星鲽染色体核型及带型的初步研究
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摘要
星鲽属鱼类包括条斑星鲽和圆斑星鲽两个种类,为冷温性大型底栖鲽类。圆斑星鲽在我国主要分布于黄渤海和东海,但多年以来,由于种种原因,圆斑星鲽自然资源衰退,捕获量极少,形不成一定的渔获量。条斑星鲽在我国黄渤海区曾有分布记录,现已多年未见渔获,接近濒危程度;星鲽具有生长速度快、抗逆能力强、耐低温等优良特性,外观形体较大,肉质细嫩鲜美,鳍边胶质厚而有弹性,营养丰富,易于烹饪,深受消费者喜爱。我国于上世纪90年代开始采捕野生圆斑星鲽亲鱼进行驯养,并逐步开展了人工育苗试验,到目前已能达到规模化繁育苗种;对条斑星鲽的研究始于2004年从日本引进条斑星鲽苗种,随后进行了养殖试验和开发,现也达到了规模化苗种繁育。星鲽属于名贵海水鱼类,具有广阔的市场前景。本文对条斑星鲽和圆斑星鲽的核型及带型进行了初步的研究和分析,并对其核型特征和演化关系作了初步探讨,比较了两者亲缘关系的远近,为其渔业资源的合理利用与保护提供参考依据,为星鲽鱼类的杂交育种、增养殖和种子工程的实施提供遗传背景资料和科学依据,对海洋鱼类多样性的保护也具有重要意义。
1.采用体内注射植物血细胞凝集素(PHA)和秋水仙素法制备星鲽染色体。实验鱼头肾细胞经体内短期培养,取出体外经低渗处理、卡诺氏固定液固定后,常规空气干燥法制作染色体标本。Giemsa染液染色后获得中期分裂相进行核型分析。结果表明,条斑星鲽和圆斑星鲽染色体数目均为2n=46,核型分别为条斑星鲽:2n=2sm+44t,NF=48;圆斑星鲽:2n=46t,NF=46。在星鲽染色体中尚未发现有多倍体的现象,也未发现异型性染色体和随体染色体。两者的核型显示其在鱼类系统上属于高位类,是比较原始的类群。条斑星鲽可能比圆斑星鲽较为特化。
2.采用Sumner(1972)的方法稍作修改后进行了星鲽染色体C带的研究。结果表明,条斑星鲽染色体显示较为复杂的C-带,有4对整条呈阳性深染,分别为3号、13号、20号和22号染色体;有3对具居间C-带,分别为1号、4号和8号;9号染色体整条为阴性,其余有大小不一的着丝粒C-带。计算其异染色质含量为54.91%。圆斑星鲽的46条染色体均有大小不一的C-带,其中第22对染色体整条呈阳性深染,第19对染色体具有端部C-带,其余均为着丝粒C-带,圆斑星鲽无居间C-带。计算得出圆斑星鲽染色体异染色质含量为30%,少于条斑星鲽的54.91%。从C-带带型看条斑星鲽和圆斑星鲽的进化关系,条斑星鲽似乎比圆斑星鲽较为进化,这与其核型特征是相一致的。
3.按Howell(1980)快速银染法进行了星鲽染色体Ag-NORs带型的研究。结果表明,条斑星鲽和圆斑星鲽均显示1对NOR,条斑星鲽的Ag-NORs位于亚中部着丝点染色体的短臂上,为端部Ag-NOR。圆斑星鲽的Ag-NORs位于2号染色体的长臂末端,也为端部Ag-NOR。条斑星鲽亚中部染色体按相对长度也属于2号染色体,这很可能说明条斑星鲽和圆斑星鲽的2号染色体具有很高的同源性,可能是经罗伯逊易位形成的,显示了两者之间较高的亲缘关系。1对端部NORs表明其核型似乎也是比较原始的,这与核型研究的结果相同。
4.采用胰酶法显带进行了星鲽染色体G-带的研究。结果表明条斑星鲽和圆斑星鲽具有差异较大的G-带带纹,但总的带纹数目相同,均为59条带。其中条斑星鲽有37条深带,22条浅带;圆斑星鲽有38条深带,21条浅带。G-带带型的研究由于受染色体收缩程度影响较大,本文得到的结果并不理想,也可能是因为鱼类染色体数目多、形态小,且与它们的染色体DNA组成有关。
5.利用谭远德、吴昌谋提出的核型似近系数和核型进化距离公式计算了星鲽的核型似近系数(λ)和核型进化距离( De )。通过计算得出λ=0.98, De =0.02,表明条斑星鲽和圆斑星鲽核型差异很小,具有很近的亲缘关系。
Verasper sp. includes two species flatfish Verasper moseri and Verasper variega tus which belong to cold-temperate macrobenthos. In china,Verasper variegatus are mainly distributed in the Yellow Sea and Bohai Sea and East China Sea,but over the years for various reasons,the natural resources of Verasper variegatus are declining,the catches are extremely few,which cannot form certain catching. Verasper moseri once had the distribution records in China’s Yellow Sea and Bohai Sea Area,but the fishing was not seen to attain for many years,closing to the extent of endangered. Verasper sp. with faster growth、high stress-resistance and excellent low-temperature -resistant characteristics,have become popular for their deliciousness,high values of nutrition and meat quality.
