光裸星虫体腔液中卵子发生的超微结构
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摘要
为探究光裸星虫卵子发生的细胞学特点,利用电镜技术观察了光裸星虫体腔液中卵子发生过程的超微结构变化。结果表明:(1)光裸星虫体腔液中存在游离的卵原细胞和卵母细胞,卵母细胞的发育分为卵黄形成前期、卵黄形成初期、卵黄旺盛合成期和成熟期4个阶段。(2)卵原细胞的细胞器少,核质比大。卵黄形成前期,卵母细胞的细胞器有所增加,大量核仁外排物进入胞质中;出现卵黄膜和胶质膜,卵黄膜遍布微孔。卵黄形成初期卵母细胞核有较多突起,细胞器大量增加,出现分散分布的卵黄粒。卵黄旺盛合成期卵母细胞迅速增大,卵黄大量积累。成熟期卵母细胞核膜突起回缩,胶质膜易于脱落。(3)卵黄分为2种。Ⅰ型卵黄电子密度高,不发生融合,为内源性卵黄;II型卵黄中等电子密度,可融合为无定形卵黄块,为外源性卵黄。(4)成熟卵母细胞卵黄膜为三层,外层为颗粒层,表面具有粒状突;中层初始为均质结构,不断增厚并纤维化;内层致密,厚度不均。胶质膜电子密度极低。(5)卵子外有滤泡细胞,其核质比很大,细胞器少,在卵母细胞成熟期发生凋亡。文章还探讨了胶质膜、卵黄膜、滤泡细胞的功能,以及细胞器在卵黄形成中的作用。研究结果积累了光裸星虫卵子发生细胞学资料,为光裸星虫生殖调控与人工繁育研究提供理论参考。
To understand the cytology underlying the oogenesis of Sipunculus mudus,scanning electron microscope and transmission electron microscope were applied to observe the morphology of oocytes in the coelomic fluid of S.nudus.The results show that the oogoniums and oocytes were free-floating in the coelomic fluid,and the development of oocyte could be divided into four stages,i.e.,pre-vitellogenic stage,primary vitellogenic stage,active vitellogenic stage,and maturation stage.Fewer organelles were found in the oogonium having large nucleo-cytoplasmic ratios.During the previtellogenic stage,more organelles appeared in cytoplasm and plenty of nucleolar exudate was released into the cytoplasm of the oogonium.At the same time,vitelline membrane and jelly coat appeared and micropore was distributed widely in the vitelline membrane.In the primary vitellogenic stage,many enations formed on surface of oocyte nucleus,the number of organelles increased,and the yolk granules distributed scatteredly in the cytoplasm of oocyte.In the active vitellogenic stage,sizes of oocyte cells increased quickly and substantial accumulation of yolk took place.During the maturation stage,enations on surface of the oocyte nucleus disappeared and jelly coat fell off easily.Two types of yolk were found during the oogenesis.Type 1 was endogenous yolk in large electron density while Type 2 was exogenous yolk in medium electron density and undergone granules fusion to form a larger yolk mass.The mature oocyte vitelline membrane could be divided into three layers.The outer layer was with granular surface and no more thickening once the layer was formed;the middle layer was homogeneous in structure at beginning and then was thickened with fibrosis post oogenesis;the inner layer was compact in texture with uneven thickness.The jelly coat was very low in electron density.Oogonia and oocytes were with follicular cells having large nucleo-cytoplasmic ratio and few organelles.The follicular cells underwent apoptosis during the maturation stage.Furthermore,the function of the jelly coat,vitelline membrane,follicular cell and organelles during the formation of yolk were discussed.The results may enrich the cytology content of the oogenesis,and provide a reference to the procreation regulation and artificial breeding of S.nudus.
