悬游双眉虫(原生动物,纤毛门)无性生殖期间的形态发生

应用蛋白银染色技术研究了悬游双眉虫青岛种群无性生殖期间皮层结构和核器的演化过程,其主要特征为:后仔虫口原基独立地出现于紧靠虫体左侧第一根横棘毛的皮层下小龛,其中毛基粒不参与其它棘毛原基的形成;老的口围带发生后半部的局部重建而非整个的由前仔虫简单继承;在前仔虫中,波动膜原基来自老结构的反分化,而在后仔虫中则来自口原基;所有棘毛原基均为独立发生并与老结构没有任何关系;在前仔虫中,口棘毛(即左侧第一根额棘毛)来自于波动膜的反分化,而在后仔虫中则为独立发生;背触毛列于老的结构当中产生,并由最右一列原基演化出3根尾棘毛;两大核片段的改组带从一端向另一端移动 ,并随着两者的融合而消失.文中同时对前人有关该属发生模式的若干存疑问题做了探讨 [动物学报 54(3):517-524,2008].     

海洋纤毛虫黄色伪角毛虫无性生殖期间的形态发生

利用蛋白银染色技术研究了海洋纤毛虫———黄色伪角毛虫Pseudokeronopsisflava (Cohn ,186 6 )Wirns berger,Larsen&Uhlig ,1987无性生殖期间的细胞发生学。其主要特征为 :1)前仔虫口原基以独立发生的方式出现并独特地形成于口前庭右侧的皮层深处 ,由其对老口围带进行完全的更新 ;2 )老口器不参与新口器的形成 ,完全被吸收 ;3)前仔虫的额 -腹 -横棘毛原基同样为独立发生 ,老结构可能不参加其随后的发育 ;4 )后仔虫的口原基、波动膜原基及额 -腹 -横棘毛原基均来自最初排列无序的毛基粒发生场 ;5 )背触毛及缘棘毛的更新发生在老的结构中 ,并向前后延伸取代老结构 ;6 )在整个发生过程中 ,无大核融合现象。文中同时对该种所表现的发生学特征及系统学意义做了探讨     

腹毛目纤毛虫皮膜演化的一种特殊形式—卵圆急纤虫无性二分裂期间的形态发

对海洋纤毛虫卵圆急纤虫无性分裂过程中的形态发生了做了跟踪观察,结果表明,该种的皮膜演化表现了一系列在腹毛目种类中所罕见的特征;１．新老口围带有一临时性汇联为一的阶段;２．左右缘棘毛原基分步出现,即明显的不同步现象;３．大核的改组在细胞分裂开始前即已完成;４．本种后仔虫之棘毛原基的分化既非独立发生又非来自于老结构的反分化,而是紧靠口原基的一侧向外分衍并极可能是孤立地发展而成;５．老的波动膜在形态发生     

四核舍太虫的细胞发生学与Helix E10-1二级结构分析

利用蛋白银染色技术,观察和研究海洋游仆虫-四核舍太虫Certesia quadrinucleata(纤毛门,游仆目)二分裂期间的形态发生学。其主要特征如下:(1)老口围带完全被前仔虫继承;(2)后仔虫口原基独立产生于皮膜深层;(3)老口侧膜参与前仔虫口侧膜原基形成,前后仔虫的口侧膜原基均发生于细胞表面, 向前贡献出第一根额腹棘毛;(4)额-腹-横棘毛以初级5原基模式产生, 且以"3:3:3:3:3"的方式分化出新的棘毛;(5)背触毛与左缘棘毛原基均来自老结构, 无尾棘毛产生。研究首次给出了背面纤毛器的发生图示,为进一步探讨舍太虫的系统地位提供了一份补足性的发生学基础资料。游仆目纤毛虫的核糖体小亚基基因HelixE10-1区域二级结构一共存在9种模式, 该区域序列长度的变异性揭示了游仆目纤毛虫在进化中可能处于比较特殊的地位。       

