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1 , 19(6): Journal of Fishery Sciences of China 研究论文 DOI: /SP.J 张颖 1, 孙慧武 2, 刘晓勇 2, 孙大江 1, 曲秋芝 1., ; 2., 摘要 : (Acipenser schrenckii) (PGCs), PGC 1.5 dph (days post-hatching) ( ) 3~7 dph, PGCs,, 17~31 dph, PGCs, ; dph, ; 90~120 dph,, 180 dph, ( ) ( )2,, d 56 d (27 ± ± ± ± ±0.5 ) 210 d,,,, (24 27 ), 27 (21 ) 50 d, d; (15 ) : ; PGCs; ; ; 中图分类号 : S963 文献标志码 : A 文章编号 : (2012) (Acipense schrenckii),, [1 3] 2006, t [2], ( ) 50% [2 3],,, t, 40.85% [4 5],,, [5], B, [6 7], [7], Devlin [8], (temperature sex determine, TSD), [9 13] 56 TSD, TSD, 收稿日期 : ; 修订日期 : 基金项目 : ( ); (2012BAD25B10-5); ( ). 作者简介 : (1977-),,,,. juletzhang@hotmail.com 通信作者 :,. sunhw@cafs.ac.cn

2 6 : 1009, 鳉 (Rivulus marmoratus),,,,, , 16 17, > 6.0 mg/l : (18±1), >6.0 mg/l, ph 7.5±0.1, 27 dph(days post-hatching), 11 mm ( 40%, 16%, 5%, 16, 12%, 2.0%) 0 dph, d 1, 7 d 1 ; 3~7 15 d 1 ; , Bouin s 24 h, 5 cm, 5 10 cm, 10 cm,,, 5 μm, H&E, Leica DM4000B 5 cm, [14] 1.3, 95 cm 50 cm 60 cm, 56 d, 3 d 1,, 5 (15, 18, 21, ), ( ) 3, 30 (±0.5 ) ph 7.5, >6.0 mg/l, 210 d,, Bouin s 24 h, 1.2, , 1.5 d, (7.32±0.16) mm,,,,,, ( I-1) 3 dph (10.58±0.25) mm, PGCs 3 ( I-2) 5 dph PGCs ( I-3) 7 dph ( I-4) 10 dph PGCs ( I-5) 17 d, (20.47±0.47) mm, PGCs ( I-6) 24 d, (24.13±1.04) mm, 1 (genital ridge), PGCs, ( I-7) 31 d, (35±2.15) mm,, PGCs ( I-8) dph,, ( I-9) 90 dph, mm, 1,, ( I-10) 120 d,

3 PGC, ( I-11) 150 d, mm,, ( I-12),, ( I-13) 180 dph, mm,,, ( I-15,18) 2 ( I-14,16,17),, dph, PGC dph, mm, ( 1, 2), ( 2),,,, ( 3) 2.1.3, 210 dph, mm, ( Ⅱ-4, -2), ( Ⅱ-5),,,, ( Ⅱ-6), ( Ⅱ-7), 2.2 1,,, (18 ), 15, 210 d, 1.. 1( -1, 2) (24 27 ),, 130 d 150 d, 1.. 1( -3, 4, 5, 6), ( -7,8; -9,10), 15 27,, (PGCs),, : 1), PGCs ( 10~22 μm),, : 2.6 [14] ; 2), 6~10 μm,,, [14 16], PGCs,,,,, (Acipenser ruthenus) (A. baerii baicalensis) (A. trasmontanus) (A. naccarii) [14, 17 18] (PGCs), 70% / temperature Tab. 1 numbers of male 表 1 培养水温对施氏鲟性腺分化和性别比率的影响 Effects of temperature on gonadal differentiation and sex ratios of Amur sturgeon numbers of female individual numbers of gonad undifferentiation n=30 sex ratio (female: male)

