EMBRYONIC AND LARVAL DEVELOPMENTAL CHARACTERISTICS OF THYMALLUS ARCTICUS GRUBEI
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摘要: 研究对人工繁殖的北极茴鱼(Thymallus arcticus grubei)胚胎发育开展系统观察, 记录分析其胚胎及仔鱼发育各时期的形态特征, 旨在为北极茴鱼的人工繁育和种质资源保护提供必要的基础数据。结果显示, 北极茴鱼受精卵呈圆球形, 金黄色, 沉性卵, 未吸水卵径(2.46±0.14) mm, 吸水卵径(3.14±0.18) mm, 卵黄质内有多个油球。油球的数量和空间分布在胚胎发育过程中发生了规律性变化。在孵化水温(11.06±0.72)℃, 溶氧8.3—9.8 mg/L条件下, 历时301h完成整个胚胎发育过程,所需积温为3384.84h·℃, 经历合子期、卵裂期、囊胚期、原肠胚期、神经胚期、器官形成期和孵化出膜7个阶段, 共26个时期。北极茴鱼仔鱼尾鳍、胸鳍的分化和眼色素沉积在受精卵胚胎发育后期就已经完成, 背鳍、腹鳍、臀鳍和脂鳍等在胚后发育过程中相继分化。其初孵仔鱼平均全长为(9.33±0.35) mm, 仔鱼卵黄囊呈圆球形, 18日龄时卵黄囊和油球被完全消耗。其早期发育阶段(0—16日龄)的生长特性符合公式: y=0.0005x4–0.0201x3+0.2264x2–0.3773x+9.6102(R2=0.9968)。研究初步阐明了北极茴鱼的胚胎发育和仔鱼发育的时序特征, 为今后的苗种规模化繁育提供了理论基础。Abstract: Thymallus arcticus grubei is only distributed in the Irtysh River basin in northern Xinjiang Uygur Autonomous Region in China. In recent years, due to overfishing and construction of water conservancy projects, its resources have declined sharply. It was listed as a second-level protected fish in Xinjiang Uygur Autonomous Region in 2004. This research carried out systematic observations on the embryonic development of artificially propagated Thymallus arcticus grubei, and recorded and analyzed the morphological characteristics of their embryos and larvae at various stages of development, aiming to provide necessary basic data for the breeding of Thymallus arcticus grubei and resource protection. The results showed that the fertilized eggs of Thymallus arcticus grubei were spherical, golden yellow, sinking eggs, unabsorbed eggs diameter (2.46±0.14) mm, water-absorbed eggs diameter (3.14±0.18) mm, and there were multiple oil droplets in the yolk. The number and spatial distribution of oil droplets changed regularly during embryonic development. Under the conditions of incubation water temperature (11.06±0.72)℃ and dissolved oxygen of 8.3—9.8 mg/L, it takes 301h to complete the entire embryonic development process, and the required accumulated temperature is 3384.84h·℃. There are 7 stages, zygote, cleavage, blastula, gastrula, neurula, organ formation and hatching, totaling 26 stages. The differentiation of caudal and pectoral fins and eye pigmentation of Thymallus arcticus grubei larvae have been completed in the late embryonic development of the fertilized egg, and the dorsal fin, pelvic fin, anal fin, adipose fin, etc. differentiated in the post-embryonic development process. The average length of the newly hatched larvae is (9.33±0.35) mm. The yolk sac and oil droplets are completely consumed at the age of 18 days. The growth characteristics of its early developmental stage (0—16 days old) conform to the formula: y=0.0005x4–0.0201x3+0.2264x2–0.3773x+9.6102 (R2=0.9968). This study preliminarily clarified the timing characteristics of the embryonic development and larval development of Thymallus arcticus grubei, and provided a theoretical basis for the future large-scale breeding of seedlings.
