红鲫繁殖周期与内分泌生理功能研究
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摘要
红鲫(Carassius auratus red variety)属鲫的变种,在我国分布甚广,是长江流域常见的小型淡水鱼类。本论文以人工养殖的红鲫为研究对象,运用组织学、免疫组织化学、激素放射免疫测定方法以及生物化学方法,对红鲫的性腺、血浆性类固醇激素及其受体和血清无机磷含量进行了较为深入的研究和分析,所得结果与结论如下:
1.红鲫性腺发育周年变化的组织学研究
1.1卵巢的发育分期和卵细胞变化
雌性红鲫具有一对卵巢,对称分布于鳔腹面两侧。非生殖期的卵巢透明,呈粉白色或肉粉色;繁殖期的卵巢体积约占体腔60～70%,呈黄色或橙黄色囊状,可清晰观察到淡黄色的卵粒;退化期的卵巢呈暗红色。组织学观察发现,红鲫卵巢属于卵母细胞发育不同步、分批产卵的类型,卵巢的周年发育分为5个时期,包括Ⅱ期、Ⅲ期、Ⅳ期、Ⅴ期和退化吸收Ⅵ期。卵细胞发育经历5个时相,滤泡膜及卵黄颗粒随着卵细胞的发育而变化。红鲫的繁殖期是在3～5月份,此时卵巢发育都在Ⅳ期到Ⅴ期;6～8月份是排卵后的退化阶段,从9月份开始进入休整期,此时的卵巢处在重复发育Ⅱ期,进入12月份以后,卵巢迅速发育至Ⅲ期,并以Ⅲ期卵巢越冬,从翌年3月份开始,卵巢迅速从Ⅲ期发育至Ⅳ期后,进入产卵高峰期。
1.2精巢的发育分期和精细胞变化
红鲫具有精巢一对,与大多数硬骨鱼类的精巢一样,红鲫的精巢属于小叶型。非繁殖期的精巢为细条状或者宽扁带状;而繁殖期精巢则肥大饱满,呈白色或乳白色。精小叶与精小叶间充满间质细胞、成纤维细胞及丰富的微血管。精子的发生经历精原细胞、初级精母细胞、次级精母细胞、精子细胞以及精子5个典型的发育阶段。从精巢发育的周年变化可以看出,其精巢发育与季节性繁殖活动同步。繁殖季节为3～5月份,6～8月份为退化吸收期,9～11月份为重复发育期,12月份开始启动生精活动。
2.红鲫性腺雌激素受体(AR)、雄激素受体(ER)、芳香化酶的免疫组织化学研究
2.1雌激素受体(ER)在性腺及性细胞中免疫组织化学定位
红鲫卵巢发育的各个时期中雌激素受体(ER)均有表达,滤泡细胞对ER呈强免疫阳性反应,ER在Ⅱ时相卵母细胞核质、胞质和大生长期的放射膜、卵黄颗粒、卵周的液泡以及Ⅵ期末退化吸收的滤泡团及排空的滤泡囊壁均有免疫阳性反应。在精巢发育的各个时期,ER主要定位在精原细胞、初级精母细胞核周、间质细胞、支持细胞,呈免疫阳性反应,早期的精原细胞和精小叶之间的间质处ER表达较强,精子细胞及精子对ER呈免疫阴性反应。
2.2雄激素受体(AR)在性腺及性细胞中免疫组织化学定位
AR在Ⅱ期卵巢的胞质、核质、滤泡膜有较强表达;卵黄发生期,AR则在滤泡膜、放射膜、卵黄颗粒表达,随着卵细胞发育表达逐渐减弱,Ⅳ时相卵母细胞晚期分布在卵周的液泡也有弱阳性反应;Ⅵ期末退化吸收的滤泡团及排空的滤泡囊壁有较强的表达。在精巢,雄激素受体(AR)在各个时期精原细胞、初级精母细胞、次级精母细胞、间质细胞表达,精子细胞和精子显免疫阴性,精小叶间质处对AR的免疫阳性反应相对较强。这些结果表明,雌激素受体、雄激素受体在生殖细胞的增殖和发育过程起作用,为证明性类固醇激素在性腺发育和成熟过程中的重要作用提供证据。
2.3性腺及性细胞中芳香化酶的免疫组织化学
芳香化酶在红鲫卵巢发育的各个时期中都有较强的免疫阳性反应。从Ⅱ时相卵母细胞到Ⅵ时相卵母细胞的卵细胞发育过程中,都对芳香化酶显免疫阳性反应。各期卵巢中Ⅱ、Ⅲ时相卵母细胞的胞质、滤泡膜;Ⅳ时相卵母细胞的胞核、卵黄颗粒、放射膜、滤泡膜以及Ⅵ卵巢中退化分解吸收的卵细胞以及排空的滤泡囊都对芳香化酶呈免疫阳性反应。芳香化酶在红鲫精细胞的表达从精母细胞成熟期(Ⅲ期)至精子成熟期(Ⅴ期)逐渐减弱,且精小叶间质细胞呈免疫阴性反应,但精子退化吸收期(Ⅵ期)精小叶间质处阳性表达较强。这些结果表明,芳香化酶的活性对红鲫性腺发育具有一定的生理效应。
3.红鲫血浆性类固醇激素水平的周年变化研究
3.1雌性红鲫血浆E2和T水平的年周期变化
采用125I标记的放射免疫测定方法(RIA)测定雌性红鲫血清睾酮(T)、雌二醇(E2)含量的季节变动情况。研究发现:繁殖期雌性红鲫血浆的T和E2含量迅速增加,呈显著性变化(P<0.05),E2含量此时高达36.8959±6.8846 pg/ml,T含量达到20.6011±3.6198ng/dl,退化期开始显著下降(P<0.05),至9～11月份进入休整期后T和E2降至全年最低,繁殖前期即12月份至翌年2月份,性激素含量开始缓慢升高。这些结果表明,雌性红鲫繁殖期血浆中雌二醇水平的提高,满足了卵黄形成和积累的需要,促进了卵巢的成熟。T含量与卵巢发育的相关性,提示它们是卵巢发育的一定阶段所必需的。
3.2雄性红鲫血浆E2和T水平的年周期变化
对雄性红鲫血浆睾酮(T)、雌二醇(E2)含量的季节变动研究发现:雄性红鲫血浆T和E2水平的周年变化规律与性腺成熟系数的变化趋势基本一致,全年最高峰值出现在繁殖季节即3～5月份,血浆T含量为37.6000±9.0260ng/dl,血浆E2含量为11.9846±3.2606pg/ml,繁殖后开始显著下降(P<0.05),至静止期降到到最低,从精巢发育Ⅳ期开始T和E2含量升高。这些结果表明,雄性红鲫繁殖期睾酮维持在较高水平,促进精子的发生和形成。
