前口虫一新种的表膜内表面结构的电镜观察
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摘要
本文利用常规透射电镜技术，改进样品制备方法的扫描电镜技术对前口虫(Frontonia sp.)的整体形态，表膜下结构和口部前庭动基列结构进行了研究。得出了以下结果：
(一)用改进的扫描电镜样品制备方法处理前口虫，使其细胞质外流后细胞膜自动外翻。从而可以在扫描电镜下从内部观察前口虫的表膜构造。
(二)在扫描电镜下可以观察到表膜下存在着三层纤维系统，分别是紧挨着表膜的由细纤维组成的网格状纤维层；第二层是出现在射出胞器周围的极细纤维束；第三层是排列规律符合右手定则的粗大纤维，故推测为动纤丝。这三层纤维有机的结合成为一个整体，推测其功能为：1、使射出体、线粒体等胞器固着在细胞表膜或细胞质中；2、作为细胞骨架的一部分，支持着细胞的正常形态。
(三)结合扫描电镜和透射电镜观察结果，在前口虫体内新看到了多种射出胞器：一种位于口部的特殊形态刺丝泡，另外还有毒素泡和粘液泡。另有其它3种类射出胞器,它们的内部组分及功能尚不清楚，所以它们归属哪一类射出体也无法确定。
(四)在高倍扫描电镜下观察到了前口虫的三排前庭动基列的微管组成。每条动基列都是由若干个粗大的柱形体规则排列而成的。并且在三条动基列间存在着数条细纤维将相邻两列连接起来。猜想这一结构可以使前口虫的胞口有力地张开和闭合，增强其摄食的能力。
This paper uses the conventional transmission electron microscope technology,improved sample preparation method for scanning electron microscopy to deal withFrontonia sp. in overall shape; surface membrane structure and export structure werestudied. Results are as follow:
(一)With improved scanning electron microscope (SEM) sample preparation methods todeal with Frontonia sp., make its cytoplasm outflow of cytomembraneautomatically. Then observe the inside structures under the SEM.
(二) There are three layers of fiber system under the surface membrane, the first one isthe fine fiber grid which next to the membrane, second layer is seen in the injectionorganelle around very fine fiber bundle, the third layer is coarse fiber, arrangedwith the right hand rule, supposedly kinetodesma. These three layers of fiber joinedas an organic whole, its functions are:1, the extrusome, mitochondria and otherorganelles fix in the cell surface membrane or cytoplasm;2, as a part of thecytoskeleton, supporting cells of the normal form.
(三) Combined with the results of scanning electron microscope and transmissionelectron microscope（TEM）, we can see multiple new kinds of extrusome inFrontonia sp. in the see injection organelles: one is a kind of trichocyst is locatedat the oral of Frontonia sp., besides that are toxicysts and mucocysts. While theother three kinds of extrusome, their internal components and functions is not clear,so they belong to which kind of extrusome is not determined.
(四) Under the high power SEM discovered that the vestibular kinety of Frontonia sp. ismade of three rows of microtubules. Each kinety base columns are composed of aplurality of thick cylinder arranged. And between three vestibular kinety are anumber of fine fibers which link the adjacent columns. Guess this structure canmake Frontonia sp. effectively to open and close its oral area, enhance its feedingcapacity.
(一)用改进的扫描电镜样品制备方法处理前口虫，使其细胞质外流后细胞膜自动外翻。从而可以在扫描电镜下从内部观察前口虫的表膜构造。
(二)在扫描电镜下可以观察到表膜下存在着三层纤维系统，分别是紧挨着表膜的由细纤维组成的网格状纤维层；第二层是出现在射出胞器周围的极细纤维束；第三层是排列规律符合右手定则的粗大纤维，故推测为动纤丝。这三层纤维有机的结合成为一个整体，推测其功能为：1、使射出体、线粒体等胞器固着在细胞表膜或细胞质中；2、作为细胞骨架的一部分，支持着细胞的正常形态。
(三)结合扫描电镜和透射电镜观察结果，在前口虫体内新看到了多种射出胞器：一种位于口部的特殊形态刺丝泡，另外还有毒素泡和粘液泡。另有其它3种类射出胞器,它们的内部组分及功能尚不清楚，所以它们归属哪一类射出体也无法确定。
(四)在高倍扫描电镜下观察到了前口虫的三排前庭动基列的微管组成。每条动基列都是由若干个粗大的柱形体规则排列而成的。并且在三条动基列间存在着数条细纤维将相邻两列连接起来。猜想这一结构可以使前口虫的胞口有力地张开和闭合，增强其摄食的能力。
This paper uses the conventional transmission electron microscope technology,improved sample preparation method for scanning electron microscopy to deal withFrontonia sp. in overall shape; surface membrane structure and export structure werestudied. Results are as follow:
(一)With improved scanning electron microscope (SEM) sample preparation methods todeal with Frontonia sp., make its cytoplasm outflow of cytomembraneautomatically. Then observe the inside structures under the SEM.
