梭鲈胚胎、仔鱼发育观察与细胞遗传学研究
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摘要
梭鲈(Sander lucioperca(Linnaeus))具有适温广、生长快、耐盐碱、抗病、肉味鲜美的优良品质,在国际水产品贸易市场非常受欢迎,是一个有着良好市场前景的水产养殖名特优新品。本研究旨在通过开展梭鲈胚胎发育和胚后发育的过程进行系统的观察,了解梭鲈胚胎及仔鱼的生物学特性,以提高人工繁殖的孵化率及苗种质量,以及今后开展多倍体育种提供理论基础;通过对梭鲈染色体核型及显带的研究为该鱼的种质资源研究、种群鉴定以及染色体原位杂交开发奠定基础,为进一步开展细胞遗传学研究及构建基因组物理图谱的精细框架提供基础依据。主要结论包括:
1.采用连续观察法通过显微观察和摄影,对梭鲈(Sander lucioperca(Linnaeus))的胚胎、仔鱼发育进行了观察和描述。结果表明:梭鲈成熟卵呈圆球状,呈淡黄色,卵径平均为0.96 mm (0.90～1.01 mm)、黏性。受精后1～2 min,吸水膨胀形成卵周隙,吸水后平均卵径为1.2 mm,卵质分布较均匀。受精卵在水温16.0～17.2℃的条件下,历时94 h完成胚胎发育过程,从受精到孵化出膜总积温为1616.8℃·h。整个胚胎发育过程分为24个期,包括受精卵、
2细胞期、4细胞期、8细胞期、16细胞期、32细胞期、64细胞期、多细胞期、囊胚早期、囊胚中期、低囊胚期、囊胚晚期、原肠早期、原肠中期、原肠晚期、神经胚期、胚孔封闭期、肌节出现期、眼基期、眼囊期、尾鳍出现期、晶体出现期、肌肉效应期、耳石期、心跳期、出膜期;依公式K=N (T-C)对不同时间和地点梭鲈胚胎发育的平均温度和孵出时间进行线性关系统计分析,得出胚胎发育速度和水温度显著相关。并计算出梭鲈受精卵发育的理论起始温度为7.55℃,发育所需的理论有效积温为903.02℃·h。梭鲈仔鱼期可分为5个时期,分别为出膜仔鱼、眼黄色素期、眼黑色素期、鳔一室期、卵黄吸尽期。2.采用RPMI l640全培养基进行梭鲈淋巴细胞体外培养,通过对培养基PHA浓度、秋水仙素浓度及低渗时间条件的摸索,建立了效果较好的培养梭鲈血淋巴细胞制备染色体标本的实验方法。结果表明:每0.2ml梭鲈外周血细胞在5mlRPMI1640培养基中(RPMI 1640体积分数80%,PHA浓度为4 ug/ml小牛血清20%,青霉素、链霉素、卡那霉素各100 IU/ml),25℃培养72 h,培养终止前3~5h加入终浓度为0.05 ug/ml左右的秋水仙素,低渗50 min后固定、滴片及染色,可以制备出分裂相较好的染色体玻片标本。
3.本试验以梭鲈外周血细胞为材料,采用PHA和秋水仙素体外培养法,经空气干燥制备了染色体并对其染色体核型进行分析。结果表明:梭鲈染色体数目为48条,包括2条中部着丝点染色体、10条亚中部着丝点染色体、12条亚端部着丝粒染色体和24条端部着丝粒染色体,未发现异型染色体对和随体。梭鲈核型公式为2n=48,2m+10sm+12st+24t,其染色体总臂数(NF)为60。
4.通过对梭鲈染色体Ag-NORs显带及C-显带的初步研究获得了Ag-NORs显带、C-显带中期染色体分裂相图谱,并进行了核型分析。结果表明:梭鲈中期分裂相中, Ag-NORs出现的位置均在亚中部着丝粒染色体短臂末端,Ag-NORs的数目在不同的细胞中表现出不同的多态性,其数目为1～2个。在间期核中,通过银染表现出1个核仁的间期核数目的频率为53.99%;2个核仁的间期核数目为45.62%,大多数间期核Ag-NoRs为圆形点状。核型分析结果表明,梭鲈在第三对亚中部着丝点染色体的短臂端部区域出现NOR,为端部Ag-NORs。梭鲈的48条染色体多数有大小不一的C-带深染,同源染色体C带的大小、位置以及着色强度基本相同,不同染色体的C带强度有一定的差异。C带显示梭鲈所有染色体上都存在阳性带,主要位置为着丝粒区、中间区和末端区。
Pike-perch (Sander lucioperca(Linnaeus)) with wide-temperature resistant, fast growth, excellent resistance to salts, disease resistance, have become popular in International aquatic products trade market for their deliciousness, high values of nutrition and meat quality. The embryonic and larval development of pike-perch was observed, in order to know its biological characteristics. The result will be used in increasing its hatchability and fry quality. The karyotype analysis and chromosome banding of pike-perch were studied for its genetic diversity, population identification and FISH. It will provide basis for Cyrogenetics and genomic physical map. The main results are as follows:
1. A successive observation was conducted to study the embryonic larval and juvenile development of pike-perch. The results showed that the eggs, measuring 0.90-1.01mm in diameter, were sticky and almost spherical in shape, with a pale yellow yolk and 1.2mm after absorbing water. Hatching occurred 94h after fertilization at 16.0-17.2℃. The embryonic development can be divided into 24 stages: fertilized egg , one cell stage, cleavage stage, 2-cell stage, 4-cell stag, 8-cell