三种腹毛目纤毛虫射出胞器的显微、亚显微结构研究
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摘要
在原生动物纤毛虫中,细胞表膜或表膜附近具有一类由膜性囊泡围着的、遇刺激时能迅速发射其内含物的特殊细胞器,即射出胞器。研究表明,在不同类纤毛虫中,射出胞器在细胞生命活动中行使防御、攻击、捕食等不同的功能。目前,对纤毛虫射出胞器的研究尚局限于较低等的类群,例如对草履虫的刺丝泡射出胞器和四膜虫的黏液泡射出胞器的了解较深入；对于较高等的类群,例如对形成明显背、腹皮层及具有纤毛结构分化的腹毛目纤毛虫其射出胞器则知之不多且仅涉及极少数种类。深入观察并阐明这类胞器的起源与发生、结构与定位、射出机制与功能,对进一步认识原生动物细胞内胞器结构的多样性与复杂性、胞器的形成与遗传调控机理,以及细胞与环境的相互作用关系等问题具有重要学术意义。
本文应用微分干涉相差显微术、蛋白银标记技术、扫描电镜术、透射电镜术以及酶细胞化学技术,以尾柱虫科(Urostylidae)念珠异列虫(Anteholosticha monilata)、尖毛虫科(Oxytrichidae)颗粒尖毛虫(Oxytricha granulifera)和印度原毛虫(Architricha indica)为材料,在显微、亚显微水平上共同显示了不同类型射出胞器的结构与特点。其中在念珠异列虫中观察到类刺丝泡射出胞器,在颗粒尖毛虫观察到黏液泡射出胞器,在印度原毛虫观察到“分泌型”射出胞器,所得结果总结如下：
1念珠异列虫的类剌丝泡射出胞器
(1)胞器形态与定位射出胞器长杆状,结构由不同电子密度片层的体部、贯穿整个胞器的中央轴杆部和多层膜的帽部组成,在营养期细胞表膜下大量成列分布,而在包囊细胞中分布较少。遇外界刺激时,该胞器突破表膜射出,其射出结构呈“金针菇”状,由一个球形头部和一个细长杆部组成。
(2)胞器起源与功能射出胞器由产生至成熟的过程中,由细胞质深处单层膜包裹的小囊泡,经历了纤维物质的聚集、内外片层的分化、致密轴杆的出现及帽部结构的形成等阶段,最终成熟定位到细胞表膜下。据结果认为,念珠异列虫的射出胞器是由高尔基体活动产生的小囊泡发育而来。本文纤毛虫所制备的电镜标本均不可避免地出现胞器射出结构,说明射出胞器具有应激或防御功能；在口区及其附近位置未见胞器的射出结构,推断胞器在细胞摄食过程中未发挥功能；在包囊细胞中,仅在自噬泡内发现一个或几个消化中的射出胞器,说明射出胞器在特殊生理条件下,可能作为细胞内利用自身物质提供能量的来源之一。
(3)与其他刺丝泡类胞器的比较与草履虫的刺丝泡比较,尽管念珠异列虫的射出胞器在细胞内的发生及胞器成熟时的形态与定位有相似之处,但草履虫的刺丝泡不太敏感,念珠异列虫的射出胞器则十分敏感,其胞器在实验过程中不可避免地发生应激反应,发射其内含物；草履虫的刺丝泡射出刺丝内含物,念珠异列虫的射出胞器则射出金针菇样内含物。与伪尾柱虫的射出胞器相比较,念珠异列虫的射出胞器在细胞内的形态及胞器射出物的形态均有相似之处。据此,将念珠异列虫的射出胞器作为一种类刺丝泡射出胞器,并认为在亲缘关系较近的纤毛虫中,射出胞器结构的分化具有相似的特征。
2颗粒尖毛虫的黏液泡射出胞器
(1)胞器形态与定位黏液泡近球形,外围一层膜结构,内含高电子密度物质,其外周有刺突或丝状结构,前端有致密的纤维状物质。该结构锚定在表膜下,在细胞质深部较少观察到。射出前,其前端的致密纤维物与表膜融合,射出过程中有微丝发射行为,部分内含物被保留在细胞外表面,粗糙不光滑,有时可见其间有丝状结构相连。
(2)胞器起源与功能由于颗粒尖毛虫细胞核附近有大量粗细相近但长度不一的内质网结构,推测黏液泡的发生可能与内质网有关。该类纤毛虫是一种土壤生纤毛虫,在不良条件下极易形成包囊,黏液泡在营养期细胞中大量存在,发射后其射出物在细胞皮层表面为不定形的黏液类物质,而在包囊细胞的皮层表面却未观察到该结构的存在。由此认为,胞器的分泌物可能与包囊壁的形成有关,同时对表膜的更新有物质贡献作用。
(3)与其他黏液泡类胞器的比较颗粒尖毛虫的黏液泡外围膜结构,内含晶状物质,遇刺激后能够发射泡内物质,这与低等纤毛虫四膜虫和腹毛类纤毛虫大尾柱虫的黏液泡有相似之处。但颗粒尖毛虫的黏液泡近球形,与四膜虫长椭球形的黏液泡和大尾柱虫可分为头、体、尾三部分的黏液泡不同；胞器前端有致密纤维物锚定在表膜下,其外围有刺突或丝状结构,这与其他黏液泡外围一层平滑的膜不一样；胞器在射出过程中有微丝发射行为,这种现象在其他黏液泡射出时并未观察到。
3印度原毛虫的“分泌型”射出胞器
(1)胞器形态与定位射出胞器长椭球形,在细胞表膜下成列分布。其前端1/4处无结构物质,形成凹陷状小窝,其余部分则充满了高电子密度物质。遇外界刺激时,前端与表膜融合,分泌物呈“杯状”,可区分为具小窝的头部、略膨大的体部和稍尖的尾部三部分。
(2)胞器起源与发生射出胞器发生早期,在细胞质深部内质网附近产生各种类型的小囊泡,且小囊泡的膜内外均呈现葡糖糖-6-磷酶反应颗粒活性；胞器成熟过程中逐渐向表膜迁移,其囊泡内包裹的纤维状物质不断聚集,葡糖糖-6-磷酶反应颗粒主要集中在囊泡的外层区域,此后泡由球形变为椭球形后定位于表膜下。据结果认为,印度原毛虫的射出胞器,可能起源于内质网的膜性囊泡。