腹毛目纤毛虫皮膜演化的一种特殊形式--卵圆急纤虫无性二分裂期间的形态发生学

对海洋纤毛虫卵圆急纤虫无性分裂过程中的形态发生做了跟踪观察,结果表明,该种的皮膜演化表现了一系列在腹毛目种类中所罕见的特征:1)新老口围带有一临时性汇联为一的阶段;2)左右缘棘毛原基分步出现,即明显的不同步现象;3)大核的改组在细胞分裂开始前即已完成;4)本种后仔虫之棘毛原基的分化既非独立发生又非来自于老结构的反分化,而是紧靠口原基的一侧向外分衍并极可能是孤立地发展而成;5)老的波动膜在形态发生过程中亦无反分化现象,而呈独特的半原基模式.     

毛尾刺虫无性生殖周期中的形态和形态发生

利用蛋白银染色法研究了毛尾刺虫的形态及无性生殖周期中的形态发生,其过程为：（１）后仔虫口原基出现在左缘棘毛内侧深层,其内的毛基体组装成整齐排列的小膜并分化成新ＡＺＭ１,ＡＺＭ２和口侧膜,（２）前仔虫口原基出现在老仔虫ＡＺＭ２之前方深处,其随后发育成前仔虫的ＡＺＭ２口侧膜及ＡＺＭ１的一部分,并更新老结构的ＡＺＭ１中第７－１１片小膜,（３）额腹横棘毛原基为５列,分别以３：３：２：２：３方式分化最终产出     

冠突伪尾柱虫的腹皮层纤毛器微管胞器及其形态发生

应用荧光紫杉醇直接荧光标记和抗α-微管蛋白抗体免疫荧光标记方法,显示冠突伪尾柱虫腹皮层纤毛器微管胞器由口围带、波动膜、额腹横棘毛和左右缘棘毛等纤毛器微管、纤毛器基部附属微管等组成。口围带基部含小膜托架及与托架相联系的肋壁微管,其中领部小膜托架间由"∧"形微管相联接;额腹横棘毛基部含前纵微管束、后纵微管束、横微管束和周围微管束,其微管在不同棘毛基部的发达程度不一,其中两列中腹棘毛基部微管紧密联系成一条粗绳索样结构,且左、右中腹棘毛基部的横微管束定向相反;左、右缘棘毛基部含前纵微管束、后纵微管束和横微管束,其中横微管束不发达。与目前已知的腹毛目纤毛虫例如贻贝棘尾虫、魏氏拟尾柱虫的纤毛器基部微管相比较,冠突伪尾柱虫腹皮层纤毛器基部微管除具有腹毛目纤毛虫纤毛器基部微管的基本特征外,也具有一些特殊的组成模式。皮层纤毛器微管形态发生中,前仔虫口围带并非全部是由老口围带更新而来的,其老口围带只有翻领部发生更新,且翻领部与领部接续处有一小段老的翻领部小膜保留,领部的小膜保留,结果其领部小膜、接续处保留的小膜与更新的翻领部小膜三部分共同组成前仔虫的新口围带。在后仔虫口原基发生的位置,其邻近的老横棘毛没有变化,此时老的横棘毛或许能起到"参照点"或定位作用;各类纤毛器发生、分化过程中,处于非原基区的老额棘毛、横棘毛及左右缘棘毛在较长时间内均未见明显的变化。它们可能是在新结构形成时仍然起到运动作用继而逐渐失去功能而退化瓦解的。     

变藓棘毛虫的形态学重描述及细胞发生学研究

通过活体观察和蛋白银染色法对采自青岛沙滩半咸水的变藓棘毛虫Sterkiella histriomuscorum(纤毛门, 腹毛目)进行了形态学及细胞发生学研究。该种群形态学与前人报道的土壤及淡水种群基本一致: 虫体近长椭圆形, 活体大小约(100-160) m (40-75) m; 无皮层颗粒; 2938片口小膜; 额棘毛3根; 额腹棘毛4根; 口后腹棘毛3根; 横前腹棘毛2根; 横棘毛3-5根; 左右缘棘毛列分别由17-23、20-24根棘毛组成; 6列背触毛; 2枚大核。其主要发生学特征如下: (1)老口围带完全保留, 老波动膜解体重建; 后仔虫口原基独立发生; (2)额腹横棘毛为5原基次级发生式, 部分原基来自老棘毛解体, 以2:3:3:4:4方式分化为新棘毛; (3)缘棘毛原基产生于老结构中, 并向两极延伸逐渐形成前后仔虫的新结构; (4)背触毛发生为典型Oxytricha模式; (5)大核在发生过程中完全融合。研究对首次在半咸水生境中发现的变藓棘毛虫种群进行了活体形态学和纤毛图式描述, 补充了显微照片、性状统计数据及发生过程的细节信息。       