4.红鲫血清无机磷含量的周年变化研究
红鲫血清无机磷的变化规律反映出其与性腺发育密切相关,无论雌性还是雄性,其周年变化规律也都与性腺成熟系数GSI变化规律一致,不同的性腺发育时期,血清无机磷的变化极为明显。繁殖期,红鲫血清无机磷含量增长迅速,呈显著性变化( P < 0.05 ) ,雌性和雄性红鲫血清无机磷含量分别为58.8975±13.6518mmol/L和47.4504±14.9158 mmol/L,6～11月份显著下降,直至繁殖前期血清无机磷含量开始缓慢升高。雌鱼和雄鱼繁殖期血清无机磷含量都为全年的最高峰,能够说明红鲫繁殖时对无机磷的需要量比较高。
Carassius auratus red variety, a variation of Crucian carp, which scatters broadly in China, is common minitype of fresh water fish in Yangtse Valley. This paper takes the artificially bred Carassius auratus red variety as study objects. Gonad, sexual steroid hormone of blood plasma and their receptors, content of serous inorganic phosphorus are studied by the methods of histological staining, immunohistochemical techniques , radioimmunoassay and biochemical methods in details. The main results and conclusions are as follows:
1. Histological study on the development and annual change in the gonad of Carassius auratus red variety
1.1 Stages and oocyte’s change during the ovary’s growth
Carassius auratus red variety has a pair of ovary, scatter symmetrically at two ventral sides of the swim bladder. The color of immature ovary is white or flesh pink; the mature ovary which looks like a yellow or the orange vesicle takes up 60 ~ 70% of body-cavity, and ovum granule can be clearly observed;the degenerate ovary assumes dark red. Histological research finds that the oocyte of Carassius auratus red variety develops out of sync and spawns in batches. The development of ovary can be divided into five stages and the oocytes be divided into five phases: phase2, phase3, phase4, phase5 and phase 6 when the oocytes degenerate. Follicle and vitellin granule are varying with the development of oocytes. The reproductive period of Carassius auratus red variety is between March and May, in which the ovaries are at phase 4 to phase 5; The ovaries degenerate after ovulation from June to August, and have a rest after September from which the ovaries are at phase 2. From the beginning of December , the ovaries live through the winter with the phase 3. From the third month of next year, the ovaries come to the peak of oviposition and went into the phase 4 immediately.
