纤毛虫重要类群的细胞发生模式研究
摘要
纤毛门原生动物由于独特的接合生殖方式、高度特化的细胞器结构以及分司不同功能的两种核型而在细胞生物学、在真核生物遗传学以及生物进化等研究领域,尤其是在“细胞分化与反分化”这一生命基本命题探讨中具有特别的研究价值和意义。因此,当今对纤毛虫原生动物的细胞发生学研究始终是广义的动物学研究中最活跃的领域之一。国际间近二十年的工作表明:即便在研究最为“彻底的”的广义“腹毛类”内,仍有大量类群处于未知、半知甚至空白状态,大量探索性研究等待深入,大量的争议、混乱和错误等待修订。
本工作选取了15个具有典型发生学代表性的种类为材料,分别隶属纤毛门内2纲、8目、13个科,研究涉及了(国际间迄今模式研究空白、发生学过程长期残缺或不详的)细胞发生模式、皮膜及核器演化、部分所代表类群的系统关系探讨以及新系统构建,揭示了一系列新的发生学现象并澄清了若干发生学上的悬疑,填补了相关类群的多项空白。同时,对前人在系统关系安排等工作中的若干不当、以及错误解读之处进行了修正和调整。主要成果包括:
1)在对代表种类凯毛虫、原盘头虫等个体发生学研究、比较以及小核糖体亚基单位(SSU rRNA)基因序列分析的基础上,将原腹毛类(亚目)升格为亚纲,将盘头类升格为目,并将其亚纲级隶属关系做了变更(移至排毛亚纲内)以及确立了尖颈虫科、双轴虫属和偏角毛虫属的系统地位等;
2)在国际间首次揭示并建立了1个亚纲(原腹毛亚纲)、1个科(尖颈虫科)、5个属(原盘头虫属、偏角毛虫属、双轴虫属、哈特曼虫属、突口虫属)的细胞发生模式,同时对迄今存在误释、缺失或不详的后尾柱虫属、齿管虫属、双眉虫属、伪小双虫属等4个属的发生模式做了修正和补足;
3)完成了对海洋凯毛虫个体发生学的详细追踪,首次阐明了本属细胞发生过程中的“所有棘毛均参与新原基的构建”、“前后仔虫波动膜来自同一由老波动膜反分化而来的原基”及“棘毛原基与背触毛原基混杂”等极其独特的发生学现象,此为迄今在旋唇类纤毛虫发生学研究中的首例报道。这一现象也揭示了凯毛虫代表着低等的异毛类向高等的腹毛类进化的一个过渡型,把原本认为没有任何联系的异毛类和腹毛类有机的联系在一起。从而为确定上述3个类群间真实的演化和隶属关系提供了重要的依据,据此并结合分子信息建立了新亚纲,原腹毛亚纲。本研究全面解决了由上世纪后半叶以来至今长期存疑的凯毛虫科系统定位及其与腹毛类等亚纲间的系统关系等问题;
4)对博罗原盘头虫的细胞发生学和系统学研究表明,盘头类纤毛虫代表了一个介于广义游仆类和排毛亚纲之间的过渡类群:其后仔虫口原基、缘棘毛原基、波动膜以及背触毛原基的发育与排毛类相类似,而尾棘毛的产生则是典型的游仆类模式。据此我们得以推断:盘头类与排毛类具有更为接近的亲缘关系,并提议将之归入排毛亚纲。此外,鉴于其特殊的发生学特征和系统地位,我们的研究建议将盘头亚目提升为一目级阶元;
5)对条形尖颈虫发生模式的研究,填补了该科、属发生学的研究空白,首次详细地给出了其无性分裂期的基本发生模式,证实了该科的细胞发生属于FVT棘毛的5-原基发生模式,表明了尖颈虫科与尖毛虫科具有较近的亲缘关系;由于其(表现型)存在前仔虫口原基、棘毛原基独立发生、老口围带完全被吸收、独特的背触毛原基以及尾棘毛发生模式等特征,证实了尖颈虫代表了一独立的科级阶元;此外,在与(经典系统中所安排的)尖颈虫科内的殖口虫属的发生学进行比较研究基础上,鉴于后者典型的尖毛虫发生模式,而明确将其重新归入尖毛虫科。
6)对厚偏角毛虫和卵圆偏角毛虫细胞发生过程中大核低度融合、缘棘毛和背触毛原基独立发生以及中腹棘毛列相互分离等特征的揭示,对于探讨相近类群(伪角毛虫属,趋角毛虫属)的发生多元性具有重要意义,并为伪角毛虫科的系统演化及科下分组提供了重要依据,本工作同时确立该偏角毛虫属(新属)的地位;
7)首次报道了双轴虫属的发生模式。鉴于本属缘棘毛原基发生及演化过程中所表现的罕见特征,我们对双轴虫属给出了新定义并且揭示了其与伪尾柱虫属较近的亲缘关系,进一步确立了双轴虫属长期争议的系统地位,将其归入伪尾柱虫科。进而探讨和整理了伪尾柱虫科的基本发生模式,并澄清了对该科在系统进化中所表现的相关趋同现象的错误理解。由此,对双轴虫属做了新定义:具明确分化的独立额棘毛及额棘毛、口棘毛各一列,具横棘毛和额前棘毛,尾棘毛缺失;右缘棘毛单列,左缘棘毛多列并由在老结构外产生的独立原基发育而来。
8)对泡状伪小双虫和拉氏伪小双虫的皮层演化过程进行了详细的重新研究,弥补了前人工作中若干发生学细节的缺失。表明本属中此两个已知种在前仔虫口原基的发育(部分更新vs.完全保留)以及额-腹-横棘毛原基的产生(初级发生型vs.次级发生型)上有显著的差异,据此证明伪小双虫属内发生模式呈较高的多态性,从不同侧面丰富了人们对属内-种间细胞结构分化以及发生学差异度的认知,也为探讨属内发生学保守性提供了新资料;
9)对中华后尾柱虫的发生学研究表明,尽管被安排入后尾柱虫属(即与属内其它种具有相同或相似的纤毛图式),但本种在发生学上具有三个独有特征:大核在发生过程中形成单一的融合体;无额外腹棘毛列的形成阶段;中腹棘毛列从未延伸至虫体末端。鉴于发生过程中的大核行为,后尾柱虫属可能系伪角毛虫科与(广义的)全列虫属之间的一个过渡类群,并与全列虫属有着较近的关系;