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Keywords:
- Embryonic development /
- Larval /
- Morphological /
- Irtysh River /
- Thymallus arcticus grubei
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图版Ⅰ 北极茴鱼胚胎发育
1. 受精卵; 2. 胚盘形成; 3. 2细胞期; 4. 4细胞期; 5. 8细胞期; 6. 16细胞期; 7. 32细胞期; 8. 64细胞期; 9. 多细胞期; 10. 囊胚早期; 11. 囊胚中期; 12. 囊胚晚期; 13. 原肠早期(箭头指示胚环); 14. 原肠中期; 15. 原肠晚期; 16. 神经胚期; 17. 胚孔封闭期; 18. 脑分化期(箭头指示脑泡); 19. 眼囊出现期(箭头指示眼囊); 20. 胸鳍芽出现期(箭头指示胸鳍芽); 21. 尾芽期(箭头指示尾芽); 22. 晶体出现期(箭头指示晶体); 23. 尾鳍期(箭头指示尾鳍); 24a. 色素沉积期; 24b. 耳石(箭头指示耳石); 25. 循环期(箭头指示心脏); 26. 出膜期。标尺表示0.4 mm
图版Ⅰ. Embryonic development of T. arcticus grubei
1. fertilized egg; 2. blastoderm; 3. 2 cell stage; 4. 4 cell stage; 5. 8 cell stage; 6. 16 cell stage; 7. 32 cell stage; 8. 64 cell stage; 9. multi-cell stage; 10. early blastula stage; 11. mid-blastula stage; 12. late blastula stage; 13. early gastrula stage (arrowhead indicates germ ring); 14. mid-gastrula stage; 15. late gastrula stage; 16. neurula stage; 17. closure of blastopore stage; 18. brain differentiation stage (arrowhead indicates cerebral vesicle); 19. appearance optic capsule (arrowhead indicates optic capsule); 20. pectoral fin appearance stage (arrowhead indicates pectoral fin); 21. tail bud period (arrowhead indicates tail bud); 22. eye lens formation stage (arrowhead indicates eye lens); 23. caudal fin appearance (arrowhead indicates caudal fin); 24a. Eye coloring material formation stage; 24b. statolith (arrowhead indicates statolith); 25. gill circulation stag (arrowhead indicates heart); 26. hatching stage. Scale bar=0.4 mm
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图版Ⅱ 北极茴鱼仔稚鱼发育
1. 1日龄; 2. 4日龄; 3. 7日龄; 4. 10日龄; 5. 13日龄; 6. 17日龄; 7. 23日龄; 8. 52日龄
图版Ⅱ. Larva development of T. arcticus grubei
1. 1d; 2. 4d; 3. 7d; 4. 10d; 5. 13d; 6. 17d; 7. 23d. 8. 52d
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图 1 北极茴鱼仔鱼卵黄囊体积与出膜天数的关系
Figure 1. Relationship between yolk sac volume and days of hatching of T. arcticus grubei
表 1 北极茴鱼发育各阶段积温
Table 1 Temperatures of T. arcticus grubei at different stages of embryonic development
发育时期
Development
stage经历时长
Elapsed
time (h)平均温度
Average
temperature
(℃)积温
Accumulative
temperature
(h·℃)图版Ⅰ