To understand the cytology underlying the oogenesis of Sipunculus mudus,scanning electron microscope and transmission electron microscope were applied to observe the morphology of oocytes in the coelomic fluid of S.nudus.The results show that the oogoniums and oocytes were free-floating in the coelomic fluid,and the development of oocyte could be divided into four stages,i.e.,pre-vitellogenic stage,primary vitellogenic stage,active vitellogenic stage,and maturation stage.Fewer organelles were found in the oogonium having large nucleo-cytoplasmic ratios.During the previtellogenic stage,more organelles appeared in cytoplasm and plenty of nucleolar exudate was released into the cytoplasm of the oogonium.At the same time,vitelline membrane and jelly coat appeared and micropore was distributed widely in the vitelline membrane.In the primary vitellogenic stage,many enations formed on surface of oocyte nucleus,the number of organelles increased,and the yolk granules distributed scatteredly in the cytoplasm of oocyte.In the active vitellogenic stage,sizes of oocyte cells increased quickly and substantial accumulation of yolk took place.During the maturation stage,enations on surface of the oocyte nucleus disappeared and jelly coat fell off easily.Two types of yolk were found during the oogenesis.Type 1 was endogenous yolk in large electron density while Type 2 was exogenous yolk in medium electron density and undergone granules fusion to form a larger yolk mass.The mature oocyte vitelline membrane could be divided into three layers.The outer layer was with granular surface and no more thickening once the layer was formed;the middle layer was homogeneous in structure at beginning and then was thickened with fibrosis post oogenesis;the inner layer was compact in texture with uneven thickness.The jelly coat was very low in electron density.Oogonia and oocytes were with follicular cells having large nucleo-cytoplasmic ratio and few organelles.The follicular cells underwent apoptosis during the maturation stage.Furthermore,the function of the jelly coat,vitelline membrane,follicular cell and organelles during the formation of yolk were discussed.The results may enrich the cytology content of the oogenesis,and provide a reference to the procreation regulation and artificial breeding of S.nudus.
引文
王庆恒,杜宇宏,林斯里等,2008.裸体方格星虫染色体组型分析.动物学杂志,43(5):95-98
王庆恒,杜晓东,李康,2006.光裸星虫遗传多样性的RAPD分析.海洋水产研究,27(3):57-61
王庆恒,杜晓东,黄红艳等,2005.湛江地区光裸星虫的生殖细胞发育和生殖周期.湛江海洋大学学报(自然科学版),25(1):5-9
兰国宝,阎冰,2002.方格星虫繁殖生物学研究.水产学报,26(6):503-509
兰国宝,阎冰,廖思明,2003.方格星虫胚胎与幼体发育的研究.热带海洋学报,22(6):70-75
成永旭,李少菁,王桂忠等,2002.锯缘青蟹卵黄发生期卵母细胞和卵泡细胞之间的结构变化.动物学报,48(1):80-92
成永旭,堵南山,赖伟,1999.中华绒螯蟹成熟卵巢的脂类及脂肪酸组成.中国水产科学,6(1):79-82
朱冬发,王桂忠,李少菁,2006.东方扁虾卵子发生的超微结构.水生生物学报,30(4):439-445
杨筱珍,王金峰,杨丽娜等,2009.日本新糠虾(Neomysis japonica卵巢发育与卵子发生.海洋与湖沼,40(3):338-346