(二) There are three layers of fiber system under the surface membrane, the first one isthe fine fiber grid which next to the membrane, second layer is seen in the injectionorganelle around very fine fiber bundle, the third layer is coarse fiber, arrangedwith the right hand rule, supposedly kinetodesma. These three layers of fiber joinedas an organic whole, its functions are:1, the extrusome, mitochondria and otherorganelles fix in the cell surface membrane or cytoplasm;2, as a part of thecytoskeleton, supporting cells of the normal form.
(三) Combined with the results of scanning electron microscope and transmissionelectron microscope（TEM）, we can see multiple new kinds of extrusome inFrontonia sp. in the see injection organelles: one is a kind of trichocyst is locatedat the oral of Frontonia sp., besides that are toxicysts and mucocysts. While theother three kinds of extrusome, their internal components and functions is not clear,so they belong to which kind of extrusome is not determined.
(四) Under the high power SEM discovered that the vestibular kinety of Frontonia sp. ismade of three rows of microtubules. Each kinety base columns are composed of aplurality of thick cylinder arranged. And between three vestibular kinety are anumber of fine fibers which link the adjacent columns. Guess this structure canmake Frontonia sp. effectively to open and close its oral area, enhance its feedingcapacity.
引文
1田果.龚大洁.闫礼.漆朝辉.原生动物在生物监测中的作用与应用.生物学通报.2009，10:11~13
2沈蕴芬.原生动物学.科学出版社(北京).1999.
3Carey, P.G. Marine interstitial ciliztes. Chapman and Hall,1992:351
4Ehrenberg, C.G. Die Infusionsthierchen als volkommene Organismen. Ein Blick in dastiefere organische Leben der Natur.1838:612
5龙红岸.山东沿海部分沙隙及内共生栖纤毛虫的分类学.中国海洋大学硕士论文.2007:1~28
6顾福康,邹士法，李艺松，倪兵.镰游仆虫腹面皮层细胞骨架的扫描电镜观察.动物学报.2003，49（4）：514~521
7顾福康,张作人.一种游仆虫皮层纤维结构的扫描电镜研究.动物学研究.1991,12(4):337~341
8周素娟,尹飞,生欣,顾福康.冠突尾柱虫皮层纤毛器微管胞器及其形态发生.动物学报.2008,54(2):299~308
9顾福康,倪兵.包囊游仆虫休眠期包囊的超微结构研究.实验生物学报.1995,28(2):163~171
10丁文乔,陈瑛,田佳玉,焦迪,邱子健.用于纤毛虫特氏前口虫皮层观察的扫描电镜样品制备的改进方法.电子显微学报.2011,30(6):574~576
11Corliss, J.O. The ciliated protozoa. Characterization, classification and guide to theliterature.2nded. Pergamon Press, Oxford.1979
12顾福康.原生动物学概论.高等教育出版社.1991:69~112
13宋微波等译《原生动物学》（Haussmann等著《Protistology》2003，Schwarzer出版社）.中国海洋大学出版社.2007
14Gortz,H.D.,Kuhlmann,H.W.,Mollenbeck,M.,Tiedtke,A.,Kusch,J.,Schmidt,H.J.,Milake,A. Intra-and intercellular communication systems in ciliates. Naturwissenschaften.1999,86:422~434
15Hausmann, K. Protozoologie. Thieme.1985
16高晶,周围,乔良,陈瑛,邱子健.一种用于观察纤毛虫表膜下结构的扫描电镜样品制备新方法.2011,9:1004~1007
17Hacke-Bell, B., Hohenberger-Bregger, R., Plattner, H. Trichocysts of Paramecium:secretory organells in search of their function. Eur.J.Protistol.1990,25:289~305
18Frankel, J. Cell biology of Terahymena thermuphila. In: Asai, D.J. Forney, J.D.(ed),Terahymena thermuphila. Academic Press, San Diego London.1999:27~108
19Giovanna, R. and Leizia, M. Extrusomes in ciliates: diversification, distribution, andphylogenetic implication. The Journal of Eukaryotic Microbiology.2003,50(6):383~402
20Fokin, S.I. Rediscovery and characterization of Frontnia fusca (Quennerstedt,1896)Kahl,1931(Ciliphora, Peniculia). Denisia23:251~259
21Harold FM et al.Microbiol Rev,1990,54:381
22MavRae TH. Eur J Biochem.1997,244:265
23Hartwell LH et al. Nature.1999,402:C47
24Hausamann, K. Extrusive organelles in Protists. Int. Rev. Qtol.1978,52:197~276
25Adoutte, A. Exocytosis: biogenesis, transport and secretion of trichocysts.1988Gortz, H.D., Paramecium.Springer-Verlag, Berlin.