PlateⅠ胚盘形成期 4h 10.80 43.20 1—2 卵裂期 10h10min 12.01 122.14 3—9 囊胚期 32h45min 12.26 401.52 10—12 原肠胚期 24h35min 11.90 292.50 13—15 神经胚期 11h30min 11.41 131.22 16—17 器官分化期 140h 10.99 1538.60 18—24 孵化出膜期 78h 10.97 855.66 25—26 
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表 2 北极茴鱼发育时序及主要特征
Table 2 Embryonic development of T. arcticus grubei
发育时期
Development stage受精后时间
Time after fertilization主要特征
Developmental characteristics图版Ⅰ
PlateⅠ受精卵 0 圆球形,微黏性,沉性卵, 金黄色, 多个分散状油球均匀分布其中 1 胚盘期 4h 受精卵吸水膨胀后形成卵周隙, 原生质集中于动物极形成胚盘 2 2细胞期 6h10min 第1次分裂, 形成两个大小相同的细胞, 油球高密度集中分布在细胞体下方 3 4细胞期 7h25min 第2次分裂, 形成2行2列排布的4个大小基本相同的细胞 4 8细胞期 8h15min 第3次分裂, 形成2行4列排布的8个大小略有差异的细胞 5 16细胞期 9h20min 第4次分裂, 形成4行4列排布的16个大小略有差异的细胞 6 32细胞期 10h50min 第5次分裂, 形成4行8列排布的32个大小略有差异的细胞 7 64细胞期 12h50min 第6次分裂, 分裂时间不完全同步, 形态大小有差异, 出现重叠排布的多个细胞 8 多细胞期 14h10min 细胞数量增多, 体积变小, 细胞界限模糊但仍可分辨 9 囊胚早期 23h 细胞界限模糊, 表面趋于平滑, 胚盘隆起达到最高 10 囊胚中期 32h15min 胚盘高度有所下降 11 囊胚晚期 46h55min 胚盘向扁平发展, 高度明显降低, 胚盘边缘的细胞开始向植物极下包,
油球开始向植物极转移12 原肠早期 53h10min 胚盘细胞延伸至卵黄直径的1/3处, 油球随着胚体下包逐渐向植物极方向转移 13 原肠中期 61h20min 胚盘下包至卵黄直径的1/2处, 胚环、胚盾形成 14 原肠晚期 71h30min 胚盘下包至卵黄直径的2/3处, 胚盾变厚, 油球随着胚体下包继续向植物极方向转移 15 神经胚期 76h30min 胚盘下包至卵黄约4/5处, 胚胎背部形成神经板, 卵黄栓出现, 油球基本均匀分布于整个卵黄囊内 16 胚孔封闭期 83h 胚孔封闭, 胚层完全覆盖卵黄栓, 外包运动结束, 胚体前部隆起 17 脑部分化期 95h 脑部开始分化 18 眼囊出现期 117h 胚体头部两侧出现一对椭圆形突起 19 胸鳍芽出现期 133h 两侧胸鳍芽出现 20 尾芽出现期 142h 尾部和卵黄囊分离, 尾芽出现 21 晶体出现期 159h 眼囊中可以看到明显的晶体轮廓 22 尾鳍出现期 166h 尾鳍出现, 胚体扭动 23 眼色素沉积期 172h 肉眼可观察到眼部有黑色素沉积, 耳囊内可观察到耳石 24 循环期 223h 可见心脏跳动和血液流动, 胚体扭动剧烈, 胚体在卵膜内转动 25 出膜期 301h 胚体在卵膜内剧烈扭动, 破膜而出 26
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表 3 北极茴鱼与其他鲑形目鱼类的比较
Table 3 Comparsion between T. arcticus grubei and other Salmoniformes
种类
Species孵化时间
Duration from fertilization to hatching (h)平均温度
Water temperature (℃)总积温
Accumulated temperature (℃·h)吸水后卵径
Egg diameter after water absorption (mm)出膜仔鱼全长
Body length at hatching (mm)北极茴鱼T. arcticus grubei 301.00 11.06 3384.84 3.14 9.33 黑龙江茴鱼T. arcticus grubei Dybowski[9] 408.00 7.35 2808.00 3.19 8.78 鸭绿江茴鱼T. arcticus yaluensis Mori[10] 600.00 7.50 4800.00 3.00 10.00 哲罗鱼H. taimen[11] 839.00 7.50 6292.56 5.55 18.45 秦岭细鳞鲑
B. lenole tsinlingensis[12]408.00 11.03 4640.40 4.25 9.64 虹鳟O. mykiss[14] 1248.00 6.30 7874.88 5.24 16.51 白点鲑S. leucomaenis [17] 1944.00 6.14 10173.00 6.17 17.89
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