海南澳洲管体星虫卵细胞发育及生殖周期研究

doi:10.11978/2021143

摘要/Abstract

摘要：

澳洲管体星虫(Siphonosoma australe)是海南当地极具特色的星虫资源, 经济价值高。近年来该资源急剧衰竭, 开展星虫资源恢复与保护十分重要。文章对海南澳洲管体星虫卵细胞的显微与超微结构进行研究。结果显示: 1) 卵细胞发育经历4个阶段: 生长前期, 体腔液中大量光滑的凹饼状红细胞出现表面凹凸不平的褶皱(直径小于30μm), 暗示这些红细胞是卵原细胞; 生长后期, 30~40μm的卵细胞表面褶皱消失, 变成光滑的圆厚饼状, 并出现厚度1μm的卵黄膜, 卵黄颗粒开始积累; 成熟前期, 卵细胞球状, 直径60~120μm, 卵黄膜增厚至5~11μm, 卵黄颗粒增多, 细胞核增大, 部分染色质形成高电子密度的团块散布于核内; 成熟后期, 直径120μm, 卵黄膜厚11~12μm, 卵黄颗粒充满整个卵细胞, 膜孔外露于膜表层。2) 成熟期澳洲管体星虫卵黄膜仅分为两层: 有一定厚度的均质内层和多层重叠膜结构组成的外层, 同时外层上覆盖有粒状突。膜孔结构简单, 主要是膜凹陷折叠形成。3) 卵黄颗粒分为Ⅰ型和Ⅱ型, 其发生途径主要由线粒体、内质网、高尔基体和溶酶体等细胞器演变而成, 亦可由卵母细胞吞饮而成。最为常见的是高尔基体包裹形成的Ⅱ型卵黄颗粒。4) 海南文昌海域澳洲管体星虫的繁殖季节为4—8月, 其中5—7月是繁殖高峰期。3月初卵母细胞开始出现, 4—8月成熟卵细胞在体腔中保持一定密度, 9月份卵细胞密度降至极低, 10月份后消失。澳洲管体星虫作为一种新的具有热带特色的水产种质资源, 其卵细胞发育及生殖周期研究将极大推动其繁育及保护的相关技术探索。

关键词: 澳洲管体星虫, 卵细胞发育, 生殖周期, 海南, 显微观察, 超微观察

Abstract:

Siphonosoma australe is an important sipuculan species and a typical local resource in Hainan with medicinal and edible value, it is distributed in tropical and subtropical coastal waters, mainly in surrounding waters of Hainan island. Due to overfishing and habitat destruction, the population size of S. australe rapidly declined in recent years. So the recovery and protection of the sipuculan resource is very important. The structures of oocytes of S. australe were observed by optical microscope and electron microscope. The results are shown as follows: 1) The oogenesis of S. australe could be divided into four stages: In the early growth stage, concave disk-shaped coelomocytes (less than 30 μm in diameter) changed from the smooth cells to lumpy and corrugated cells, which suggested that concave coelomocytes were the oogonia; in the late growth stage, the surface of 30 ~ 40 μm oocytes became smooth and unfolded, and the vitelline membrane of 1 μm in thickness appeared with the accumulation of yolk granules; in the early mature stage, oocytes were 60 ~ 120 μm in diameter and transformed into spherical shape. The yolk granules increased. The nucleus was enlarged. Some of the chromatin formed clumps with high electron density and dispersed in the nucleus; in the late mature stage, oocytes were 120 μm in diameter with the yolk granules filling the whole cells, and the membrane pores were exposed on the membrane surface. 2) The vitelline membrane of mature oocyte can be divided into two layers: the homogeneous inner layer with a thin thickness and the multi-folded outer layer with granular protuberances. The structure of membrane micropore was simple and formed by the invagination of the folded membranes. 3) During oogenesis, a variety of organelles increased and the yolk granules (type Ⅰ and Ⅱ) were combined with organelles, e.g. mitochondria, endoplasmic reticulum, Golgi apparatus, lysosome, evenly transformed by phagocytic vesicle. The type Ⅱ yolk-granules enveloped by Golgiosomes were common. 4) The reproductive season of S. australe in Wenchang, Hainan is from April to August and the reproductive peak is in May and July. Oocytes begin to appear in early March, quantitatively drop to a very low level in September and disappear in October. The mature oocytes keep a certain density in the coelom from April to August. It is speculated that the reproduction period is related to the monthly water-temperature and the season of highest water-temperature is the period of reproductive peak. As a new Hainan fishery resource with tropical characteristics, this research on oocyte development and reproductive cycle of S. australe is expected to greatly promote the development on related technologies for reproduction and protection.