26Ehret C F, Haller G. Origin development and maturation of organelle system of thecell surface in Paramecium. Ultrasture.1963,6:1~42
27Raikov, I.B., Kovaleva, V.G. Fine structure of circumoral ciliature and pharyngealapparatus of the marine ciliate Helicoprodon gigas (Gymnostomata). Protistologica,1980,16:177~198
28Kruger. Die Ttichocysten der Ciliaten im Dunkelfedbild. Zoological,1936,34:1~83
29Wessenberg, H., Antipa,G. Capture and ingestion of Paramecium by Disiniumnasustum. Protozool.1970,17:240~270
30Frankel,J. Cell biology of Tetrahymena thermuphila. San Diego, London.1999:27~108
31周瑶.腹毛目纤毛虫新尾柱虫微管胞器和射出胞器的研究.华东师范大学博士论文.2010:56~104
32Kalatenbach A. Nahbrungsauswshl and Nahrungsaufnahme holotricher Zilaten derDonaulitioralzone and im Myriophyllum-Aufwuchs des DonausltwassersGansechaufel. Wass.Abwass. Wien.1962:157-185
33Kahl A.Urtier oder Protozoa. Tierweld Dtl.1931,21:181~398
34Poter E D. The buccal organelles in Paramecium awrelia during fission andconjugation with special reference to the kinetosome. Protozool.1960,7:211~217
35史新柏.草履虫口器的形态和形态发生的意义.动物学报.1980,26:289~300
36陈晶，邱子健，王瑞清，秦宇虹，2005运用激光扫描共聚焦显微技术研究大草履虫口胞器与消化胞器动物学报51（4）：718~722
37宋微波.尖前口虫的口器发生研究(纤毛门，膜口目).水生生物学报.1995,19(3):257~262
38邹士法,张作人.银白前口虫Frontonia leucas Ehrenberg切割口器再生研究.华东师范大学学报(自然科学版).1990,3:89~93
39Long H.,Song W., Gong J., Hu X.,Ma H.,Zhu M., Wang M. Frontonia lynni n. sp., anew marine ciliate(Protozoa, Ciliophpra,Hymenostomatida)from Qingdao, China.Zootaxa.2005,1003:57~64
40Foissner, W., Leipe, D. Morphology and ecology of Siroloxophyllum utriculariae(penard,1922) n.g., n.comb.(Ciliophora, Pleurostormatida) and an improvesclassification of plurostomatid ciliates. Eukaryot. Microbiol.1995,42:476~490
41Berger, H. Monograph of the oxytrichidae (Ciliophora, Hypotrophy).1999, KluwerAcademic Publishers Netherlands
42Estbve, J.C. Cytochimie ultrastructurale du trichocyste in situ chez Parameciumcaudatum. Protistologica.1974,10:371~378
43Allman, G.J. On the occurrence among Infusoria of peculiar organs resemblingthread cell. Q.J.Microse. Sci.1855,3:177~179
2沈蕴芬.原生动物学.科学出版社(北京).1999.
3Carey, P.G. Marine interstitial ciliztes. Chapman and Hall,1992:351
4Ehrenberg, C.G. Die Infusionsthierchen als volkommene Organismen. Ein Blick in dastiefere organische Leben der Natur.1838:612
5龙红岸.山东沿海部分沙隙及内共生栖纤毛虫的分类学.中国海洋大学硕士论文.2007:1~28
6顾福康,邹士法，李艺松，倪兵.镰游仆虫腹面皮层细胞骨架的扫描电镜观察.动物学报.2003，49（4）：514~521
7顾福康,张作人.一种游仆虫皮层纤维结构的扫描电镜研究.动物学研究.1991,12(4):337~341
8周素娟,尹飞,生欣,顾福康.冠突尾柱虫皮层纤毛器微管胞器及其形态发生.动物学报.2008,54(2):299~308
9顾福康,倪兵.包囊游仆虫休眠期包囊的超微结构研究.实验生物学报.1995,28(2):163~171
10丁文乔,陈瑛,田佳玉,焦迪,邱子健.用于纤毛虫特氏前口虫皮层观察的扫描电镜样品制备的改进方法.电子显微学报.2011,30(6):574~576
11Corliss, J.O. The ciliated protozoa. Characterization, classification and guide to theliterature.2nded. Pergamon Press, Oxford.1979
12顾福康.原生动物学概论.高等教育出版社.1991:69~112
13宋微波等译《原生动物学》（Haussmann等著《Protistology》2003，Schwarzer出版社）.中国海洋大学出版社.2007
14Gortz,H.D.,Kuhlmann,H.W.,Mollenbeck,M.,Tiedtke,A.,Kusch,J.,Schmidt,H.J.,Milake,A. Intra-and intercellular communication systems in ciliates. Naturwissenschaften.1999,86:422~434
15Hausmann, K. Protozoologie. Thieme.1985
