广西北部湾海域织锦巴非蛤卵巢发育、卵子及卵黄发生的研究

doi:10.11978/2023070

摘要/Abstract

摘要：

采用解剖观察、卵巢组织切片和透射电镜技术对广西北部湾海域织锦巴非蛤(Paphia textile)的卵巢发育、卵子发育及卵黄粒合成过程和特点进行探讨。结果表明, 织锦巴非蛤在1周年内具有1个繁殖周期, 根据卵巢发育时期划分为5个时期, 分别为增殖期(5月—7月)、生长期(7月—9月)、成熟期(9月—11月)、排放期(11月—翌年1月)、休止期(翌年1月— 4月)。根据织锦巴非蛤卵子发育和卵黄合成的相关特点, 可将卵子发育划分4个主要阶段, 第一阶段为卵原细胞, 第二阶段为卵黄合成前期卵母细胞, 第三阶段为卵黄合成期卵母细胞, 第四阶段为卵黄合成后期卵母细胞, 其中卵黄合成期是卵黄粒生成的重要时期, 在此期线粒体、内质网、高尔基体、溶酶体等细胞器通过不同的方式合成卵黄粒。此外, 海区水温与织锦巴非蛤卵巢发育密切相关, 是影响织锦巴非蛤卵巢发育的重要因素。广西北部湾海域织锦巴非蛤为雌雄异体, 未发现雌雄同体现象。

关键词: 织锦巴非蛤, 卵巢发育, 卵子发生, 超微结构, 组织切片观察

Abstract:

In this study, ovarian development, reproductive cycle, egg development and its ultrastructure of Paphia textile in Beibu Gulf of Guangxi were investigated using anatomical observation, ovarian tissue sectioning and transmission electron microscopy techniques. The results showed that P. textile has one reproductive cycle in one year, and the ovarian development of P. textile can be divided into five stages: proliferating stage (May to July), growing stage (July to September), maturing stage (September to November), spawning stage (November to January of the next year), and resting stage (January to April of the next year). According to the relevant characteristics of the egg development and vitellogenesis of P. textile, it can be divided into four main stages: the first stage is the oocytes stage, the second stage is the phase of previtellogenesis oocytes stage, the third stage is the phase of vitellogenesis oocytes stage, and the fourth stage is the phase of postvitellogenic oocytes stages. The phase of vitellogenesis is an important stage of vitellogenesis, during which mitochondria, endoplasmic reticulum, golgiosome, lysosomes and other organelles form yolk granule in different ways. The water temperature in the sea area was closely related to the ovarian development of P. textile and was an important factor affecting the ovarian development of P. textile. P. textile is hermaphroditic, and no hermaphroditism has been found. The results of the study provide providing basic data for reproductive biology, artificial breeding, breeding and germplasm resources protection of P. textile.

Key words: Paphia textile, ovarian development, oogenesis, ultrastructure, histological section observation

徐炳杰, 刘一鸣, 连昌朋, 吴韬, 潘英. 广西北部湾海域织锦巴非蛤卵巢发育、卵子及卵黄发生的研究[J]. 热带海洋学报, 2024, 43(2): 48-58.

XU Bingjie, LIU Yiming, LIAN Changpeng, WU Tao, PAN Ying. Ovarian development, oocyte and yolk production of Paphia textile in Beibu Gulf, Guangxi[J]. Journal of Tropical Oceanography, 2024, 43(2): 48-58.

图/表 6

图 1

表1

图2

表2

织锦巴非蛤各期卵母细胞特征"

