河南省驴寄生圆线虫的研究
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摘要
圆线虫隶属于杆形目(Rhabditida)圆线亚目(Strongylina)圆线科(Strongylidae),是马属动物体内寄生线虫的一个重要类群。圆线虫主要寄生在大肠内,对宿主的危害较大,往往引起疾病甚至死亡。在国外,关于圆线虫的分类、生活史、致病机理、疾病控制、流行病学、DNA分子诊断及系统发育学等方面的研究比较多,而在国内,除了关于形态学分类的报道较多以外,其它方面较少,甚至仍是空白。本论文首次对河南省驴体内寄生圆线虫的种类、形态描述、流行病学以及分子系统发育进行详细地报道。
从2006年2月至2007年1月调查了河南省34头驴体内寄生线虫的感染情况。通过解剖驴的大肠,共采集到22种圆线虫,分属于2个亚科8个属。此次调查发现,寄生数量较大的种类有普通戴拉风线虫Delafondia vulgaris、鼻状杯环线虫Cylicocyclus nassatus、小唇片冠环线虫Coronocyclus labratus、四刺盅口线虫Cyathostomum tetracanthum、耳状杯环线虫Cylicocylus auriculatus等;而长形杯环线虫Cylicocyclus elongatus、小杯杯冠线虫Cylicostephanus calicatus、艾氏杯环线虫Cylicocyclus adersi、真臂副杯口线虫Parapoteriostomum euproctus、日本三齿线虫Triodontophorus nipponicus、锯齿状三齿线虫Triodontophorus serratus、无齿阿尔夫线虫Alfortia edentatus等的寄生数量相对较小。
本研究在光学显微镜下对每一种圆线虫进行仔细地观察和测量,同时对数量较多的种类做了扫描电镜观察,根据观察结果详细地描述了每种圆线虫的形态特征。运用扫描电镜技术观察和描述圆线虫的形态和结构,在我国尚属首次,同时为该类线虫的分类鉴定提供了新的依据。作者所采的标本与国内外的文献报道相比,其形态、结构基本一致。虽然有些种类在体长、外叶冠数、食道长、交合刺长、阴道长以及尾长等方面存在差异,但是作者认为,这可能是由于调查的对象以及采集的数量、采集的时间和地点不同造成的种内变异。
圆线虫的流行病学调查结果显示:每头驴感染圆线虫的数量变化范围是120条～1939条,平均498.6条;被解剖的驴肠道内,有的仅发现1种圆线虫,有的多达15种,大多数是多种圆线虫混合感染。在此次调查中,普通戴拉风线虫(88.2%)、鼻状杯环线虫(73.5%)、小唇片冠环线虫(70.6%)和大唇片冠环线虫Coronocyclus labiatus(67.6%)是感染率较高的种类,为本次调查的核心种;小唇片冠环线虫(175.9±291.3)、四刺盅口线虫(156.0±251.7)、耳状杯环线虫(127.2±221.7)和鼻状杯环线虫(110.1±124.4)是平均感染强度较高的种类;平均丰度较高的种类是小唇片冠环线虫(124.2±256.4)、四刺盅口线虫(96.4±210.5)和鼻状杯环线虫(80.9±117.1)。此次调查还发现,季节对圆线虫种类的分布和感染情况也有一定的影响。在一年内,寄生种类最多的季节是秋季,其次是春季和夏季,冬季种类最少。不同种类的圆线虫其感染率和平均丰度的季节变化明显不一致,大多数种类的变化趋势是:感染率和平均丰度在春季和秋季出现两个高峰,冬季和夏季往往较低。圆线虫的流行病学调查在国内尚属首次报道。
利用聚合酶链式反应(PCR)技术对7种杯环线虫(辐射杯环线虫Cylicocylus radiatus、艾氏杯环线虫、阿氏杯环线虫Cylicocylus ashworthi、耳状杯环线虫、长形杯环线虫、显形杯环线虫Cylicocylus insignis和鼻状杯环线虫)的核糖体DNA(rDNA)的内转录间隔区(Internal transcribed spacer,ITS)及5.8S rDNA进行扩增、测序。序列分析结果表明,所测样品的ITS1-5.8S-ITS2序列的长度范围为829-843bp,变异位点(variable sites)为88个,简约信息位点(parsimony informative sites)为56个;其中ITS1和ITS2的长度范围分别为366-370bp和307-321bp,变异位点为22个和66个,简约信息位点为14个和42个;7种样品的5.8S序列完全一样,长度为153bp。另外,ITS1区的G+C含量(45.8-47.7%)明显高于ITS2区(39.2-41.9%)。通过序列两两比对,7种杯环线虫ITS1区和ITS2区的种间差异性分别为1.4-3.8%和3.1-14.3%。
基于上述7种线虫和Genebank中已登录的5种杯环线虫(阿氏杯环线虫、短口囊杯环线虫Cylicocylus brevicapsulatus、显形杯环线虫、细口杯环线虫Cylicocylus leptostomus和鼻状杯环线虫)的ITS1和ITS2序列的综合数据,以长伞杯冠线虫作为外群,使用PAUP软件构建NJ(neighbor–joining)和MP(maximum parsimony)系统发育树。结果发现,在NJ树和MP树中,短口囊杯环线虫都被聚在内群的最外支,说明短口囊杯环线虫与其它种类的亲缘关系比较远;阿氏杯环线虫、细口杯环线虫、辐射杯环线虫和鼻状杯环线虫总是聚在一起,说明它们之间的亲缘关系比较近。需要指出的是,阿氏杯环线虫与细口杯环线虫的种间差异和阿氏杯环线虫的种内差异相似,聚类分析结果表明这两个种难以分开。出现这种结果的原因可能与阿氏杯环线虫具有较大的种内变异,而两个种的种间变异较小有关。也有可能阿氏杯环线虫存在隐含种或者细口杯环线虫和阿氏杯环线虫是同一个种。当然这还需要进一步对细口杯环线虫和阿氏杯环线虫进行形态学和分子生物学的深入研究。
Strongyloid nematodes belong to family Strongylidae, suborder Strongylina, order Rhabditida. Infections of equids (inculding horse, donkey, mule and zebra) with strongyloid nematodes are of major veterinary importance. Adult strongyles inhabit in the large intestine of equids, leading to severe disease or death. Many studies on equine parasites have been carried out in various parts of the world, but there is no detailed survey except taxonomy in China. This study is the first report about the species, morphological description, epidemiology and phylogeny of strongyloids nematodes in the large intestine of donkeys in Henan Province, China.