吴明灿,张立,潘英等,2015.糙海参卵子发生及卵黄发生的超微结构.热带海洋学报,34(3):68-74
邹杰,彭慧婧,蒋艳等,2010.方格星虫亲体培育试验.渔业现代化,37(3):30-33
应雪萍,2002.文蛤卵母细胞卵黄发生的超微结构.中国水产科学,9(2):125-128
张琴,童潼,童万平等,2012.饲料蛋白水平对方格星虫稚虫日增重和消化酶活性的影响.渔业现代化,39(2):41-47
张永泉,尹家胜,杜佳等,2013.哲罗鱼(Hucho taimen)卵巢滤泡细胞发生的超微结构研究.海洋与湖沼,44(1):262-266
陈振国,班廷辉,2015.方格星虫捕捉机:中国,CN201420615340.0
竺俊全,王伟,丁理法,2012.可口革囊星虫(Phascolosoma esculenta)卵黄合成期卵母细胞发育及卵黄发生与卵膜形成的超微结构.海洋与湖沼,43(4):870-876
金春华,竺俊全,许式见等,2011.可口革囊星虫(Phascolosoma esculenta)胚胎及幼虫发育研究.海洋与湖沼,42(1):94-100
姜永华,颜素芬,宋振荣,2005.凡纳滨对虾卵母细胞卵黄发生的超微结构.动物学报,51(1):133-141
洪水根,1992.长毛对虾卵子发生的研究II.卵黄发生.海洋与湖沼,23(3):280-284
顾晓英,竺俊全,许式见等,2009.可口革囊星虫(Phascolosoma esculenta)卵子发生的组织学研究.海洋与湖沼,40(3):283-288
顾海龙,林志华,沈伟良等,2013.泥蚶初级卵母细胞发育及卵黄发生的超微结构研究.海洋科学,37(1):49-53
郭学武,李复雪,1993.光裸星虫生殖周期的研究.热带海洋,12(2):69-76
堵南山,赖伟,陈鹏程等,1999.中华绒螯蟹卵黄形成的研究.动物学报,45(1):88-92
蒋艳,蔡德建,邹杰等,2010.方格星虫苗种池塘中间培育试验研究.广西科学,17(2):175-177
曾志南,李复雪,1991.青蛤卵母细胞发育过程和卵黄发生的超微结构研究.海洋学报,13(5):679-683
操向文,王庆恒,黄若男等,2009.光裸星虫消化系统的解剖学和组织学研究.广东海洋大学学报,29(3):6-10
Du X D,Chen Z,Deng Y W et al,2008.Genetic Diversity and Population Structure of the Peanut Worm(Sipunculus nudus)in Southern China as Inferred from Mitochondrial 16S r RNASequences.The Israeli Journal of Aquaculture-Bamidgeh,60(4):237-242
Du X D,Chen Z,Deng Y W et al,2009.Comparative analysis of genetic diversity and population structure of Sipunculus nudus as revealed by mitochondrial COI sequences.Biochemical Genetics,47(11-12):884-891
Gonse P H,1956.L’ovogenese chez Phascolosoma vulgare:Ⅱ.Recherches biométriques sur les ovocytes.Acta Zoologica,37(3):225-233
Rice M E,1974.Gametogenesis in three species of Sipuncula:Phascolosoma agassizi,Golfingia pugettensis,and Themiste pyroides.La Cellule,79(2-3):259-313
Rice M E,1988.Observations on development and metamorphosis of Siphonosoma cumanense with comparative remarks on Sipunculus nudus(Sipuncula,Sipunculidae).Bulletin of Marine Science,42(1):1-15
Rice M E,1989.Comparative observations of gametes,fertilization,and maturation in sipunculans.In:Ryland J Sed.Reproduction,Genetics and Distributions of Marine Organisms:23rd European Marine Biology Symposium.Swansea:School of Biological Sciences,University of Wales,167-183
Rice M E,Reichardt H F,Piraino J et al,2012.Reproduction,development,growth,and the length of larval life of Phascolosoma turnerae,a wood-dwelling deep-sea sipunculan.Invertebrate Biology,131(3):204-215
Sawada N,1975.An electron microscopic study on the oogenesis of Sipunculan worms.In:Mary E,Todorovic M,eds.Proceedings of the International Symposium on the Biology of the Sipuncula and Echiura.Belgrade:Institute for Biological Research,169-176
王庆恒,杜晓东,李康,2006.光裸星虫遗传多样性的RAPD分析.海洋水产研究,27(3):57-61
王庆恒,杜晓东,黄红艳等,2005.湛江地区光裸星虫的生殖细胞发育和生殖周期.湛江海洋大学学报(自然科学版),25(1):5-9
兰国宝,阎冰,2002.方格星虫繁殖生物学研究.水产学报,26(6):503-509
兰国宝,阎冰,廖思明,2003.方格星虫胚胎与幼体发育的研究.热带海洋学报,22(6):70-75
成永旭,李少菁,王桂忠等,2002.锯缘青蟹卵黄发生期卵母细胞和卵泡细胞之间的结构变化.动物学报,48(1):80-92