1.2 Stages and the spermatid change during the testicle’s growth
Like other teleost fishes, Carassius auratus red variety has a pair of testicles. Carassius auratus red variety’s testicle is lobular type in appearance. The immature testicle b looks like pinstriped shape or flat tape shape while the mature one is big and assume white color. Be skill and accurate, foliole is full of the inferstitial cells, mechanocyte and lots of blood capillaries. Sperm experience 5 steps during its life: spermatogonium, spermatocyte, spermatocyteⅡ,spoermatid, sperm. The anniversary change from the testicle is it can be seen, testicle growth and seasonal characteristic breed the synchronism. From the beginning of March the testicle start to breed, and it degenerate after May, September to December is convalescent period, after which the sperm happens.
2. Immunohistochemical study on the gonad of Carassius auratus red variety including AR, ER and aromatase.
2.1 Immunohistochemical study on ER in gonadal and sexual cells
The estrogen receptor (ER) is express in each period of development, follicle cells show strong immunopositive reaction to ER, and immunopositive reaction is observed in phase 2 oocyte’s karyon as well as cytoplast, high phase oocyte’s zona radiate, vitellin granule, vacuole around ovary, follicle in the phase 6 and vacuous follicle wall. During the testicle’s development, immunopositive reaction of ER happens in spermatogonium, nucleolus of spermatocyte, inferstitial cells, sertoli cell. Strong expression of ER is found in early spermatogonium and space between folioles. Spermid and sperm assume immunonegative reaction to ER.
2.2 Immunohistochemical study on AR in gonadal and sexual cells
Androgen (AR) is expressed strongly in ovary’s cytoplasm, karyoplasms, follicular membrane; During the vitellus period, AR is expressed in follicular membrane, zona radiate, vitellin granule, and the reaction is weakened by the development of ovary. The vacuole around ovary in the late phase 4 also finds weak immunopositive reaction. AR is strongly expressed in follicle in the phase 6 and vacuous follicle wall. As to testicle, spermatogonium, spermatocyte, spermatocyte, inferstitial cells, sperm and spermid in all phase assume immunonegative reaction to AR. Strong expression of AR is found in the space between folioles. These results indicate ER and AR play an important role in the multiplication and development of germ cell, as well as a proof of the important function of sexual steroid hormone in gonadal growth and matureness.
2.3 Immunohistochemical study on aromatase in gonadal and sexual cells
Strong immunopositive reaction is found in ovary’s development from the phase 2 oocytes to the phase 6 ones. The phase 2 and phase 3 oocytes’cytoplast, follicular membrane; the phase 4 oocytes’nucleus, vitellin granule, zona radiate, follicular membrane and follicle ,vacuous follicle wall in the phase 6 are all found immunopositive reaction. As for testicle, the immunopositive reaction is weakened from the phase 3 to the phase 5, even it assume immunonegative reaction to aromatase in the space between folioles, but strong expression is found in the phase 6. All these results indicate aromatase is important for Carassius auratus red variety’s gonadal growth.