Thirty-four donkeys from Henan Province were examined at necropsy for strongyloid nematodes in the large intestine from February 2006 to January 2007. Twenty-two species belonging to 8 genus and 2 subfamilies were identified. Of all strongyloid nematodes, some species are of large number, for example, Delafondia vulgaris, Cylicocyclus nassatus, Coronocyclus labratus, Cyathostomum tetracanthum and Cylicocyclus auriculatus. However, other species are of small number, for example, Cylicocyclus elongatus, Cylicostephanus calicatus, Cylicocyclus adersi, Parapoteriostomum euproctus, Triodontophorus nipponicus, Triodontophorus serratus and Alfortia edentatus.
Twenty-two species of strongyloid nematodes were measured, redescribed and photographed through observation under light microscopy and scanning electron microscopy (SEM). SEM observations of strongyles were made at first time in China and the results will provide new characters for the identification and taxonomy. Morphological characters of strongyles in this study were similar to that reported in previously reported, but differences were also found in the numbers of external leaf crown and the length of body, esophagus, spicules, vagina and tail. These differences were considered as intraspecific variation, which can be related to the host, geographical location and climatic conditions.
Epidemiological investigation showed that total numbers of strongyles ranged from 120 to 1939 worms, with a mean of 499 worms per donkey. The numbers of species ranged from 1 to 15 worms per donkey. Only 5.9% of donkeys were infected by a single species, the other were infected with multiple species. Of 22 strongyles, the four most prevalent species are D. vulgaris (88.2%) C. nassatus (73.5%), C. labratus (70.6%) and Coronocyclus labiatus (67.6%). Whereas C. labratus (175.9±291.3), C. tetracanthum (156.0±251.7), C. auriculatus (127.2±221.7) and C. nassatus (110.1±124.4) are the species with the greatest mean intensity. The three most abundant species are C. labratus (124.2±256.4), C.tetracanthum (96.4±210.5) and C. nassatus (80.9±117.1). In addition, season was a significant factor on the effect of species distribution, the prevalence and mean abundance of 22 strongyles. The species richness of strongyles examined are the most in autumn, followed by spring, summer and winter. The major of the species has much higher prevalence and mean abundance in spring and autumn than in summer and winter. Epidemiological investigation on strongyloid nematodes was the first study in China.