(3)胞器射出机制与功能胞器射出前,泡外层膜与泡内结构脱离,形成较大的空隙,此后其前端膜与表膜融合,泡内结构从泡、膜融合处的缺口“射出”或者排出,胞器“射出”物几乎原封不动地残留在细胞表面,呈“杯状”的形态。胞器“射出”过程可能与细胞行使某种应激或防御功能有关,胞器“射出”时细胞内线粒体大量聚集,结果表明胞器的活动不仅对促进细胞内膜结构的更新有作用,并与线粒体的功能活动相联系。
(4)与其他类射出胞器的比较印度原毛虫的射出胞器明显区别于目前已报道的类型：胞器在发生过程中泡内纤维状物质是从泡外周向中心积累的；在射出前,胞器的膜结构先与内含物脱离,其前端膜再与表膜融合；在“射出”过程中,泡内结构似乎是逐渐或缓慢地排出的,结果使射出物几乎原封不动地残留在细胞表面,对这一过程与其说是射出的过程,还不如说是分泌的过程。由此认为,印度原毛虫的射出胞器是一种“分泌型”射出胞器,这是未报道的又一类射出胞器。
4总结
本文应用显微和亚显微方法,显示并探索了三种腹毛类纤毛虫射出胞器的形态与定位、起源与发生、射出机制与功能；比较了念珠异列虫射出胞器与草履虫刺丝泡及伪尾柱虫射出胞器的异同,将念珠异列虫的射出胞器作为类刺丝泡射出胞器；通过将颗粒尖毛虫的黏液泡与四膜虫、大尾柱虫的黏液泡的比较,提出了颗粒尖毛虫的黏液泡不同于其他纤毛虫黏液泡的特征；通过对印度原毛虫射出胞器的射出过程和射出物的观察,认为印度原毛虫的射出胞器是一种“分泌型”射出胞器。所得结果,对深入认识腹毛类纤毛虫射出胞器这类特殊细胞器提供了形态学资料,并对进一步认识及阐明原生动物细胞的结构、结构形成与进化、结构功能与细胞调控等提供了基础资料。
Extrusomes are membrane-bound peculiar organelles in ciliates of protists and generally found in their cortical cytoplasm or just beneath the pellicle. They are considered to play an important role in the life activity of different kinds of ciliates, such as defensive, offensive and predation by discharging their internal content to the outside of the cell in response to stimulus. The previous researches mainly focused on some extrusomes of lower forms of ciliates, particularly trichocyst extrusomes in Paramecium and mucocyst extrusomes in Tetrahymena. To date, the research data are very limited in the hypotrichous ciliates showing differentiation of the dorsoventral cortex and functional ciliature. To study the origin and morphogenesis, morphology and localization, extrusion mechanism and functions of extrusomes is of great academic significance for the further understanding the diversity and complexity, the genetics and regulation mechanism of the organelles in protozoa cells, and the interaction relationship between the cells and the external environment.
By using differential interference contrast microscopy, protargol staining, scanning and transmission electron microscopy and enzymo-cytochemistry, the present paper demonstrates the different types of extrusomes at microscopic and submicroscopic level:trichocyst-like extrusomes in Anteholosticha monilata of Urostylidae, mucocyst extrusomes in Oxytricha granulifera of Oxytrichidae and excretion extrusomes in Architricha indica of Oxytrichidae. The results are as follows:
1Trichocyst-like extrusomes in Anteholosticha monilata
(1) Morphology and localization Extrusomes are long rod-shaped and consist of three parts: a body of stratiform structures of uneven electron density, an elongated shaft located at the center of the body and a cap including multilayer membrane-structure. A great number of extrusomes are found to arrange in rows just beneath the pellicle of vegetative cells but less in cyst. The organelles break through the pellicle and discharge their internal substances when suffered from external stimulus. The extruded structures are needle mushroom shaped including a spherical head and a long rod.
(2) Origin and function During the process of maturation, small vesicles undergo the collection of the filamentous materials, the differentiation of inner and outer layers, the appearance of dense shaft, the formation of cap structure and finally positioned in their functional areas. So these organelles may originate from membrane-limited vesicle related to Golgi apparatus. The present study reveals that the extrusomes in Anteholosticha monilata discharge inevitably during the experimental process, which suggests the organelles are involved in defense against chemical stimulus; however, no extrusomes are found near the oral region, indicating that the organelles do not play a role in the process of prey capture. The extrusomes are digested inside autophagic vacuole of cyst, suggesting the organelles may provide energy as material in the special physiological conditions.
(3) Compared with trichocysts extrusomes Although the extrusomes in Anteholosticha monilata have similarities in morphogenesis, morphology and localization with trichocysts in Paramecium, the extrusomes are sensitive to external stimulus and discharge inevitably lots of mushroom-shaped structure while trichocysts are insensitive and eject needle-shaped inclusion. In addition, these characteristics are consistent with the trichocysts-like extrusomes of Urostylidae ciliates. The results show that the extrusomes in Anteholosticha monilata are identified as trichocysts-like extrusomes and their structure may have similar differentiation characteristics in the close relationship of ciliates.
