车轮虫齿体特征的量化及其种内种间系统发育的研究
QUANTITATIVE ANALYSIS OF TRICHODINA DENTICLATING CHARACTERS AND PHYLOGENETIC RELATIONSHIP STUDIES ON INTERSPECIES AND INTRASPECIES
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摘要: 齿体特征是车轮虫物种鉴定的一个重要依据.本文首次提出一种齿体结构面积比例量化法,即通过对齿钩、齿锥、齿棘、齿体环各部分反口面观面积的测量,分别计算出前三者与所在齿体的面积比以及所有齿体总面积与齿体环的面积比,从而将齿体特征转换为量化指标.依据所测量化值,采用相似性聚类法,我们首先探讨了显著车轮虫(Trichodina nobbillis Chen,163)的三个种群的亲缘关系,然后以劳牧小车轮虫(Trichodinella lomi Xu,Song & Warren, 1)为外类群,选择显著车轮虫(Trichodina nobillis Chen,163)、异齿车轮虫(Trichodina heterodentata Duncan,177)、杜氏车轮虫(Trichodina dormergui Wallengren,187)和Trichodina fahaka(Khaled A.S.Al-Rasheid,2000)为研究对象,对车轮虫种间亲缘关系进行了分析.结果表明,Pdr值可以有效地表现种群间的差异;同时还发现齿棘面积比例较高的杜氏车轮虫和异齿车轮虫进化较晚,而齿棘面积比例较低的显著车轮虫进化较早,并得出了齿棘越发达,车轮虫越进化这一结论,支持了徐奎栋等所提出的游走亚目六种基本齿型的进化顺序.这种量化研究方法不但可对车轮虫齿体的各部分结构以及它们之间的相互关系进行定量描述,而且还可以这些数据为基础对车轮虫种群间及物种间亲缘关系进行研究.Abstract: Since Ehrenberg(1838)firstly reported Tricodina pediculus,there were two milestones in the research work on trichodinids. The first was the widespread acceptance of the uniform specific characteristics proposed by Lom(1958),which resulted in a standardization of the taxonomy of trichodinids. The second was the appliance of guidelines for elucidating the shape of denticles advanced by Van As Basson(1989),which can also be used to re evaluate previous work on trichodinids and especially to distinguish denticles between closely related species. As an important diagnosis marker,denticlating character has been widely used in species identification of the trichodine ciliates. In this paper,we firstly introduced a quantitative method of area percentage to describe denticlating. Quantitative description was achieved by measuring the percentages of the respective area of blade,central part and thorn to the denticle’s and the total area of denticles to the denticlating ring’s from aboral view. Basing on these quntitative values,first,we discussed the phylogenetic relationship of intraspecies of Trichodina nobillis from three populations; then we selected Trichodinella lomi as an outer group,and tentatively performed fuzzy clustering analysis on Trichodina nobillis, Trichodina heterodentata, Trichodina dormergui, and Trichodina fahaka, and studied the phylogenetic relationship of intraspecies among the four trichodinids. The results revealed the Pdr(Percentage of the area of denticles to the denticlating ring’s) showed distinct diference in interspecies, and we also found Trichodina heterodentata and Trichodina dormergui with higher Ptd(Percentage of thorn area to dentice’s) formed a sister group, and evolved lately, but Trichodina nobillis with lower Ptd evolved more early relatively. So, we concluded that the higher the Ptd was, the more lately Trichodina evolved. The conclusion approved the theory of evolutionary order of 6 types of denticles of Mobilina that Kuidong Xu et al. had put forward. It may not only describe quantitatively the structure of the three parts of the denticle and their interrelation, but also provide a method for the research of Trichodina phylogeny by this quantitative mean.
