中国13种海参系统发育和骨片演化分析
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摘要
运用扫描电子显微镜观察了中国沿海13种代表性海参的骨片,并获得了8种海参的线粒体16SrDNA基因序列,通过整合网上序列联合分析以期了解13种海参的系统发育关系和骨片演化规律。结果表明, 13种海参的骨片包括桌形体、假桌形体、扣状体、杆状体、穿孔板、花纹样体、颗粒体7种类型; 8种海参的16S rDNA片段长度在465 bp左右,富含AT碱基, A+T的含量平均为55.8%,符合无脊椎动物线粒体DNA序列的特征。从GenBank中获取11种海参16S rDNA基因序列,以杂色角孔海胆(Salmacis sphaeroides)为外群,联合本研究中所测序列构建了NJ和ME系统树。结果表明,NJ、ME系统发育树基本一致,海参科(Holothuriidae)内,柄体参属(Labidodemas)与海参属(Holothuria)亲缘关系极近且骨片形态均相似,可划分为同一个属;与传统形态学分类不同,刺参科(Stichopodidae)同尻参科(Caudinidae)、沙鸡子科(Phyllophoridae)、硬瓜参科(Sclerodactylidae)、瓜参科(Cucumariidae)聚成一簇,而与海参科亲缘关系较远,符合最新分类系统。本研究根据NJ和ME系统发育树结果,结合海参骨片的形态学资料,对海参科内骨片的演化进行了分析,认为虽然桌形体的进化史尚不清晰,但在海参科内,海参骨片由复杂的桌形体、扣状体向简单的花纹样体和颗粒体演化。
To fully determine the phylogenetic relationship and ossicle evolution of holothurians, the ossicles of 13 representative sea cucumber species in the China Sea were observed by scanning electronic microscopy and the mitochondrial 16 S rDNA gene from eight specimens were amplified and sequenced. The results demonstrated that ossicles of the 13 sea cucumbers could be classified into seven types, which were termed as table, pseudo-table,button, rod, plate, rosette, and oblate grain. The length of 16 S rDNA fragments amplified from eight sea cucumbers was approximately 465 bp. These fragments were enriched in AT bases with an average A+T content of 55.8%,which is in accordance with the characteristics of mitochondrial DNA in invertebrates. 16 S rDNA sequences of 11 sea cucumbers obtained from GenBank were combined with sequences from our study to construct a phylogenetic tree. Salmacis sphaeroides was regarded as a control group. Neighbor-joining(NJ) and minimum evolution(ME)methods provided almost identical results. Genetic relationships between Labidodemas and Holothuria in Holothuriidae were rather close and shapes of their ossicles were also similar, thus they could be divided into the same genus; Caudinidae, Phyllophoridae, Sclerodactylidae, and Cucumariidae were clustered with Stichopodidae, and separated from Holothuriidae. Contrary to traditional taxonomy, our study met the criteria of the newest taxonomy system, wherein Aspidochirotida was removed and Holothuroidea was redivided into seven orders, including Dendrochirotida, Synallactida, Molpadida, Persicalida, Holothuriida, Elasipodida, and Apodida. Based on molecular phylogenetic trees and morphological data of ossicles, we hypothesized that ossicles of sea cucumbers in Holothuriidae underwent slow evolution from a complex shape, such as table and button, to simple patterns, such as rosette and oblate grain. Furthermore, considering that the rod in Holothuria pervicax and the button in Holothuria fuscocinerea, we concluded that ossicles in Holothuriidae underwent slow transition rather than direct evolution.
To fully determine the phylogenetic relationship and ossicle evolution of holothurians, the ossicles of 13 representative sea cucumber species in the China Sea were observed by scanning electronic microscopy and the mitochondrial 16 S rDNA gene from eight specimens were amplified and sequenced. The results demonstrated that ossicles of the 13 sea cucumbers could be classified into seven types, which were termed as table, pseudo-table,button, rod, plate, rosette, and oblate grain. The length of 16 S rDNA fragments amplified from eight sea cucumbers was approximately 465 bp. These fragments were enriched in AT bases with an average A+T content of 55.8%,which is in accordance with the characteristics of mitochondrial DNA in invertebrates. 16 S rDNA sequences of 11 sea cucumbers obtained from GenBank were combined with sequences from our study to construct a phylogenetic tree. Salmacis sphaeroides was regarded as a control group. Neighbor-joining(NJ) and minimum evolution(ME)methods provided almost identical results. Genetic relationships between Labidodemas and Holothuria in Holothuriidae were rather close and shapes of their ossicles were also similar, thus they could be divided into the same genus; Caudinidae, Phyllophoridae, Sclerodactylidae, and Cucumariidae were clustered with Stichopodidae, and separated from Holothuriidae. Contrary to traditional taxonomy, our study met the criteria of the newest taxonomy system, wherein Aspidochirotida was removed and Holothuroidea was redivided into seven orders, including Dendrochirotida, Synallactida, Molpadida, Persicalida, Holothuriida, Elasipodida, and Apodida. Based on molecular phylogenetic trees and morphological data of ossicles, we hypothesized that ossicles of sea cucumbers in Holothuriidae underwent slow evolution from a complex shape, such as table and button, to simple patterns, such as rosette and oblate grain. Furthermore, considering that the rod in Holothuria pervicax and the button in Holothuria fuscocinerea, we concluded that ossicles in Holothuriidae underwent slow transition rather than direct evolution.