At the beginning of the last century 90's Chinese started to catch the wild Vera- sper variegatus for domestication,the experiments of artificial breeding had started progressively. To the present it has been able to achieve large-scale breedings. The stu -dies to Verasper moseri began in 2004 with the introduction of seedlings from Japan,followed by experiments and developments of aquaculture. The breeding of Verasper moseri have reached large-scales now. Verasper sp. belong to valuable marine fish which have broad market prospects.
In this paper,the chromosomal conventional karyotypes and banding patterns on Verasper sp. were studied. The author has made the preliminary discussion to their kar yotypes and the evolved relations,compared the relationship between the two speci- es,which could provide the reference for reasonable use and protect of their fishery resources,and the genetic background information and scientific basis for the flound- er fish’s crossbreeding and culturing. It also has the important meanings for marine fi- shes diversity protection.
1. The karyotype of Verasper sp. was studied through the chromosome preparation obtained from head kidney by the method of injecting PHA and colchicines. After low permeating and Carnoy′s fixation,air drying method was used to make chromosome samples,which were dyed by Giemsa afterwards. The results showed that the diploid chromosomal number of V. moseri and V. variegatus is 2n=46 and their respective kar- yotype formulae are 2n=2sm+44t,NF=48 and 2n=46t,NF=46. It means that there are
1 pair of submetacentric chromosomes and 22 pairs of telocentric chromosomes in V. moseri and 23 pairs of telocentric chromosomes in V. variegatus. There are no visible evidence of polyploidy chromosome and sex chromosome. According to the karyotyp e of the 2 species,Verasper sp. belong to height classification in system classify and they are the more primitive groups. V. moseri may be more evolutional than V. variega tus.
2. Using the methods of Sumner (1972) slightly modified,V. moseri shows more complicated C-bands. There are 4 pairs of chromosomes which have the whole chromosomes C-bands,their number is 3、13、20、22;the number of 1、4 and 8 chromosomes have interstitial C-bands;the No.9 chromosome don’t have C-bands,and the rest of the chromosomes have the centromere C-bands. The heterochromatin content of V. moseri is 54.91%. C-bands of V. variegates are located mainly in the centromere regions of chromosomes except the No.19 and 22 chromosomes,the No.19 chromosome of V. variegates have terminal C-bands and the No.22 chromosome of V. variegates have the whole chromosomes C-bands. The heterochromatin content of V. variegates is 30% which less than V. moseri. The characteristics of the number and sites of C-bands in Verasper sp. chromosomes reveal that V. moseri is more evolutional than V. variegates which is consistent to their karyotype.
3. The chromosomes of Verasper sp. were thus stained according to the method of Howell (1980) that was slightly modified. The study showed that 2 Ag-NORs bands were observed in these 2 species in their metaphases respectively. The position of nucleolus organizer regions of V. moseri was situated on the terminal region of short arm of the submetacentric chromosomes. The position of nucleolus organizer regions of V. variegates was situated on the terminal region of No.2 chromosomes. This may explain that No.2 chromosomes of V. moseri and V. variegates are highly homologous because the submetacentric chromosomes of V. moseri also belong to No.2 chromosomes according to the relative length of it’s chromosomes. It shows that the 2 species of the Verasper sp. have very close relationship. One pair of telomeric NORs reveal that Verasper sp. are the more primitive groups.