10)对中华哈特曼虫的细胞发生进行了详细的追踪研究,建立了属级发生模式,并证明了管口类科属间口器发生的高度稳定性,基本特征为:后仔虫的口前和围口动基列均来自口原基,虫体中部3列口后动基列参与口原基的形成,而亲体口前和围口动基列完全保留;老的胞口和咽杆在发生中期会被新结构所取代;在发生晚期,异质大核的主体和副体会发生核质融合;
11)对摩涅齿管虫细胞发生过程进行了补足性重描述,为本类群的发生学了解添加了新的资料;
12)首次揭示和描述了异毛目中叶状突口虫发生期间前仔虫口原基及波动膜原基的反分化以及体动基列中毛基粒对无规则增殖的现象,并建立了本属的细胞发生模式;
13)详细追踪了棕色尖毛虫、卡龙游仆虫以及偏寡毛双眉虫三个种的细胞分裂和皮层演化过程,为研究相应属内发生学保守性研究提供了补足性资料。
成果此外还包括:根据发生学和形态学所给出的重要指征建立了若干新的分类阶元,包括1新属(偏角毛虫属)、2新种(中华哈特曼虫、中华后尾柱虫)和3新组合(厚偏角毛虫、南极偏角毛虫和卵圆偏角毛虫)。
对新建立的偏角毛虫属做了定义:具冠状排列的额棘毛双列,左右缘棘毛各一列,口棘毛2根以上;两列中腹棘毛列明显相互分离,即中腹棘毛不呈典型的锯齿状排布,横棘毛高度发达,数目众多;无尾棘毛,具有额前棘毛,无触毛区棘毛;在细胞发生的过程中大核融合并形成多个融合体,而非单一大团;缘棘毛和背触毛原基为独立发生式。
本研究同时对趋角虫属进行了修订:具冠状排列额棘毛,左右缘棘毛各一列;两列中腹棘毛列明显相互分离;额前棘毛存在,横棘毛不发达,具有触毛区棘毛;在细胞发生过程中,缘棘毛及背触毛原基独立发生。
Ciliated protozoa is the most complex and highly differentiated among single-celled organisms,of which some groups are closely related to occurrence of red-tide, bait-animal culture in aquaculture, disease of animal, water quality purification in polysaprobic environment, material cycle and energy flow in“micro-food-loop”. Moreover, owe to its special nuclear dualism, complicated morphogenetic process and unique sexual reproduction (conjugation), ciliates play important roles in molecular biology, cell biology and researches on relationship between nucleus and cytoplasm.
(1) Main stages of general cortical development during binary division of Kiitricha marina which are characterized by“numerous”unique features that are never observed in all other known spirotrichs were investigated, the features are as follows: (1) no new UM-anlage in the opisthe is formed, hence the undulating membranes in the opisthe derive from the division of the de-differentiated parental ones; (2) no marginal cirral rows are formed; (3) no typical dorsal kineties (DK) are generated (i.e. several rows of pre-DK consisting of dikinetids with both basal bodies ciliated appear to mix with either isolated small cirri or cirral fragments; (4) all cirral anlagen are formed interkinetally from the de-differentiated old cirral rows; and (5) unlike all other known“hypotrichs”(s. l.), the differentiation of ciliature from