2Mucocyst extrusomes in Oxytricha granulifera
(1) Morphology and localization The mature extrusomes are suborbicular vesicles and arrange in rows beneath the pellicle. These organelles are surrounded with a plasma membrane with spike or filament structure and containing high electron dense materials. When suffered from the external stimulus, the inclusions of mucocyst are extruded and stay outside of the cell surface. The structure of the ejection seems to be not smooth and some filamentous structures are often found.
(2) Origin and function Due to lots of similar thickness but different length of endoplasmic reticulum structure distribute around the membrane of nuclear, we speculate that the formation of these organelles is connection with endoplasmic reticulum. Oxytricha granulifera is a kind of soil ciliate, which will becomes cyst easily under adverse conditions. The extruded structure are often found on the vegetative cell surface, but not on the cyst surface, which suggests the extruded structure is beneficial to the formation of the cyst wall and contributes to the pellicle retrieval or renewal.
(3) Compared with reported mucocyst extrusomes Mucocyst of Oxytricha granulifera, which are membrane-bound vesicles and filled with high election dense materials and can discharge their content when suffered from stimulus, is similar to mucocyst of lower formers of ciliates Tetrahymena and hypotrichous ciliates Urostyla grandis. However, the mucocyst in Oxytricha granulifera are suborbicular with spike or filament structure, which is different from the oval shaped mucocyst in Tetrahymena and from mucocyst Urostyla grandis which has three recognizable parts: head body and tail. In addition, the extrusomes in Oxytricha granulifera have the phenomenon of microfilament occurrence during ejection while the phenomenon is not to be observed in reported mucocyst during the process.
3Excretion extrusomes in Architricha indica
(1) Morphology and localization The mature extrusomes are ellipsoidal vesicles and arrange in rows beneath the pellicle. Two parts can be recognized according to its internal structure:a spherical cavity possibly containing structureless substance, occupies about1/4of the extrusomes volume; while the rest part is full of different electron dense materials. When suffered from the external stimulus, the extrusomes are extruded and present a cyathiform-shaped structure including a head with a cavity, a slightly swollen body and a pointed tail.
(2) Origin and morphogenesis The observation showed small vesicles are closely bounded up with the endoplasmic reticulum and developing extrusomes migrate to the cell pellicle. Moreover, the results reveal that glucose-6-phosphatase reactive granules are found not only in the endoplasmic reticulum and its nearby vesicles, but in those developing extrusomes in the cytoplasm. We can speculate that the extrusomes of Architricha indica might originate from endoplasmic reticulum.
(3) Extrusion mechanism and function Before discharge, the extrusomes expand gradually to separate its membrane from internal content forming a lager space, then the membrane fuse with the pellicle and the internal content break through the pellicle and eject out. During ejection, the entire internal content of extrusomes has not obviously morphological change and the extruded structures of extrusomes are cyathiform and remain outside of the cell surface intactly. The process of ejection of extrusomes in Architricha indica may perform stress response or defensive function, which associates with the activity of mitochondria and contributes to the structure of membrane retrieval or renewal.
(4) Compared with reported extrusomes The extrusomes of Architricha indica clearly differ from that of previously reported extrusomes in other ciliates. In their morphogenetic processes, the filamentous substances in vesicles cluster gradually from outer to center area. Before discharge, the extrusomes expand gradually to separate its membrane from internal content and then the membrane fuse with the pellicle. After extrusion, the extruded structure remain outside of the cell surface intactly appear not to "eject" but "secrete" gradually. Therefore, the extrusomes in Architricha indica are excretion extrusomes and are a new type of extrusomes without report in ciliates.