引文
[1]Smiley S,Harrison F W,Chia F S.Microscopic Anatomy of Invertebrates:Echinodermata[M].New York:Wiley-Liss Press,1994:401-471.
[2]Liao Y L,Xiao N.Species composition and faunal characteristics of echinoderms in China seas[J].Biodiversity Science,2011,19(6):729-736.[廖玉麟,肖宁.中国海棘皮动物的种类组成及区系特点[J].生物多样性,2011,19(6):729-736.]
[3]Cui G Y,Zhao L.Names and species identifying of edible sea cucumber in China[J].Cuisine Journal of Yangzhou University,2000,17(3):13-18.[崔桂友,赵廉.食用海参的名称与种类鉴别[J].扬州大学烹饪学报,2000,17(3):13-18.]
[4]Wang L,Lu H X,Chen Q.Analysis and evaluation of the nutritional components of sea cucumber(Acaudina molpadioidea)[J].Food and Fermentation Industries,2014,40(8):215-218.[王磊,陆海霞,陈青.东海海参(Acaudina molpadioidea)营养成分分析及评价[J].食品与发酵工业,2014,40(8):215-218.]
[5]Guo Y Y,Ding Y,Xu F F,et al.Progress in research on main bioactive constituents of sea cucumber[J].Food Science,2014,35(15):335-344.[郭盈莹,丁燕,徐飞飞,等.海参中主要生物活性成分研究进展[J].食品科学,2014,35(15):335-344.]
[6]Liao Y L.Fauna Sinica:Phylum Echinodermata,Class Holothuroidea[M].Beijing:Science Press,1997.[廖玉麟.中国动物志:棘皮动物门,海参纲[M].北京:科学出版社,1997.]
[7]Zhao S M.Shallow-Water Sea Cucumber of Taiwan[M].Taichung:Museum of Natural Science,1998.[赵世民.台湾礁岩海岸的海参[M].台中:自然科学博物馆,1998.]
[8]Li Y,Fei L H,Chen J X.The morphology of ossicles of 15commercial holothurians[J].Periodical of Ocean University of China,2008,38(2):211-216.[李赟,费来华,陈家鑫.十五种海参骨片的形态学研究[J].中国海洋大学学报(自然科学版),2008,38(2):211-216.]
[9]Miller A K,Kerr A M,Paulay G,et al.Molecular phylogeny of extant Holothuroidea(Echinodermata)[J].Molecular Phylogenetics and Evolution,2017,111:110-131.
[10]Wen J,Hu C Q,Zhang L P,et al.PCR-RFLP identification of 16 commercial sea cucumber species on the basis of 16Sr RNA gene[J].Journal of Fishery Sciences of China,2011,18(2):451-457.[文菁,胡超群,张吕平,等.16种商品海参16S r RNA的PCR-RFLP鉴定方法[J].中国水产科学,2011,18(2):451-457.]
[11]Kerr A M,Janies D A,Clouse R M,et al.Molecular phylogeny of coral-reef sea cucumbers(Holothuriidae:Aspidochirotida)based on 16S mitochondrial ribosomal DNA sequence[J].Marine Biotechnology,2005,7(1):53-60.
[12]Fei L H,Li Y,Chen J X.16S rDNA sequence diversity and phylogenetic relationship of ten sea cucumbers[J].Journal of Fishery Sciences of China,2008,15(5):755-765.[费来华,李赟,陈家鑫.10种海参16S r DNA序列多样性及其亲缘关系分析[J].中国水产科学,2008,15(5):755-765.]
[13]Wen J,Hu C Q,Fan S G.Molecular phylogeny of 15 sea cucumbers and the inferences about the evolution of ossicles[J].Marine Sciences,2011,35(5):66-72.[文菁,胡超群,范嗣刚.中国15种海参的分子系统发育和骨片演化的分析[J].海洋科学,2011,35(5):66-72.]
[14]Yan J X.The phylogeny of six species sea cucumbers in South China Sea and studies on embryonic and larval development of sea cucumber Holothria scabra and Stichopus variegates[D].Haikou:Hainan University,2012.[严俊贤.六种南海海参的系统发育与糙海参和花刺参的胚胎与幼体发育研究[D].海口:海南大学,2012.]
[15]Kerr A M,Kim J.Phylogeny of Holothuroidea(Echinodermata)inferred from morphology[J].Zoological Journal of the Linnean Society,2001,133(1):63-81.
[16]Chen L M,Li Q,Li Y.Sequence analysis of mitochondrial16S rRNA and COI gene and molecular phylogeny of four species of sea cucumber[J].Journal of Fishery Sciences of China,2008,15(6):935-942.[陈丽梅,李琪,李赟.4种海参16S r RNA和COI基因片段序列比较及系统学研究[J].中国水产科学,2008,15(6):935-942.]
