology of Triparma laevis NIES-2565(Parmales,Heterokontophyta).Plos One,9:e103289.
Yamada K,Yoshikawa S,Ohki K,et al,2016.Ultrastructural analysis of siliceous cell wall regeneration in the stramenopile Triparma laevis(Parmales,Bolidophyceae).Phycologia,55:602-609.
Yang E C,Boo G H,Kim H J,et al,2012.Supermatrix data highlight the phylogenetic relationships of photosynthetic Stramenopiles.Protist,163:217-231.
高月鑫,江志兵,曾江宁,等,2018.春季长江口北支邻近海域浮游植物群落及其影响因子.海洋通报,37(4):430-439.
郭术津,李彦翘,张翠霞,等,2014.渤海浮游植物群落结构及与环境因子的相关性分析.海洋通报,33(1):95-105.
李冠国,范振刚,等,2011.海洋生态学(第二版),北京:高等教育出版社,74-82.
李洪武,宋培学,等,2012.海洋浮游生物学,合肥:中国科学技术大学出版社,7-14.
李佳俊,谭烨辉,周林滨,等,2016.南海东北部贫营养海区营养盐对浮游植物生长的限制.海洋通报,35(5):562-570.
刘涛,等,2017.藻类系统学,北京:海洋出版社,255-259.
钱罡,魏玉秋,孙军,等,2017.2015年春季桑沟湾浮游植物群落研究.海洋科学,41(2):44-52.
孙军,魏玉秋,2018.聚球藻硅质化作用初探.生态学报,38(14):5234-5243.
杨阳,孙军,关翔宇,等,2016.渤海网采浮游植物群集的季节变化.海洋通报,35(2):121-131.
Bailey J C,Bldigare R R,Christensen S J,et al,1995.Phaeothamniophyceae classis nova:a new lineage of chromophytes based upon photosynthetic pigments,rbc L sequence analysis and ultrastructure.Protist,149:245-263.
Booth B C,Marchant H J,1987.Parmales,a new order of marine Chrysophytes,with descriptions of three new genera and seven new species.Joural of Phycology,23:245-260.
Booth B C,Marchant H J,1988.Triparmaceae,a substitute name for a family in the order Parmales(Chrysophyceae).Joural of Phycology,24:124.
Bravo-Sierra E,Hernández-Becerril D U,2003.Parmales(Chrysophyceae)from the Gulf of Tehuantepec,Mexico,including the description of a new species,Tetraparma insecta sp Nov.,and a proposal to the taxonomy of the group.Joural of Phycology,39:577-583.
Daugbjerg N,Guillou L,2001.Phylogenetic analyses of Bolidophyceae(Heterokontophyta)using rbc L gene sequences support their sister group relationship to diatoms.Phycologia,40(2):153-161.
Derelle R,López-García P,Timpano H,et al,2016.A phylogenomic framework to study the diversity and evolution of stramenopiles(=Heterokonts).Molecular.Biology.Evolution,33:2890-2898.
Drebes G,1997.“Sexuality,”in The Biology of Diatoms,Oxford:Blackwell Scientific Publications,250-283.
Edgar L A,Pickett-Heaps J D,1984.Valve morphogenesis in the pennate diatom Navicula cuspidata.Joural of Phycology,,20:47-61.
Guillou L,2011.Characterization of the Parmales:much more than the resolution of a taxonomic enigma.Joural of Phycology,,47:2-4.
Guillou L,Chrétiennot-Dinet M J,Medlin L K,et al,1999a.Bolidomonas:a new genus with two species belonging to a new algal class,the Bolidophyceae(Heterokonta).Joural of Phycology,,35:368-381.
Guillou L,Claustre H,Vaulot D,et al,1999b.Diversity and Abundance of Bolidophyceae(Heterokonta)in Two Oceanic Regions.Environmental.Microbiology,4528-4536.
Ichinomiya M,Dos Santos A L,Gourvil P,et al,2016.Diversity and oceanic distribution of the Parmales(Bolidophyceae),a picoplanktonic group closely related to diatoms.Microbial.Ecology,10:2419-2434.
Ichinomiya M,Yoshikawa S,Kamiya M,et al,2011.Isolation and characterization of Parmales(Heterokonta/Heterokontophyta/Stramenopiles)from the Oyashio region,Western North Pacific.Joural of Phycology,47:144-151.
Idei M,Osada K,Sato S,et al,2012.Gametogenesis and auxospore development in Actinocyclus(Bacillariophyta).Plos One,7:e41890.
Kawachi M,Inouye I,Honda D,et al,2000.The Pinguiophyceae classis nova,chromophytes algae producing large amounts of omega-3 fatty acids.Joural of Phycology,36:35.
Kessenich C R,Ruck E C,SchurkoA M,et al,2014.Transcriptomic Insights into the Life History of Bolidophytes,the Sister Lineage to Diatoms.Joural of Phycology,50:977-983.
Konno S,Jordan R W,2007a.An amended terminology for the Parmales(Chrysophyceae).Phycologia,46:612-616.
Konno S,Ohira R,Harada N,et al,2007b.Six new taxa of subarctic Parmales(Chrysophyceae).Joural of Nannoplankton Research,29:108-128.