4Conclusion
By using the microscopic and submicroscopic methods, the present paper observed and explored the morphology and localization, origin and morphogenesis, extrusion mechanism and functions of different extrusomes in three kinds of hypotrichous ciliate. Based on the comparison of difference and similarities of extrusomes between Anteholosticha monilata and Paramecium and Pseudourostyla ciliates, the extrusomes in Anteholosticha monilata are identified as trichocysts-like extrusomes. The mucocyst extrusomes in Oxytricha granulifera have some obvious characteristics distinguishing from that of Tetrahymena and Urostyla grandis. The extrusomes in Architricha indica are excretion extrusomes by ejected process and extruded structure observation. The present paper supplies the morphological data in detail for the investigation on extrusomes in hypotrichous ciliates. Meanwhile, the results also provide basic information for the further understanding the structure, formation, evolution and regulation of protozoa cells.
本文应用微分干涉相差显微术、蛋白银标记技术、扫描电镜术、透射电镜术以及酶细胞化学技术,以尾柱虫科(Urostylidae)念珠异列虫(Anteholosticha monilata)、尖毛虫科(Oxytrichidae)颗粒尖毛虫(Oxytricha granulifera)和印度原毛虫(Architricha indica)为材料,在显微、亚显微水平上共同显示了不同类型射出胞器的结构与特点。其中在念珠异列虫中观察到类刺丝泡射出胞器,在颗粒尖毛虫观察到黏液泡射出胞器,在印度原毛虫观察到“分泌型”射出胞器,所得结果总结如下：
1念珠异列虫的类剌丝泡射出胞器
(1)胞器形态与定位射出胞器长杆状,结构由不同电子密度片层的体部、贯穿整个胞器的中央轴杆部和多层膜的帽部组成,在营养期细胞表膜下大量成列分布,而在包囊细胞中分布较少。遇外界刺激时,该胞器突破表膜射出,其射出结构呈“金针菇”状,由一个球形头部和一个细长杆部组成。
(2)胞器起源与功能射出胞器由产生至成熟的过程中,由细胞质深处单层膜包裹的小囊泡,经历了纤维物质的聚集、内外片层的分化、致密轴杆的出现及帽部结构的形成等阶段,最终成熟定位到细胞表膜下。据结果认为,念珠异列虫的射出胞器是由高尔基体活动产生的小囊泡发育而来。本文纤毛虫所制备的电镜标本均不可避免地出现胞器射出结构,说明射出胞器具有应激或防御功能；在口区及其附近位置未见胞器的射出结构,推断胞器在细胞摄食过程中未发挥功能；在包囊细胞中,仅在自噬泡内发现一个或几个消化中的射出胞器,说明射出胞器在特殊生理条件下,可能作为细胞内利用自身物质提供能量的来源之一。
(3)与其他刺丝泡类胞器的比较与草履虫的刺丝泡比较,尽管念珠异列虫的射出胞器在细胞内的发生及胞器成熟时的形态与定位有相似之处,但草履虫的刺丝泡不太敏感,念珠异列虫的射出胞器则十分敏感,其胞器在实验过程中不可避免地发生应激反应,发射其内含物；草履虫的刺丝泡射出刺丝内含物,念珠异列虫的射出胞器则射出金针菇样内含物。与伪尾柱虫的射出胞器相比较,念珠异列虫的射出胞器在细胞内的形态及胞器射出物的形态均有相似之处。据此,将念珠异列虫的射出胞器作为一种类刺丝泡射出胞器,并认为在亲缘关系较近的纤毛虫中,射出胞器结构的分化具有相似的特征。
2颗粒尖毛虫的黏液泡射出胞器
(1)胞器形态与定位黏液泡近球形,外围一层膜结构,内含高电子密度物质,其外周有刺突或丝状结构,前端有致密的纤维状物质。该结构锚定在表膜下,在细胞质深部较少观察到。射出前,其前端的致密纤维物与表膜融合,射出过程中有微丝发射行为,部分内含物被保留在细胞外表面,粗糙不光滑,有时可见其间有丝状结构相连。
(2)胞器起源与功能由于颗粒尖毛虫细胞核附近有大量粗细相近但长度不一的内质网结构,推测黏液泡的发生可能与内质网有关。该类纤毛虫是一种土壤生纤毛虫,在不良条件下极易形成包囊,黏液泡在营养期细胞中大量存在,发射后其射出物在细胞皮层表面为不定形的黏液类物质,而在包囊细胞的皮层表面却未观察到该结构的存在。由此认为,胞器的分泌物可能与包囊壁的形成有关,同时对表膜的更新有物质贡献作用。