4. The G-banding was carried out by the trypsin method. The study showed that V. moseri and V. variegates have more difference in G-bands of chromosomes,but they have the same total numbers of G-bands which are 59. V. moseri have 37 deeply stained bands and 22 shallow stained bands,while V. variegates have 38 deeply stained bands and 21 shallow stained bands,but the region of G-bands in Verasper sp. chromosomes are different. G-banding is susceptible to interference by variations such as the degree of chromosome contraction,so the results in this paper are not satisfactory. It also may be due to the larger number and the tiny shape of fish chromosome and their DNA composition of chromosomes.
5. Calculated the karyotype resemblance near coefficients (λ) and the distance of karyotype evolution ( De ) of Verasper sp.,the resultλ=0.98, De =0.02 indicate that the difference between V. moseri and V. variegates are very little,they have very close relationship.
1.采用体内注射植物血细胞凝集素(PHA)和秋水仙素法制备星鲽染色体。实验鱼头肾细胞经体内短期培养,取出体外经低渗处理、卡诺氏固定液固定后,常规空气干燥法制作染色体标本。Giemsa染液染色后获得中期分裂相进行核型分析。结果表明,条斑星鲽和圆斑星鲽染色体数目均为2n=46,核型分别为条斑星鲽:2n=2sm+44t,NF=48;圆斑星鲽:2n=46t,NF=46。在星鲽染色体中尚未发现有多倍体的现象,也未发现异型性染色体和随体染色体。两者的核型显示其在鱼类系统上属于高位类,是比较原始的类群。条斑星鲽可能比圆斑星鲽较为特化。
2.采用Sumner(1972)的方法稍作修改后进行了星鲽染色体C带的研究。结果表明,条斑星鲽染色体显示较为复杂的C-带,有4对整条呈阳性深染,分别为3号、13号、20号和22号染色体;有3对具居间C-带,分别为1号、4号和8号;9号染色体整条为阴性,其余有大小不一的着丝粒C-带。计算其异染色质含量为54.91%。圆斑星鲽的46条染色体均有大小不一的C-带,其中第22对染色体整条呈阳性深染,第19对染色体具有端部C-带,其余均为着丝粒C-带,圆斑星鲽无居间C-带。计算得出圆斑星鲽染色体异染色质含量为30%,少于条斑星鲽的54.91%。从C-带带型看条斑星鲽和圆斑星鲽的进化关系,条斑星鲽似乎比圆斑星鲽较为进化,这与其核型特征是相一致的。
3.按Howell(1980)快速银染法进行了星鲽染色体Ag-NORs带型的研究。结果表明,条斑星鲽和圆斑星鲽均显示1对NOR,条斑星鲽的Ag-NORs位于亚中部着丝点染色体的短臂上,为端部Ag-NOR。圆斑星鲽的Ag-NORs位于2号染色体的长臂末端,也为端部Ag-NOR。条斑星鲽亚中部染色体按相对长度也属于2号染色体,这很可能说明条斑星鲽和圆斑星鲽的2号染色体具有很高的同源性,可能是经罗伯逊易位形成的,显示了两者之间较高的亲缘关系。1对端部NORs表明其核型似乎也是比较原始的,这与核型研究的结果相同。
4.采用胰酶法显带进行了星鲽染色体G-带的研究。结果表明条斑星鲽和圆斑星鲽具有差异较大的G-带带纹,但总的带纹数目相同,均为59条带。其中条斑星鲽有37条深带,22条浅带;圆斑星鲽有38条深带,21条浅带。G-带带型的研究由于受染色体收缩程度影响较大,本文得到的结果并不理想,也可能是因为鱼类染色体数目多、形态小,且与它们的染色体DNA组成有关。
5.利用谭远德、吴昌谋提出的核型似近系数和核型进化距离公式计算了星鲽的核型似近系数(λ)和核型进化距离( De )。通过计算得出λ=0.98, De =0.02,表明条斑星鲽和圆斑星鲽核型差异很小,具有很近的亲缘关系。
Verasper sp. includes two species flatfish Verasper moseri and Verasper variega tus which belong to cold-temperate macrobenthos. In china,Verasper variegatus are mainly distributed in the Yellow Sea and Bohai Sea and East China Sea,but over the years for various reasons,the natural resources of Verasper variegatus are declining,the catches are extremely few,which cannot form certain catching. Verasper moseri once had the distribution records in China’s Yellow Sea and Bohai Sea Area,but the fishing was not seen to attain for many years,closing to the extent of endangered. Verasper sp. with faster growth、high stress-resistance and excellent low-temperature -resistant characteristics,have become popular for their deliciousness,high values of nutrition and meat quality.
