广西北部湾施氏獭蛤卵巢发育、卵子和卵黄发生的研究

doi:10.11978/2023027

摘要/Abstract

摘要：

为探究广西北部湾海域施氏獭蛤卵巢发育、卵子和卵黄发生, 采用组织切片分析卵巢周年变化, 并利用光学和电子显微镜观察雌性生殖细胞的变化过程。结果表明, 广西北部湾海域施氏獭蛤卵巢发育周期为一年, 每年12月至翌年4月性腺饱满, 为繁殖盛期, 产卵高峰期略滞后于性腺成熟期, 每期5%~15%个体发育滞后于群体。卵子发生过程中, 雌性生殖细胞逐渐脱离滤泡壁, 进入滤泡腔。卵子发育过程中, 卵径由6.9~8.3μm变大至70.0~74.9μm, 细胞质内细胞器数量增加, 出现卵黄粒, 自噬泡吞噬脂滴和线粒体。卵黄发生期间, 核仁经染色后分为两部分, 颜色深浅不一, 核质间出现物质交换, 卵母细胞质膜内出现多泡小体, 在靠近卵周隙处形成微吞饮泡, 分别从卵周隙和滤泡吸收外源性卵黄物质。此外, 本实验还观察到施氏獭蛤存在滤泡混合型和滤泡共存型雌雄同体, 性转换方向表现为雌性向雄性转换。本研究结果为施氏獭蛤的人工繁育提供基础资料。

关键词: 施氏獭蛤, 卵巢发育, 卵子发生, 卵子发育, 超微结构, 雌雄同体

Abstract:

For understanding the ovarian development, oogenesis and vitellogenesis of Lutraria sieboldii in the Beibu Gulf of Guangxi, we used tissue section, optical and electronic microscope observed the annual change process of female germ cells. The results showed that the ovary development of L. sieboldii in the Beibu Gulf of Guangxi was a reproductive cycle every year, and the peak period of reproduction was from December to April of the next year when the gonads were full, 5%~15% individual development lagged behind the population in each stage. During oogenesis, female reproductive cell gradually detached from the follicle walls, and entered the follicular cavity. In the process of ovum development, the oocyte diameter changed from 6.9~8.3 μm to 70.0~74.9 μm, the number of organelles in the cytoplasm increased, and appeared yolk granules, autophagy vesicles engulf lipid droplets and mitochondria. During vitellogenesis, the nucleolus turned into two parts with different staining depth, material exchange occurred between nucleoplasm. Oocytes formed polyvesicular bodies through plasma membrane depression and formed microphagulation vesicles near the perioval space, which absorb exogenous yolk substances from perioval space and follicles, respectively. In addition, we observed that there were follicular mixed type and follicular coexisting type hermaphroditism, and the direction of sexual transformation was from female to male. The results provide basic data for the artificial breeding of L. sieboldii.

Key words: Lutraria sieboldii, ovarian development, oogenesis, ovum development, ultrastructure, hermaphrodite

吴韬, 潘英, 连昌朋, 刘一鸣, 徐炳杰, 王超奇, 杨凌. 广西北部湾施氏獭蛤卵巢发育、卵子和卵黄发生的研究[J]. 热带海洋学报, 2023, 42(6): 137-149.

WU Tao, PAN Ying, LIAN Changpeng, LIU Yiming, XU Bingjie, WANG Chaoqi, YANG Ling. The study on ovarian development, oogenesis and vitellogenesis of Lutraria sieboldii in Beibu Gulf of Guangxi[J]. Journal of Tropical Oceanography, 2023, 42(6): 137-149.

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数值	壳长/mm	壳高/mm	壳宽/mm	体质量/g
平均值	94.12	46.03	32.61	98.89
最大值	116.80	57.74	43.83	199.96
最小值	80.12	39.53	24.72	55.30

发育分期	各期特点
增殖期(Ⅰ)	结缔组织包裹至软体部上缘, 性腺颜色透明, 雌雄外观无区别, 呈细线分布。结缔组织间形状不规则且狭长的滤泡构成性腺, 滤泡末端分支由少变多, 滤泡由小变大形成滤泡腔, 滤泡间隙较大。此时期滤泡腔大多为空腔, 滤泡壁厚, 滤泡内大部分为卵原细胞, 卵原细胞松散分布在滤泡壁上。此期为8月下旬—9月
生长期(Ⅱ)	性腺增殖至软体部上缘, 仍无法从性腺颜色分辨雌雄。滤泡不断增大, 数量不断增多, 结缔组织减少, 滤泡间存在间隙。滤泡壁变薄, 滤泡壁上雌性生殖细胞数量增加, 部分次级卵母细胞生成卵柄。小部分成熟卵母细胞从滤泡壁脱落进入滤泡腔, 大量未发育成熟的卵母细胞占据着滤泡。此期为10月—11月下旬
成熟期(Ⅲ)	性腺增殖至软体部腹缘和足基部, 呈白色, 肉眼可分辨雌雄。此期滤泡最大, 结缔组织退化, 滤泡相互挤压呈不规则形。滤泡壁变为单层, 其上残留少数初级卵母细胞。次级卵母细胞卵柄断裂进入滤泡腔内, 发育为成熟卵子, 滤泡腔内卵子形状不规则。此期为12月—次年3月底
排放期(Ⅳ)	性腺仍饱满, 大部分成熟卵子排出体外, 腔内残留着少量成熟卵子与滤泡壁相连。滤泡壁变厚, 滤泡逐渐缩小, 呈现出大小不一的空腔, 滤泡间结缔组织开始增多。此期为次年4月
休止期(Ⅴ)	软体部消瘦, 性腺变薄, 无法从软体部颜色分辨雌雄。滤泡间隙进一步加大, 结缔组织增长挤压滤泡。滤泡内已无成熟卵子, 滤泡壁上可见少量处于重吸收的卵原细胞, 滤泡壁变厚呈粗线条状。此期为5月下旬—8月中旬