3. Study on the seasonal changes of sexual steroid hormone in blood plasma of Carassius auratus red variety.
3.1 Female Carassius auratus red variety’s 17β-Estradiol and testosterone seasonal changes.
Plasma testosterone(T) and 17β-Estradiol(E2) seasonal changes in female Carassius auratus red variety was detected with radio immunoassay(RIA). The results show that Plasma testosterone and 17β-Estradiol are significantly increased during the reproductive period, 17β-Estradiol’s contents are up to 36.8959±6.8846 pg/ml, testosterone’s contents reach 20.6011±3.6198ng/dl.E2 significantly descend and fall to the annual minimum after November, then the sexual steroid hormone will slowly ascend next year. All these results show the increase of female Carassius auratus red variety’s plasma 17β-Estradiol contents, have satisfied the vitellus’formation and accumulation, even accelerate the ovary’s maturation. The correlativity of testosterone and ovary also reveal they are essential for ovary’s development.
3.2 Male Carassius auratus red variety’s 17β-Estradiol and testosterone seasonal changes.
Studies on the seasonal change of testosterone(T) and 17β-Estradiol(E2) finds that the anniversary change law of testosterone and 17β-Estradiol is consistent with GSI change trend, the peak of the year is reproductive season, in which testosterone’s content is 37.6000±9.0260ng/dl. Plasma 17β-Estradiol content is 11.9846±3.2606 pg/ml, and descend after propagation, then touch the lowest point till the phase 4. All these results indicate testosterone maintain high level to boost sperm’s formation in the reproductive period of Carassius auratus red variety.
4. Study on the anniversary change of serous inorganic phosphorus in Carassius auratus red variety.
Serous inorganic phosphorus change law reflects it’s correlativity to gonadal development and GSI. Significant variation of serous inorganic phosphorus is found among different periods. At reproductive phases, serous inorganic phosphorus contents increase rapidly, male fishes’serous inorganic phosphorus content is 47.4504±14.9158 mmol/L and the female’s is 58.8975±13.6518mmol/L. Significant descendant happens after June, until early reproductive period serous inorganic phosphorus starts to increase. Both female and male fishes reach the peak in reproductive period, which show the phosphorus need when Carassius auratus red variety breeds.
1.红鲫性腺发育周年变化的组织学研究
1.1卵巢的发育分期和卵细胞变化
雌性红鲫具有一对卵巢,对称分布于鳔腹面两侧。非生殖期的卵巢透明,呈粉白色或肉粉色;繁殖期的卵巢体积约占体腔60～70%,呈黄色或橙黄色囊状,可清晰观察到淡黄色的卵粒;退化期的卵巢呈暗红色。组织学观察发现,红鲫卵巢属于卵母细胞发育不同步、分批产卵的类型,卵巢的周年发育分为5个时期,包括Ⅱ期、Ⅲ期、Ⅳ期、Ⅴ期和退化吸收Ⅵ期。卵细胞发育经历5个时相,滤泡膜及卵黄颗粒随着卵细胞的发育而变化。红鲫的繁殖期是在3～5月份,此时卵巢发育都在Ⅳ期到Ⅴ期;6～8月份是排卵后的退化阶段,从9月份开始进入休整期,此时的卵巢处在重复发育Ⅱ期,进入12月份以后,卵巢迅速发育至Ⅲ期,并以Ⅲ期卵巢越冬,从翌年3月份开始,卵巢迅速从Ⅲ期发育至Ⅳ期后,进入产卵高峰期。