Key words: Siphonosoma australe, Hainan, oogenesis, microstructural observation, ultrastructural observation, reproductive cycle

中图分类号:

S917

乔立君, 姚雪梅, 余巧驰. 海南澳洲管体星虫卵细胞发育及生殖周期研究[J]. 热带海洋学报, 2022, 41(5): 161-169.

QIAO Lijun, YAO Xuemei, YU Qiaochi. Development of oocytes and reproductive cycle of Siphonosoma australe in Hainan[J]. Journal of Tropical Oceanography, 2022, 41(5): 161-169.

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参考文献 22

[1]	陈慧, 林国文, 陈武, 等, 2009. 可口革囊星虫生殖周期的观察[J]. 海洋渔业, 31(2): 139-145.
	CHEN HUI, LIN GUOWEN, CHEN WU, et al, 2009. Observation on the reproductive cycle of Phascolosoma esculenta[J]. Marine Fisheries, 31(2): 139-145. (in Chinese with English abstract)
[2]	陈子安, 杜晓东, 王庆恒, 等, 2007. 3种星虫线粒体16S rRNA、CO I和Cyt b基因片段的序列比较[J]. 广东海洋大学学报, 27(4): 3-10.
	CHEN ZI’AN, DU XIAODONG, WANG QINGHENG, et al, 2007. Sequence comparison and phylogenetic analysis of mtDNA 16SrRNA and CO I and Cyt b gene fragments in three species of Sipuncula[J]. Journal of Guangdong Ocean University, 27(4): 3-10. (in Chinese with English abstract)
[3]	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[J]. 海洋与湖沼, 40(3): 283-288.
	GU XIAOYING, ZHU JUNQUAN, XU SHIJIAN, et al, 2009. Oogenesis of Phascolosoma esculenta: histological study[J]. Oceanologia et Limnologia Sinica, 40(3): 283-288. (in Chinese with English abstract)
[4]	郭学武, 李复雪, 1993. 光裸星虫生殖周期的研究[J]. 热带海洋学报, 12(2): 69-76.
	GUO XUEWU, LI FUXUE, 1993. Studies on the reproductive cycle of Sipunculus nudus linnaeus[J]. Tropic Oceanology, 12(2): 69-76. (in Chinese with English abstract)
[5]	兰国宝, 阎冰, 2002. 方格星虫繁殖生物学研究[J]. 水产学报, 26(6): 503-509.
	LAN GUOBAO, YAN BING, 2002. The reproductive biology of peanut worm, Sipunculus nudus[J]. Journal of Fisheries of China, 26(6): 503-509. (in Chinese with English abstract)
[6]	李凤鲁, 周红, 王玮, 1992. 中国沿海管体星虫属(星虫动物门)的研究[J]. 青岛海洋大学学报, 22(1): 97-102.
	LI FENGLU, ZHOU HONG, WANG WEI, 1992. Studies on the genus Siphonosoma (sipuncula) off the China coasts[J]. Journal of Ocean University of Qingdao, 22(1): 97-102. (in Chinese with English abstract)
[7]	刘懂, 龙玲利, 曾海祥, 等, 2014. 可口革囊星虫体腔中卵细胞大小组成的周年变化[J]. 生物学杂志, 31(4): 29-32.
	LIU DONG, LONG LINGLI, ZENG HAIXIANG, et al, 2014. Annual variation of egg size in the coelom of Phascolosoma esculenta[J]. Journal of Biology, 31(4): 29-32. (in Chinese with English abstract)
[8]	王庆恒, 杜晓东, 黄洪艳, 等, 2005. 湛江地区光裸星虫的生殖细胞发育和生殖周期[J]. 湛江海洋大学学报, 25(1): 5-9.
	WANG QINGHENG, DU XIAODONG, HUANG HONGYAN, et al, 2005. Development of germ cells and reproductive cycle of Sipunculus nudus L in Zhanjiang[J]. Journal of Zhanjiang Ocean University, 25(1): 5-9. (in Chinese with English abstract)
[9]	王庆恒, 张家炜, 郝瑞娟, 等, 2017. 光裸星虫体腔液中卵子发生的超微结构[J]. 海洋与湖沼, 48(1): 57-66.
	WANG QINGHENG, ZHANG JIAWEI, HAO RUIJUAN, et al, 2017. Ultrastructure of oogenesis in coelomic fluid of Sipunculus nudus[J]. Oceanologia et Limnologia Sinica, 48(1): 57-66. (in Chinese with English abstract)