the primordia occurs mostly after cell division. Based on all the morphogenetic information showing above, we conclude that Kiitricha may represent an intermediate taxon between heterotrichs (s. l.) and the Stichotrichia-Hypotrichia-complex and could be similar to the ancestor form of the latter group. So, Kiitricha might be placed in the class Spirotrichea at about subclass level, next to the Phaconidiidia, Hypotrichia, and Stichotrichia. We support the establishment of a new subclass Protohypotrichia Shi et al., 1999 within the class Spirotrichea and we transfer all taxa assigned to the order Kiitrichida to it.
(2) The morphogenetic data reveals that the discocephalines have both stichotrich and hypotrich characteristics. Five morphogenetic characters are highly characteristic of the stichotrichs and only two morphogenetic features that are shared by discocephalines and euplotids to the exclusion of stichotrichs. On balance, therefore, the morphogenetic data suggest that the discocephalines are more closely related to the stichotrichs than the hypotrichs. Based on the data all above, we suggest that the discocephalines should have the taxonomic rank of order (i.e. Discocephalida Wicklow, 1982) as suggested by Puytorac et al. (1993), within the subclass Stichotrichia.
(3) In general, the developmental pattern of Trachelostyla pediculiformis conforms to that of oxytrichids, i.e. typical 5-FVT-anlagen mode. It differs conspicuously in some other morphogenetic features from typical oxytrichids in some key points, e.g. the old AZM is renewed completely in the proter; the generation mode of dorsal kineties is of“one group type”, and all three caudal cirri derive from the right-most 3 DK-anlagen (or kineties), and only two old dorsal kineties are involved in the construction of the new anlagen and the DK-anlage fragmentation occurs only in the single, left-most primordium. We conclude from the available morphogenetic evidence that: (1) Trachelostylidae, represented by the genus Trachelostyla, might be a lower but sister group to the closely-related family Oxytrichidae; (2) Gonostomum should be assigned to Oxytrichidae (s. l.).