stage, 16-cell stage, morula stage, early blastula stage, mid blastula stage, late blastula stage, early gastrula stage, mid gastrula stage, late gastrula stage, neurula stage, closure of blastopore stage, body segment appearance stage, optic rudiment stage, optic vesicle stage, caudal fin formation stage, lens formation stage, muscular contraction stage, otolithes formation stage, heart pulsation stage, hatching-out stage. According to the formula K=N (T-C) , the average temperature and hatch time were analysis with statistics. Speed of embryonic development and water temperature was very closely related. It was calculated that the newly hatched larvae’s lowest temperature is about 7.55℃, and the sum calculated temperature 903.02℃·h is necessary. The larval development of pike-perch can be divided into 5 stages: hatching stage, xanthic eye stage, melanoid eye stage, one chamber air bladder stage, exhaustion of yolk stage.
2. Use the whole culture medium of RPMI1640 to culture the blood cell in vitro. With researching the content of colchicines and permeating time, establish a set of mature experimental method to cultivate the blood lymphocytic cell and prepare chromosome of pike-perch. The result showed that the blood cell 0.2ml was cultivated in culture medium 5ml of RPMI1640 (RPMI 1640 : FBS=4:1; PHA concentration is 4 ug/ml; Penicillin, streptomycin, kanamycin each concentration is 100 IU/ml ) under a temperature of 25℃for 72h. Colchicines were added until it reached concentration of about 0.2ug/ml working for 3-5h before the end of cell cultivation. And the lymphocytes were permeated for 50min with KCl. The good chromosome specimens were prepared by the routine methods of hypotonicity, fixing and coloration.
3. Metaphase chromosome specimens of pike-perch were prepared from short-term culture of peripheral blood lymphocytes with air-drying technique. It diploid chromosome number is 2n=48. The karyotype is compose of 2m, 10sm, 10st,12t, NF=60. The C-banding, NOR banding distribution were analyzed in chromosomes of pike-perch.The chromosomes of pike-perch were stained according to the method of Howell (1980) that was slightly modified. The result showed that 2 Ag-NORs bands were observed in these 2 species in their metaphases respectively. The position of nucleolus organizer regions of pike-perch was situated on the terminal region of short arm of the subteloeentrie chromosomes. The number of Ag-NORs showed polymorphism in different cells.The nucleus were stained by the Ag-staining, 53.99% of then have one nucleolus and 45.62% of nucleus have two nucleoli. All the chromosomes of pike-perch have C-bands. The C-bands of homologous chromosome are approximate the same in size, location and intensity of colouring. C-banding of pike-perch mainly distributed at centromeres, interstitial and terminal regions.