Internal transcribed spacer (ITS) and 5.8S of ribosomal DNA (rDNA) of 7 species of the genus Cylicocyclus (C. radiatus, C. adersi, C. ashworthi, C. auriculatus, C. elongatus, C. insignis and C. nassatus) were amplified and sequenced using polymerase chain reaction (PCR) techniques. The length of the ITS1-5.8S-ITS2 sequences ranged from 829 to 843bp, and there are 88 variable sites and 56 parsimony informative sites. The length of the ITS1 sequences ranged form 366 to 370bp, with 22 variable sites and 14 parsimony informative sites. The length of the ITS2 sequences ranged from 307 to 321bp, with 66 variable sites and 42 parsimony informative sites. The 5.8S gene sequences of all 7 species are identical. For all 7 species, the G+C content is higher in the ITS1 (45.8-47.7%) than in the ITS2 (39.2-41.9%). Pairwise comparison revealed differences in sequences ranging from 1.4 to 3.8% in the ITS1 and from 3.1 to 14.3% in the ITS2.
Based on ITS1 and ITS2 sequences from the above 7 species and 5 species of the genus Cylicocyclus (C. ashworthi, C. brevicapsulatus, C. insignis, C. leptostomus and C. nassatus) from Genebank, neighbor-joining (NJ) tree and maximum-parsimony (MP) tree were built using Cylicostephanus longibursatus as outgroup employing PAUP 4.0 (Phylogenetic Analysis Using Parsimony. 4.0 beta version) software. The results showed that C. brevicapsulatus was always on a branch external to the other species in the NJ and MP tree, as indicated that there was a distant relationship between C. brevicapsulatus and the other species. While C. nassatus, C. radiatus, C. ashworthi and C. leptostomus always clustered together, and this demonstrated they have a closed relationship. However, C. leptostomus always clustered with 3 individuals of C. ashworthi, and they could not be completely separated in the NJ and MP tree. This suggested that cryptic species may be existent within C. ashworthi or C. ashworthi and C. leptostomus are the same species. Of course, this problem still need to be further studied.
从2006年2月至2007年1月调查了河南省34头驴体内寄生线虫的感染情况。通过解剖驴的大肠,共采集到22种圆线虫,分属于2个亚科8个属。此次调查发现,寄生数量较大的种类有普通戴拉风线虫Delafondia vulgaris、鼻状杯环线虫Cylicocyclus nassatus、小唇片冠环线虫Coronocyclus labratus、四刺盅口线虫Cyathostomum tetracanthum、耳状杯环线虫Cylicocylus auriculatus等;而长形杯环线虫Cylicocyclus elongatus、小杯杯冠线虫Cylicostephanus calicatus、艾氏杯环线虫Cylicocyclus adersi、真臂副杯口线虫Parapoteriostomum euproctus、日本三齿线虫Triodontophorus nipponicus、锯齿状三齿线虫Triodontophorus serratus、无齿阿尔夫线虫Alfortia edentatus等的寄生数量相对较小。
