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寄生于黄颡鱼鳃上的流行小车轮虫研究

2014-07-16胡银亨

江苏农业科学 2014年3期

摘要:流行小车轮虫是一种寄主广泛的寄生虫,对四川省泸州市鲇类寄生车轮虫进行调查,用国际间统一的干银法染色显示车轮虫的附着盘结构和口围绕度,用Lom“统一的特定方法”进行测量,用Basson等方法描述车轮虫的齿体。结果表明,在黄颡鱼鳃上发现的2个种群车轮虫是流行小车轮虫(Trichodinella epzootica),是黄颡鱼的首记录寄主。

关键词:黄颡鱼;流行小车轮虫;卡普小车轮虫;周丛小车轮虫

中图分类号: S941.5 文献标志码: A 文章编号:1002-1302(2014)03-0168-03

1950年Raabe发现了Brachyspira epzootica,1953年Sˇramek-Huek 建立了小车轮虫属,因此把Brachyspira epzootic重命名为流行小车轮虫(Trichodinella epzootica)。1937年,Mueller发现了眉溪车轮虫(Trichodina myakkae),后来重命名为Trichodinella myakkae,它也是流行小车轮虫的同种异名。

流行小车轮虫是一种广泛性寄生车轮虫,欧亚大陆、非洲、北美和澳洲都有报道,它在世界上存在着许多同种异名,在我国,1961年陈启鎏报道的江鳕小车轮虫(T. lotae)就是流行小车轮虫,近年来,许多学者描述的卡普小车轮虫(Trichodinella carpi)和周丛小车轮虫(Trichodinella epzootica)都是流行小车轮虫。

1 材料与方法

寄主黄颡鱼于2009—2012年分别采集于四川省泸县和泸州市江阳区,用黄颡鱼的鳃作涂片,自然干燥后,用国际间统一的干银法染色以显示车轮虫的附着盘结构和口围绕度[1];显影后自然干燥,用高浓度乙醇脱水,用中性胶封片制成永久装片,用凤凰PH50-3A43L-A显微镜观察,借助 MC-D500U(C) 在100倍油镜下拍摄显微照片。

车轮虫形态学特征的描述依据Lom倡导的“统一的特定方法”(Uniform Specific Characteristic System)[2]进行。车轮虫附着盘中齿体定位描述依据Basson等从车轮虫附着盘中央引线至齿钩边缘的“定位描述法”[3]进行;齿体定位图(图1)借助于CorelDRAW X4绘制,并用CorelDRAW X4的标注工具进行数据测量[4]。用Microsoft Excel 2010进行数据统计,一般数据统计模式为:最小值~最大值(平均值±标准差),幅线数和齿体数统计模式为:最小值~最大值(众数)。

小核相对于大核的位置,采用Lom引用Dogiel于1940年提出的标准,即:+y位:小核在大核一臂的外侧;-y位:小核在大核臂端的对立侧面;-y1位:小核在大核一臂的内侧[2]。

2 结果与分析

2.1 种群1

由图2、图3可见,虫体直径21.9~26.1(23.9±1.9) μm,附着盘直径17.9~20.4(19.1±1.1) μm,齿环直径8.5~10.5(9.2±0.8) μm,缘膜2.9 μm,辐线数5根/齿体,齿体数21~22个。齿体纵长5.2~6.0(5.5±0.2) μm,齿长2.5~3.5(3.1±0.3) μm,齿钩长3.2~3.9(3.6±0.2) μm,齿锥宽0.9~1.4(1.2±0.2) μm,齿棘长0.4~0.9(0.8±0.1) μm。齿钩外切缘几乎和缘膜平行,骨突约90°,位置略高于外切缘。齿钩前缘向下弯曲,向前、向后,然后向前形成明显的钩突,钩突超过y-1轴。齿钩连接短而宽,后缘向下行并向前凹,和后面的钩突相嵌。齿锥不超过y轴,x轴上锥形略大于x轴下锥形。齿棘细小,呈弯钩状,染色效果好的标本方可见到,染色效果差的标本不可见。

大核马蹄形,外直径25.1~31.7(28.6±2.9) μm,厚90~9.7(9.4±0.3) μm;小核球形,与大核的位置-y1,直径27~3.0(2.9±0.1) μm。口围绕度159.0~166.0(1625±3.5)°。