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Keywords:
- Trichodina /
- Quantitative analysis /
- Phylogeny relationship
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analysis[J].J.Agricul.Eng.,1997,12:278-281.[吕志南、吕艳燕.模糊聚类分析在研究生物进化系统中的应用.农业工程学报,1997,12:278-281.][26] Miao W.Molecular Phylogeny of Peritrichous Ciliates[D].Wuhan: Institute of Hydrobiology,The Chinese Academy of Sciences,2002.[缪炜.缘毛类纤毛虫分子系统发育学研究.武汉:中国科学院水生生物研究所,2002.][27] Lei A P,Shi Z X,Wei X Y.Diversity of the phyloplankton in Donghu Lake,Wuhan[J].Acta Hydrobiologica Sinica,2003,27:179-184.[雷安平施之新魏心印.武汉东湖浮游藻类物种多样性的研究.水生生物学报,2003,27:179-184.] Asmat G S M.Trichodina canningensis sp.n.(Ciliophora:Trichodinidae)from an Indian Estuarine Fish,Mystus gulio(Hamilton)(Bagridae)[J].Acta protozool.,2001 40:147-151[2] Xu K D,Song W B,Warren.Trichodinid ectoparasites(Ciliophora:Peritricha)from the gills of maricultue molluscs in China,with the descriptions of four new species of Trichodina Ehrenberg,1838[J].Syst.Parasitol.,1999b,42:229-237[3] Xu K D,Song W B,Warren.Observations on trichodinid ectoparasites(Ciliophora:Peritricha) from the gills of maricultue molluscs in China,with the descriptions of three new species of Trichodina Ehrenberg,1838[J].Syst.Parasitol., 2000,45:17-24[4] Li L X.Descriptions of two new species of Trichodinids from freshwater fishes, Ctenogobius giurnus Retter [J].Acta Hydrobiologica Sinica,2001,25:503-507.[李连祥,（鱼假）虎鱼寄生车轮虫两新种的记述.水生生物学报,2001,25:503-507.][5] Lom J.A contribution to the systematics and morphology of endoparasitic trichodinids from amphibians with a proposal of uniform specific characteristics[J].J.Protozool.,1958,5:251-263[6] Stein G A.New data on Parasitic ciliates of the family Urceolariidae(Peritricha,Mobilina) from some marine invertebrates [J].Zool.Zhurn.,1979,53:965-973[7] Chen Q L.Studies on the ectoparasitic Trichodinids from fishes,tadpoles and crustaceas[J].Acta Hydrobiologica Sinica,1963,3:99-111.[陈启鎏.鱼类、蝌蚪和甲壳动物体外寄生车轮虫(Trichodina)的研究.水生生物学集刊,1963,3:99-111.][8] Van As J G,Basson L.A further contribution to the taxonomy of the Trichodinidae (Ciliophhora: Peritricha)and a review of the taxonomic status of some fish ectoparasitic trichodinids[J].Syst.Parasitol.,1989,14: 157-179[9] Lom J.The adhesive disc of Trichodinella epizootica-ultrastructure and injury to the host tissue[J].Folia Parasitol.,1973b,20:913-202[10] Hausmann K,Hausmann E.Structural studies on Trichodina pediculus(Ciliophora,Peritricha).Ⅰ.The locomotor fringe and the oral apparatus[J].Journal of Ultrastructure Research,1981a,74:131-143[11] Hausmann K,Hausmann E.Structural studies on Trichodina pediculus (Ciliophora,Peritricha).