[10]	吴斌, 1999. 光裸方格星虫(Sipunculus nudus L.)生殖细胞及胚胎发育[J]. 广西科学, 6(3): 222-226.
	WU BIN, 1999. Development of genital cells and embryo of Sipunculus nudus L.[J]. Guangxi Sciences, 6(3): 222-226. (in Chinese with English abstract)
[11]	徐海娇, 2012. AFM研究高尔基体和内质网的结构[D]. 长春: 东北师范大学.
	XU HAIJIAO, 2012. Studying the structure of Golgi complex and endoplasmic reticulum by AFM[D]. Changchun: Northeast Normal University. (in Chinese with English abstract)
[12]	周红, 李凤鲁, 王玮, 2007. 中国动物志无脊椎动物第四十六卷星虫动物门螠虫动物门[M]. 北京: 科学出版社.
	ZHOU HONG, LI FENGLU, WANG WEI, 2007. Fauna sinica:invertebrata vol. 46. sipuncula echiura[M]. Beijing: Science Press. (in Chinese)
[13]	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[J]. 海洋与湖沼, 43(4): 870-876.
	ZHU JUNQUAN, WANG WEI, DING LIFA, 2012. Ultrastructural observation on oogenesis, vitellogenesis and vitelline membrane formation during yolk-synthesizing stage in Phascolosoma esculenta[J]. Oceanologia et Limnologia Sinica, 43(4): 870-876. (in Chinese with English abstract)
[14]	AMOR A, 1993. Reproductive cycle of Golfingia margaritacea, a bipolar sipunculan, in subantarctic water[J]. Marine Biology, 117(3): 409-414. doi: 10.1007/BF00349316
[15]	ANDREWS E A, 1889. The reproductive organs of Phascolosoma gouldii[J]. Zoologischer Anzeiger, 12: 140-142.
[16]	CATALAN M A B, YAMAMOT M, 1994. Annual reproductive cycle of two Japanese species of sipunculans: Siphonosoma cumanense (Sipunculidae) and Phascolosoma scolops (Phascolosomatidae)[J]. Pacific Science, 48(2): 145-157.
[17]	GONSE P H, 1956. L’ovogenese chez Phascolosoma vulgare: Ⅱ. Recherches biométriques sur les ovocytes[J]. Acta Zoologica, 37(3): 225-233. doi: 10.1111/j.1463-6395.1956.tb00045.x
[18]	RICE M E, 1974. Gametogenesis in three species of sipuncula: Phascolosoma agassizii, Golfingia pugettensis, and Themiste pyroides[J]. La Cellule, 79(2-3): 259-313.
[19]	RICE M E, 1988. Observations on development and metamorphosis of Siphonosoma cumanense with comparative remarks on Sipunculus nudus (Sipuncula, Sipunculidae)[J]. Bulletin of Marine Science, 42(1): 1-15.
[20]	RICE M E, 1989. Comparative observations of gametes, fertilization, and maturation in sipunculans[C]// Proceedings of the 23^rd European Marine Biology Symposium. Fredensborg: Olsen and Olsen: 167-183.
[21]	SAWADA N, 1975. An electron microscopic study on the oogenesis of Sipunculan worms[C]// Proceedings of International Symposium on the Biology of the Sipuncula and Echiura. Belgrade: Nauč-no Delo Press: 169-176.
[22]	YING XUEPING, DAHMS H U, LIU XINMEI, et al, 2009. Development of germ cells and reproductive biology in the sipunculid Phascolosoma esculenta[J]. Aquaculture Research, 40(3): 305-314. doi: 10.1111/j.1365-2109.2008.02093.x