缘毛类和盾纤类纤毛虫的分类学与分子系统学研究
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摘要
纤毛虫作为形态分化最复杂的单细胞真核生物，是微食物网中不可或缺的组成部分，具有重要的生态学研究意义。而且以缘毛类和盾纤类为代表的纤毛虫病原已给水产养殖业带来了重大的经济损失。尽管这些类群具有生态和经济的双重重要性，人们对其分类学和系统学等基础资料还知之甚少。主要原因在于：这些类群的形态分类学基础研究还不足，许多早期的资料存在大量鉴定错误，且近年来有关其系统关系的界定又存在着巨大争议。
鉴于上述问题，本工作于2008—2012年间采集了大连、青岛、福州和海南等地的寄生性和自由生的缘毛类和盾纤类纤毛虫，开展了形态分类学和分子系统学为主的研究。结合基于活体观察和现代银染技术的形态学方法，对疑难种和误鉴定种进行了重描述；通过增加序列样品量、多基因测序、改进数据分析方法等手段，来提高分子系统分析的可靠性。并结合形态学和分子系统学的信息，来对这些类群的分类系统进行修订。
主要成果包括：
1）对隶属于12属19种纤毛虫进行了重描述，其中14种为国内新纪录物种。
2）运用多种序列比对和校正方法、多种限制性假设检验方法以及不同样品量的小核糖体亚基基因信息对缘毛类纤毛虫的单系性问题进行综合探讨，发现缘毛类单系群的支持与否，取决于分析方法的不同。检测了Alpha微管蛋白基因序列在缘毛类纤毛虫系统关系研究中的适用性，发现基于该基因信息的系统发育分析结果比较混乱，并不能很好解决缘毛类的单系性问题。
3）结合形态学和分子系统学数据，对缘毛亚纲的5科，6属纤毛虫的系统地位进行了调整和确认。并对游走目下的4科，4属做了修订：把Leiotrocha，Polycycla和Trichodinopsis合并到壶形虫属Urceolaria；把Leiotrochidae，Polycyclidae和Trichodinopsidae合并到壶形虫科Urceolariidae中。此外，确定了Urceolaria，Trichodinella和Trichodina的系统关系。
4）通过增加盾纤类23个种/种群的核糖体小亚基基因信息，并对盾纤亚纲的系统地位进行探讨，结果表明盾纤类纤毛虫的单系性被拒绝，并且分为两大分枝：一个是具有“经典”盾纤类特征的类群，包含嗜污目、帆口目和触毛目；另一个则是具有盾纤类和膜口类共同特征的斜头类。而后者的单系性并未能得到很好地支持，且与前口亚纲的系统关系较近。同时，对盾纤类纤毛虫长期系统地位混乱或不明的11个属的系统地位做了调整和确认，分别为： Ancistrum， Biggaria， Boveria，Conchophthirus，Cyclidium，Madsenia，Myxophyllum，Mytilophilus，Paralembus，Pseudocyclidium和Protophyra。
5）首次针对高害性盾纤类纤毛虫—水滴伪康纤虫Pseudocohnilembuspersalinus开发了特异性寡核苷酸探针，并结合荧光原位杂交技术实现对其的快速鉴定。并与另外一种快速鉴定方法—DNA条形码技术进行比较，探讨了它们在病害纤毛虫诊断中的应用前景。此外，本文总共完成了对缘毛亚纲和盾纤亚纲内部35个种/种群的核糖体亚基基因和3个种的Alpha微管蛋白基因以及1个种的线粒体细胞色素C氧化酶Ⅰ亚基基因的序列测定并提交到GenBank。
Ciliated protozoa are morphologically highly differentiated single-celledeukaryotes. They are essential components in microbial food webs, and play akey role in various ecosystems. In world aquaculture, the pathogenic diseasescaused by Peritrichia and Scuticociliatia have resulted in serious economicloss. Despite their economic and ecological significance, there are limitedstudies on the diversity and pathogenic mechanism of these groups. Thereasons include:1) there are many uncertainties in the taxonomy of thesegroups, and2) there are still many phylogentic discrepancies even based onrecent molecular data.