1.2精巢的发育分期和精细胞变化
红鲫具有精巢一对,与大多数硬骨鱼类的精巢一样,红鲫的精巢属于小叶型。非繁殖期的精巢为细条状或者宽扁带状;而繁殖期精巢则肥大饱满,呈白色或乳白色。精小叶与精小叶间充满间质细胞、成纤维细胞及丰富的微血管。精子的发生经历精原细胞、初级精母细胞、次级精母细胞、精子细胞以及精子5个典型的发育阶段。从精巢发育的周年变化可以看出,其精巢发育与季节性繁殖活动同步。繁殖季节为3～5月份,6～8月份为退化吸收期,9～11月份为重复发育期,12月份开始启动生精活动。
2.红鲫性腺雌激素受体(AR)、雄激素受体(ER)、芳香化酶的免疫组织化学研究
2.1雌激素受体(ER)在性腺及性细胞中免疫组织化学定位
红鲫卵巢发育的各个时期中雌激素受体(ER)均有表达,滤泡细胞对ER呈强免疫阳性反应,ER在Ⅱ时相卵母细胞核质、胞质和大生长期的放射膜、卵黄颗粒、卵周的液泡以及Ⅵ期末退化吸收的滤泡团及排空的滤泡囊壁均有免疫阳性反应。在精巢发育的各个时期,ER主要定位在精原细胞、初级精母细胞核周、间质细胞、支持细胞,呈免疫阳性反应,早期的精原细胞和精小叶之间的间质处ER表达较强,精子细胞及精子对ER呈免疫阴性反应。
2.2雄激素受体(AR)在性腺及性细胞中免疫组织化学定位
AR在Ⅱ期卵巢的胞质、核质、滤泡膜有较强表达;卵黄发生期,AR则在滤泡膜、放射膜、卵黄颗粒表达,随着卵细胞发育表达逐渐减弱,Ⅳ时相卵母细胞晚期分布在卵周的液泡也有弱阳性反应;Ⅵ期末退化吸收的滤泡团及排空的滤泡囊壁有较强的表达。在精巢,雄激素受体(AR)在各个时期精原细胞、初级精母细胞、次级精母细胞、间质细胞表达,精子细胞和精子显免疫阴性,精小叶间质处对AR的免疫阳性反应相对较强。这些结果表明,雌激素受体、雄激素受体在生殖细胞的增殖和发育过程起作用,为证明性类固醇激素在性腺发育和成熟过程中的重要作用提供证据。
2.3性腺及性细胞中芳香化酶的免疫组织化学
芳香化酶在红鲫卵巢发育的各个时期中都有较强的免疫阳性反应。从Ⅱ时相卵母细胞到Ⅵ时相卵母细胞的卵细胞发育过程中,都对芳香化酶显免疫阳性反应。各期卵巢中Ⅱ、Ⅲ时相卵母细胞的胞质、滤泡膜;Ⅳ时相卵母细胞的胞核、卵黄颗粒、放射膜、滤泡膜以及Ⅵ卵巢中退化分解吸收的卵细胞以及排空的滤泡囊都对芳香化酶呈免疫阳性反应。芳香化酶在红鲫精细胞的表达从精母细胞成熟期(Ⅲ期)至精子成熟期(Ⅴ期)逐渐减弱,且精小叶间质细胞呈免疫阴性反应,但精子退化吸收期(Ⅵ期)精小叶间质处阳性表达较强。这些结果表明,芳香化酶的活性对红鲫性腺发育具有一定的生理效应。
3.红鲫血浆性类固醇激素水平的周年变化研究
3.1雌性红鲫血浆E2和T水平的年周期变化
采用125I标记的放射免疫测定方法(RIA)测定雌性红鲫血清睾酮(T)、雌二醇(E2)含量的季节变动情况。研究发现:繁殖期雌性红鲫血浆的T和E2含量迅速增加,呈显著性变化(P<0.05),E2含量此时高达36.8959±6.8846 pg/ml,T含量达到20.6011±3.6198ng/dl,退化期开始显著下降(P<0.05),至9～11月份进入休整期后T和E2降至全年最低,繁殖前期即12月份至翌年2月份,性激素含量开始缓慢升高。这些结果表明,雌性红鲫繁殖期血浆中雌二醇水平的提高,满足了卵黄形成和积累的需要,促进了卵巢的成熟。T含量与卵巢发育的相关性,提示它们是卵巢发育的一定阶段所必需的。
3.2雄性红鲫血浆E2和T水平的年周期变化
对雄性红鲫血浆睾酮(T)、雌二醇(E2)含量的季节变动研究发现:雄性红鲫血浆T和E2水平的周年变化规律与性腺成熟系数的变化趋势基本一致,全年最高峰值出现在繁殖季节即3～5月份,血浆T含量为37.6000±9.0260ng/dl,血浆E2含量为11.9846±3.2606pg/ml,繁殖后开始显著下降(P<0.05),至静止期降到到最低,从精巢发育Ⅳ期开始T和E2含量升高。这些结果表明,雄性红鲫繁殖期睾酮维持在较高水平,促进精子的发生和形成。
4.红鲫血清无机磷含量的周年变化研究
红鲫血清无机磷的变化规律反映出其与性腺发育密切相关,无论雌性还是雄性,其周年变化规律也都与性腺成熟系数GSI变化规律一致,不同的性腺发育时期,血清无机磷的变化极为明显。繁殖期,红鲫血清无机磷含量增长迅速,呈显著性变化( P < 0.05 ) ,雌性和雄性红鲫血清无机磷含量分别为58.8975±13.6518mmol/L和47.4504±14.9158 mmol/L,6～11月份显著下降,直至繁殖前期血清无机磷含量开始缓慢升高。雌鱼和雄鱼繁殖期血清无机磷含量都为全年的最高峰,能够说明红鲫繁殖时对无机磷的需要量比较高。
Carassius auratus red variety, a variation of Crucian carp, which scatters broadly in China, is common minitype of fresh water fish in Yangtse Valley. This paper takes the artificially bred Carassius auratus red variety as study objects. Gonad, sexual steroid hormone of blood plasma and their receptors, content of serous inorganic phosphorus are studied by the methods of histological staining, immunohistochemical techniques , radioimmunoassay and biochemical methods in details. The main results and conclusions are as follows:
1. Histological study on the development and annual change in the gonad of Carassius auratus red variety
1.1 Stages and oocyte’s change during the ovary’s growth