成永旭,堵南山,赖伟,1999.中华绒螯蟹成熟卵巢的脂类及脂肪酸组成.中国水产科学,6(1):79-82
朱冬发,王桂忠,李少菁,2006.东方扁虾卵子发生的超微结构.水生生物学报,30(4):439-445
杨筱珍,王金峰,杨丽娜等,2009.日本新糠虾(Neomysis japonica卵巢发育与卵子发生.海洋与湖沼,40(3):338-346
吴明灿,张立,潘英等,2015.糙海参卵子发生及卵黄发生的超微结构.热带海洋学报,34(3):68-74
邹杰,彭慧婧,蒋艳等,2010.方格星虫亲体培育试验.渔业现代化,37(3):30-33
应雪萍,2002.文蛤卵母细胞卵黄发生的超微结构.中国水产科学,9(2):125-128
张琴,童潼,童万平等,2012.饲料蛋白水平对方格星虫稚虫日增重和消化酶活性的影响.渔业现代化,39(2):41-47
张永泉,尹家胜,杜佳等,2013.哲罗鱼(Hucho taimen)卵巢滤泡细胞发生的超微结构研究.海洋与湖沼,44(1):262-266
陈振国,班廷辉,2015.方格星虫捕捉机:中国,CN201420615340.0
竺俊全,王伟,丁理法,2012.可口革囊星虫(Phascolosoma esculenta)卵黄合成期卵母细胞发育及卵黄发生与卵膜形成的超微结构.海洋与湖沼,43(4):870-876
金春华,竺俊全,许式见等,2011.可口革囊星虫(Phascolosoma esculenta)胚胎及幼虫发育研究.海洋与湖沼,42(1):94-100
姜永华,颜素芬,宋振荣,2005.凡纳滨对虾卵母细胞卵黄发生的超微结构.动物学报,51(1):133-141
洪水根,1992.长毛对虾卵子发生的研究II.卵黄发生.海洋与湖沼,23(3):280-284
顾晓英,竺俊全,许式见等,2009.可口革囊星虫(Phascolosoma esculenta)卵子发生的组织学研究.海洋与湖沼,40(3):283-288
顾海龙,林志华,沈伟良等,2013.泥蚶初级卵母细胞发育及卵黄发生的超微结构研究.海洋科学,37(1):49-53
郭学武,李复雪,1993.光裸星虫生殖周期的研究.热带海洋,12(2):69-76
堵南山,赖伟,陈鹏程等,1999.中华绒螯蟹卵黄形成的研究.动物学报,45(1):88-92
蒋艳,蔡德建,邹杰等,2010.方格星虫苗种池塘中间培育试验研究.广西科学,17(2):175-177
曾志南,李复雪,1991.青蛤卵母细胞发育过程和卵黄发生的超微结构研究.海洋学报,13(5):679-683
操向文,王庆恒,黄若男等,2009.光裸星虫消化系统的解剖学和组织学研究.广东海洋大学学报,29(3):6-10
Du X D,Chen Z,Deng Y W et al,2008.Genetic Diversity and Population Structure of the Peanut Worm(Sipunculus nudus)in Southern China as Inferred from Mitochondrial 16S r RNASequences.The Israeli Journal of Aquaculture-Bamidgeh,60(4):237-242
Du X D,Chen Z,Deng Y W et al,2009.Comparative analysis of genetic diversity and population structure of Sipunculus nudus as revealed by mitochondrial COI sequences.Biochemical Genetics,47(11-12):884-891
Gonse P H,1956.L’ovogenese chez Phascolosoma vulgare:Ⅱ.Recherches biométriques sur les ovocytes.Acta Zoologica,37(3):225-233
Rice M E,1974.Gametogenesis in three species of Sipuncula:Phascolosoma agassizi,Golfingia pugettensis,and Themiste pyroides.La Cellule,79(2-3):259-313
Rice M E,1988.Observations on development and metamorphosis of Siphonosoma cumanense with comparative remarks on Sipunculus nudus(Sipuncula,Sipunculidae).Bulletin of Marine Science,42(1):1-15
Rice M E,1989.Comparative observations of gametes,fertilization,and maturation in sipunculans.In:Ryland J Sed.Reproduction,Genetics and Distributions of Marine Organisms:23rd European Marine Biology Symposium.Swansea:School of Biological Sciences,University of Wales,167-183
Rice M E,Reichardt H F,Piraino J et al,2012.Reproduction,development,growth,and the length of larval life of Phascolosoma turnerae,a wood-dwelling deep-sea sipunculan.Invertebrate Biology,131(3):204-215
Sawada N,1975.An electron microscopic study on the oogenesis of Sipunculan worms.In:Mary E,Todorovic M,eds.Proceedings of the International Symposium on the Biology of the Sipuncula and Echiura.Belgrade:Institute for Biological Research,169-176
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