时期	细胞特点	细胞器特点	卵黄特点
卵原细胞期	卵原细胞紧靠滤泡壁, 由泡壁上的生殖上皮细胞增殖而来, 形状呈椭圆形或圆形。细胞卵径约为0.94~8.96μm, 细胞间界限不清晰, 但可观察到细胞核。	细胞核内核质聚集, 核仁明显, 细胞在此期整体染色较深。电镜下可观察到卵原细胞的胞质呈颗粒状, 胞质存在较多的囊泡, 线粒体等细胞器较少, 核仁不位于细胞核的中央。	此期未开始卵黄合成。
卵黄合成前期	在滤泡内壁周围。细胞卵径约为9.51~15.43μm, 前期与滤泡壁相贴合, 后期逐渐朝向滤泡中心, 可见卵质有一部分为变窄狭长, 变为滤泡壁相连的卵柄。形状为椭圆形或梨形。细胞核的染色质有聚集成团块的趋势, 同时卵母细胞胞质内存在许多大小、形状不一的囊泡。	细胞核中核质膨胀成泡状, 逐渐透亮, 形成一个生发泡, 卵母细胞胞质中可观察到线粒体等细胞器, 高尔基体复合体开始形成。粗面内质网形成少量囊泡。脂滴的数量开始增多。	此时有少量卵黄粒在合成, 卵黄粒无膜结构, 体积较小, 呈球状。同时胞质周围有大量的糖原颗粒分布。
卵黄合成期	在滤泡周围和靠中央位置, 细胞卵径约15.06~57.20μm, 细胞形态从扁平或不规则逐渐变为梨形或者葡萄状, 卵柄变窄, 与滤泡壁脱离或仅剩一点柄状连接。细胞核偏向一边, 细胞内卵黄粒增多。卵母细胞表面微绒毛发达。卵母细胞胞质内有大量内质网、线粒体等细胞器。	卵母细胞中细胞器数量多。内质网发达, 分为层状和环状结构, 环状结构还会包围脂滴进行卵黄粒的合成。高尔基体复合体在边缘形成囊泡, 包囊包裹线粒体等进行卵黄粒沉淀。	此时卵黄大量合成, 卵黄的来源多种, 形状与大小不一, 不同来源的卵黄粒沉积程度有差别。
卵黄合成后期	成熟卵子形状为球形, 卵子间相互挤压, 分布在滤泡中央位置。卵径比大生长期的卵母细胞稍大一些, 细胞卵径约为 48.78~62.54μm, 呈饱满状, 核膜清晰, 核仁靠在细胞核的一边、卵黄粒多, 大量分布在核质中。胞质内充满卵黄粒, 卵黄饱满, 成熟卵细胞相互挤压, 细胞核清晰、有少部分可观察到存在2个核仁。	卵母细胞的细胞器数量大幅减少。内质网和高尔基体复合体等细胞器特征已经不明显, 形成包囊。Ⅱ型线粒体内部脊消失, 出现空泡化, 合成卵黄粒。	大量卵黄分布在成熟卵子内, 多为球形, 大部分发育成熟。卵黄粒在卵母细胞内呈现饱满状, 数量多、体积大。