Carassius auratus red variety has a pair of ovary, scatter symmetrically at two ventral sides of the swim bladder. The color of immature ovary is white or flesh pink; the mature ovary which looks like a yellow or the orange vesicle takes up 60 ~ 70% of body-cavity, and ovum granule can be clearly observed;the degenerate ovary assumes dark red. Histological research finds that the oocyte of Carassius auratus red variety develops out of sync and spawns in batches. The development of ovary can be divided into five stages and the oocytes be divided into five phases: phase2, phase3, phase4, phase5 and phase 6 when the oocytes degenerate. Follicle and vitellin granule are varying with the development of oocytes. The reproductive period of Carassius auratus red variety is between March and May, in which the ovaries are at phase 4 to phase 5; The ovaries degenerate after ovulation from June to August, and have a rest after September from which the ovaries are at phase 2. From the beginning of December , the ovaries live through the winter with the phase 3. From the third month of next year, the ovaries come to the peak of oviposition and went into the phase 4 immediately.
1.2 Stages and the spermatid change during the testicle’s growth
Like other teleost fishes, Carassius auratus red variety has a pair of testicles. Carassius auratus red variety’s testicle is lobular type in appearance. The immature testicle b looks like pinstriped shape or flat tape shape while the mature one is big and assume white color. Be skill and accurate, foliole is full of the inferstitial cells, mechanocyte and lots of blood capillaries. Sperm experience 5 steps during its life: spermatogonium, spermatocyte, spermatocyteⅡ,spoermatid, sperm. The anniversary change from the testicle is it can be seen, testicle growth and seasonal characteristic breed the synchronism. From the beginning of March the testicle start to breed, and it degenerate after May, September to December is convalescent period, after which the sperm happens.
2. Immunohistochemical study on the gonad of Carassius auratus red variety including AR, ER and aromatase.
2.1 Immunohistochemical study on ER in gonadal and sexual cells
The estrogen receptor (ER) is express in each period of development, follicle cells show strong immunopositive reaction to ER, and immunopositive reaction is observed in phase 2 oocyte’s karyon as well as cytoplast, high phase oocyte’s zona radiate, vitellin granule, vacuole around ovary, follicle in the phase 6 and vacuous follicle wall. During the testicle’s development, immunopositive reaction of ER happens in spermatogonium, nucleolus of spermatocyte, inferstitial cells, sertoli cell. Strong expression of ER is found in early spermatogonium and space between folioles. Spermid and sperm assume immunonegative reaction to ER.