2.2 种群2

由图4、图5可见,虫体直径19.7~23.0(21.2±1.2) μm,附着盘直径15.5~18.9(17.1±1.4) μm,齿环直径7.0~9.4(8.4±0.9) μm,缘膜1.7~1.8 μm,辐线数4~5根/齿体,齿体数19~22个。齿体纵长3.8~5.0(4.4±0.3) μm,齿长 1.9~3.2(2.6±0.4) μm,齿钩长 2.6~2.9

(3.2±0.3) μm,齿锥宽06~1.2(0.9±0.2) μm,齿棘长06~0.9(0.6±03) μm。齿钩外切缘几乎和缘膜平行,骨突略高于外节缘。齿钩前缘向下行,略向后,再急剧向前,超过y-1轴,形成较长的钩突。齿钩后缘下行,形成齿钩后突起。齿钩后缘向前凹,和后面的钩突相嵌。齿钩连接短而细。齿锥不超过y轴,x轴上锥形略小于x轴下锥形。齿棘细小,呈弯钩状。

大核马蹄形,外直径17.2~21.7(19.1±1.9) μm,厚44~8.4(6.5±1.7) μm;小核球形,与大核的位置-y1,直径3.1~3.4(3.3±0.1) μm。口围绕度145.0~160.0(1525±75)°。

3 小结和讨论

流行小车轮虫是一种寄主广泛的车轮虫,广泛寄生于河

鲈(Perca fluviatilus)、上口欧鳊(Abramis ballerus,Acerina cernua)、白斑狗鱼(Esox lucius)、江鳕(Lota lota)、鲤(Cyprinus carpio)、鲫(Carassius auratus)、银鲫(C. auratus gibelio)、鲃(Barbus barbus)、粗鳞鳊(Blicca bjoerkna)、暗斑梭鲈(Stizostedion lucioperca)、纵带泥鳅(Misgurnus fossilis)、红大麻哈鱼(Oncorhynchus nerka)、欧飘鱼(Pelecus cultratus)、鳟(Salmo trutta)、丝鳑鲏(Rhodeus sericeus)、红眼鱼(Scardinius erythophthalmus)、丁(Tinca tinca)、罗非鱼(Oncorhynchus nerka)、伦氏罗非鱼(Tilapia rendalli)、褶唇丽鱼(Pseudocrenilabrus philander)、鲃属之Barbus trimaculatus及B.paludinosus上[5-6],在黄颡鱼上还没有报道。本研究在黄颡鱼鳃上发现流行小车轮虫,是黄颡鱼的首记录寄主。

由于流行小车轮虫是一种寄主广泛的车轮虫,存在许多同种异名,有必要进行澄清。Lom等指出“Trichodinella epzootica Raabe,1950”“Trichodinella f. prcarum Lom,1959”“Trichodinella percarum (Dogiel,1940)Chan,1961”“Trichodinella percarum f. lotae Chan,1961”“Trichodinella lotae (Chan,1961)Stein,1962”“Trichodinella maior Reicenbac Klinke,1962”和“Trichodinella carassii (Dogiel,1940)Kostenko,1969 pro parte”均是流行小车轮虫的同种异名[6]。1937年Mueller发现了眉溪车轮虫(Trichodina myakkae)[7],后来重命名为Trichodinella myakkae,我国1973年陈启鎏也报道了此小车轮虫,另外,陈启鎏还报道的江鳕小车轮虫“Trichodinella lotae”“Trichodinella percarum (Dogiel,1940)”“Trichodinella percarum f. lotae”[6]也都是流行小车轮虫[8]。

Stein指出“Trichodina domerguel f. percarum Dogiel,1940,part”“Trichodina percarum sensu Chan,1961”“Brachyspira epzootica Raabe,1950”“B. baltica Stryjecka-Trembaczowska,1953”“Trichodinella epzootica f. prcarum Lom,1959,part”“Trichodinella percarum sensu Chan,1961”“Trichodinella baltica Stein,1962”“Trichodinella carassii Haider,1964,part”“Trichodinella carassii Kostenko,1969,part”和“Trichodinella cyprinid Kaschkovsky,1969”也均为流行小车轮虫的同种异名[10]。