1.采用连续观察法通过显微观察和摄影,对梭鲈(Sander lucioperca(Linnaeus))的胚胎、仔鱼发育进行了观察和描述。结果表明:梭鲈成熟卵呈圆球状,呈淡黄色,卵径平均为0.96 mm (0.90～1.01 mm)、黏性。受精后1～2 min,吸水膨胀形成卵周隙,吸水后平均卵径为1.2 mm,卵质分布较均匀。受精卵在水温16.0～17.2℃的条件下,历时94 h完成胚胎发育过程,从受精到孵化出膜总积温为1616.8℃·h。整个胚胎发育过程分为24个期,包括受精卵、
2细胞期、4细胞期、8细胞期、16细胞期、32细胞期、64细胞期、多细胞期、囊胚早期、囊胚中期、低囊胚期、囊胚晚期、原肠早期、原肠中期、原肠晚期、神经胚期、胚孔封闭期、肌节出现期、眼基期、眼囊期、尾鳍出现期、晶体出现期、肌肉效应期、耳石期、心跳期、出膜期;依公式K=N (T-C)对不同时间和地点梭鲈胚胎发育的平均温度和孵出时间进行线性关系统计分析,得出胚胎发育速度和水温度显著相关。并计算出梭鲈受精卵发育的理论起始温度为7.55℃,发育所需的理论有效积温为903.02℃·h。梭鲈仔鱼期可分为5个时期,分别为出膜仔鱼、眼黄色素期、眼黑色素期、鳔一室期、卵黄吸尽期。2.采用RPMI l640全培养基进行梭鲈淋巴细胞体外培养,通过对培养基PHA浓度、秋水仙素浓度及低渗时间条件的摸索,建立了效果较好的培养梭鲈血淋巴细胞制备染色体标本的实验方法。结果表明:每0.2ml梭鲈外周血细胞在5mlRPMI1640培养基中(RPMI 1640体积分数80%,PHA浓度为4 ug/ml小牛血清20%,青霉素、链霉素、卡那霉素各100 IU/ml),25℃培养72 h,培养终止前3~5h加入终浓度为0.05 ug/ml左右的秋水仙素,低渗50 min后固定、滴片及染色,可以制备出分裂相较好的染色体玻片标本。
3.本试验以梭鲈外周血细胞为材料,采用PHA和秋水仙素体外培养法,经空气干燥制备了染色体并对其染色体核型进行分析。结果表明:梭鲈染色体数目为48条,包括2条中部着丝点染色体、10条亚中部着丝点染色体、12条亚端部着丝粒染色体和24条端部着丝粒染色体,未发现异型染色体对和随体。梭鲈核型公式为2n=48,2m+10sm+12st+24t,其染色体总臂数(NF)为60。
4.通过对梭鲈染色体Ag-NORs显带及C-显带的初步研究获得了Ag-NORs显带、C-显带中期染色体分裂相图谱,并进行了核型分析。结果表明:梭鲈中期分裂相中, Ag-NORs出现的位置均在亚中部着丝粒染色体短臂末端,Ag-NORs的数目在不同的细胞中表现出不同的多态性,其数目为1～2个。在间期核中,通过银染表现出1个核仁的间期核数目的频率为53.99%;2个核仁的间期核数目为45.62%,大多数间期核Ag-NoRs为圆形点状。核型分析结果表明,梭鲈在第三对亚中部着丝点染色体的短臂端部区域出现NOR,为端部Ag-NORs。梭鲈的48条染色体多数有大小不一的C-带深染,同源染色体C带的大小、位置以及着色强度基本相同,不同染色体的C带强度有一定的差异。C带显示梭鲈所有染色体上都存在阳性带,主要位置为着丝粒区、中间区和末端区。
Pike-perch (Sander lucioperca(Linnaeus)) with wide-temperature resistant, fast growth, excellent resistance to salts, disease resistance, have become popular in International aquatic products trade market for their deliciousness, high values of nutrition and meat quality. The embryonic and larval development of pike-perch was observed, in order to know its biological characteristics. The result will be used in increasing its hatchability and fry quality. The karyotype analysis and chromosome banding of pike-perch were studied for its genetic diversity, population identification and FISH. It will provide basis for Cyrogenetics and genomic physical map. The main results are as follows:
1. A successive observation was conducted to study the embryonic larval and juvenile development of pike-perch. The results showed that the eggs, measuring 0.90-1.01mm in diameter, were sticky and almost spherical in shape, with a pale yellow yolk and 1.2mm after absorbing water. Hatching occurred 94h after fertilization at 16.0-17.2℃. The embryonic development can be divided into 24 stages: fertilized egg , one cell stage, cleavage stage, 2-cell stage, 4-cell stag, 8-cell stage, 16-cell stage, morula stage, early blastula stage, mid blastula stage, late blastula stage, early gastrula stage, mid gastrula stage, late gastrula stage, neurula stage, closure of blastopore