2.2 Immunohistochemical study on AR in gonadal and sexual cells
Androgen (AR) is expressed strongly in ovary’s cytoplasm, karyoplasms, follicular membrane; During the vitellus period, AR is expressed in follicular membrane, zona radiate, vitellin granule, and the reaction is weakened by the development of ovary. The vacuole around ovary in the late phase 4 also finds weak immunopositive reaction. AR is strongly expressed in follicle in the phase 6 and vacuous follicle wall. As to testicle, spermatogonium, spermatocyte, spermatocyte, inferstitial cells, sperm and spermid in all phase assume immunonegative reaction to AR. Strong expression of AR is found in the space between folioles. These results indicate ER and AR play an important role in the multiplication and development of germ cell, as well as a proof of the important function of sexual steroid hormone in gonadal growth and matureness.
2.3 Immunohistochemical study on aromatase in gonadal and sexual cells
Strong immunopositive reaction is found in ovary’s development from the phase 2 oocytes to the phase 6 ones. The phase 2 and phase 3 oocytes’cytoplast, follicular membrane; the phase 4 oocytes’nucleus, vitellin granule, zona radiate, follicular membrane and follicle ,vacuous follicle wall in the phase 6 are all found immunopositive reaction. As for testicle, the immunopositive reaction is weakened from the phase 3 to the phase 5, even it assume immunonegative reaction to aromatase in the space between folioles, but strong expression is found in the phase 6. All these results indicate aromatase is important for Carassius auratus red variety’s gonadal growth.
3. Study on the seasonal changes of sexual steroid hormone in blood plasma of Carassius auratus red variety.
3.1 Female Carassius auratus red variety’s 17β-Estradiol and testosterone seasonal changes.
Plasma testosterone(T) and 17β-Estradiol(E2) seasonal changes in female Carassius auratus red variety was detected with radio immunoassay(RIA). The results show that Plasma testosterone and 17β-Estradiol are significantly increased during the reproductive period, 17β-Estradiol’s contents are up to 36.8959±6.8846 pg/ml, testosterone’s contents reach 20.6011±3.6198ng/dl.E2 significantly descend and fall to the annual minimum after November, then the sexual steroid hormone will slowly ascend next year. All these results show the increase of female Carassius auratus red variety’s plasma 17β-Estradiol contents, have satisfied the vitellus’formation and accumulation, even accelerate the ovary’s maturation. The correlativity of testosterone and ovary also reveal they are essential for ovary’s development.
3.2 Male Carassius auratus red variety’s 17β-Estradiol and testosterone seasonal changes.
Studies on the seasonal change of testosterone(T) and 17β-Estradiol(E2) finds that the anniversary change law of testosterone and 17β-Estradiol is consistent with GSI change trend, the peak of the year is reproductive season, in which testosterone’s content is 37.6000±9.0260ng/dl. Plasma 17β-Estradiol content is 11.9846±3.2606 pg/ml, and descend after propagation, then touch the lowest point till the phase 4. All these results indicate testosterone maintain high level to boost sperm’s formation in the reproductive period of Carassius auratus red variety.
4. Study on the anniversary change of serous inorganic phosphorus in Carassius auratus red variety.
Serous inorganic phosphorus change law reflects it’s correlativity to gonadal development and GSI. Significant variation of serous inorganic phosphorus is found among different periods. At reproductive phases, serous inorganic phosphorus contents increase rapidly, male fishes’serous inorganic phosphorus content is 47.4504±14.9158 mmol/L and the female’s is 58.8975±13.6518mmol/L. Significant descendant happens after June, until early reproductive period serous inorganic phosphorus starts to increase. Both female and male fishes reach the peak in reproductive period, which show the phosphorus need when Carassius auratus red variety breeds.
引文
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