(3)与其他黏液泡类胞器的比较颗粒尖毛虫的黏液泡外围膜结构,内含晶状物质,遇刺激后能够发射泡内物质,这与低等纤毛虫四膜虫和腹毛类纤毛虫大尾柱虫的黏液泡有相似之处。但颗粒尖毛虫的黏液泡近球形,与四膜虫长椭球形的黏液泡和大尾柱虫可分为头、体、尾三部分的黏液泡不同；胞器前端有致密纤维物锚定在表膜下,其外围有刺突或丝状结构,这与其他黏液泡外围一层平滑的膜不一样；胞器在射出过程中有微丝发射行为,这种现象在其他黏液泡射出时并未观察到。
3印度原毛虫的“分泌型”射出胞器
(1)胞器形态与定位射出胞器长椭球形,在细胞表膜下成列分布。其前端1/4处无结构物质,形成凹陷状小窝,其余部分则充满了高电子密度物质。遇外界刺激时,前端与表膜融合,分泌物呈“杯状”,可区分为具小窝的头部、略膨大的体部和稍尖的尾部三部分。
(2)胞器起源与发生射出胞器发生早期,在细胞质深部内质网附近产生各种类型的小囊泡,且小囊泡的膜内外均呈现葡糖糖-6-磷酶反应颗粒活性；胞器成熟过程中逐渐向表膜迁移,其囊泡内包裹的纤维状物质不断聚集,葡糖糖-6-磷酶反应颗粒主要集中在囊泡的外层区域,此后泡由球形变为椭球形后定位于表膜下。据结果认为,印度原毛虫的射出胞器,可能起源于内质网的膜性囊泡。
(3)胞器射出机制与功能胞器射出前,泡外层膜与泡内结构脱离,形成较大的空隙,此后其前端膜与表膜融合,泡内结构从泡、膜融合处的缺口“射出”或者排出,胞器“射出”物几乎原封不动地残留在细胞表面,呈“杯状”的形态。胞器“射出”过程可能与细胞行使某种应激或防御功能有关,胞器“射出”时细胞内线粒体大量聚集,结果表明胞器的活动不仅对促进细胞内膜结构的更新有作用,并与线粒体的功能活动相联系。
(4)与其他类射出胞器的比较印度原毛虫的射出胞器明显区别于目前已报道的类型：胞器在发生过程中泡内纤维状物质是从泡外周向中心积累的；在射出前,胞器的膜结构先与内含物脱离,其前端膜再与表膜融合；在“射出”过程中,泡内结构似乎是逐渐或缓慢地排出的,结果使射出物几乎原封不动地残留在细胞表面,对这一过程与其说是射出的过程,还不如说是分泌的过程。由此认为,印度原毛虫的射出胞器是一种“分泌型”射出胞器,这是未报道的又一类射出胞器。
4总结
本文应用显微和亚显微方法,显示并探索了三种腹毛类纤毛虫射出胞器的形态与定位、起源与发生、射出机制与功能；比较了念珠异列虫射出胞器与草履虫刺丝泡及伪尾柱虫射出胞器的异同,将念珠异列虫的射出胞器作为类刺丝泡射出胞器；通过将颗粒尖毛虫的黏液泡与四膜虫、大尾柱虫的黏液泡的比较,提出了颗粒尖毛虫的黏液泡不同于其他纤毛虫黏液泡的特征；通过对印度原毛虫射出胞器的射出过程和射出物的观察,认为印度原毛虫的射出胞器是一种“分泌型”射出胞器。所得结果,对深入认识腹毛类纤毛虫射出胞器这类特殊细胞器提供了形态学资料,并对进一步认识及阐明原生动物细胞的结构、结构形成与进化、结构功能与细胞调控等提供了基础资料。
Extrusomes are membrane-bound peculiar organelles in ciliates of protists and generally found in their cortical cytoplasm or just beneath the pellicle. They are considered to play an important role in the life activity of different kinds of ciliates, such as defensive, offensive and predation by discharging their internal content to the outside of the cell in response to stimulus. The previous researches mainly focused on some extrusomes of lower forms of ciliates, particularly trichocyst extrusomes in Paramecium and mucocyst extrusomes in Tetrahymena. To date, the research data are very limited in the hypotrichous ciliates showing differentiation of the dorsoventral cortex and functional ciliature. To study the origin and morphogenesis, morphology and localization, extrusion mechanism and functions of extrusomes is of great academic significance for the further understanding the diversity and complexity, the genetics and regulation mechanism of the organelles in protozoa cells, and the interaction relationship between the cells and the external environment.
