微生物学报

小袋科纤毛虫分类与遗传多态性的研究进展

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闫文朝 ¹ , 李明 ² , 胡苏辉 ¹ , 王天奇 ¹

微生物学报 | 综述 2026,66(7): 3250-3259

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微生物学报 |综述 2026 , 66 (7) : 3250 -3259

小袋科纤毛虫分类与遗传多态性的研究进展

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闫文朝¹, 李明², 胡苏辉¹, 王天奇¹

作者信息

^1.河南科技大学动物科技学院，河南洛阳

^2.中国科学院水生生物研究所，湖北武汉

作者简介:

作者贡献声明

闫文朝：论文构思、撰写和基金项目支持；李明：论文撰写与修改；胡苏辉：系统发育树构建和参考文献整理；王天奇：论文构思和修改。

Advancements in the classification and genetic diversity of Balantidiidae ciliates

Wenchao YAN¹, Ming LI², Suhui HU¹, Tianqi WANG¹

Affiliations

^1.College of Animal Science and Technology, Henan University of Science and Technology, Luoyang, Henan, China

^2.Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan, Hubei, China

出版时间: 2026-07-04 doi: 10.13343/j.cnki.wsxb.20260042

文章导航

摘要

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小袋科纤毛虫是一类广泛寄生于多种无脊椎动物和脊椎动物消化系统的纤毛虫。其中，结肠小袋纤毛虫呈世界性分布，是一种重要的人兽共患病原。长期以来，由于不同宿主来源的小袋科纤毛虫滋养体形态相似，且其形态学特征会受虫体不同生长阶段和宿主肠道微环境的影响而发生变化，导致其分类地位、属级划分及命名在国内外文献中存在分歧与混乱。近年来，小袋科纤毛虫的系统发育关系和遗传多态性研究取得了重要进展，为其分类学重构提供了新的证据。本文在系统梳理国内外研究文献的基础上，结合作者团队的相关研究成果，从形态学特征、成囊能力、宿主特异性及分子系统发育等方面，综述并比较不同宿主源小袋科纤毛虫的分类地位与遗传多态性。基于现有证据，本文支持将小袋科纤毛虫划分为肠袋属(Balantidium)和小袋属(Balantioides)，并明确指出感染人和多种哺乳动物的小袋科纤毛虫应统一归为结肠小袋纤毛虫(Balantioides coli)。这些研究成果和认识有助于澄清小袋科纤毛虫的分类与命名体系，为其生物学特性研究以及人和动物小袋虫病的诊断与防控提供重要参考。

关键词

小袋科 / 肠袋属 / 小袋属 / 结肠小袋纤毛虫 / 分类与命名 / 遗传多态性分析

Abstract

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Ciliates of the family Balantidiidae widely parasitize the digestive systems of various invertebrates and vertebrates. Among them, Balantioides coli with cosmopolitan distribution is a major zoonotic pathogen. For a long time, due to the similar morphology of trophozoites of Balantidiidae ciliates from various hosts and the influences of various growing stages and host intestinal microenvironments on morphological characteristics, there have been some inconsistencies and confusions in their classification and nomenclature in literature. Recently, significant progress has been achieved in the study of the phylogenetic relationship and genetic diversity of Balantidiidae ciliates, providing new evidences for their taxonomic reconstruction. On the basis of a systematic review of literature and our findings, this article summarizes the taxonomic status and genetic diversity of Balantidiidae ciliates from different hosts in terms of the morphological characteristics, encystation ability, host specificity, and phylogenetic relationships. The existing evidence supports the division of the family Balantidiidae into two genera Balantidium and Balantioides and indicates that the ciliate infecting humans and various mammals should be classified as Balantioides coli. The related findings and understanding help clarify the taxonomy and nomenclature system of Balantidiidae ciliates and provide an important reference for studying the biological characteristics of Balantidiidae ciliates as well as the diagnosis and control of balantiosis in humans and animals.