Kosman C A,Thomsen H A,and覫stergaard J B,1993.Parmales(Chrysophyceae)from Mexican,Californian,Baltic,Arctic and Antarctic waters with the description of a new subspecies and several new forms.Phycologia,32:116-128.
Kuwata A,Yamada K,Ichinomiya M,2018.Bolidophyceae,a sister picoplanktonic group of diatoms-a review.Marine.Biogeochemistry,5.
Lee M,Li C W,1992.The origin of the silica deposition vesicle of diatoms.Bot.Bull.Acad.Sin,33:317-325.
Mann D,Marchant H,1989.The origins of the diatom and its life cycle.The Chromophyte Algae:Problems and Perspectives.Oxford,Clarendon Press,307-323.
Marchant H J,McEldowney A,1986.Nanoplanktonic siliceous cysts from Antarctica are algae.Marine Biology,92:53-57.
Moestrup覫,1982.Flagellar structure in algae:a review,with new observations particularly on the Chrysophyceae,Phaeophyceae(Fucophyceae),Euglenophyceae,and Reckertia.Phycologia,21:427-528.
Preisig H R,1989.The flagellar base ultrastructure and phylogeny of chromophytes.Tn:The chromophyte algae:problems and perspectives.167-187.
Preisig H R,1994.Siliceous structures and silicification in flagellated protists.Protoplasma,181:29-42.
Riisberg I,Orr R J S,Kluge R,et al,2009.Seven gene phylogeny of Heterokonts.Protist,160:191-204.
Sara L.Thomas,Douglas A.Campbell,2013.Photophysiology of Bolidomonas pacifica.J.Plankton Res,35(2):260-269.
Saunders G W,Poter D,Paskind M P,et al,1995.Cladistic analyses of combined traditional and molecular data sets reveal an algal lineage.Proceedings of the National Academy of Sciences of the United States of America,92:244-248.
Secq M P O Le,Green B R,2011.Complex repeat structures and novel features in the mitochondrial genomes of the diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana.Gene,476:20-26.
譒evèíkováT,Horák A,Klime觢V,et al,2015.Updating algal evolutionary relationships through plastid genome sequencing:did alveolate plastids emerge through endosymbiosis of an ochrophyte?Scientific Reports,5:10134.
Silver M W,Mitchell J G,Ringo D L,1980.Siliceous nanoplankton.II.Newly discovered cysts and abundant choanoflagellates from the Weddell Sea,Antarctica.Marine Biology,58:211-217.
Simpson T L,Volcani B E,1981.Silicon and Siliceous Structures in Biological Systems,Hamburg:Springer Verlag Gmbh.
Stoermer E F,Pankratz H S,and Bowen C C,1965.Fine structure of the diatom Amphipleura pellucida.II.Cytoplasmic fine structure and frustule formation.American Joural of Botany,52,1067-1078.
Tajima N,Saitoh K,Sato S,et al,2016.Sequencing and analysis of the complete organellar genomes of Parmales,a closely related group to Bacillariophyta(diatoms).Current.Genetics,62:887-896.
Vaulot D,Eikrem W,Viprey M,et al,2008.The diversity of small eukaryotic phytoplankton(≤3μm)in marine ecosystems.FEMS Microbiology.Reviews,32:795-820.
Véronique Martin-Jézéquel,2000.Silicon metabolism in diatoms:implications for growth.Journal of Phycology,36:821-840.
Waterbury J B,Watson S W,Guillard R R L,et al,1979.Widespread occurrence of a unicellular,marine,planktonic,cyanobacterium.Nature,277:293-294.
Yamada K,Yoshikawa S,Ichinomiya M,et al,2014.Effects of siliconlimitation on growth and morphology of Triparma laevis NIES-2565(Parmales,Heterokontophyta).Plos One,9:e103289.
Yamada K,Yoshikawa S,Ohki K,et al,2016.Ultrastructural analysis of siliceous cell wall regeneration in the stramenopile Triparma laevis(Parmales,Bolidophyceae).Phycologia,55:602-609.
Yang E C,Boo G H,Kim H J,et al,2012.Supermatrix data highlight the phylogenetic relationships of photosynthetic Stramenopiles.Protist,163:217-231.
高月鑫,江志兵,曾江宁,等,2018.春季长江口北支邻近海域浮游植物群落及其影响因子.海洋通报,37(4):430-439.
郭术津,李彦翘,张翠霞,等,2014.渤海浮游植物群落结构及与环境因子的相关性分析.海洋通报,33(1):95-105.
李冠国,范振刚,等,2011.海洋生态学(第二版),北京:高等教育出版社,74-82.
李洪武,宋培学,等,2012.海洋浮游生物学,合肥:中国科学技术大学出版社,7-14.
李佳俊,谭烨辉,周林滨,等,2016.南海东北部贫营养海区营养盐对浮游植物生长的限制.海洋通报,35(5):562-570.
刘涛,等,2017.藻类系统学,北京:海洋出版社,255-259.
钱罡,魏玉秋,孙军,等,2017.2015年春季桑沟湾浮游植物群落研究.海洋科学,41(2):44-52.
孙军,魏玉秋,2018.聚球藻硅质化作用初探.生态学报,38(14):5234-5243.
杨阳,孙军,关翔宇,等,2016.渤海网采浮游植物群集的季节变化.海洋通报,35(2):121-131.
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