4 6 : ~100, 5~23, 40~50, 79~109 [14], PGCs, 25~74, (A. gueldenstaedtii) 32~80, 5~50 2~24 7~33 (A. stellatus Pallas) 23~27 [14 15, 19], PGCs 20~30, PGCs, [14 19] PGCs PGCs, [14 19], PGCs 1.5 d, 1 d, 1~2 h, (A. baerii) 3 d PGCs 6.5 d [14 20] PGCs, PGC [11 18, 21 26], PGCs, PGC, [11 16] (Brachydanio rerio)pgc, PGCs 24 h 4,, 24 h [23 24], PGC, 鳉 (Oryzias latipes) (Micropterus salnmides) [27 29], 1.5 d PGCs PGCs, [14], PGCs : 1) ; 2) ; 3) [15 16,21,24] PGCs, [14], PGCs,,, (A. naccari) [18] 3.1.3,, 180 dph, [14,17 20,25 26],,, (Huso huso A. ruthen)(6 ), 3, (8 ) (A.brevirostrum)(7 ) (A.sinensis) (9 ) [17 20,25 26,30 31], 2 : 1), ; 2), [14, 17 20, 25 26],, [25],,,,,,,, [14 32],,,,, [16], 3.2 [33],,

5 (TSD) [34], TSD, (Oreochromis niloticus) (Dicentrar chllslabrar L.) [35 36],,,, TSD,, [37], Navarro-Martín [38] cyp19a, cyp19a, SF-1 Foxl2, cyp19a DNA TSD 参考文献 : [1],,,. [J]., 2010, 17(4): [2],,,. [J]., 2011, 24(4): [3],,,. [J], 2004, 28(5): [4] [D]. :, 2011 [5],,. [J]., 2011(7): [6],,,. [J]., 2009, 31(2): [7],,,. [J]., 2011, 42(12): [8] Guerrero-Estévez S., Moreno-Mendoza N. Sexual determination and differentiation in teleost fish[j]. Rev Fish Biol Fisheries, 2009, DOI /s [9] Ospina-Álvarez N, Piferrer F. Temperature-Dependent Sex Determination in Fish Revisited: Prevalence, a Single Sex Ratio Response Pattern, and Possible Effects of Climate Change. PLoS ONE, 2008,3 (7): e2837 DOI: /journal. pone [10] Wang L H, Tsai CL. Effects of temperature on the deformity and sex differentiation of tilapia, Oreochromis mossambicus[j]. J Exp Zool, 2000, 286: [11],,. [J]., 2007, 14 (1): [12] Pavlidis M, Koumoundouros G, Sterioti A, et al. Evidence of temperaturedependent sex determination in the European sea bass (Dicentrarchus labrax L.) [J]. J Exp Zool, 2000, 287(3): [13] Harrington RW, Crossman RA. Effects of temperature and sex genotype on meristic counts of the gonochoristic cyprinodontid fish Rivulus cylindraceus Poey[J]. Can J Zool, 1976, 54: [14]. [M]. :, 1981: [15],,,. [J]. :, 2010, 40(2): [16]. [J]., 1991, 15(1): 1 7. [17] Akhundov M M, Fedorov K Y. Effect of exogenous estradiol on ovarian development in juvenile sterlet, Acipenser ruthenus[j]. J Ichthyol, 1995, 35: [18] Grandi G, Giovannini S, Chicca M. Gonadogenesis in early developmental stages of Acipenser naccarii and influence of estrogen immersion on feminization [J]. Appl Ichthyol, 2007, 23: 3 8. [19] Akhundov M M, Fedorov KY. Early gametogenesis and gonadogenesis in sturgeons. On criteria for comparative assessment of juvenile gonadal development in the example of the Russian sturgeon, Acipenser gueldenstaedtii [J]. J Ichthyol, 1991, 31: [20] Grandi G, Chicca M. Histological and ultrastructural investigation of early gonad development and sex differentiation in Adriatic sturgeon (Acipenser naccarii, Acipenseriformes, Chondrostei) [J]. J Morphol, 2008, 269(10): [21] Saito T, Goto-Kazeto R, Kawakami Y, et al. The mechanism for primordial germ-cell migration is conserved between Japanese eel and zebrafish[j]. PLoS ONE, 2011, 6(9): e24460, DOI: /journal.pone [22] Kurokawa H, Aoki Y, Nakamura S, et al. Time-lapse analysis reveals different modes of primordial germ cell migration in the medaka Oryzias latipes [J]. Dev Growth Differ, 2006, 48:

6 6 : 1013 [23] Weidinger G, Wolke U, Köprunner M, et al. Identification of tissues and patterning events required for distinct steps in early migration of zebrafish primordial germ cells[j]. Development, 1999, 126: [24] Richardson B E, Lehmann R. Mechanisms guiding primordial germ cell migration: strategies from different organisms[j]. Nat Rev Mol Cell Biol, 2010, 11: [25],,, [J]., 2004, 28(6): [26],,,. [J]., 2010, 17(3): [27] Sanchez-Sanchez AV, Camp E, Garcia-Espana A, et al. Medaka oct4 is expressed during early embryo development, and in primordial germ cells and adult gonads[j]. Dev Dyn, 2010, 239(2): [28] Shinomiya A, Tanaka M, Kobayashi T, et al. The vasa-like gene, olvas, identifies the migration path of primordial germ cells during embryonic body formation stage in the medaka, Oryzias latipes[j]. Dev Growth Different, 2000, 42(4): [29] Johnston P M. The embryonic history of the germ cells of the largemouth black bass, Micropterus salmoides salmoides (Lacépède) [J]. J Morphol, 1951, 88(3): [30] Flynn S R, Benfey T J. Sex differentiation and aspects of gametogenesis in shortnose sturgeon Acipenser brevirostrum Lesueur [J]. J Fish Biol, 2007, 70: [31] Kijima T, Maruyama T. Histological research for the development of the gonad of hybrid sturgeon, bester (Acipenser ruthenus L. male Huso huso L. female) [J]. Bull Nat Res Ins Aqu, 1985, 8: [32], [J], 2004(4): [33] Devlin R H, Nagahama Y. Sex determination and sex differentiation in fish:an overview of genetic, physiobgica, and environmental influences[j]. Aquaculture, 2002, 208: [34] Baroiller J F, D Cotta H, Saillant E. Environmental effects on fish sex determination and differentiation[j]. Sex Dev, 2009, 3(2 3): [35] Baroiller J F, Nakayama I, Fostier A, et a1. Sex determenation studies in two species of teleost fish, Oreochromis niloticus and Leporinus elongates[j]. Zool Stud, 1996, 35: [36] Blazquez M, Zanuy S, Carillo M, et a1.effects of rearing temperature on sex differentiation in the European sea bass (Dicentrar chllslabrar L.) [J]. J Exp Zool, 1998, 281: [37] Spotila R J, Spotila D L, Kaufer F N. Molecular mechanisms of TSD in reptiles: A search for the magic bullet[j]. J Exp Zool, 1994, 270(1): [38] Navarro-Martín L, Viñas J, Ribas L, et al. DNA Methylation of the gonadal Aromatase (cyp19a) promoter is involved in temperature-dependent sex ratio shifts in the European sea bass[j]. PLoS Genet, 7(12): 2011, e DOI: / journal.pgen

7 Histological observation of gonadal differentiation and effect of rearing temperature on sex differentiation in Amur sturgeon Acipenser schrenckii ZHANG Ying 1, SUN Huiwu 2, LIU Xiaoyong 2, QU Qiuzhi 1, SUN Dajiang 1 1. Heilongjiang River Fisheries Research Institute, Chinese Academy of Fishery Sciences, Harbin , China; 2. Chinese Academy of Fishery Sciences, Beijing , China Abstract: Temperature-dependent sex determination has been demonstrated in some species of fish and temperature during the period of sex differentiation typically produces male or female-dominant population. This research investigated the gonadal sex differentiation and effect of rearing temperature on the sex ratio in larval Amur sturgeon Acipenser schrenckii. In 1.5 days post-hatch, a PGC is visible along the dorsal epithelium of the body cavity, between the mesentery and the kidney. Later, in 3-7 days post-hatch, the numbers of the primordial germ cells (PGC) increased, and gathered into cell cable. At dph, PGCs along the mesenteric migrate to below the kidney area and cells together to form the gonadal ridges. When dph, the original gonadal development, groups of small cells from kidney area migrate to original gonad, original gonadal columnar epithelial cells. At dph, germinal epithelium and cells appears in the original gonad, fat cells and blood vessels increase. Putative ovaries with notches in the germinal epithelium and presumed testes with smooth germinal epithelium appeared in 180 days post-hatch, signs of juvenile Acipenser schrenckii already on the level of differentiation of gonads in anatomy. Ovaries with proliferating oogonia and early meiotic oocytes clusters were observed in 210-day-old juveniles. In a temperature controlled experiment, the proportions of females were 50% at 21, 43.3% at 15, 43.3% at 18, 50% at 24, and 46.7% (the same as 1:1 sex ratio) at 27. These results suggest that morphological sex differentiation in Amur sturgeon occurs at approximately 180 days after hatch, and though the high temperatures can not change the 1:1 sex ratio in this species, but it can help to promote gonadal development of juvenile Amur sturgeon. Key words: Acipenser schrenckii; PGCs; gonadal differentiation; temperature; sex ratio Corresponding author: SUN Huiwu. sunhw@cafs.ac.cn