阔口尖毛虫无性生殖和生理改组过程的比较

阔口尖毛虫无性生殖中先后发生后口围带原基,前、后的波动膜原基,额腹横棘毛原基和左、右缘棘毛原基,老口围带也在此期间更新,结果形成2套新纤毛结构,原老结构瓦解消失;生理改组时按同样顺序产生口围带原基等几类腹面纤毛原基,结果形成1套新纤毛结构,替换老结构。在这两个截然不同的过程中,新纤毛结构的分化和老纤毛结构退化时也表现出某些相似的特征。作者据此推测,这种纤毛虫无性生殖和生理改组中,纤毛原基的发生、发育和定位在细胞控制机理上可能是相同的。     

小腔游仆虫形态学、个体发育与分子系统学研究

游仆类是纤毛虫中进化最为复杂和高等的一大类群,为了进一步探索和完善游仆类的多样性,本研究利用活体观察、蛋白银和银浸法染色技术对采自青岛小西湖的小腔游仆虫(Euplotes aediculatus)的形态学及细胞发生学进行了详尽的研究,并在完整的形态学及发生学研究基础上,测定了小腔游仆虫的核糖体小亚基基因(SSU r DNA)序列,通过序列比较和分子系统树构建等方法,对小腔游仆虫的系统地位进行了分析。结果表明:本种鉴别特征为9根额腹棘毛,5根横棘毛,2根缘棘毛,2根尾棘毛,8列背触毛,double-eurystomus型银线系。发生学特征包括:(1)后仔虫口原基在表皮下独立发生,前仔虫完全继承老口围带;(2)额–腹–横棘毛原基从左向右按照3:3:3:2:2的模式形成额腹棘毛和横棘毛;(3)前后仔虫最左侧额腹棘毛分别由独立产生的原基形成;(4)缘棘毛原基独立发生;(5)初级背触毛原基来自虫体中部老结构的反分化;(6)前后仔虫尾棘毛分别来自最右侧2列背触毛原基和老背触毛列末端;这些特征显示出游仆虫属个体发生模式的高度保守性。分子系统分析与形态学数据一致,即游仆虫属为单元发生,且小腔游仆虫与艾美游仆虫(Euplotes amieti)、阔口游仆虫(E.eurystomus)和伍氏游仆虫(E.woodruffi)聚在一起。     

Morphologie und Morphogenese des Bodenciliaten Oxytricha granulifera sp.n. (CiIiophora, Oxytrichidae)

Oxytricha granulifera sp.n. differs from other members of the genus by its subpellicular granules and the strongly shortened dorsal kinety 4. The overall pattern of the morphogenetic events is similar to that known from other Oxytrichidae. However, the oral primordium evolves de novo between the left marginal cirral row and the postoral cirri. The six anlagen of the frontoventral cirri are of different origin. Two anlagen of the proter evolve from parental frontal cirri, two from the opisthe, and one includes basal bodies of the proter and opisthe. Two anlagen of the opisthe evolve from the oral primordium, and three primordia originate from the postoral cirri. Frontal cirrus 1 evolves from the paroral membrane in the proter, and from the oral primordium and the anlagen of the frontoventral cirri in the opisthe. The genus Oxytricha can be subdivided into several groups with regard to the origin of its oral primordium and the development of the frontoventral cirri. The morphogenesis of the dorsal kineties in the Hypotrichida is reviewed. Seven different modes of origin are distinguished. We conclude that morphogenetic features cannot be used in the classification of the Hypotrichida at the generic level, because we have too little information to decide whether special morphogenetic features are important at the generic or species level.     