近年来,我国学者描述的卡普小车轮虫[9,11-13](图2)和周丛小车轮虫[9、11](图4),都是流行小车轮虫的不同种群。

Duncan 在菲律宾的鲤(Cyprinus carpio)鳃上发现Trichodinella carpi[14],Duncan以寄主(鲤)命名,我国学者把它翻译成卡普小车轮虫,应该说翻译为鲤小车轮虫更准确。Lom等指出,Trichodinella carpi除齿环直径和齿体数不同于流行小车轮虫外,其余完全相同[6],仅限于齿环直径和齿体数不足以建立一个新的物种,该小车轮虫是流行小车轮虫的1个种群。

Raabe 1950年发现Brachyspira epzootica[15],Sˇ ramek-Huek 1953年建立小车轮虫属,把它重命名为流行小车轮虫(Trichodinella epzootica)[16]。此后,欧洲、非洲、北美洲、澳洲和亚洲都报道了流行小车轮虫[17-27]。“Brachyspira epzootica”“Trichodinella epzootica Raabe,1950”、“Trichodinella epzootica (Raabe,1950)Sˇramek-Huekek,1953”是同一物种,即流行小车轮虫,而我国许多学者把“Trichodinella epzootica (Raabe,1950)Sˇramek-Huek,1953”说成是周丛小车轮虫[9,11-12],当成是小车轮虫的另一个物种,这显然是不正确的。

参考文献:

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[6]Lom J,Haldar D P. Ciliates of the cenera Trichodinella,Tripartiella and Paratrichodina(Peritricha,Mobilina) invading fish gills[J]. Folia Parasitol,1977,24(3):193-210.

[7]Mueller J F. Some species of Trichodina(Ciliata:Peritricha)parasitic on fishes[J]. Trans Amer Micros Soc,1937,56:177-184.

[8]陈启鎏. 病原体分类[M]. 北京:科学出版社,1973:101-110.

[9]赵元莙,唐发辉,唐安科. 重庆地区淡水车轮虫研究 Ⅰ. 小车轮虫、三分虫种类及周丛小车轮虫种群周年动态[J]. 重庆师范大学学报:自然科学版,2007,24(1):1-6.

[10]Stein G A. Guide to the paratites of the freshwater fish fauna of the USSR[M]//Schulman S S. Suborder Mobilina:Vol.1.Parasitic protozoa. Nauka:Bauer O N.1984:322-389.

[11]唐发辉,赵元乔,唐安科.重庆地区鲫鱼外寄生车轮虫的记述及一新种描述[J]. 动物分类学报,2005,30(1):35-40.

[12]李文会,管越强,张耀红,等. 河北白洋淀鱼类寄生小车轮虫属和三分虫属分类学研究[J]. 动物医学进展,2012,33(1):48-52.

[13]Yu S,Tang F,zhao Y. Geographical distribution and diversity of Trichodinid ectoparasites(Ciliophora,Oligohymenophorea,Mobilia)from the gills of fresh and estuarine fishes in Zhejiang Province,China and coastal regions of the East China sea[J]. European Journal of Scientific Research,2011,64:58-70.

[14]Duncan B L. Urceolariid ciliates,including three new species,from cultured Philippine fishes[J]. Trans Amer Micros Soc,1977,96:76-81.

[15]Raabe Z. Uwagio Urceolariidae(ciliate-Peritricha) Skrzel ryb[J]. Annal Univ M Curie-Sklodowska Lublin,1950,5:292-310.

[16]Sˇramek-Huek R. Zur frage der taxonomie und der pathogenitat unserer ektoparasitischen Unceolariiden[J]. Folia Zoologica et Entomologie,1953,2:167-180 .

[17]Lom J. The ciliates of the family Urceolariidae inhabiting gills of fishes(Trichodinella group)[J]. Věst Cˇs Spole cˇ Zool,1963,27:7-19.

[18]Lom J. The adhesive disc of Trichodinella epizootica-ultrastructure and injury to the host tissue[J]. Folia Parasitol,1973,20:193-202.

[19]Lom J,Golemansky V,Grupcheva G. Protozoan parasites of carp(Cyprinus carpio L.):a comparative study of their occurrence in Bulgaria and Czechoslovakia,with the description of Trichodina perforata sp. n.[J]. Folia Parasitologica,1976,23(4):289-300.