To investigate the issues mentioned above, we collected the parasitc andfree-living peritrich and scuticociliatid ciliates from sites located at Dalian,Qingdao, Fuzhou and Hainan in China during2008-2012. Using modernmorphological and molecular methods, we aimed to solve those confusions inmorphological taxonomy and systematology of deep subclades of thesegroups. We also analyzed sequences of various genetic markers to investigatethe disagreements between different researchers.
The results are summarized as following:
(1)19ambigous species belonging to12genera were redescribed. Amongthese ciliates,14species are first recorded in China.
(2) Based on large number of SSU-Rdna sequences of Peritrichia, andcomparison of different alignment and masking methods, we found thatsupport for and against monophyly of the Peritrichia were highlydependent upon alignment and masking methodologies.
(3) The support for and against monophyly of the Peritrichia s.l. were not significant basing on alpha tubulin gene sequences. And the resultshowed alpha tubulin gene was dubious help when inferringphylogenetic relationships.
(4) Based on the morphological and molecular data, we redefined thesystemic position of six peritrichous genera. Four families and fourgenera in Mobilida were emended: the genera Leiotrocha, Polycycla andTrichodinopsis were transferred to Urceolaria; and consequently thefamily Leiotrochidae, Polycyclidae and Trichodinopsidae weretransferred to Urceolariidae.
(5) The monophyly of the subclass Scuticociliatia was not supported withadding new23scuticociliatid SSU-rDNA sequences. The phylogeneticpositions of11genera, whose assignments were historically unclear,have been studied. These11genera are: Ancistrum, Biggaria, Boveria,Conchophthirus, Cyclidium, Madsenia, Myxophyllum, Mytilophilus,Paralembus, Pseudocyclidium, and Protophyra.
(6) A fluorescent probe targeting small subunit ribosomal RNA (SSU-rRNA)was designed and optimized in a fluorescence in situ hybridization (FISH)to detect the pathogenic ciliate Pseudocohnilembus persalinus inaquaculture.
Additionally, small subunit rDNA sequences of35peritichous andscuticociliatid species/populations, alpha tubulin gene sequences of threeperitichs, and mitochondrial cytochrome c oxidase subunit Ⅰ gene sequence ofone scuticociliatid species were submitted to the GenBank.
鉴于上述问题，本工作于2008—2012年间采集了大连、青岛、福州和海南等地的寄生性和自由生的缘毛类和盾纤类纤毛虫，开展了形态分类学和分子系统学为主的研究。结合基于活体观察和现代银染技术的形态学方法，对疑难种和误鉴定种进行了重描述；通过增加序列样品量、多基因测序、改进数据分析方法等手段，来提高分子系统分析的可靠性。并结合形态学和分子系统学的信息，来对这些类群的分类系统进行修订。
主要成果包括：
1）对隶属于12属19种纤毛虫进行了重描述，其中14种为国内新纪录物种。
2）运用多种序列比对和校正方法、多种限制性假设检验方法以及不同样品量的小核糖体亚基基因信息对缘毛类纤毛虫的单系性问题进行综合探讨，发现缘毛类单系群的支持与否，取决于分析方法的不同。检测了Alpha微管蛋白基因序列在缘毛类纤毛虫系统关系研究中的适用性，发现基于该基因信息的系统发育分析结果比较混乱，并不能很好解决缘毛类的单系性问题。
3）结合形态学和分子系统学数据，对缘毛亚纲的5科，6属纤毛虫的系统地位进行了调整和确认。并对游走目下的4科，4属做了修订：把Leiotrocha，Polycycla和Trichodinopsis合并到壶形虫属Urceolaria；把Leiotrochidae，Polycyclidae和Trichodinopsidae合并到壶形虫科Urceolariidae中。此外，确定了Urceolaria，Trichodinella和Trichodina的系统关系。
4）通过增加盾纤类23个种/种群的核糖体小亚基基因信息，并对盾纤亚纲的系统地位进行探讨，结果表明盾纤类纤毛虫的单系性被拒绝，并且分为两大分枝：一个是具有“经典”盾纤类特征的类群，包含嗜污目、帆口目和触毛目；另一个则是具有盾纤类和膜口类共同特征的斜头类。而后者的单系性并未能得到很好地支持，且与前口亚纲的系统关系较近。同时，对盾纤类纤毛虫长期系统地位混乱或不明的11个属的系统地位做了调整和确认，分别为： Ancistrum， Biggaria， Boveria，Conchophthirus，Cyclidium，Madsenia，Myxophyllum，Mytilophilus，Paralembus，Pseudocyclidium和Protophyra。