本研究在光学显微镜下对每一种圆线虫进行仔细地观察和测量,同时对数量较多的种类做了扫描电镜观察,根据观察结果详细地描述了每种圆线虫的形态特征。运用扫描电镜技术观察和描述圆线虫的形态和结构,在我国尚属首次,同时为该类线虫的分类鉴定提供了新的依据。作者所采的标本与国内外的文献报道相比,其形态、结构基本一致。虽然有些种类在体长、外叶冠数、食道长、交合刺长、阴道长以及尾长等方面存在差异,但是作者认为,这可能是由于调查的对象以及采集的数量、采集的时间和地点不同造成的种内变异。
圆线虫的流行病学调查结果显示:每头驴感染圆线虫的数量变化范围是120条～1939条,平均498.6条;被解剖的驴肠道内,有的仅发现1种圆线虫,有的多达15种,大多数是多种圆线虫混合感染。在此次调查中,普通戴拉风线虫(88.2%)、鼻状杯环线虫(73.5%)、小唇片冠环线虫(70.6%)和大唇片冠环线虫Coronocyclus labiatus(67.6%)是感染率较高的种类,为本次调查的核心种;小唇片冠环线虫(175.9±291.3)、四刺盅口线虫(156.0±251.7)、耳状杯环线虫(127.2±221.7)和鼻状杯环线虫(110.1±124.4)是平均感染强度较高的种类;平均丰度较高的种类是小唇片冠环线虫(124.2±256.4)、四刺盅口线虫(96.4±210.5)和鼻状杯环线虫(80.9±117.1)。此次调查还发现,季节对圆线虫种类的分布和感染情况也有一定的影响。在一年内,寄生种类最多的季节是秋季,其次是春季和夏季,冬季种类最少。不同种类的圆线虫其感染率和平均丰度的季节变化明显不一致,大多数种类的变化趋势是:感染率和平均丰度在春季和秋季出现两个高峰,冬季和夏季往往较低。圆线虫的流行病学调查在国内尚属首次报道。
利用聚合酶链式反应(PCR)技术对7种杯环线虫(辐射杯环线虫Cylicocylus radiatus、艾氏杯环线虫、阿氏杯环线虫Cylicocylus ashworthi、耳状杯环线虫、长形杯环线虫、显形杯环线虫Cylicocylus insignis和鼻状杯环线虫)的核糖体DNA(rDNA)的内转录间隔区(Internal transcribed spacer,ITS)及5.8S rDNA进行扩增、测序。序列分析结果表明,所测样品的ITS1-5.8S-ITS2序列的长度范围为829-843bp,变异位点(variable sites)为88个,简约信息位点(parsimony informative sites)为56个;其中ITS1和ITS2的长度范围分别为366-370bp和307-321bp,变异位点为22个和66个,简约信息位点为14个和42个;7种样品的5.8S序列完全一样,长度为153bp。另外,ITS1区的G+C含量(45.8-47.7%)明显高于ITS2区(39.2-41.9%)。通过序列两两比对,7种杯环线虫ITS1区和ITS2区的种间差异性分别为1.4-3.8%和3.1-14.3%。
基于上述7种线虫和Genebank中已登录的5种杯环线虫(阿氏杯环线虫、短口囊杯环线虫Cylicocylus brevicapsulatus、显形杯环线虫、细口杯环线虫Cylicocylus leptostomus和鼻状杯环线虫)的ITS1和ITS2序列的综合数据,以长伞杯冠线虫作为外群,使用PAUP软件构建NJ(neighbor–joining)和MP(maximum parsimony)系统发育树。结果发现,在NJ树和MP树中,短口囊杯环线虫都被聚在内群的最外支,说明短口囊杯环线虫与其它种类的亲缘关系比较远;阿氏杯环线虫、细口杯环线虫、辐射杯环线虫和鼻状杯环线虫总是聚在一起,说明它们之间的亲缘关系比较近。需要指出的是,阿氏杯环线虫与细口杯环线虫的种间差异和阿氏杯环线虫的种内差异相似,聚类分析结果表明这两个种难以分开。出现这种结果的原因可能与阿氏杯环线虫具有较大的种内变异,而两个种的种间变异较小有关。也有可能阿氏杯环线虫存在隐含种或者细口杯环线虫和阿氏杯环线虫是同一个种。当然这还需要进一步对细口杯环线虫和阿氏杯环线虫进行形态学和分子生物学的深入研究。
Strongyloid nematodes belong to family Strongylidae, suborder Strongylina, order Rhabditida. Infections of equids (inculding horse, donkey, mule and zebra) with strongyloid nematodes are of major veterinary importance. Adult strongyles inhabit in the large intestine of equids, leading to severe disease or death. Many studies on equine parasites have been carried out in various parts of the world, but there is no detailed survey except taxonomy in China. This study is the first report about the species, morphological description, epidemiology and phylogeny of strongyloids nematodes in the large intestine of donkeys in Henan Province, China.