Key words

Balantidiidae / Balantidium / Balantioides / Balantioides coli / taxonomy and nomenclature / genetic diversity analysis

引用本文

闫文朝, 李明, 胡苏辉, 王天奇. 小袋科纤毛虫分类与遗传多态性的研究进展. 微生物学报, 2026 , 66 (7) : 3250 -3259 . DOI: 10.13343/j.cnki.wsxb.20260042

Wenchao YAN, Ming LI, Suhui HU, Tianqi WANG. Advancements in the classification and genetic diversity of Balantidiidae ciliates[J]. Acta Microbiologica Sinica, 2026 , 66 (7) : 3250 -3259 . DOI: 10.13343/j.cnki.wsxb.20260042

正文

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小袋科(Balantidiidae Doflein & Reichenow, 1929)纤毛虫隶属于纤毛门(Ciliophora Doflein, 1901)线口纲(Litostomatea Small & Lynn, 1981)毛口亚纲(Trichostomatia Bütschli, 1889)前庭目(Vestibuliferida de Puytorac et al., 1974)，是一类广泛寄生于多种无脊椎动物和脊椎动物消化系统的纤毛虫，其宿主范围广泛，从低等的腔肠动物、扁形动物、环节动物、软体动物、节肢动物，到较高等的鱼类、两栖类、爬行类、鸟类，直至哺乳类和人类均有报道^[1-3]。寄生于无脊椎动物、鱼类、两栖类和爬行类动物体内的小袋科纤毛虫(Balantidium spp.)，一般认为是共生物，对宿主无致病性，且能提高宿主肠道微生物的多样性和有益菌群的丰度^[2,4-6]。寄生于哺乳动物、鸟类和人盲肠和结肠内的结肠小袋纤毛虫(Balantioides coli，简称小袋虫)是一种机会性病原，当宿主处于应激或免疫抑制状态，或长期采食富含淀粉的饲料或食物时，滋养体数量激增，入侵肠黏膜，继而进入黏膜下组织和淋巴组织，引发小袋虫性腹泻、盲肠和结肠溃疡，甚至穿孔^[1,7-9]。小袋虫感染会降低肠道微生物的多样性和有益菌群的丰度^[10]。此外，人的病例中还存在泌尿生殖道、肝脏和肺脏等肠外感染的报道^[11-14]。

据记载，不同宿主源的小袋科纤毛虫种类多达100余种，其中一些种类形态结构极为相似，在显微镜下难以辨别^[2,15-17]。近年来，随着分子遗传数据的不断积累，出现了小袋科纤毛虫的分子鉴定结果与经典形态分类学结果不一致的问题^[3,15,18]。在教材和论文等文献资料中存在拉丁文名称和中文名称不统一的现象，给同行交流和临床诊断带来了困扰^[3,18-20]。本文查阅了国内外相关文献，结合作者长期的研究成果，从经典形态分类学和分子遗传学2个方面总结和阐述小袋科纤毛虫分类和遗传多态性的研究进展，澄清感染不同宿主的小袋科纤毛虫的分类地位，以期为同行和临床上人和动物小袋虫病的诊断和防控提供参考。