Cortical structure in non-dividing and dividing Diophrys japonica spec. nov. (Ciliophora, Euplotida) with notes on morphological variation

The morphology and morphogenesis of Diophrys japonica spec. nov., isolated from the Mie Port, Nagasaki, Japan, were investigated from life and following impregnation with protargol. The new species is recognized by the following characters: Body elliptical in outline and slightly greyish to yellowish in color; size in vivo about 80-120 x 50-70 microm; pellicle flexible, with underlying granules densely arranged in lines; ciliature comprising about 30-46 adoral membranelles, 4-7 frontal, 1-4 ventral and 4-7 transverse cirri, always 1 left marginal and 3 caudal cirri, and 4 dorsal kineties; usually two macronuclear nodules; fragment kinety with 2-5 dikinetids; marine habitat. The main morphogenetic events are: (1) the opisthe's oral primordium develops de novo in a subsurface pouch near the left transverse cirri; (2) the proter retains the parental AZM except for reorganization of some proximal membranelles; (3) cirral anlagen for the frontal, ventral and transverse cirri in both dividers develop separately from the oral primordium or parental cirri, and are derived from the separation of primary primordia that originate de novo; (4) the anlagen for the left marginal cirrus and fragment kinety also form de novo and separately; (5) dorsal kinety anlagen occur within the parental structures at mid-body and posterior end of the cell, of which the right-most one contributes three caudal cirri from its posterior portion. Based on available ontogenetic data, the author proposes that the numbers of left marginal and caudal cirri can be regarded as reliable characters for species identification, while the numbers of frontal, ventral and transverse cirri are not consistent enough for species distinction. A key to the eleven adequately known species of Diophrys is presented.     

Morphogenesis of the marine ciliate, Pseudoamphisiella alveolata (Kahl, 1932) Song & Warren, 2000 (Ciliophora, Stichotrichia, Urostylida) during binary fission

Morphogenesis during the binary fission of the stichotrich ciliate Pseudoamphisiella alveolata, isolated from Jiaozhou Bay near Qingdao, China, was investigated using protargol silver impregnation. The process is characterized as follows: (1) in the proter, only the posterior part of the parental adoral zone of membranelles is renewed, where the membranelles dedifferentiate and then rebuild the UM-anlage and the missing membranelles, (2) the oral primordium in the opisthe and the FVT-anlagen in both dividers are formed de novo on the cell surface, (3) an "extra" anlage, which is generated on the right of the right marginal anlage, develops into three or four "extra" marginal cirri that connect the caudal cirri with the marginal rows, (4) the right marginal anlage is formed within the old structure, (5) the FVT-cirri develop in a primary mode, and (6) unlike most stichotrichs, the right marginal anlagen in both dividers generate closely together. As an additional contribution, the diversity of morphogenetic patterns within the genus Pseudoamphisiella is discussed. Based on both morphogenetic and SS rRNA gene sequencing data, the systematic position of the genus Pseudoamphisiella as well as the family Pseudoamphisiellidae Song et al. 1997 is briefly analyzed. The results indicate that they should very possibly represent a higher evolved group in the order Urostylida.     

Morphogenetic characters of the model ciliate Euplotes vannus (Ciliophora,Spirotrichea): Notes on cortical pattern formation during conjugational and postconjugational reorganization

Ciliated protists represent a morphologically and genetically distinct group of single-celled eukaryotes which can reproduce asexually and sexually. Morphogenesis occurs in both asexual and sexual modes of reproduction which is of interest for researchers investigating cell differentiation, regeneration, systematics and evolution. However, studies of morphogenesis have concentrated almost entirely on the asexual mode. Here we use protargol staining to investigate the morphogenetic processes during sexual reproduction in the model species Euplotes vannus (Müller). The major events include: (1) two rounds of morphogenesis occur during sexual reproduction, i.e., conjugational and postconjugational reorganization; (2) in both processes the oral primordium is generated de novo in a pouch beneath the cortex; (3) the frontoventral-transverse cirri anlagen are formed de novo and fragment in a 3:3:3:3:2 pattern; (4) the leftmost cirrus and the paroral membrane do not change during conjugational morphogenesis, but reorganize de novo during postconjugational morphogenesis; (5) marginal cirral anlagen are formed de novo in both morphogenetic processes; (6) two or three caudal cirri are formed at the ends of the rightmost two or three old dorsal kineties; (7) the dorsal kineties are retained entirely. These results can serve as reference to investigate the morphogenetic events in the different stages of sexual reproduction.     