(4) The origin of marginal and dorsal kineties anlagen in Apokeronopsis crassa and Apokeronopsis ovalis, derived de novo, that means have no business with the old structure, and the developmental mode of the macronuclear nodules of this species (fuse into many masses) were shown for the first time, and Apokeronopsis nov. gen. was established based on morphogenetic data combined. Additionally we conclude that Apokeronopsis is a transitional form between the Pseudokeronopsinae and Thigmokeronopsis as it shares morphogenetic features with both.
(5) The morphogenetic process of Diaxonella trimarginata almost follows a typical urostylid mode, one of the most significant morphogenetic features in D. trimarginata is the mode of the formation of the left marginal rows: only a single anlage develops intrakinetally to the right of the old left marginal rows, and then develops to replace the old structures, which is similar to that described in Pseudourostyla cristata and Pseudourostyla nov. The morphogenetic data suggest a close relationship between the genera Diaxonella and Pseudourostyla and we place the genus Diaxonella in the family Pseudourostylidae Jankowski, 1979.
(6) Morphogenesis of Pseudoamphisiella alveolata was investigated and compared with its congener P. lacazei. With reference to the dissimilarities, we judged that morphogenetic patterns in the genus Pseudoamphisiella are not conservative. Moreover, on the same basis we suggest that the family Pseudoamphisiellidae might likewise be a highly evolved taxon within Urostylida and the assertion that pseudoamphisiellids at family level is justified.
(7) The morphogenetic process of Metaurostylopsis sinica was thoroughly investigated. Compared with its congener M. marina, whose morphogenetic features was known, M. sinica shows some unique characteristics: the macronuclear nodules fuse into a single sausage-like mass; FVT-anlage streak n forms two frontoterminal cirri, an extra ventral cirrus and a transverse cirrus, i.e. typical urostylid mode but not a Metaurostylopsis mode. Metaurostylopsis was considered to be the sister group of the Pseudokeronopsidae because the macronucleus already shows the tendency not to fuse to a compact, globular mass and M. sinica is considered to be closer to the lower urostylids while M. marina is closer to the pseudokeronopsids.
(8) The complete morphogenetic process of Chlamydodon mnemosyne (including the development of terminal and equtorial fragmnets) was reported and contributed new information to the understandability of morphogenesis of cyrtophorids.
(9) As for morphogenesis, Hartmannula sinica correspond well with its congeners. This suggests that the mode of cell development among congeners could be very stable. The basic morphogenetic characteristics of H. sinica are in accordance with most other cyrtophorids (e.g. Hartmannulopsis, Thigmogaster, Chlamydonyx, Trochilioides, Brooklynella, Chilodonella and Trithigmostoma) in two main aspects: (1) the parental oral ciliature (circumoral and preoral kineties) is retained for the proter, and (2) the new oral primordium for the opisthe originates from several inner left somatic kineties.
(10) Morphogenesis of Condylostoma spatiosum proceeds basically as described in a previous report and can be summarized as follows: (1) the parental adoral zone of membranelles is partly dedifferentiated and then renewed in the posterior portion; (2) in the proter, both the frontal cirri and the paroral membrane are newly formed by anlagen derived from the disaggregated old structures; (3) in the opisthe the anlagen of the paroral membrane and the frontal cirri develop from the right margin of the oral primordium; (4) two frontal cirri are formed one after the other by the frontal cirral anlage; (5) during morphogenesis, no recognizable duplication of basal bodies takes place in somatic kineties; and (6) macronucleus divides after prior fusion.
(11) The morphogenetic process of Oxytricha, Euplotes charon and Diophrys apoligothrix thoroughly described and compared with their congeners. The result shows that the morphogenesis of these 4 genera is highly conservative.
(12) A new genus Apokeronopsis nov. gen. was established and other two Thigmokeronopsis Wicklow, 1981 and Diaxonella Jankowski, 1979 are redefined.