1 小袋科纤毛虫经典形态学分类

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小袋科纤毛虫滋养体一般呈卵圆形或梨形，前端稍尖，后端钝圆(图1)。其内部存在双核结构，即大核和小核。大核多呈肾形或香肠状，为营养核，高度多倍体，富含调控虫体生理生化代谢活动的基因；小核呈球形，多位于大核凹陷处，为生殖核，二倍体，负责传递遗传信息^[1,19]。虫体体表布满斜纵向排列的纤毛(图2A-2C)，这些纤毛具有正常的“9+2”型微管结构，纤毛基部与表膜下纤维系统(即由动基体及其附属纤维结构组成)相连，是滋养体的运动器官；每排纤毛之间有纵向排列的突起，即动基列(图2B、2D)^[21-24]。滋养体前腹侧有一倾斜内陷的胞口，相对于体表纤毛，胞口内表面覆盖更长更密的纤毛，胞口具有摄取淀粉、细菌等食物的作用(图2C、2D)^[25-26]。一些种类的滋养体胞口右侧体表存在Villeneuve-Brachon’s field (V-B区)，该区纤毛相对其他体表区较短，由4-20排动基列组成，该区域与滋养体接合生殖时的黏附结合相关^[6]。胞口向下形成管状结构，即胞咽，其盲端终止于细胞质。滋养体后端有一胞肛，具有排泄虫体不能消化的残渣废物的作用(图2B)^[19,26]。另外，滋养体细胞质内还存在数量不等的粗面内质网、食物泡、伸缩泡等细胞器，食物泡具有暂时储存食物的作用，伸缩泡具有调节细胞内渗透压的功能^[1-2,19]。早期透射电镜观察结果显示，小袋科纤毛虫滋养体细胞质内存在线粒体^[22]，但后来的透射电镜和基因组分析结果显示，小袋科纤毛虫滋养体内无线粒体和线粒体相关基因，而是在动基体周围和动基列内部有产氢体(又称退化的线粒体结构)，为滋养体运动提供能量，这与小袋科纤毛虫适应宿主肠道的厌氧环境相一致^[4,6,27]。以上结构特征是小袋科纤毛虫的共同特征。

虽然有学者提出可将滋养体的大小和形态、胞口形状和长度、伸缩泡数量和位置、大核形状、V-B区纤毛长度和动基列数量等参数作为小袋科不同种属分类的依据，但由于滋养体所处宿主肠道微环境和生长阶段不同，上述参数会出现较大变化，因此多数学者认为以这些滋养体参数作为小袋科不同种属的分类依据可靠性较低^[1,15,26]。另外，在显微镜和扫描电镜下，小袋科不同种属纤毛虫滋养体的形态结构较为相似(图1)，常难以准确区分^[6,15]。

根据经典形态学分类标准，宿主种类也是小袋科纤毛虫分类的判定指标之一^[1,19]。目前，寄生于猪、牛、羊、非人灵长类、豚鼠等哺乳动物以及鸵鸟和人结肠和盲肠的小袋纤毛虫归到Balantioides属，为同一个种Balantioides coli^[1,15,28]。来自鱼类、两栖类、爬行类和昆虫的小袋科纤毛虫则归到Balantidium属，其模式种为内生肠袋虫(Balantidium entozoon)；Balantidium属种类繁多，仅在鱼类和两栖类肠道中发现的就有50余种，如鱼源的鲩肠袋虫(B. ctenopharyngodonis)、多泡肠袋虫(B. polyvacuolum)、B. jocularum等^[24-26]；蛙源的B. entozoon、十二指肠肠袋虫(B. duodeni)、格里姆肠袋虫(B. grimi)、洪湖肠袋虫(B. honghuensis)等^[6,29-30]；大鲵源的中华肠袋虫(B. sinensis)、大鲵肠袋虫(B. andianusis)等^[31]。另外，小袋科纤毛虫回归感染模型试验鲜有成功报道，尚不足以支撑其作为分类的标准^[32]。

不同小袋科纤毛虫物种的成囊能力不同。例如，寄生于哺乳动物、鸵鸟和人的Balantioides coli滋养体具有成囊能力，虫体表膜下的黏液囊泡参与成囊过程^[1,18]。来自鱼类、两栖类、爬行类的Balantidium纤毛虫在体内或体外培养过程中以滋养体形式存在，通常认为其不具成囊能力^[18,26]。也有学者认为，它们应该具有成囊能力，只是由于它们的宿主经常生活在水中，缺乏干燥等逆环境来迫使滋养体形成包囊^[7,30]。此外，不同纤毛虫物种成囊所需的条件不同，如牛源和猴源的巴克斯顿纤毛虫(Buxtonella spp.)滋养体在蒸馏水和自来水中就容易形成包囊，而猪源Balantioides coli滋养体在蒸馏水和自来水中不能形成包囊反而崩解，说明Balantioides coli的成囊条件更为苛刻^[18,32-33]。