8 第6期张颖等: 施氏鲟的性腺分化及养殖水温对其性腺分化的影响图版 I Plate I 施氏鲟 PGCs 的出现迁移及性腺原基的形成 PGCs arise, migration and gonad primordia formation of Amur sturgeon 1015

1016 中国水产科学第 19 卷图版 I 说明 1. 孵化后 1.5 d 仔鱼的横切, 箭头示单个 PGC 细胞; 2. 孵化后 3 d, 仔鱼的横切, 箭头示 3 个 PGC 细胞; 3. 孵化后 5 d, 示多个 PGCs 在肾管和肠管之间; 4. 孵化后 7 d, PGCs 形成背细胞索; 5. 孵化后 10 d, PGCs 从侧板的中胚层时其沿肠系膜向肾管区迁移; 6.

9 1016 中国水产科学第 19 卷图版 I 说明 1. 孵化后 1.5 d 仔鱼的横切, 箭头示单个 PGC 细胞; 2. 孵化后 3 d, 仔鱼的横切, 箭头示 3 个 PGC 细胞; 3. 孵化后 5 d, 示多个 PGCs 在肾管和肠管之间; 4. 孵化后 7 d, PGCs 形成背细胞索; 5. 孵化后 10 d, PGCs 从侧板的中胚层时其沿肠系膜向肾管区迁移; 6. 孵化后 17 d, PGCs 沿腹膜上皮迁移到中肾管下方肠壁上方; 7. 孵化后 24 d, 示 1 对生殖嵴; 8. 孵化后 31 d, 示原始性腺, 箭头示生殖脊和 PGCs; 9. 孵化后 d, 成群的小细胞团由肾区向原始性腺迁移, 原始性腺上皮细胞柱状; 10. 孵化后 90 d, 原始性腺背面出现生殖上皮细胞, 腹侧出现脂肪细胞, 靠近柄部出现血管; d, PGCs 增多, 核仁染色清晰; 12. 孵化后 150 d, 原始性腺继续增大, 切面呈叶片状; 13. 孵化后 150 d, 游离末端脂肪组织变大, 血管丰富; 14. 孵化后 180 d, 卵巢已经分化; 15. 孵化后 180 d, 精巢已经分化; 16. 孵化后 180 d, 卵巢已经分化; 17. 孵化后 180 d, 分化卵巢的生殖上皮形成皱褶; 18. 孵化后 180 d, 分化精巢的生殖上皮. B:血管, ET:生殖上皮, F:脂肪, GO:性腺, GR:生殖脊, GV:生殖沟, IN:肠, K:肾脏, OV:卵巢, PGC:原始生殖细胞,TE:精巢. Explanation of Plate I d after hating, a PGC appeared in mesoderm, the arrow indicated PGC; 2. 3 d after hating, three PGCs appeared in Virginia Woolf's tube, the arrow indicated PGCs; 3. 5 d after hating, multiple PGCs appeared between the renal tube and bowel; 4. 7d after hating, PGCs alignment in dorsal side; d after hating, PGCs migrated from lateral plate mesoderm along the mesenteric and renal districts; d after hating, PGCs along the peritoneum epithelial migration and the renal tube in the intestinal wall; d after hating, form a pair of genital Ridge, the arrow indicated the genital Ridge and kindey; d after hating, PGCs has moved into genital ridge, constitute the original gonad, the arrow indicated PGCs and genital ridge; d after hating, flocks of small cells from kidney area migrated to the original gonad, the original sexual gland epithelial cells cylindrical; d after hating, epithelial cells of original gonadal occurs on the back, abdominal fat cells appear, blood vessel appears near the shank vascular; d after hating, PGCs proliferation and nucleolus stain clear; d after hating, original gonads grow, slices are leaf-shaped; d after hating, adipose tissue and blood vessel-rich; d after hating, ovary has differentiated; d after hating, testis has differentiated; d after hating, differentiated ovary; d after hating, germinal epithelium formed folds or groove of the ovary; d after hating, germinal epithelium of differentiated testis. B: Vascular, ET: Germinal epithelium, F: fat, GO: gonad, GR: gonadal ridge, GV: genital groove, IN: intestinal, K: kidney, OV: ovarian, PGC: primordial germ cells, TE: testes. 图版 II 施氏鲟卵巢和精巢的分化 1. 孵化后 210 d, 示已分化卵巢, 卵巢内出现产卵板, 箭头示血管; 2. 孵化后 210 d, 示已分化卵巢, 每个初级卵母细胞由卵泡细胞和颗粒细胞包围; 3. 孵化后 210 d, 箭头示正在进行第 1 次成熟分裂的卵母细胞; 4. 孵化后 210 d, 示已分化的精巢, 箭头示脂肪; 5. 孵化后 210 d, 示已分化精巢的生殖上皮; 6. 孵化后 210 d, 箭头示初级精母细胞 AC: 脂肪细胞, BV: 血管, ET: 生示产卵板. 殖上皮, F: 脂肪, FC: 卵泡细胞, GC: 颗粒细胞, OO: 卵母细胞, OV: 卵巢, SC: 初级精母细胞, TE: 精巢, Plate II Ovarian and testicular differentiation of Amur sturgeon d after hatching, showing differentiated ovary, and ovarian board appeared in ovarian, arrow indicating blood vessels; d after hatching, showing the differentiated ovary, and the primary oocyte surrounded by follicular cells and granule cells; 3. primary oocyte, arrow indicating oocyte; d after hatching, showing the differentiated testis, arrow indicating fat; d after hatching, showing germinal epithelium of primary spermatocytes; d after hatching, showing primary spermatocytes. AC: Adipose Cell, BV: Vascular, ET: Germinal epithelium, F: Fat, FC: Follicular cells, GC: Granulosa cells, OO:Oocyte, OV: ovarian, SC: primary eggs plates. spermatocytes, TE: testes,