[20]Grigoryan D A,Stein G A. Parasitic ciliates(Peritricha:Urceolariidae)infecting fishes of the water boodles of the Armenian SSR[J]. Parazitol,1981,15:305-312.

[21]Basson L,Van As J G,Paperna I. Trichodinid parasites of cichlids and cyprinid fishes of South Africa and Israel[J]. Sys Parasitol,1983,5:245-257.

[22]Al-Rasheid K A S,Ali M A,Sakran T,et al. Trichodinid ectoparasites(Ciliophora:Peritrichida) of some River Nile fish,Egypt[J]. Parasitology International,2000,49:131-137

[23]Lom J,Laird M. Parasitic protozoa from marine and euryhaline fish of Newfoundland and New Brunswick. Ⅰ. Peritrichous ciliates[J]. Can J Zool,1969,47:1367-1380

[24]Basson L. First records of Trichodinid ectoparasites(Ciliophora:Peritrichia)from introduced freshwater fishes in Tasmania,Australia,with comments on pathogenicity[J]. Acta Protozoologica,2010,49:253-265.

[25]Albaladejo J D,Arthur J R. Some trichodinids(Protozoa:Ciliophora:Pertrichida)from freshwater fishes imported into the Philippines[J]. Asian Fisheries Science,1989,3:1-25.

[26]Asmat G S M. Occurrence and morphology of Trichodinella epizootica(Raabe,1950)Sˇrámek-Huek,1953 in India[J]. The Chittagong Univ J Sc,2001,25(2):7-43.

[27]Habib M M A,Asmat G S M. Record of Trichodinella epizootica(reebe,1950) Sˇrámek-Huek(Ciliophora:Trchodinidae) from a major carp,Labeo rohita from Tanguar haorin in Sunamganj[J]. J Asiat Soc Bangladesh Sci,2008,34(1):89-92.

[7]Mueller J F. Some species of Trichodina(Ciliata:Peritricha)parasitic on fishes[J]. Trans Amer Micros Soc,1937,56:177-184.

[8]陈启鎏. 病原体分类[M]. 北京:科学出版社,1973:101-110.

[9]赵元莙,唐发辉,唐安科. 重庆地区淡水车轮虫研究 Ⅰ. 小车轮虫、三分虫种类及周丛小车轮虫种群周年动态[J]. 重庆师范大学学报:自然科学版,2007,24(1):1-6.

[10]Stein G A. Guide to the paratites of the freshwater fish fauna of the USSR[M]//Schulman S S. Suborder Mobilina:Vol.1.Parasitic protozoa. Nauka:Bauer O N.1984:322-389.

[11]唐发辉,赵元乔,唐安科.重庆地区鲫鱼外寄生车轮虫的记述及一新种描述[J]. 动物分类学报,2005,30(1):35-40.

[12]李文会,管越强,张耀红,等. 河北白洋淀鱼类寄生小车轮虫属和三分虫属分类学研究[J]. 动物医学进展,2012,33(1):48-52.

[13]Yu S,Tang F,zhao Y. Geographical distribution and diversity of Trichodinid ectoparasites(Ciliophora,Oligohymenophorea,Mobilia)from the gills of fresh and estuarine fishes in Zhejiang Province,China and coastal regions of the East China sea[J]. European Journal of Scientific Research,2011,64:58-70.

[14]Duncan B L. Urceolariid ciliates,including three new species,from cultured Philippine fishes[J]. Trans Amer Micros Soc,1977,96:76-81.

[15]Raabe Z. Uwagio Urceolariidae(ciliate-Peritricha) Skrzel ryb[J]. Annal Univ M Curie-Sklodowska Lublin,1950,5:292-310.

[16]Sˇramek-Huek R. Zur frage der taxonomie und der pathogenitat unserer ektoparasitischen Unceolariiden[J]. Folia Zoologica et Entomologie,1953,2:167-180 .

[17]Lom J. The ciliates of the family Urceolariidae inhabiting gills of fishes(Trichodinella group)[J]. Věst Cˇs Spole cˇ Zool,1963,27:7-19.

[18]Lom J. The adhesive disc of Trichodinella epizootica-ultrastructure and injury to the host tissue[J]. Folia Parasitol,1973,20:193-202.

