Histological observation on gonadal differentiation and first annual gonadal development of cobia (Rachycentron canadum)
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摘要: 本研究采用石蜡组织切片和H.E.染色法对军曹鱼原始性腺的形成、分化及精巢和卵巢首周年发育的组织结构变化进行观察。结果显示，军曹鱼原始生殖细胞在7孵化日龄(days post hatching，dph)时迁移到达生殖嵴，随后体细胞出现聚集和分裂，并于15 dph将原始生殖细胞完全包绕形成原始性腺。军曹鱼卵巢分化的时间要早于精巢，34 dph的稚鱼性腺组织尚未分化，但可观察到两种存在明显组织学差异的性腺类型，其中一种类型的性腺出现卵原细胞群，而另一种类型的性腺横切面较狭长，生殖细胞数量明显较少，因此推定为未分化精巢；卵巢的解剖学分化开始于44 dph，其标志为卵巢腔的形成；50 dph时精巢开始细胞学上的分化，此时精原细胞由基底膜包被形成囊泡状的细胞团。在首周年发育期间 (60～360 dph)，军曹鱼的精巢发育包含I、II、III、IV、V 5个时期，而卵巢发育只包含I、II、III 3个时期。60 dph时，精巢和卵巢均处于I期；90 dph时，精巢发育至II期，卵巢仍处于I期；120 dph时，超过半数的精巢已发育至 III期，仅少部分卵巢发育至II期；150 dph时，精巢已发育至III期，而大部分卵巢发育至II期；185 dph时，精巢仍为III期，卵巢均已发育至II期；210 dph时，大部分精巢发育至IV期，卵巢仍处于II期；360 dph时，精巢已发育至V期，大部分卵巢发育至III期。上述研究结果可丰富军曹鱼的繁殖生物学研究基础，阐明其早期性腺发育规律，还可为其人工繁殖提供理论依据。Abstract: In this study, histological observation of gonadogenesis, gonadal differentiation and first annual gonadal development in cobia (Rachycentron canadum) was performed based on paraffin section and H.E. staining method. The results of histological observations showed that the primordial germ cells (PGCs) arrived at the genital ridge at 7 dph (days post hatching), then the somatic cells began to aggregate around the PGCs. At 15 dph, the PGCs were enclosed entirely by somatic cells and formed the primary gonad. Two types of primary gonads with noticeable histological differences could be observed at 34 dph. The presence of clusters of oogonia characterized a part of primary gonads. The other primary gonads showed a narrower cross-section with fewer germ cells, which were presumed to be undifferentiated testis. Similar to other fishes, ovarian differentiation of cobia was anterior to testicular differentiation. The anatomical indication of ovarian differentiation was the ovarian cavity formation, which occurred at 44 dph. The cytological differentiation of testis began at about 50 dph and was characterized by the presence of spermatogonial acinar-clusters. The first annual development of testes could be divided into five stages (I, II, III, IV, V), while the development of the ovaries was divided into three stages (I, II, III). Testes and ovaries were both developed to early Stage I at 60 dph. All of the testes developed to Stage II at 90 dph; the ovaries were still at Stage I. More than half of the testes developed to Stage III and part of the ovaries developed to Stage II at 120 dph. The testes were mainly developed to Stage III at 150 dph; most of the ovaries were at Stage II. The testes were still at stage III, and the ovaries were developed to Stage II at 185 dph. Most of the testes developed to Stage IV at 210 dph, but the ovaries were still at Stage II. At 360 dph, the testes entered Stage V, and most of the ovaries developed to Stage III. These findings could clarify gonadal development characteristics and provide a theoretical basis for the artificial breeding of cobia.