2 小袋科纤毛虫分子遗传学鉴定

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2.1 基于18S rRNA基因序列的分子鉴定

18S rRNA基因是小袋科纤毛虫种类鉴定中应用最广泛的遗传标记基因，基于18S rRNA基因序列的分析结果将小袋科纤毛虫分为3个群系(图3)，分别为蛙源群系、鱼源群系和哺乳动物源群系(该群系除鸵鸟外，其他宿主均为哺乳动物)^[18,34-37]。在GenBank中，蛙源群系目前有3个种具有18S rRNA基因序列数据，分别为B. entozoon、B. duodeni、B. grimi，该群系内种间序列相似性为98.6%-99.0%；鱼源群系目前有2个种具有18S rRNA基因序列数据，分别为B. polyvacuolum和B. ctenopharyngodoni，该群系内种间序列相似性为96.9%；哺乳动物源群系目前只有1个种，即Balantioides coli，有猪、大猩猩、鹿、鸵鸟、大象等不同宿主源的虫株，种内序列相似性为99.0%-100.0%。3种群系间比较显示，蛙源群系与鱼源群系间序列相似性为93.9%-94.5%；蛙源群系与哺乳动物源群系间序列相似性为95.8%-96.2%；鱼源群系与哺乳动物源群系间序列相似性为94.8%-96.1%。鱼源群系与蛙源群系、哺乳动物源群系遗传距离均较远，而与毛口亚纲寄生于有袋类动物消化道内的Amylovorax属纤毛虫遗传距离更近^[30,34]，并处于种系发育树同一分支上(图3)。鱼源群系是否单列为一个新属，有待更多生物学特征和遗传数据的积累与验证。

2.2 基于ITS1-5.8S rDNA-ITS2序列的遗传多态性分析

小袋科纤毛虫ITS1-5.8S rDNA-ITS2序列公开发表的有多种宿主源的Balantioides coli、鱼源的B. ctenopharyngodoni和蛙源的B. entozoon 3个虫种，其中B. ctenopharyngodoni和B. entozoon均仅有1个虫株的序列，而Balantioides coli具有来自不同宿主和不同国家地区的多个虫株的序列^[18,36-39]。

基于ITS1-5.8S rDNA-ITS2序列比对分析结果显示，B. entozoon与B. ctenopharyngodoni的序列相似性为83.3%，与Balantioides coli为84.1%-88.0%；B. ctenopharyngodoni与Balantioides coli为83.5%-87.4%，Balantioides coli种内不同虫株间序列相似性为91.5%-99.5%。基于5.8S rRNA基因序列比对分析结果显示，B. entozoon与B. ctenopharyngodoni的序列相似性为96.6%，与Balantioides coli为94.6%-96.0%；B. ctenopharyngodoni与Balantioides coli为97.4%，Balantioides coli种内不同虫株间序列相似性为100%。基于5.8S rRNA基因序列分析的结果与基于18S rRNA基因序列分析结果相似，说明5.8S rRNA基因序列可以有效区分小袋科不同种属纤毛虫，但无法区分种内不同虫株之间的遗传差异。在ITS1-5.8S rDNA-ITS2序列中，小袋科纤毛虫的5.8S rRNA基因序列相对保守，序列变异性最高的是内转录间隔区ITS1，其次是ITS2区^[18,38-39]。