[19]Lom J,Golemansky V,Grupcheva G. Protozoan parasites of carp(Cyprinus carpio L.):a comparative study of their occurrence in Bulgaria and Czechoslovakia,with the description of Trichodina perforata sp. n.[J]. Folia Parasitologica,1976,23(4):289-300.

[20]Grigoryan D A,Stein G A. Parasitic ciliates(Peritricha:Urceolariidae)infecting fishes of the water boodles of the Armenian SSR[J]. Parazitol,1981,15:305-312.

[21]Basson L,Van As J G,Paperna I. Trichodinid parasites of cichlids and cyprinid fishes of South Africa and Israel[J]. Sys Parasitol,1983,5:245-257.

[22]Al-Rasheid K A S,Ali M A,Sakran T,et al. Trichodinid ectoparasites(Ciliophora:Peritrichida) of some River Nile fish,Egypt[J]. Parasitology International,2000,49:131-137

[23]Lom J,Laird M. Parasitic protozoa from marine and euryhaline fish of Newfoundland and New Brunswick. Ⅰ. Peritrichous ciliates[J]. Can J Zool,1969,47:1367-1380

[24]Basson L. First records of Trichodinid ectoparasites(Ciliophora:Peritrichia)from introduced freshwater fishes in Tasmania,Australia,with comments on pathogenicity[J]. Acta Protozoologica,2010,49:253-265.

[25]Albaladejo J D,Arthur J R. Some trichodinids(Protozoa:Ciliophora:Pertrichida)from freshwater fishes imported into the Philippines[J]. Asian Fisheries Science,1989,3:1-25.

[26]Asmat G S M. Occurrence and morphology of Trichodinella epizootica(Raabe,1950)Sˇrámek-Huek,1953 in India[J]. The Chittagong Univ J Sc,2001,25(2):7-43.

[27]Habib M M A,Asmat G S M. Record of Trichodinella epizootica(reebe,1950) Sˇrámek-Huek(Ciliophora:Trchodinidae) from a major carp,Labeo rohita from Tanguar haorin in Sunamganj[J]. J Asiat Soc Bangladesh Sci,2008,34(1):89-92.

[7]Mueller J F. Some species of Trichodina(Ciliata:Peritricha)parasitic on fishes[J]. Trans Amer Micros Soc,1937,56:177-184.

[8]陈启鎏. 病原体分类[M]. 北京:科学出版社,1973:101-110.

[9]赵元莙,唐发辉,唐安科. 重庆地区淡水车轮虫研究 Ⅰ. 小车轮虫、三分虫种类及周丛小车轮虫种群周年动态[J]. 重庆师范大学学报:自然科学版,2007,24(1):1-6.

[10]Stein G A. Guide to the paratites of the freshwater fish fauna of the USSR[M]//Schulman S S. Suborder Mobilina:Vol.1.Parasitic protozoa. Nauka:Bauer O N.1984:322-389.

[11]唐发辉,赵元乔,唐安科.重庆地区鲫鱼外寄生车轮虫的记述及一新种描述[J]. 动物分类学报,2005,30(1):35-40.

[12]李文会,管越强,张耀红,等. 河北白洋淀鱼类寄生小车轮虫属和三分虫属分类学研究[J]. 动物医学进展,2012,33(1):48-52.

[13]Yu S,Tang F,zhao Y. Geographical distribution and diversity of Trichodinid ectoparasites(Ciliophora,Oligohymenophorea,Mobilia)from the gills of fresh and estuarine fishes in Zhejiang Province,China and coastal regions of the East China sea[J]. European Journal of Scientific Research,2011,64:58-70.

[14]Duncan B L. Urceolariid ciliates,including three new species,from cultured Philippine fishes[J]. Trans Amer Micros Soc,1977,96:76-81.

[15]Raabe Z. Uwagio Urceolariidae(ciliate-Peritricha) Skrzel ryb[J]. Annal Univ M Curie-Sklodowska Lublin,1950,5:292-310.

[16]Sˇramek-Huek R. Zur frage der taxonomie und der pathogenitat unserer ektoparasitischen Unceolariiden[J]. Folia Zoologica et Entomologie,1953,2:167-180 .

[17]Lom J. The ciliates of the family Urceolariidae inhabiting gills of fishes(Trichodinella group)[J]. Věst Cˇs Spole cˇ Zool,1963,27:7-19.

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