(13) Two new combinations, Apokeronopsis crassa (Claparède & Lachmann, 1858) Shao et al. 2008 and Apokeronopsis antarctica (Petz, 1995) Shao et al. 2008 are proposed. Two new species, Hartmannula sinica and Metaurostylopsis sinica, are also suggested.
本工作选取了15个具有典型发生学代表性的种类为材料,分别隶属纤毛门内2纲、8目、13个科,研究涉及了(国际间迄今模式研究空白、发生学过程长期残缺或不详的)细胞发生模式、皮膜及核器演化、部分所代表类群的系统关系探讨以及新系统构建,揭示了一系列新的发生学现象并澄清了若干发生学上的悬疑,填补了相关类群的多项空白。同时,对前人在系统关系安排等工作中的若干不当、以及错误解读之处进行了修正和调整。主要成果包括:
1)在对代表种类凯毛虫、原盘头虫等个体发生学研究、比较以及小核糖体亚基单位(SSU rRNA)基因序列分析的基础上,将原腹毛类(亚目)升格为亚纲,将盘头类升格为目,并将其亚纲级隶属关系做了变更(移至排毛亚纲内)以及确立了尖颈虫科、双轴虫属和偏角毛虫属的系统地位等;
2)在国际间首次揭示并建立了1个亚纲(原腹毛亚纲)、1个科(尖颈虫科)、5个属(原盘头虫属、偏角毛虫属、双轴虫属、哈特曼虫属、突口虫属)的细胞发生模式,同时对迄今存在误释、缺失或不详的后尾柱虫属、齿管虫属、双眉虫属、伪小双虫属等4个属的发生模式做了修正和补足;
3)完成了对海洋凯毛虫个体发生学的详细追踪,首次阐明了本属细胞发生过程中的“所有棘毛均参与新原基的构建”、“前后仔虫波动膜来自同一由老波动膜反分化而来的原基”及“棘毛原基与背触毛原基混杂”等极其独特的发生学现象,此为迄今在旋唇类纤毛虫发生学研究中的首例报道。这一现象也揭示了凯毛虫代表着低等的异毛类向高等的腹毛类进化的一个过渡型,把原本认为没有任何联系的异毛类和腹毛类有机的联系在一起。从而为确定上述3个类群间真实的演化和隶属关系提供了重要的依据,据此并结合分子信息建立了新亚纲,原腹毛亚纲。本研究全面解决了由上世纪后半叶以来至今长期存疑的凯毛虫科系统定位及其与腹毛类等亚纲间的系统关系等问题;
4)对博罗原盘头虫的细胞发生学和系统学研究表明,盘头类纤毛虫代表了一个介于广义游仆类和排毛亚纲之间的过渡类群:其后仔虫口原基、缘棘毛原基、波动膜以及背触毛原基的发育与排毛类相类似,而尾棘毛的产生则是典型的游仆类模式。据此我们得以推断:盘头类与排毛类具有更为接近的亲缘关系,并提议将之归入排毛亚纲。此外,鉴于其特殊的发生学特征和系统地位,我们的研究建议将盘头亚目提升为一目级阶元;
5)对条形尖颈虫发生模式的研究,填补了该科、属发生学的研究空白,首次详细地给出了其无性分裂期的基本发生模式,证实了该科的细胞发生属于FVT棘毛的5-原基发生模式,表明了尖颈虫科与尖毛虫科具有较近的亲缘关系;由于其(表现型)存在前仔虫口原基、棘毛原基独立发生、老口围带完全被吸收、独特的背触毛原基以及尾棘毛发生模式等特征,证实了尖颈虫代表了一独立的科级阶元;此外,在与(经典系统中所安排的)尖颈虫科内的殖口虫属的发生学进行比较研究基础上,鉴于后者典型的尖毛虫发生模式,而明确将其重新归入尖毛虫科。
6)对厚偏角毛虫和卵圆偏角毛虫细胞发生过程中大核低度融合、缘棘毛和背触毛原基独立发生以及中腹棘毛列相互分离等特征的揭示,对于探讨相近类群(伪角毛虫属,趋角毛虫属)的发生多元性具有重要意义,并为伪角毛虫科的系统演化及科下分组提供了重要依据,本工作同时确立该偏角毛虫属(新属)的地位;
7)首次报道了双轴虫属的发生模式。鉴于本属缘棘毛原基发生及演化过程中所表现的罕见特征,我们对双轴虫属给出了新定义并且揭示了其与伪尾柱虫属较近的亲缘关系,进一步确立了双轴虫属长期争议的系统地位,将其归入伪尾柱虫科。进而探讨和整理了伪尾柱虫科的基本发生模式,并澄清了对该科在系统进化中所表现的相关趋同现象的错误理解。由此,对双轴虫属做了新定义:具明确分化的独立额棘毛及额棘毛、口棘毛各一列,具横棘毛和额前棘毛,尾棘毛缺失;右缘棘毛单列,左缘棘毛多列并由在老结构外产生的独立原基发育而来。