基于ITS1/2序列的高度变异特性，将Balantioides coli分为遗传变异型A、B和C 3种，其中遗传变异型A为人兽共患型，可感染人、猪、豚鼠、奶牛、绵羊、灵长类和鸵鸟等宿主；遗传变异型B是猪群感染的优势种群，同时存在于豚鼠、奶牛、鸵鸟、绵羊、灵长类等动物，目前尚未见感染人的报道；遗传变异型C目前仅见于感染豚鼠的报道^[38-40]。另外，有学者基于ITS1-5.8S rDNA-ITS2序列建立了聚合酶链反应-限制性片段长度多态性(polymerase chain reaction-restriction fragment length polymorphism, PCR-RFLP)分型方法，该方法可以有效区分遗传变异型A和B，并且将遗传变异型B细分为B-c和B-t 2个亚型^[17,41]。

相关研究显示，在同一份样品或同一个滋养体中存在Balantioides coli 2个或2个以上ITS遗传变异型，且同一遗传变异型内也存在多个亚型^[18,36,38]。这说明Balantioides coli的ITS1/2区为非基因编码区，碱基突变频率极高，使其作为遗传标记分子时分析结果的代表性有一定局限性。

2.3 基于β-微管蛋白基因的遗传多态性分析

目前β-微管蛋白基因在小袋科纤毛虫中仅有Balantioides coli有报道^[40]。β-微管蛋白基因是Balantioides coli的结构基因，碱基突变可能改变基因的功能^[42-43]。Balantioides coli与嗜热四膜虫和草履虫β-微管蛋白基因序列相似性分别为81.8%和80.0%，Balantioides coli种内不同分离株之间序列相似性为88.1%-100.0%。基于56份样品的β-微管蛋白基因分析结果显示，来源于猪、豚鼠、奶牛和绵羊的Balantioides coli被分为基因型I、Ⅱ和Ⅲ 3种基因型，其中基因型Ⅱ为中国猪群感染的优势种群，基因型Ⅲ为豚鼠源虫株，目前暂无其他宿主感染该基因型的报道。3种基因型之间序列相似性为88.1%-92.9%，基因型内不同分离株之间序列相似性为94.4%-100.0%。这说明β-微管蛋白基因是Balantioides coli一个变异性较强、分辨率高的遗传标记基因，可用于Balantioides coli的基因型分析，相较于ITS1/2序列，其分析结果更具代表性和可信度。

3 结肠小袋纤毛虫的分类与命名

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1857年，Malmsten从2位痢疾患者体内检测到纤毛虫，并将其认定为草履虫，命名为Paramecium coli；1858年，Claparède和Lachmann 2位学者提议在前庭目(Vestibuliferida)下设立Balantidiidae科Balantidium属，并将蛙源Balantidium entozoon作为Balantidium属的模式种；1861年，Leuckart从猪的大肠中分离到与Paramecium coli形态相似的纤毛虫；1863年，Stein通过滋养体形态学观察和比较，将来自人和猪的Paramecium coli划归到Balantidium属，并命名为Balantidium coli^[1,28,46]。由于猪、人和蛙遗传进化关系较远，结合虫体形态结构特征，1931年，Alexeieff^[44]建议将来自人和猪的Balantidium coli归为一个新属——Balantioides，种名为Balantioides coli。遗憾的是，该建议未引起寄生虫学界的关注，大家仍然沿用Balantidium coli，中文学名为结肠小袋纤毛虫^[19,28]。后来在豚鼠、鸵鸟、非人灵长类等多种动物体内均发现与Balantidium属形态相似的纤毛虫，学者建议以宿主种类来命名来自不同宿主的Balantidium属纤毛虫，如猪源小袋纤毛虫命名为Balantidium suis，豚鼠源的为Balantidium caviae，鸵鸟源的为Balantidium struthionis。后来基于分子鉴定、滋养体形态结构特征和寄生部位等数据证实，Balantidium suis、Balantidium caviae、Balantidium struthionis等多种宿主源Balantidium属纤毛虫均为Balantidium coli的同物异名^[1,15]。