Divisional morphogenesis in the marine ciliate, Hemigastrostyla enigmatica (Dragesco & Dragesco-Kernéis, 1986) and redefinition of the genus Hemigastrostyla Song & Wilbert, 1997 (Protozoa, Ciliophora)

Morphogenetic events during the division of the marine hypotrichous ciliate, Hemigastrostyla enigmatica (Dragesco & Dragesco-Kernéis, 1986) Song & Wilbert, 1997 are described. The morphogenesis is characterized by:(1) 5 frontoventral-transverse cirral anlagen develop into 8 frontal, 5 ventral and 5 transverse cirri after Oxytricha-pattern;(2) there may be 6 FVT-anlagen in some individuals giving rise to more cirri which, however, will be resorbed after division;(3) anlage of the right marginal row at least in the opisthe occurs de novo right to the parental structures instead of within them;(4) according to the origin, the two extra ventral cirri right to transverse ones are not ventral or transverse cirri, which are from the retained old structure;(5) dorsal kineties originate from one group of DK-anlagen in both dividers with, very uniquely, an additional fragmentation of DK1, and(6) oral primordia will be formed in both dividing parts, from which the newly-built membranelles in the proter replace the posterior part of the parental AZM with a particular 'piecing together mode.Some features during the morphogenesis (e.g. variable number of cirral anlagen, presence of primary primordia, the mode of rebuilding of the proter's adoral zone of membranelles, origin of dorsal kineties and caudal cirri etc.) indicate that the genus Hemigastrostyla might present a intermediate form between oxytrichids and other related higher taxa. Based on our new observations, an improved diagnosis for genus Hemigastrostyla is given: marine or brackish water Oxytrichidae with slightly to conspicuously cephalized body shape; mostly 8–10 frontal, 5 ventral, 5 transverse and two to several extra ventral cirri to the right of the transverse ones, which are from the retained parental structure; caudal cirri present.     

Morphogenesis in the marine spirotrichous ciliate Apokeronopsis crassa (Claparède & Lachmann, 1858) n. comb. (Ciliophora: Stichotrichia), with the establishment of a new genus, Apokeronopsis n. g., and redefinition of the genus Thigmokeronopsis

Morphogenetic events during the division of the marine spirotrichous ciliate, Apokeronopsis crassa (Claparède & Lachmann 1858) n. comb. were investigated. Compared with members of the well-known genera Thigmokeronopsis, Uroleptopsis, and Pseudokeronopsis, A. crassa has one row of buccal cirri, high number of transverse cirri, clearly separated midventral rows, lacks thigmotactic cirri and a gap in adoral zone, its undulating membranes (UMs) anlage forms one cirrus and marginal rows and dorsal kineties form apokinetally during division. All these characteristics indicate that this organism represents a new taxon at the generic level, and hence a new genus is suggested, Apokeronopsis n. g. It is defined as thus: Pseudokeronopsidae with Pseudokeronopsis-like bicorona of frontal cirri and one marginal row on each side; one row of two or more buccal cirri in ordinary position; two midventral rows distinctly separated, hence of cirri that are not in a typical zig-zag pattern; high number of transverse cirri, caudal cirri absent, and frontoterminal cirri present; thigmotactic cirri absent, many macronuclear nodules fuse into many masses as well as marginal and dorsal kineties form apokinetally during morphogenesis. At the same time, the genus ThigmokeronopsisWicklow, 1981 is redefined, and one new combination, Apokeronopsis antarctica (Petz, 1995) n. comb. is proposed. The morphogenetic events of A. crassa are characterized as follows: (1) In the proter, the adoral zone of membranelles and UMs are completely renewed by the oral primordium. The UM anlage is formed apokinetally on the dorsal wall of the buccal cavity and is hence clearly separated from the frontoventral-transverse (FVT) cirral anlagen in the proter. (2) Frontoventral-transverse cirral anlagen are generated de novo in the outermost region of the cortex to the right of the old UMs. (3) A row of buccal cirri arises from FVT cirral streak I. (4) The marginal rows and dorsal kineties originate de novo in both dividers; no caudal cirri are formed. (5) The last FVT-streak contributes two frontoterminal cirri. (6) The many macronuclear nodules fuse into many masses (about 50 segments) during division, unlike a singular or branched mass as described in other urostylids.     