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图 1 军曹鱼原始生殖细胞的胚后迁移
a. 原始生殖细胞分布位置示意图（红点表示原始生殖细胞）；b, c. 3 dph和5 dph单个游离存在的原始生殖细胞；d, e. 7 dph和9 dph原始生殖细胞迁移至生殖嵴；f. 11 dph原始生殖细胞周围开始出现体细胞聚集；g. 15 dph原始生殖细胞被体细胞包围形成原始性腺雏形；PGC. 原始生殖细胞；GC. 生殖细胞；SC. 体细胞；PG. 原始性腺；MD. 中肾管；G. 肠
Fig. 1 Migration of primordial germ cells of Rachycentron canadum during early development
a. Schematic diagram of PGCs distribution (red dots indicated primordial germ cells); b, c. individual PGC at 3 dph and 5 dph; d, e. PGCs arrive at the genital ridge at 7 dph and 9 dph; f. somatic cells began to aggregate around PGCs at 11 dph; g. somatic cells encompassed PGCs and the prototype primary gonad was formed at 15 dph; PGC. primordial germ cells; GC. germ cell; SC. somatic cell; PG. primary gonad; MD. mesonephric duct; G. gut
图 2 军曹鱼原始性腺的分化
a−e. 18 dph、21 dph、24 dph、27 dph和30 dph尚未出现分化特征的原始性腺；f. 34 dph早期卵巢，形成卵原细胞群；g. 34 dph推定的精巢组织；h. 40 dph早期精巢，精原细胞出现；i. 50 dph早期精巢，形成精原细胞囊；j. 44 dph早期卵巢，出现卵巢腔；k. 48 dph早期卵巢，卵巢腔扩大；l. 52 dph早期卵巢，卵原细胞开始向初级卵母细胞过渡；BC. 血细胞；BV. 血管；GC. 生殖细胞；PG. 原始性腺；O. 卵巢；OG. 卵原细胞；PT. 推定的精巢组织；T. 精巢；SG. 精原细胞
Fig. 2 Differentiation of primary gonad of Rachycentron canadum
a−e. Undifferentiated primary gonad at 18 dph, 21 dph, 24 dph, 27 dph and 30 dph; f. early differentiated ovary at 34 dph, clusters of oogonia formed; g. presumptive testis at 34 dph; h. early differentiated testis at 40 dph, clusters of spermatogonia formed; i. early differentiated testis at 50 dph, cysts of spermatogonia formed; j. early differentiated ovary at 44 dph, ovarian cavity formed; k. early differentiated ovary at 48 dph, ovarian cavity increased in size; l. early differentiated ovary at 52 dph, oogonia began to transform into primary oocytes; BC. blood cell; BV. blood vessel; GC. germ cell; PG. primary gonad; O. ovary; OG. oogonia; PT. presumptive testis; T. testis; SG: spermatogonia
图 3 军曹鱼精巢首周年发育的组织学变化
A, a. 60 dph精巢切片；B, b. 90 dph 精巢切片；C, c. 120 dph精巢切片；D, d. 150 dph精巢切片；E, e. 185 dph 精巢切片；F, f. 210 dph精巢切片；G, g. 360 dph精巢切片；SG. 精原细胞；PSC. 初级精母细胞；SSC. 次级精母细胞；ST. 精细胞；SP. 精子；BC. 血细胞
Fig. 3 The histologic changes of first annual testicular development of Rachycentron canadum
A, a. Section of testis at 60 dph; B, b. section of testis at 90 dph; C, c. section of testis at 120 dph; D, d. section of testis at 150 dph; E, e. section of testis at 185 dph; F, f. section of testis at 210 dph; G, g. section of testis at 360 dph; SG.spermatogonia; PSC. primary spermatocytes; SSC. secondary spermatocytes; ST. spermatids; SP. sperms; BC. blood cells
图 4 军曹鱼卵巢首周年发育的组织学变化
A, a. 60 dph卵巢切片；B, b. 90 dph卵巢切片；C, c. 120 dph卵巢切片；D, d. 150 dph卵巢切片；E, e. 185 dph卵巢切片；F, f. 210 dph卵巢切片；G, g. 360 dph卵巢切片；OG. 卵原细胞；I. 第I时相卵母细胞；II. 第II时相卵母细胞；III. 第III时相卵母细胞；NU. 核仁；OD. 油滴；YN. 卵黄核
Fig. 4 The histologic changes of first annual ovarian development of Rachycentron canadum
A, a. Section of ovary at 60 dph; B, b. section of ovary at 90 dph; C, c. section of ovary at 120 dph; D, d. section of ovary at 150 dph; E, e. section of ovary at 185 dph; F, f. section of ovary at 210 dph; G, g. section of ovary at 360 dph; OG. oogonium; I. oocyte at Stage I; II. oocyte at Stage II; III. oocyte at Stage III; NU. nucleolus; OD. oil droplet; YN. yolk nucleus
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