表2

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图4

参考文献 40

[1]	白胡木吉力图, 马汝河, 高悦勉, 等, 2008. 大连海区青蛤的性腺发育和生殖周期[J]. 大连水产学院学报, 23(3): 196-199.
	BAIHUMU JILITU, MA RUHE, GAO YUEMIAN, et al, 2008. The gonadal development and reproductive cycle of clam Cyclina sinensis in coastal Dalian[J]. Journal of Dalian Fisheries University, 23(3): 196-199 (in Chinese with English abstract).
[2]	曹伏君, 刘永, 张春芳, 等, 2012. 施氏獭蛤(Lutraria sieboldii)性腺发育和生殖周期的研究[J]. 海洋与湖沼, 43(5): 976-982.
	CAO FUJUN, LIU YONG, ZHANG CHUNFANG, et al, 2012. Studies on the sex gonad development and reproductive cycle of Lutraria sieboldii[J]. Oceanologia et Limnologia Sinica, 43(5): 976-982 (in Chinese with English abstract).
[3]	邓正华, 翟子钦, 魏海军, 等, 2022. 织锦巴非蛤幼虫对不同种类单胞藻的摄食和消化效果[J]. 南方农业学报, 53(5): 1448-1556.
	DENG ZHENGHUA, ZHAI ZIQIN, WEI HAIJUN, et al, 2022. Effects of Paphia textile larvae on ingestion and digestion of different microalgae species[J]. Journal of Southern Agriculture, 53(5): 1448-1556 (in Chinese with English abstract).
[4]	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[J]. 海洋科学, 37(1): 49-53.
	GU HAILONG, LIN ZHIHUA, SHEN WEILIANG, et al, 2013. Ultrastructural studies on the development of primary oocytes and vitellogenesis of Tegillarca granosa[J]. Marine Sciences, 37(1): 49-53 (in Chinese with English abstract).
[5]	柯巧珍, 李琪, 闫红伟, 等, 2012. 山东北部沿海四角蛤蜊性腺发育年周期研究[J]. 中国海洋大学学报(自然科学版), 42(11): 28-34.
	KE QIAOZHEN, LI QI, YAN HONGWEI, et al, 2012. Annual observation on the reproduction of surf clam (Mactra veneriformis) inhabiting the north coast of Shandong peninsula[J]. Periodical of Ocean University of China, 42(11): 28-34 (in Chinese with English abstract).
[6]	赖胜琪, 尹聪, 邱炬维, 等, 2022. 北部湾沿海织锦巴非蛤不同地理群体形态差异研究[J]. 广东农业科学, 49(7): 105-112.
	LAI SHENGQI, YIN CONG, QIU JUWEI, et al, 2022. Morphological variation analysis of different Paphia textile geographical populations along the coast of Beibu Gulf[J]. Guangdong Agricultural Sciences, 49(7): 105-112 (in Chinese with English abstract).
[7]	李庆昌, 刘坦, 陈小明, 等, 2016. 织锦巴非蛤斧足颜色与总类胡萝卜素含量相关分析[J]. 海洋科学, 40(10): 120-125.
	LI QINGCHANG, LIU TAN, CHEN XIAOMING, et al, 2016. Correlation analysis of the color and total carotenoid content in Paphia textile foot tissue[J]. Marine Sciences, 40(10): 120-125 (in Chinese with English abstract).
[8]	李霞, 梁峻, 闫喜武, 等, 2007. 人工控温促熟条件下不同海域菲律宾蛤仔的性腺发育[J]. 水产学报, 31(6): 853-859.
	LI XIA, LIANG JUN, YAN XIWU, et al, 2007. The relationship between water temperature and periodic variation of histology with gonad development of Manila clam Ruditapes philippinarum in different regions[J]. Journal of Fisheries of China, 31(6): 853-859 (in Chinese with English abstract).
[9]	栗志民, 刘志刚, 韩伟贤, 2011. 织锦巴非蛤稚贝盐度适应性研究[J]. 海洋科学, 35(10): 96-102.
	LI ZHIMIN, LIU ZHIGANG, HAN WEIXIAN, et al, 2011. Salinity adaptability of Paphia textile spats at different salinity levels[J]. Marine Sciences, 35(10): 96-102 (in Chinese with English abstract).