Thirty-four donkeys from Henan Province were examined at necropsy for strongyloid nematodes in the large intestine from February 2006 to January 2007. Twenty-two species belonging to 8 genus and 2 subfamilies were identified. Of all strongyloid nematodes, some species are of large number, for example, Delafondia vulgaris, Cylicocyclus nassatus, Coronocyclus labratus, Cyathostomum tetracanthum and Cylicocyclus auriculatus. However, other species are of small number, for example, Cylicocyclus elongatus, Cylicostephanus calicatus, Cylicocyclus adersi, Parapoteriostomum euproctus, Triodontophorus nipponicus, Triodontophorus serratus and Alfortia edentatus.
Twenty-two species of strongyloid nematodes were measured, redescribed and photographed through observation under light microscopy and scanning electron microscopy (SEM). SEM observations of strongyles were made at first time in China and the results will provide new characters for the identification and taxonomy. Morphological characters of strongyles in this study were similar to that reported in previously reported, but differences were also found in the numbers of external leaf crown and the length of body, esophagus, spicules, vagina and tail. These differences were considered as intraspecific variation, which can be related to the host, geographical location and climatic conditions.
Epidemiological investigation showed that total numbers of strongyles ranged from 120 to 1939 worms, with a mean of 499 worms per donkey. The numbers of species ranged from 1 to 15 worms per donkey. Only 5.9% of donkeys were infected by a single species, the other were infected with multiple species. Of 22 strongyles, the four most prevalent species are D. vulgaris (88.2%) C. nassatus (73.5%), C. labratus (70.6%) and Coronocyclus labiatus (67.6%). Whereas C. labratus (175.9±291.3), C. tetracanthum (156.0±251.7), C. auriculatus (127.2±221.7) and C. nassatus (110.1±124.4) are the species with the greatest mean intensity. The three most abundant species are C. labratus (124.2±256.4), C.tetracanthum (96.4±210.5) and C. nassatus (80.9±117.1). In addition, season was a significant factor on the effect of species distribution, the prevalence and mean abundance of 22 strongyles. The species richness of strongyles examined are the most in autumn, followed by spring, summer and winter. The major of the species has much higher prevalence and mean abundance in spring and autumn than in summer and winter. Epidemiological investigation on strongyloid nematodes was the first study in China.
Internal transcribed spacer (ITS) and 5.8S of ribosomal DNA (rDNA) of 7 species of the genus Cylicocyclus (C. radiatus, C. adersi, C. ashworthi, C. auriculatus, C. elongatus, C. insignis and C. nassatus) were amplified and sequenced using polymerase chain reaction (PCR) techniques. The length of the ITS1-5.8S-ITS2 sequences ranged from 829 to 843bp, and there are 88 variable sites and 56 parsimony informative sites. The length of the ITS1 sequences ranged form 366 to 370bp, with 22 variable sites and 14 parsimony informative sites. The length of the ITS2 sequences ranged from 307 to 321bp, with 66 variable sites and 42 parsimony informative sites. The 5.8S gene sequences of all 7 species are identical. For all 7 species, the G+C content is higher in the ITS1 (45.8-47.7%) than in the ITS2 (39.2-41.9%). Pairwise comparison revealed differences in sequences ranging from 1.4 to 3.8% in the ITS1 and from 3.1 to 14.3% in the ITS2.
Based on ITS1 and ITS2 sequences from the above 7 species and 5 species of the genus Cylicocyclus (C. ashworthi, C. brevicapsulatus, C. insignis, C. leptostomus and C. nassatus) from Genebank, neighbor-joining (NJ) tree and maximum-parsimony (MP) tree were built using Cylicostephanus longibursatus as outgroup employing PAUP 4.0 (Phylogenetic Analysis Using Parsimony. 4.0 beta version) software. The results showed that C. brevicapsulatus was always on a branch external to the other species in the NJ and MP tree, as indicated that there was a distant relationship between C. brevicapsulatus and the other species. While C. nassatus, C. radiatus, C. ashworthi and C. leptostomus always clustered together, and this demonstrated they have a closed relationship. However, C. leptostomus always clustered with 3 individuals of C. ashworthi, and they could not be completely separated in the NJ and MP tree. This suggested that cryptic species may be existent within C. ashworthi or C. ashworthi and C. leptostomus are the same species. Of course, this problem still need to be further studied.
引文
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