By using differential interference contrast microscopy, protargol staining, scanning and transmission electron microscopy and enzymo-cytochemistry, the present paper demonstrates the different types of extrusomes at microscopic and submicroscopic level:trichocyst-like extrusomes in Anteholosticha monilata of Urostylidae, mucocyst extrusomes in Oxytricha granulifera of Oxytrichidae and excretion extrusomes in Architricha indica of Oxytrichidae. The results are as follows:
1Trichocyst-like extrusomes in Anteholosticha monilata
(1) Morphology and localization Extrusomes are long rod-shaped and consist of three parts: a body of stratiform structures of uneven electron density, an elongated shaft located at the center of the body and a cap including multilayer membrane-structure. A great number of extrusomes are found to arrange in rows just beneath the pellicle of vegetative cells but less in cyst. The organelles break through the pellicle and discharge their internal substances when suffered from external stimulus. The extruded structures are needle mushroom shaped including a spherical head and a long rod.
(2) Origin and function During the process of maturation, small vesicles undergo the collection of the filamentous materials, the differentiation of inner and outer layers, the appearance of dense shaft, the formation of cap structure and finally positioned in their functional areas. So these organelles may originate from membrane-limited vesicle related to Golgi apparatus. The present study reveals that the extrusomes in Anteholosticha monilata discharge inevitably during the experimental process, which suggests the organelles are involved in defense against chemical stimulus; however, no extrusomes are found near the oral region, indicating that the organelles do not play a role in the process of prey capture. The extrusomes are digested inside autophagic vacuole of cyst, suggesting the organelles may provide energy as material in the special physiological conditions.
(3) Compared with trichocysts extrusomes Although the extrusomes in Anteholosticha monilata have similarities in morphogenesis, morphology and localization with trichocysts in Paramecium, the extrusomes are sensitive to external stimulus and discharge inevitably lots of mushroom-shaped structure while trichocysts are insensitive and eject needle-shaped inclusion. In addition, these characteristics are consistent with the trichocysts-like extrusomes of Urostylidae ciliates. The results show that the extrusomes in Anteholosticha monilata are identified as trichocysts-like extrusomes and their structure may have similar differentiation characteristics in the close relationship of ciliates.
2Mucocyst extrusomes in Oxytricha granulifera
(1) Morphology and localization The mature extrusomes are suborbicular vesicles and arrange in rows beneath the pellicle. These organelles are surrounded with a plasma membrane with spike or filament structure and containing high electron dense materials. When suffered from the external stimulus, the inclusions of mucocyst are extruded and stay outside of the cell surface. The structure of the ejection seems to be not smooth and some filamentous structures are often found.