8)对泡状伪小双虫和拉氏伪小双虫的皮层演化过程进行了详细的重新研究,弥补了前人工作中若干发生学细节的缺失。表明本属中此两个已知种在前仔虫口原基的发育(部分更新vs.完全保留)以及额-腹-横棘毛原基的产生(初级发生型vs.次级发生型)上有显著的差异,据此证明伪小双虫属内发生模式呈较高的多态性,从不同侧面丰富了人们对属内-种间细胞结构分化以及发生学差异度的认知,也为探讨属内发生学保守性提供了新资料;
9)对中华后尾柱虫的发生学研究表明,尽管被安排入后尾柱虫属(即与属内其它种具有相同或相似的纤毛图式),但本种在发生学上具有三个独有特征:大核在发生过程中形成单一的融合体;无额外腹棘毛列的形成阶段;中腹棘毛列从未延伸至虫体末端。鉴于发生过程中的大核行为,后尾柱虫属可能系伪角毛虫科与(广义的)全列虫属之间的一个过渡类群,并与全列虫属有着较近的关系;
10)对中华哈特曼虫的细胞发生进行了详细的追踪研究,建立了属级发生模式,并证明了管口类科属间口器发生的高度稳定性,基本特征为:后仔虫的口前和围口动基列均来自口原基,虫体中部3列口后动基列参与口原基的形成,而亲体口前和围口动基列完全保留;老的胞口和咽杆在发生中期会被新结构所取代;在发生晚期,异质大核的主体和副体会发生核质融合;
11)对摩涅齿管虫细胞发生过程进行了补足性重描述,为本类群的发生学了解添加了新的资料;
12)首次揭示和描述了异毛目中叶状突口虫发生期间前仔虫口原基及波动膜原基的反分化以及体动基列中毛基粒对无规则增殖的现象,并建立了本属的细胞发生模式;
13)详细追踪了棕色尖毛虫、卡龙游仆虫以及偏寡毛双眉虫三个种的细胞分裂和皮层演化过程,为研究相应属内发生学保守性研究提供了补足性资料。
成果此外还包括:根据发生学和形态学所给出的重要指征建立了若干新的分类阶元,包括1新属(偏角毛虫属)、2新种(中华哈特曼虫、中华后尾柱虫)和3新组合(厚偏角毛虫、南极偏角毛虫和卵圆偏角毛虫)。
对新建立的偏角毛虫属做了定义:具冠状排列的额棘毛双列,左右缘棘毛各一列,口棘毛2根以上;两列中腹棘毛列明显相互分离,即中腹棘毛不呈典型的锯齿状排布,横棘毛高度发达,数目众多;无尾棘毛,具有额前棘毛,无触毛区棘毛;在细胞发生的过程中大核融合并形成多个融合体,而非单一大团;缘棘毛和背触毛原基为独立发生式。
本研究同时对趋角虫属进行了修订:具冠状排列额棘毛,左右缘棘毛各一列;两列中腹棘毛列明显相互分离;额前棘毛存在,横棘毛不发达,具有触毛区棘毛;在细胞发生过程中,缘棘毛及背触毛原基独立发生。
Ciliated protozoa is the most complex and highly differentiated among single-celled organisms,of which some groups are closely related to occurrence of red-tide, bait-animal culture in aquaculture, disease of animal, water quality purification in polysaprobic environment, material cycle and energy flow in“micro-food-loop”. Moreover, owe to its special nuclear dualism, complicated morphogenetic process and unique sexual reproduction (conjugation), ciliates play important roles in molecular biology, cell biology and researches on relationship between nucleus and cytoplasm.