[10]	连昌朋, 吴韬, 王超奇, 等, 2022. 广西北海营盘海域钝缀锦蛤(Tapes conspersus)卵巢发育、卵子和卵黄发生的研究[J]. 热带海洋学报, 41(5): 170-179. doi: 10.11978/2021185
	LIAN CHANGPENG, WU TAO, WANG CHAOQI, et al, 2022. Ovarian development, histology of oogenesis and yolk formation of Tapes conspersus from the Beihai Yingpan, Guangxi[J]. Journal of Tropical Oceanography, 41(5): 170-179 (in Chinese with English abstract).
[11]	廖承义, 徐应馥, 王远隆, 等, 1983. 栉孔扇贝的生殖周期[J]. 水产学报, 7(1): 1-13.
	LIAO CHENGYI, XU YINGFU, WANG YUANLONG, et al, 1983. Reproductive cycle of the scallop Chlamys farreri (Jones et Preston) at Qingdao[J]. Journal of Fisheries of China, 7(1): 1-13 (in Chinese with English abstract).
[12]	刘海娟, 陈瑞芳, 曾梦清, 等, 2022. 织锦巴非蛤人工养殖技术研究[J]. 科学养鱼, 38(11): 61-63.
	LIU HAIJUAN, CHEN RUIFANG, ZENG MENGQING, et al, 2022. Study on the artificial culture technology of Paphia textile[J]. Scientific Fish Farming, 38(11): 61-63 (in Chinese with English abstract).
[13]	刘相全, 方建光, 包振民, 等, 2007. 中国蛤蜊繁殖生物学的初步研究[J]. 中国海洋大学学报(自然科学版), 37(1): 89-92.
	LIU XIANGQUAN, FANG JIANGUANG, BAO ZHENMIN, et al, 2007. A preliminary study on the reproductive biology of Mactra Chinensis Philippi[J]. Periodical of Ocean University of China, 37(1): 89-92 (in Chinese with English abstract).
[14]	吕敏, 李琪, 2022. 山东田横岛海域长牡蛎性腺发育及生化成分周年变化[J]. 中国海洋大学学报(自然科学版), 52(2): 33-40.
	LYU MIN, LI QI, 2022. Seasonal variations of gonadal development and biochemical components of Crassostrea gigas in Tianheng island sea area, Shandong[J]. Periodical of Ocean University of China, 52(2): 33-40 (in Chinese with English abstract).
[15]	宁军号, 常亚青, 宋坚, 等, 2015. 偏顶蛤的性腺发育和生殖周期[J]. 中国水产科学, 22(3): 469-477.
	NING JUNHAO, CHNAG YAQING, SONG JIAN, et al, 2015. Gonadal development and the reproductive cycle of Modiolus modiolus[J]. Journal of Fishery Sciences of China, 22(3): 469-477 (in Chinese with English abstract).
[16]	上官步敏, 刘正琮, 1995. 缢蛏卵母细胞卵黄发生过程的超微结构研究[J]. 海洋与湖沼, 26(1): 48-52.
	SHANGGUAN BUMIN, LIU ZHENGCONG, 1995. Ultrastructural studies on the formation of yolk globule of the oocyte in Sinonovacula constricta[J]. Oceanologia et Limnologia Sinica, 26(1): 48-52 (in Chinese with English abstract).
[17]	闻海波, 孙光兴, 丁图强, 等, 2020. 淮河橄榄蛏蚌繁殖类型与性腺发育观察[J]. 中国水产科学, 27(10): 1156-1160.
	WEN HAIBO, SUN GUANGXING, DING TUQIANG, et al, 2020. Reproductive type and gonad development in the threatened freshwater mussel Solenaia oleivora (Heude) from the Huaihe River[J]. Journal of Fishery Sciences of China, 27(10): 1156-1160 (in Chinese with English abstract).
[18]	吴加莹, 戴明姝, 刘志刚, 等, 2023. 温度对织锦巴非蛤稚贝生存和生长的影响[J]. 南方水产科学, 19(2): 62-69.
	WU JIAYING, DAI MINGSHU, LIU ZHIGANG, et al, 2023. Effects of temperature on survival and growth of Paphia textile juveniles[J]. South China Fisheries Science, 19(2): 62-69 (in Chinese with English abstract).
[19]	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[J]. 热带海洋学报, 34(3): 68-74. doi: 10.11978/j.issn.1009-5470.2015.03.009