取样日期	盐度/‰	水温/℃	总数	雌雄同体数	卵巢发育分期
取样日期	盐度/‰	水温/℃	总数	雌雄同体数	Ⅰ	Ⅱ	Ⅲ	Ⅳ	Ⅴ
2020-11-26	27.4	22.0	9			8	1
2020-12-21	27.5	17.0	10		2	2	5		1
2021-01-08	29.4	15.0	9			1	8
2021-02-23	30.9	26.3	14			4	10
2021-03-20	30.0	21.0	11			3	8
2021-04-22	30.0	27.5	12					10	2
2021-05-26	28.0	29.5	12						12
2021-06-21	28.4	30.5	7	1			1	1	5
2021-07-14	29.0	29.7	11		2			4	6
2021-08-17	30.0	29.8	10	1	1	1	1	3	4
2021-09-15	31.0	29.8	9	1	4	1		1	3
2021-10-20	29.0	25.5	9	2		5	2

发生分期	各期发生特点
增殖期	出现在Ⅰ—Ⅳ期卵巢。体积小、染色浅的卵原细胞大量增殖, 与滤泡细胞一起平行排列于滤泡壁基膜上
小生长期	出现在Ⅰ—Ⅳ期卵巢。卵原细胞体积增大, 分化形成生长较缓慢的初级卵母细胞, 细胞的大部分仍紧贴滤泡壁基膜, 细胞核内有一核仁常偏位, 细胞周围围绕着一层滤泡细胞。细胞内较为空虚, 染色质数量较少且不清晰, 细胞核染色较浅
大生长期	初级卵母细胞迅速生长, 细胞核内卵黄粒大量形成, 细胞质内细胞器大量形成, 生长期可延续到第一次减数分裂形成次级卵母细胞前, 出现在Ⅰ—Ⅳ期卵巢
成熟期	出现在施氏獭蛤卵巢发育的Ⅲ—Ⅳ期。此期成熟卵子体积达到最大, 成熟卵子呈不规则圆形。初级卵母细胞经过小、大生长期, 胞质内物质大量累积, 发育到次级卵母细胞, 逐渐脱离滤泡壁基膜, 后期卵柄断裂萎缩, 卵母细胞发育成熟进入滤泡腔内, 少数成熟卵子退化被滤泡重吸收

细胞器	细胞器变化情况
线粒体	线粒体为细胞内部活动提供能量, 卵黄合成前期胞质中线粒体数量较少, 内部嵴少且清晰。随着卵黄的合成, 线粒体数量逐渐增多, 内部嵴模糊, 卵黄物质在线粒体基质中的内膜沉积并逐渐充满整个线粒体, 线粒体内膜逐渐溶解消失, 最终形成单层膜结构的卵黄粒(图6e—f)
粗面内质网	卵黄合成期粗面内质网呈环形状和片条状, 片条状粗面内质网两端形成囊泡, 环状粗面内质网内部包裹成椭圆形囊泡, 胞质中高电子密度物质沉积逐渐形成卵黄粒。环状粗面内质网中央盘绕成球状, 胞质中的脂滴、蛋白质等物质沉积在中央, 随着沉淀物质的增加体积不断扩大最终形成卵黄粒(图6h)
高尔基复合体	扁平囊两端发育形成囊泡, 部分囊泡内部被脂类物质沉积形成电子密度较浅的脂滴, 囊泡通过内质网和核糖体的加工, 内部沉积卵黄蛋白形成卵黄粒(图6j)
溶酶体	溶酶体体积较小, 约0.5μm, 电子密度高, 基质均匀, 吞噬胞质内的核糖体、线粒体等细胞器和蛋白质等物质并将其分解沉积形成卵黄(图6e—f)
核糖体	核糖体在施氏獭蛤卵黄发生期间大量游离在细胞质中并被各细胞器及囊泡吸收, 间接参与合成卵黄(图6e、j)