(1) Main stages of general cortical development during binary division of Kiitricha marina which are characterized by“numerous”unique features that are never observed in all other known spirotrichs were investigated, the features are as follows: (1) no new UM-anlage in the opisthe is formed, hence the undulating membranes in the opisthe derive from the division of the de-differentiated parental ones; (2) no marginal cirral rows are formed; (3) no typical dorsal kineties (DK) are generated (i.e. several rows of pre-DK consisting of dikinetids with both basal bodies ciliated appear to mix with either isolated small cirri or cirral fragments; (4) all cirral anlagen are formed interkinetally from the de-differentiated old cirral rows; and (5) unlike all other known“hypotrichs”(s. l.), the differentiation of ciliature from the primordia occurs mostly after cell division. Based on all the morphogenetic information showing above, we conclude that Kiitricha may represent an intermediate taxon between heterotrichs (s. l.) and the Stichotrichia-Hypotrichia-complex and could be similar to the ancestor form of the latter group. So, Kiitricha might be placed in the class Spirotrichea at about subclass level, next to the Phaconidiidia, Hypotrichia, and Stichotrichia. We support the establishment of a new subclass Protohypotrichia Shi et al., 1999 within the class Spirotrichea and we transfer all taxa assigned to the order Kiitrichida to it.
(2) The morphogenetic data reveals that the discocephalines have both stichotrich and hypotrich characteristics. Five morphogenetic characters are highly characteristic of the stichotrichs and only two morphogenetic features that are shared by discocephalines and euplotids to the exclusion of stichotrichs. On balance, therefore, the morphogenetic data suggest that the discocephalines are more closely related to the stichotrichs than the hypotrichs. Based on the data all above, we suggest that the discocephalines should have the taxonomic rank of order (i.e. Discocephalida Wicklow, 1982) as suggested by Puytorac et al. (1993), within the subclass Stichotrichia.
(3) In general, the developmental pattern of Trachelostyla pediculiformis conforms to that of oxytrichids, i.e. typical 5-FVT-anlagen mode. It differs conspicuously in some other morphogenetic features from typical oxytrichids in some key points, e.g. the old AZM is renewed completely in the proter; the generation mode of dorsal kineties is of“one group type”, and all three caudal cirri derive from the right-most 3 DK-anlagen (or kineties), and only two old dorsal kineties are involved in the construction of the new anlagen and the DK-anlage fragmentation occurs only in the single, left-most primordium. We conclude from the available morphogenetic evidence that: (1) Trachelostylidae, represented by the genus Trachelostyla, might be a lower but sister group to the closely-related family Oxytrichidae; (2) Gonostomum should be assigned to Oxytrichidae (s. l.).
(4) The origin of marginal and dorsal kineties anlagen in Apokeronopsis crassa and Apokeronopsis ovalis, derived de novo, that means have no business with the old structure, and the developmental mode of the macronuclear nodules of this species (fuse into many masses) were shown for the first time, and Apokeronopsis nov. gen. was established based on morphogenetic data combined. Additionally we conclude that Apokeronopsis is a transitional form between the Pseudokeronopsinae and Thigmokeronopsis as it shares morphogenetic features with both.