	WU MINGCAN, ZHANG LI, PAN YING, et al, 2015. Ultrastructure of oogenesis and vitellogenesis of sea cucumber Holothuria scabra[J]. Journal of Tropical Oceanography, 34(3): 68-74 (in Chinese with English abstract).
[20]	巫旗生, 文宇, 曾志南, 等, 2017. 钝缀锦蛤繁殖周期和胚胎发育[J]. 中国水产科学, 24(3): 488-496.
	WU QISHENG, WEN YU, ZENG ZHINAN, et al, 2017. The reproductive cycle and embryonic development of the bivalve mollusk Tapes conspersus[J]. Journal of Fishery Sciences of China, 24(3): 488-496 (in Chinese with English abstract). doi: 10.3724/SP.J.1118.2017.16280
[21]	徐凤山, 张素萍, 2008. 中国海产双壳类图志[M]. 北京: 科学出版社:244- 245.
	XU FENGSHAN, ZHANG SUPING, 2008. An illustrated bivalvia mollusca fauna of China Seas[M]. Beijing: Science Press: 244-245 (in Chinese with English abstract).
[22]	杨林, 李琪, 闫红伟, 等, 2010. 山东沿海中国蛤蜊的繁殖生物学特性[J]. 中国水产科学, 17(3): 514-520.
	YANG LIN, LI QI, YAN HONGWEI, et al, 2010. Reproductive characteristics of clam (Mactra chinensis) along coast of Shandong Province[J]. Journal of Fishery Sciences of China, 17(3): 514-520 (in Chinese with English abstract).
[23]	应雪萍, 杨万喜, 2001. 泥螺卵黄发生过程中线粒体的变化[J]. 动物学研究, 22(5): 379-382.
	YING XUEPING, YANG WANXI, 2001. Mitochondrial changes during vitellogenesis in oocytes of Bullacta exarata[J]. Zoological Research, 22(5): 379-382 (in Chinese with English abstract).
[24]	曾庆民, 赵虹博, 邓素贞, 等, 2022. 织锦巴非蛤不同颜色斧足的转录组[J]. 水产学报, 46(9): 1562-1571.
	ZENG QINGMIN, ZHAO HONGBO, DENG SUZHEN, et al, 2022. Transcriptome analysis of Paphia textile with different foot colors[J]. Journal of Fisheries of China, 46(9): 1562-1571 (in Chinese with English abstract).
[25]	翟子钦, 喻达辉, 白丽蓉, 2022. 织锦巴非蛤形态性状对体质量的影响[J]. 广东农业科学, 49(7): 113-119.
	ZHAI ZIQIN, YU DAHUI, BAI LIRONG, 2022. Effects of morphological traits on body mass of Paphia textile[J]. Guangdong Agricultural Sciences, 49(7): 113-119 (in Chinese with English abstract).
[26]	赵雯, 车宗豪, 刘括, 等, 2022. 日本海神蛤(Panopea japonica)性腺发育生物学零度及有效积温研究[J]. 海洋科学, 46(10): 78-84.
	ZHAO WEN, CHE ZONGHAO, LIU KUO, et al, 2022. Biological zero degree and effective accumulated temperature of the geoduck clam Panopea japonica[J]. Marine Sciences, 46(10): 78-84 (in Chinese with English abstract).
[27]	朱星海, 孙红振, 杨祖晶, 等, 2019. 风信标扇贝的性腺发育与繁殖周期规律研究[J]. 中国海洋大学学报(自然科学版), 49(2): 52-58.
	ZHU XINGHAI, SUN HONGZHEN, YANG ZUJING, et al, 2019. Studies on gonadal development and the reproductive cycle of Patinopecten caurinus[J]. Periodical of Ocean University of China, 49(2): 52-58 (in Chinese with English abstract).
[28]	庄启谦, 2001. 中国动物志: 软体动物门、双壳纲、帘蛤科[M]. 北京: 科学出版社:53- 54.
	ZHUANG QIQIAN, 2001. Zoology of China: Mollusca, Bivalvia, Veneridae[M]. Beijing: Science Press: 53-54 (in Chinese with English abstract).
[29]	邹杰, 彭慧婧, 杨家林, 2019. 织锦巴非蛤人工种苗培育及浅海养殖试验[J]. 科学养鱼, 35(10): 57-58.
	ZOU JIE, PENG HUIJING, YANG JIALIN, 2019. Experiment on artificial propagation and culture in shallow sea of Paphia textile[J]. Scientific Fish Farming, 35(10): 57-58 (in Chinese with English abstract).