5）首次针对高害性盾纤类纤毛虫—水滴伪康纤虫Pseudocohnilembuspersalinus开发了特异性寡核苷酸探针，并结合荧光原位杂交技术实现对其的快速鉴定。并与另外一种快速鉴定方法—DNA条形码技术进行比较，探讨了它们在病害纤毛虫诊断中的应用前景。此外，本文总共完成了对缘毛亚纲和盾纤亚纲内部35个种/种群的核糖体亚基基因和3个种的Alpha微管蛋白基因以及1个种的线粒体细胞色素C氧化酶Ⅰ亚基基因的序列测定并提交到GenBank。
Ciliated protozoa are morphologically highly differentiated single-celledeukaryotes. They are essential components in microbial food webs, and play akey role in various ecosystems. In world aquaculture, the pathogenic diseasescaused by Peritrichia and Scuticociliatia have resulted in serious economicloss. Despite their economic and ecological significance, there are limitedstudies on the diversity and pathogenic mechanism of these groups. Thereasons include:1) there are many uncertainties in the taxonomy of thesegroups, and2) there are still many phylogentic discrepancies even based onrecent molecular data.
To investigate the issues mentioned above, we collected the parasitc andfree-living peritrich and scuticociliatid ciliates from sites located at Dalian,Qingdao, Fuzhou and Hainan in China during2008-2012. Using modernmorphological and molecular methods, we aimed to solve those confusions inmorphological taxonomy and systematology of deep subclades of thesegroups. We also analyzed sequences of various genetic markers to investigatethe disagreements between different researchers.
The results are summarized as following:
(1)19ambigous species belonging to12genera were redescribed. Amongthese ciliates,14species are first recorded in China.
(2) Based on large number of SSU-Rdna sequences of Peritrichia, andcomparison of different alignment and masking methods, we found thatsupport for and against monophyly of the Peritrichia were highlydependent upon alignment and masking methodologies.
(3) The support for and against monophyly of the Peritrichia s.l. were not significant basing on alpha tubulin gene sequences. And the resultshowed alpha tubulin gene was dubious help when inferringphylogenetic relationships.
(4) Based on the morphological and molecular data, we redefined thesystemic position of six peritrichous genera. Four families and fourgenera in Mobilida were emended: the genera Leiotrocha, Polycycla andTrichodinopsis were transferred to Urceolaria; and consequently thefamily Leiotrochidae, Polycyclidae and Trichodinopsidae weretransferred to Urceolariidae.
(5) The monophyly of the subclass Scuticociliatia was not supported withadding new23scuticociliatid SSU-rDNA sequences. The phylogeneticpositions of11genera, whose assignments were historically unclear,have been studied. These11genera are: Ancistrum, Biggaria, Boveria,Conchophthirus, Cyclidium, Madsenia, Myxophyllum, Mytilophilus,Paralembus, Pseudocyclidium, and Protophyra.
(6) A fluorescent probe targeting small subunit ribosomal RNA (SSU-rRNA)was designed and optimized in a fluorescence in situ hybridization (FISH)to detect the pathogenic ciliate Pseudocohnilembus persalinus inaquaculture.
Additionally, small subunit rDNA sequences of35peritichous andscuticociliatid species/populations, alpha tubulin gene sequences of threeperitichs, and mitochondrial cytochrome c oxidase subunit Ⅰ gene sequence ofone scuticociliatid species were submitted to the GenBank.
引文
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