(5) The morphogenetic process of Diaxonella trimarginata almost follows a typical urostylid mode, one of the most significant morphogenetic features in D. trimarginata is the mode of the formation of the left marginal rows: only a single anlage develops intrakinetally to the right of the old left marginal rows, and then develops to replace the old structures, which is similar to that described in Pseudourostyla cristata and Pseudourostyla nov. The morphogenetic data suggest a close relationship between the genera Diaxonella and Pseudourostyla and we place the genus Diaxonella in the family Pseudourostylidae Jankowski, 1979.
(6) Morphogenesis of Pseudoamphisiella alveolata was investigated and compared with its congener P. lacazei. With reference to the dissimilarities, we judged that morphogenetic patterns in the genus Pseudoamphisiella are not conservative. Moreover, on the same basis we suggest that the family Pseudoamphisiellidae might likewise be a highly evolved taxon within Urostylida and the assertion that pseudoamphisiellids at family level is justified.
(7) The morphogenetic process of Metaurostylopsis sinica was thoroughly investigated. Compared with its congener M. marina, whose morphogenetic features was known, M. sinica shows some unique characteristics: the macronuclear nodules fuse into a single sausage-like mass; FVT-anlage streak n forms two frontoterminal cirri, an extra ventral cirrus and a transverse cirrus, i.e. typical urostylid mode but not a Metaurostylopsis mode. Metaurostylopsis was considered to be the sister group of the Pseudokeronopsidae because the macronucleus already shows the tendency not to fuse to a compact, globular mass and M. sinica is considered to be closer to the lower urostylids while M. marina is closer to the pseudokeronopsids.
(8) The complete morphogenetic process of Chlamydodon mnemosyne (including the development of terminal and equtorial fragmnets) was reported and contributed new information to the understandability of morphogenesis of cyrtophorids.
(9) As for morphogenesis, Hartmannula sinica correspond well with its congeners. This suggests that the mode of cell development among congeners could be very stable. The basic morphogenetic characteristics of H. sinica are in accordance with most other cyrtophorids (e.g. Hartmannulopsis, Thigmogaster, Chlamydonyx, Trochilioides, Brooklynella, Chilodonella and Trithigmostoma) in two main aspects: (1) the parental oral ciliature (circumoral and preoral kineties) is retained for the proter, and (2) the new oral primordium for the opisthe originates from several inner left somatic kineties.
(10) Morphogenesis of Condylostoma spatiosum proceeds basically as described in a previous report and can be summarized as follows: (1) the parental adoral zone of membranelles is partly dedifferentiated and then renewed in the posterior portion; (2) in the proter, both the frontal cirri and the paroral membrane are newly formed by anlagen derived from the disaggregated old structures; (3) in the opisthe the anlagen of the paroral membrane and the frontal cirri develop from the right margin of the oral primordium; (4) two frontal cirri are formed one after the other by the frontal cirral anlage; (5) during morphogenesis, no recognizable duplication of basal bodies takes place in somatic kineties; and (6) macronucleus divides after prior fusion.
(11) The morphogenetic process of Oxytricha, Euplotes charon and Diophrys apoligothrix thoroughly described and compared with their congeners. The result shows that the morphogenesis of these 4 genera is highly conservative.
(12) A new genus Apokeronopsis nov. gen. was established and other two Thigmokeronopsis Wicklow, 1981 and Diaxonella Jankowski, 1979 are redefined.
(13) Two new combinations, Apokeronopsis crassa (Claparède & Lachmann, 1858) Shao et al. 2008 and Apokeronopsis antarctica (Petz, 1995) Shao et al. 2008 are proposed. Two new species, Hartmannula sinica and Metaurostylopsis sinica, are also suggested.
引文
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