第6期张颖等: 施氏鲟的性腺分化及养殖水温对其性腺分化的影响图板 III 1017 温度对施氏鲟性别比率的影响 1. 15 组精巢; 2. 15 组卵巢; 3. 27 组卵巢; 4. 27 组精巢; 5. 24 组卵巢; 6. 24 组精巢; 7. 18 组卵巢; 8. 18 组精巢; 9. 15 组卵巢; 10. 15 组精巢. AC:脂肪细胞, BV:血管, ET:生殖上皮, OO:卵母细胞, OV:卵巢, TE:精巢.

10 第6期张颖等: 施氏鲟的性腺分化及养殖水温对其性腺分化的影响图板 III 1017 温度对施氏鲟性别比率的影响组精巢; 组卵巢; 组卵巢; 组精巢; 组卵巢; 组精巢; 组卵巢; 组精巢; 组卵巢; 组精巢. AC:脂肪细胞, BV:血管, ET:生殖上皮, OO:卵母细胞, OV:卵巢, TE:精巢. Plate III Effect of temperature on sex ratio of Amur sturgeon 1.Testis of 15 group; 2. Ovary of 15 group; 3. Ovary of 27 group; 4. estis of 27 group; 5. Ovary of 24 group; 6. Testis of 24 group; 7. Ovary of 18 group; 8. Testis of 18 group; 9. Ovary of 15 group; 10. Testis of 15 group. AC: adipose cell, BV: vascular, ET: germinal epithelium, OO:oocyte, OV: ovarian, TE: Testes.

<4D6963726F736F667420506F776572506F696E74202D2032303135D0D4B1F0BEF6B6A8D3EBC9FAD6B3CFB8B0FBB7A2C9FA205BBCE6C8DDC4A3CABD5D>

<4D6963726F736F667420506F776572506F696E74202D2032303135D0D4B1F0BEF6B6A8D3EBC9FAD6B3CFB8B0FBB7A2C9FA205BBCE6C8DDC4A3CABD5D> 性别决定与生殖细胞发育历史观点公元前 335 年,Aristotle 指出性别决定于热 (heat) 1600s/1700s, 已认识到 female 可以产生能够传递亲本性状的卵子 19 世纪末之前, 人们认为温度营养年龄等环境因素决定了性别有利于能量和营