16高晶,周围,乔良,陈瑛,邱子健.一种用于观察纤毛虫表膜下结构的扫描电镜样品制备新方法.2011,9:1004~1007
17Hacke-Bell, B., Hohenberger-Bregger, R., Plattner, H. Trichocysts of Paramecium:secretory organells in search of their function. Eur.J.Protistol.1990,25:289~305
18Frankel, J. Cell biology of Terahymena thermuphila. In: Asai, D.J. Forney, J.D.(ed),Terahymena thermuphila. Academic Press, San Diego London.1999:27~108
19Giovanna, R. and Leizia, M. Extrusomes in ciliates: diversification, distribution, andphylogenetic implication. The Journal of Eukaryotic Microbiology.2003,50(6):383~402
20Fokin, S.I. Rediscovery and characterization of Frontnia fusca (Quennerstedt,1896)Kahl,1931(Ciliphora, Peniculia). Denisia23:251~259
21Harold FM et al.Microbiol Rev,1990,54:381
22MavRae TH. Eur J Biochem.1997,244:265
23Hartwell LH et al. Nature.1999,402:C47
24Hausamann, K. Extrusive organelles in Protists. Int. Rev. Qtol.1978,52:197~276
25Adoutte, A. Exocytosis: biogenesis, transport and secretion of trichocysts.1988Gortz, H.D., Paramecium.Springer-Verlag, Berlin.
26Ehret C F, Haller G. Origin development and maturation of organelle system of thecell surface in Paramecium. Ultrasture.1963,6:1~42
27Raikov, I.B., Kovaleva, V.G. Fine structure of circumoral ciliature and pharyngealapparatus of the marine ciliate Helicoprodon gigas (Gymnostomata). Protistologica,1980,16:177~198
28Kruger. Die Ttichocysten der Ciliaten im Dunkelfedbild. Zoological,1936,34:1~83
29Wessenberg, H., Antipa,G. Capture and ingestion of Paramecium by Disiniumnasustum. Protozool.1970,17:240~270
30Frankel,J. Cell biology of Tetrahymena thermuphila. San Diego, London.1999:27~108
31周瑶.腹毛目纤毛虫新尾柱虫微管胞器和射出胞器的研究.华东师范大学博士论文.2010:56~104
32Kalatenbach A. Nahbrungsauswshl and Nahrungsaufnahme holotricher Zilaten derDonaulitioralzone and im Myriophyllum-Aufwuchs des DonausltwassersGansechaufel. Wass.Abwass. Wien.1962:157-185
33Kahl A.Urtier oder Protozoa. Tierweld Dtl.1931,21:181~398
34Poter E D. The buccal organelles in Paramecium awrelia during fission andconjugation with special reference to the kinetosome. Protozool.1960,7:211~217
35史新柏.草履虫口器的形态和形态发生的意义.动物学报.1980,26:289~300
36陈晶，邱子健，王瑞清，秦宇虹，2005运用激光扫描共聚焦显微技术研究大草履虫口胞器与消化胞器动物学报51（4）：718~722
37宋微波.尖前口虫的口器发生研究(纤毛门，膜口目).水生生物学报.1995,19(3):257~262
38邹士法,张作人.银白前口虫Frontonia leucas Ehrenberg切割口器再生研究.华东师范大学学报(自然科学版).1990,3:89~93
39Long H.,Song W., Gong J., Hu X.,Ma H.,Zhu M., Wang M. Frontonia lynni n. sp., anew marine ciliate(Protozoa, Ciliophpra,Hymenostomatida)from Qingdao, China.Zootaxa.2005,1003:57~64
40Foissner, W., Leipe, D. Morphology and ecology of Siroloxophyllum utriculariae(penard,1922) n.g., n.comb.(Ciliophora, Pleurostormatida) and an improvesclassification of plurostomatid ciliates. Eukaryot. Microbiol.1995,42:476~490
41Berger, H. Monograph of the oxytrichidae (Ciliophora, Hypotrophy).1999, KluwerAcademic Publishers Netherlands
42Estbve, J.C. Cytochimie ultrastructurale du trichocyste in situ chez Parameciumcaudatum. Protistologica.1974,10:371~378
43Allman, G.J. On the occurrence among Infusoria of peculiar organs resemblingthread cell. Q.J.Microse. Sci.1855,3:177~179