Ⅱ.The adhesive disc[J].Journal of Ultrastructure Research,1981b,74:144-155[12] Van As J G,Basson L.An articulated internal skeleton resembling a spinal column in a ciliated protozoan[J].Syst.Parasit.,1990,14:157-179[13] Urawa S,Arthur J R.First record of the parasitic ciliate Trichodina truttae Mueller,1937 on chum salumon fry (Oncorhynchus keta )from Japan[J].Gyobyo Kenkyu,1991,26:83-89[14] Kruger J,Basson L,Van As J G.On the ultrastructure of the adhesive disc of Trichodina xenofodos Fantham,1924 and T.heterodentata Duncan,1977 (Ciliophhora:Peritricha)[J]Acta Protozool., 1993,32:245-253[15] Gaze W H,Wooten R.An SEM study of adhesive disc skeletal structures isolated from trichodinids (Ciliophora:Peritrichida) of the genera Trichodina Ehrenberg,1838 and Paratrichodina Lom, 1963[J].Syst.Parasit.,1999,43:167-174[16] Kruger J,Basson L,Van As J.Observations on the adhesive disc of Trichodina xenofodos Fantham,1924 and T.heterodentata Duncan,1977 during binary fission (Ciliophhora: Peritricha)[J].Acta Protozool.,1995, 34:203-209[17] Li LX.Observations on Ultrastructre of Trichodina algonquiensis Li et Desser,1983 and its location of classification[C].Proceedings of the 11th symposium of the Chinese Protozoological society(abstracts).Wuhan:2001,134-139.[李连祥.阿冈车轮虫的超微结构观察及其分类位置.中国原生动物学学会第十一次学术讨论会(论文摘要汇编),武汉:2001,134-139.][18] Song W B.Progress in Protozoology[M].Qingdao:Qingdao Ocean University Press,1999.[宋微波,原生动物学专论.青岛:青岛海洋大学出版社,1999.][19] Shen Y F.Protozoo1ogy[M].Beijing:Science Press,1999.[沈蕴芬.原生动物学.北京:科学出版社,1999.][20] Institute of Hydrobiology,The Chinese Academy of Sciences.The pathogeny graphy of fishes in Hubei province[M].Beijing:Science Press,1973,194-202[中国科学院水生生物研究所.湖北省鱼病病原区系图志.北京:科学出版社,1973,194-202][21] Xu K D,Song W B.Parasitic Trichodinid Ciliates(Ciliophora,Peritrichida) from Mariculture Fishes Ⅱ, Genara Paratrichodina,Trichodinella and Dipartiella[J].J.of Ocean Univ.Qingdao,2000,30(3):413-417.[徐奎栋宋微波.海水养殖鱼类外寄生车轮虫Ⅱ.拟车轮虫属、小车轮虫属、及两分虫属.青岛海洋大学学报,2000,30(3):413-417[22] Al-Rasheid K A S,Ali M A,Sakran T,et al.Trichodinid ectoparasites(Ciliophora:Peritrichida) of some River Nile fish,Egypt[J].Parasitol.Internat.,2000,49:131-137[23] Ahmet ?iZER.The Occurrence of Trichodina domergui Wallengren,1897 and Trichodina tenuidens Faur é-Fremiet 1944(Peritrichia) on Three-spined Stickleback,Gasterosteus aculeatus L.,1758 Found in a Brackish and Freshwater Environment[J].Acta Protozool.,2003, 42:41-46[24] Xu K D.Parasitic and Commensal Ciliated Protozoa from Marine Molluscs and Fishes off the Coast of the Yellow Sea and the Bohai Bay, with the Review of Mobiline Peritrichous Ciliates(Protozoa,Ciliophora)[D].Qingdao: Ocean University of Qingdao.1999.[徐奎栋.黄渤海常见经济贝类及鱼类的危害性纤毛虫原生动物.青岛:青岛海洋大学,1999.][25] Lv Z N,Lv Y Y.Studies on biology evolution system with fuzzy clustering analysis[J].J.Agricul.Eng.,1997,12:278-281.[吕志南、吕艳燕.模糊聚类分析在研究生物进化系统中的应用.农业工程学报,1997,12:278-281.][26] Miao W.Molecular Phylogeny of Peritrichous Ciliates[D].Wuhan: Institute of Hydrobiology,The Chinese Academy of Sciences,2002.[缪炜.缘毛类纤毛虫分子系统发育学研究.武汉:中国科学院水生生物研究所,2002.][27] Lei A P,Shi Z X,Wei X Y.Diversity of the phyloplankton in Donghu Lake,Wuhan[J].Acta Hydrobiologica Sinica,2003,27:179-184.[雷安平施之新魏心印.武汉东湖浮游藻类物种多样性的研究.水生生物学报,2003,27:179-184.]
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