[17]Massin C,Mercier A,Hamel J F.Ossicle change in Holothuria scabra with a discussion of ossicle evolution within the Holothuriidae(Echinodermata)[J].Acta Zoologica(Stockholm),2000,81(1):77-91..
[2]Liao Y L,Xiao N.Species composition and faunal characteristics of echinoderms in China seas[J].Biodiversity Science,2011,19(6):729-736.[廖玉麟,肖宁.中国海棘皮动物的种类组成及区系特点[J].生物多样性,2011,19(6):729-736.]
[3]Cui G Y,Zhao L.Names and species identifying of edible sea cucumber in China[J].Cuisine Journal of Yangzhou University,2000,17(3):13-18.[崔桂友,赵廉.食用海参的名称与种类鉴别[J].扬州大学烹饪学报,2000,17(3):13-18.]
[4]Wang L,Lu H X,Chen Q.Analysis and evaluation of the nutritional components of sea cucumber(Acaudina molpadioidea)[J].Food and Fermentation Industries,2014,40(8):215-218.[王磊,陆海霞,陈青.东海海参(Acaudina molpadioidea)营养成分分析及评价[J].食品与发酵工业,2014,40(8):215-218.]
[5]Guo Y Y,Ding Y,Xu F F,et al.Progress in research on main bioactive constituents of sea cucumber[J].Food Science,2014,35(15):335-344.[郭盈莹,丁燕,徐飞飞,等.海参中主要生物活性成分研究进展[J].食品科学,2014,35(15):335-344.]
[6]Liao Y L.Fauna Sinica:Phylum Echinodermata,Class Holothuroidea[M].Beijing:Science Press,1997.[廖玉麟.中国动物志:棘皮动物门,海参纲[M].北京:科学出版社,1997.]
[7]Zhao S M.Shallow-Water Sea Cucumber of Taiwan[M].Taichung:Museum of Natural Science,1998.[赵世民.台湾礁岩海岸的海参[M].台中:自然科学博物馆,1998.]
[8]Li Y,Fei L H,Chen J X.The morphology of ossicles of 15commercial holothurians[J].Periodical of Ocean University of China,2008,38(2):211-216.[李赟,费来华,陈家鑫.十五种海参骨片的形态学研究[J].中国海洋大学学报(自然科学版),2008,38(2):211-216.]
[9]Miller A K,Kerr A M,Paulay G,et al.Molecular phylogeny of extant Holothuroidea(Echinodermata)[J].Molecular Phylogenetics and Evolution,2017,111:110-131.
[10]Wen J,Hu C Q,Zhang L P,et al.PCR-RFLP identification of 16 commercial sea cucumber species on the basis of 16Sr RNA gene[J].Journal of Fishery Sciences of China,2011,18(2):451-457.[文菁,胡超群,张吕平,等.16种商品海参16S r RNA的PCR-RFLP鉴定方法[J].中国水产科学,2011,18(2):451-457.]
[11]Kerr A M,Janies D A,Clouse R M,et al.Molecular phylogeny of coral-reef sea cucumbers(Holothuriidae:Aspidochirotida)based on 16S mitochondrial ribosomal DNA sequence[J].Marine Biotechnology,2005,7(1):53-60.
[12]Fei L H,Li Y,Chen J X.16S rDNA sequence diversity and phylogenetic relationship of ten sea cucumbers[J].Journal of Fishery Sciences of China,2008,15(5):755-765.[费来华,李赟,陈家鑫.10种海参16S r DNA序列多样性及其亲缘关系分析[J].中国水产科学,2008,15(5):755-765.]
[13]Wen J,Hu C Q,Fan S G.Molecular phylogeny of 15 sea cucumbers and the inferences about the evolution of ossicles[J].Marine Sciences,2011,35(5):66-72.[文菁,胡超群,范嗣刚.中国15种海参的分子系统发育和骨片演化的分析[J].海洋科学,2011,35(5):66-72.]
[14]Yan J X.The phylogeny of six species sea cucumbers in South China Sea and studies on embryonic and larval development of sea cucumber Holothria scabra and Stichopus variegates[D].Haikou:Hainan University,2012.[严俊贤.六种南海海参的系统发育与糙海参和花刺参的胚胎与幼体发育研究[D].海口:海南大学,2012.]
[15]Kerr A M,Kim J.Phylogeny of Holothuroidea(Echinodermata)inferred from morphology[J].Zoological Journal of the Linnean Society,2001,133(1):63-81.
[16]Chen L M,Li Q,Li Y.Sequence analysis of mitochondrial16S rRNA and COI gene and molecular phylogeny of four species of sea cucumber[J].Journal of Fishery Sciences of China,2008,15(6):935-942.[陈丽梅,李琪,李赟.4种海参16S r RNA和COI基因片段序列比较及系统学研究[J].中国水产科学,2008,15(6):935-942.]
[17]Massin C,Mercier A,Hamel J F.Ossicle change in Holothuria scabra with a discussion of ossicle evolution within the Holothuriidae(Echinodermata)[J].Acta Zoologica(Stockholm),2000,81(1):77-91..
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