2013年，Pomajbíková等^[18]通过18S rRNA基因和ITS1-5.8S rDNA-ITS2序列的系统分析发现，来自人、猪、鸵鸟和非人灵长类的Balantidium coli与蛙源Balantidium entozoon遗传差异明显，因此认为Balantidium coli和Balantidium entozoon应该分别属于不同的属，建议新属名为Neobalantidium，将Balantidium coli改为Neobalantidium coli。2014年，Chistyakova等^[30]学者认为Balantioides作为属名比Neobalantidium更早，具有优先权。2018年，Jirků-Pomajbíková和Stensvold^[45]对前期研究结果和文献进行重新评估后，承认Balantioides作为属名比Neobalantidium具有优先权，建议将Neobalantidium coli更名为Balantioides coli。这一建议后来被更多学者接受和认可，并将Balantioides coli引起宿主的疾病命名为balantiosis^[46-48]。

虽然来源于人、猪、豚鼠和鸵鸟等宿主的Balantioides coli的拉丁文学名已确定下来，但是其中文学名尚未定论。从构词上讲，“Balantio-”是Balantioides的词根，希腊语“balantídion”意思是“小的袋子”，“-ides”是后缀，意思是“类/拟”^[7,19]，按照字面意思应该将Balantioides coli翻译为结肠类小袋纤毛虫或结肠拟小袋纤毛虫。然而，长期以来，寄生虫学界同行已习惯将Balantidium coli翻译为结肠小袋纤毛虫，简称小袋虫^[19,22]。按照约定俗成和便于交流的原则，Balantioides coli应该沿用原来的中文学名即结肠小袋纤毛虫，简称小袋虫，不宜改为“结肠类小袋纤毛虫”或“结肠拟小袋纤毛虫”。另外，“Balantidium”在鱼源和蛙源小袋科纤毛虫中通常被翻译为“肠袋属”，如Balantidium ctenopharyngodoni和Balantidium polyvacuolum中文学名分别为鲩肠袋虫和多泡肠袋虫^[2-3,19,24]。因此，建议将“Balantidium”和“Balantioides”分别译为“肠袋属”和“小袋属”，也恰好将小袋科的这2个属在拉丁文名称和中文学名上区分开来。该建议已与相关领域专家(中国海洋大学宋微波院士、中国农业大学索勋教授和河南农业大学张龙现教授)讨论，并达成共识。

4 总结与展望

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基于滋养体形态结构特征、成囊能力、感染宿主种类、寄生部位和分子遗传关系等数据，将前庭目小袋科纤毛虫分为肠袋属(Balantidium)和小袋属(Balantioides)。肠袋属包括感染两栖动物类、鱼类、昆虫类、爬行动物类的小袋科纤毛虫；小袋属包括感染猪、牛、羊、豚鼠、非人灵长类、鸵鸟等动物和人的小袋科纤毛虫，目前只有一个种即Balantioides coli，它是一种机会性病原，存在较高的遗传多态性，具有重要的公共卫生意义。18S rRNA和β-微管蛋白基因是小袋科纤毛虫2个重要的遗传标记基因，是其分类和基因型分析的重要工具。

鱼源肠袋纤毛虫是否独立为一个新属，有待更多基因组、形态特征对比等数据来验证。对小袋科不同种属纤毛虫进行分类澄清和遗传多态性研究有助于深入了解和研究其生物学特性，对结肠小袋纤毛虫分子流行病学研究具有重要的理论指导意义。未来应对结肠小袋纤毛虫更多遗传标记基因的开发、跨宿主传播机制、致病机理等方面进行深入研究。

基金

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国家自然科学基金(31772733)
国家自然科学基金(32302898)

参考文献

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文献

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2026年第66卷第7期

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doi: 10.13343/j.cnki.wsxb.20260042

接收时间：2026-01-16
首发时间：2026-07-06
出版时间：2026-07-04

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收稿日期：2026-01-16
录用日期：2026-03-04

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The National Natural Science Foundation of China(31772733)

国家自然科学基金(31772733)

国家自然科学基金(32302898)

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^1.河南科技大学动物科技学院，河南洛阳

^2.中国科学院水生生物研究所，湖北武汉

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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