The morphology and morphogenesis of a marine ciliate,Epiclintes auricularis rarisetus nov. sspec. (Ciliophora,Epiclintidae), from the Yellow Sea

The morphology and morphogenesis of a marine ciliate, Epiclintes auricularis rarisetus nov. sspec., collected from Qingdao, northern China were studied on live and protargol-impregnated specimens. This isolate can be recognized by its elongate, contractile and tripartite body with a size of 200-400×20-40 μm in vivo, about 30 ventrally located adoral membranelles, short undulating membranes, 2-3 frontal and 10-18 transverse cirri as well as 8-9 fronto-midventral rows, 22-31 left and 35-54 right marginal cirri, but no caudal cirri, invariably 3 dorsal kineties, of which kineties 1 and 3 link together anteriorly, 24-70 macronuclear nodules, marine habitat, and contribution of almost all frontal-midventral transverse cirral anlagen (FVT anlagen) to the formation of fronto-midventral rows. In its divisional morphogenesis, this species demonstrates some features rarely if ever found in any other urostyloids, e.g. partial replacement of the old adoral zone of membranelles, de novo formation of the oral primordium and the anlagen for marginal rows and dorsal kineties, contribution of almost all FVT anlagen to the transverse cirri, lack of frontoterminal cirri. In addition, its morphogenesis appears to differ in detail from that of E. auricularis auricularis nov. stat., based on published data. These remarkable morphogenetic traits of Epiclintes suggests a questionable systematic position for this genus in the Urostyloidea, therefore, further researches are urgently needed.     

Morphogenesis of the marine spirotrichous ciliate, Trachelostyla pediculiformis (Cohn, 1866) (Ciliophora, Stichotrichia), with consideration of its phylogenetic position

The cortical development during binary fission of the relatively poorly known stichotrich ciliate, Trachelostyla pediculiformis (Cohn, 1866) Borror, 1972, found in coastal waters near Qingdao, China, was investigated using the protargol impregnation method. The morphogenetic process reveals some pretty unusual characteristics, which do not follow the Oxytricha-pattern: (1) the parental oral apparatus is entirely renewed from an oral primordium formed de novo in the proter; (2) in the proter, the parental undulating membranes are not involved in the formation of the newly formed oral primordium; both undulating membrane-anlagen (UM-anlage) and frontoventral-transverse cirral anlagen (FVT-anlagen) develop from the oral primordium in the proter; (3) the dorsal kineties (DK) are generated in a unique way, that is, in both dividers, two separate groups of DK-anlagen develop in the right- and left-most DK, generate all the DK and evolve to replace the old structures; (4) three caudal cirri are formed at the posterior ends of three right-most dorsal kinety anlagen; (5) eight frontal, five ventral and five transverse cirri are derived from six streaks, namely, the UM-anlage and 5 FVT-anlagen; the cirri are segregated from these anlagen in the pattern 1:3:3:3:4:4 (from left to right) in the Oxytricha mode. Based on both SSrRNA gene sequencing and morphogenetic data, the systematic positions of the genus Trachelostyla Borror, 1972 as well as the family Trachelostylidae Small and Lynn, 1985 are briefly analyzed. The results indicate that this genus/family could be a highly isolated lineage and might be ancestral to other well-known oxytrichids.