[30]	AEGERTER S, JALABERT B, 2004. Effects of post-ovulatory oocyte ageing and temperature on egg quality and on the occurrence of triploid fry in rainbow trout, Oncorhynchus mykiss[J]. Aquaculture, 231(1-4): 59-71. doi: 10.1016/j.aquaculture.2003.08.019
[31]	AGNESE M, VERDERAME M, MEO E D, 2013. A network system for vitellogenin synthesis in the mussel Mytilus galloprovincialis (L.)[J]. Journal of Cellular Physiology, 228(3): 547-555. doi: 10.1002/jcp.v228.3
[32]	BI JINHONG, LI QI, ZHANG XINJUN, et al, 2016. Seasonal variation of biochemical components in clam (Saxidomus purpuratus Sowerby 1852) in relation to its reproductive cycle and the environmental condition of Sanggou Bay, China[J]. Oceanic and Coastal Sea Research, 15(2): 341-350.
[33]	CHEN JIAN, DENG ZHENHUA, WEI HAIJUN, et al, 2021. Spermatozoa morphology and embryo development of four species of bivalves from Beibu Gulf[J]. Tuikish Journal of Fisheries and Aquatic Sciences, 21(2): 51-61.
[34]	CHEN XIAOMING, LI JIAKAI, XIAO SHIJUN, et al, 2016. De novo assembly and characterization of foot transcriptome and microsatellite marker development for Paphia textile[J]. Gene, 576(1): 537-543. doi: 10.1016/j.gene.2015.11.001
[35]	ERKAN E, 2009. Ultrastructure of ovary and oogenesis in Chamelea gallina (Linné, 1758) (Bivalvia, Veneridae)[J]. Invertebrate Reproduction & Development, 53(4): 201-209.
[36]	LIU WENGUANG, LI QI, GAO FENGXIANG, et al, 2010. Effect of starvation on biochemical composition and gametogenesis in the Pacific oyster Crassostrea gigas[J]. Fisheries Science, 76(5): 737-745. doi: 10.1007/s12562-010-0274-y
[37]	NI JIANBIN, ZENG ZHEN, KONG DEZHENG, et al, 2014. Vitellogenin of Fujian oyster, Crassostrea angulata: synthesized in the ovary and controlled by estradiol-17β[J]. General and Comparative Endocrinology, 202: 35-43. doi: 10.1016/j.ygcen.2014.03.034
[38]	NIE HONGTAO, LU YAO, LIU HAI, et al, 2016. Seasonal variations in biochemical composition of the clam Dosinia corrugate in relation to the reproductive cycle and environmental conditions[J]. Journal of Shellfish Research, 35(2): 369-377. doi: 10.2983/035.035.0211
[39]	QIN YANPING, LI XINGYOU, LI JUN, et al, 2021. Seasonal variations in biochemical composition and nutritional quality of Crassostrea hongkongensis, in relation to the gametogenic cycle[J]. Food Chemistry, 356: 129736. doi: 10.1016/j.foodchem.2021.129736
[40]	YURCHENKO O V, KALACHEV A V, 2019. Morphology of nutrient storage cells in the gonadal area of the Pacific oyster, Crassostrea gigas (Thunberg, 1793)[J]. Tissue and Cell, 56: 7-13. doi: 10.1016/j.tice.2018.11.004

取样日期/(年-月-日)	卵巢发育各期内的个体数					总数/个	盐度/‰	水温/℃
取样日期/(年-月-日)	Ⅰ	Ⅱ	Ⅲ	Ⅳ	Ⅴ	总数/个	盐度/‰	水温/℃
2020-11-29			2	4		6	27.4	22.0
2020-12-21			5	6		11	27.5	17.2
2021-01-08			2	9	1	12	29.4	15.6
2021-02-24				2	8	10	30.9	17.3
2021-03-18				1	8	9	30.0	21.0
2021-04-22					12	12	30.0	27.5
2021-05-20	2	1			4	7	28.0	29.5
2021-06-23	2	4			4	10	28.4	30.5
2021-07-15	5	6	2			13	29.0	29.7
2021-08-18	3	6	1			10	30.0	29.8
2021-09-17	2	3	5	2		12	31.0	29.8
2021-10-20			13	1		14	29.0	25.5