(2) Origin and function Due to lots of similar thickness but different length of endoplasmic reticulum structure distribute around the membrane of nuclear, we speculate that the formation of these organelles is connection with endoplasmic reticulum. Oxytricha granulifera is a kind of soil ciliate, which will becomes cyst easily under adverse conditions. The extruded structure are often found on the vegetative cell surface, but not on the cyst surface, which suggests the extruded structure is beneficial to the formation of the cyst wall and contributes to the pellicle retrieval or renewal.
(3) Compared with reported mucocyst extrusomes Mucocyst of Oxytricha granulifera, which are membrane-bound vesicles and filled with high election dense materials and can discharge their content when suffered from stimulus, is similar to mucocyst of lower formers of ciliates Tetrahymena and hypotrichous ciliates Urostyla grandis. However, the mucocyst in Oxytricha granulifera are suborbicular with spike or filament structure, which is different from the oval shaped mucocyst in Tetrahymena and from mucocyst Urostyla grandis which has three recognizable parts: head body and tail. In addition, the extrusomes in Oxytricha granulifera have the phenomenon of microfilament occurrence during ejection while the phenomenon is not to be observed in reported mucocyst during the process.
3Excretion extrusomes in Architricha indica
(1) Morphology and localization The mature extrusomes are ellipsoidal vesicles and arrange in rows beneath the pellicle. Two parts can be recognized according to its internal structure:a spherical cavity possibly containing structureless substance, occupies about1/4of the extrusomes volume; while the rest part is full of different electron dense materials. When suffered from the external stimulus, the extrusomes are extruded and present a cyathiform-shaped structure including a head with a cavity, a slightly swollen body and a pointed tail.
(2) Origin and morphogenesis The observation showed small vesicles are closely bounded up with the endoplasmic reticulum and developing extrusomes migrate to the cell pellicle. Moreover, the results reveal that glucose-6-phosphatase reactive granules are found not only in the endoplasmic reticulum and its nearby vesicles, but in those developing extrusomes in the cytoplasm. We can speculate that the extrusomes of Architricha indica might originate from endoplasmic reticulum.
(3) Extrusion mechanism and function Before discharge, the extrusomes expand gradually to separate its membrane from internal content forming a lager space, then the membrane fuse with the pellicle and the internal content break through the pellicle and eject out. During ejection, the entire internal content of extrusomes has not obviously morphological change and the extruded structures of extrusomes are cyathiform and remain outside of the cell surface intactly. The process of ejection of extrusomes in Architricha indica may perform stress response or defensive function, which associates with the activity of mitochondria and contributes to the structure of membrane retrieval or renewal.
(4) Compared with reported extrusomes The extrusomes of Architricha indica clearly differ from that of previously reported extrusomes in other ciliates. In their morphogenetic processes, the filamentous substances in vesicles cluster gradually from outer to center area. Before discharge, the extrusomes expand gradually to separate its membrane from internal content and then the membrane fuse with the pellicle. After extrusion, the extruded structure remain outside of the cell surface intactly appear not to "eject" but "secrete" gradually. Therefore, the extrusomes in Architricha indica are excretion extrusomes and are a new type of extrusomes without report in ciliates.
4Conclusion
By using the microscopic and submicroscopic methods, the present paper observed and explored the morphology and localization, origin and morphogenesis, extrusion mechanism and functions of different extrusomes in three kinds of hypotrichous ciliate. Based on the comparison of difference and similarities of extrusomes between Anteholosticha monilata and Paramecium and Pseudourostyla ciliates, the extrusomes in Anteholosticha monilata are identified as trichocysts-like extrusomes. The mucocyst extrusomes in Oxytricha granulifera have some obvious characteristics distinguishing from that of Tetrahymena and Urostyla grandis. The extrusomes in Architricha indica are excretion extrusomes by ejected process and extruded structure observation. The present paper supplies the morphological data in detail for the investigation on extrusomes in hypotrichous ciliates. Meanwhile, the results also provide basic information for the further understanding the structure, formation, evolution and regulation of protozoa cells.
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