stage, body segment appearance stage, optic rudiment stage, optic vesicle stage, caudal fin formation stage, lens formation stage, muscular contraction stage, otolithes formation stage, heart pulsation stage, hatching-out stage. According to the formula K=N (T-C) , the average temperature and hatch time were analysis with statistics. Speed of embryonic development and water temperature was very closely related. It was calculated that the newly hatched larvae’s lowest temperature is about 7.55℃, and the sum calculated temperature 903.02℃·h is necessary. The larval development of pike-perch can be divided into 5 stages: hatching stage, xanthic eye stage, melanoid eye stage, one chamber air bladder stage, exhaustion of yolk stage.
2. Use the whole culture medium of RPMI1640 to culture the blood cell in vitro. With researching the content of colchicines and permeating time, establish a set of mature experimental method to cultivate the blood lymphocytic cell and prepare chromosome of pike-perch. The result showed that the blood cell 0.2ml was cultivated in culture medium 5ml of RPMI1640 (RPMI 1640 : FBS=4:1; PHA concentration is 4 ug/ml; Penicillin, streptomycin, kanamycin each concentration is 100 IU/ml ) under a temperature of 25℃for 72h. Colchicines were added until it reached concentration of about 0.2ug/ml working for 3-5h before the end of cell cultivation. And the lymphocytes were permeated for 50min with KCl. The good chromosome specimens were prepared by the routine methods of hypotonicity, fixing and coloration.
3. Metaphase chromosome specimens of pike-perch were prepared from short-term culture of peripheral blood lymphocytes with air-drying technique. It diploid chromosome number is 2n=48. The karyotype is compose of 2m, 10sm, 10st,12t, NF=60. The C-banding, NOR banding distribution were analyzed in chromosomes of pike-perch.The chromosomes of pike-perch were stained according to the method of Howell (1980) that was slightly modified. The result showed that 2 Ag-NORs bands were observed in these 2 species in their metaphases respectively. The position of nucleolus organizer regions of pike-perch was situated on the terminal region of short arm of the subteloeentrie chromosomes. The number of Ag-NORs showed polymorphism in different cells.The nucleus were stained by the Ag-staining, 53.99% of then have one nucleolus and 45.62% of nucleus have two nucleoli. All the chromosomes of pike-perch have C-bands. The C-bands of homologous chromosome are approximate the same in size, location and intensity of colouring. C-banding of pike-perch mainly distributed at centromeres, interstitial and terminal regions.
引文
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