At the beginning of the last century 90's Chinese started to catch the wild Vera- sper variegatus for domestication,the experiments of artificial breeding had started progressively. To the present it has been able to achieve large-scale breedings. The stu -dies to Verasper moseri began in 2004 with the introduction of seedlings from Japan,followed by experiments and developments of aquaculture. The breeding of Verasper moseri have reached large-scales now. Verasper sp. belong to valuable marine fish which have broad market prospects.
In this paper,the chromosomal conventional karyotypes and banding patterns on Verasper sp. were studied. The author has made the preliminary discussion to their kar yotypes and the evolved relations,compared the relationship between the two speci- es,which could provide the reference for reasonable use and protect of their fishery resources,and the genetic background information and scientific basis for the flound- er fish’s crossbreeding and culturing. It also has the important meanings for marine fi- shes diversity protection.
1. The karyotype of Verasper sp. was studied through the chromosome preparation obtained from head kidney by the method of injecting PHA and colchicines. After low permeating and Carnoy′s fixation,air drying method was used to make chromosome samples,which were dyed by Giemsa afterwards. The results showed that the diploid chromosomal number of V. moseri and V. variegatus is 2n=46 and their respective kar- yotype formulae are 2n=2sm+44t,NF=48 and 2n=46t,NF=46. It means that there are
1 pair of submetacentric chromosomes and 22 pairs of telocentric chromosomes in V. moseri and 23 pairs of telocentric chromosomes in V. variegatus. There are no visible evidence of polyploidy chromosome and sex chromosome. According to the karyotyp e of the 2 species,Verasper sp. belong to height classification in system classify and they are the more primitive groups. V. moseri may be more evolutional than V. variega tus.
2. Using the methods of Sumner (1972) slightly modified,V. moseri shows more complicated C-bands. There are 4 pairs of chromosomes which have the whole chromosomes C-bands,their number is 3、13、20、22;the number of 1、4 and 8 chromosomes have interstitial C-bands;the No.9 chromosome don’t have C-bands,and the rest of the chromosomes have the centromere C-bands. The heterochromatin content of V. moseri is 54.91%. C-bands of V. variegates are located mainly in the centromere regions of chromosomes except the No.19 and 22 chromosomes,the No.19 chromosome of V. variegates have terminal C-bands and the No.22 chromosome of V. variegates have the whole chromosomes C-bands. The heterochromatin content of V. variegates is 30% which less than V. moseri. The characteristics of the number and sites of C-bands in Verasper sp. chromosomes reveal that V. moseri is more evolutional than V. variegates which is consistent to their karyotype.
3. The chromosomes of Verasper sp. were thus stained according to the method of Howell (1980) that was slightly modified. The study showed that 2 Ag-NORs bands were observed in these 2 species in their metaphases respectively. The position of nucleolus organizer regions of V. moseri was situated on the terminal region of short arm of the submetacentric chromosomes. The position of nucleolus organizer regions of V. variegates was situated on the terminal region of No.2 chromosomes. This may explain that No.2 chromosomes of V. moseri and V. variegates are highly homologous because the submetacentric chromosomes of V. moseri also belong to No.2 chromosomes according to the relative length of it’s chromosomes. It shows that the 2 species of the Verasper sp. have very close relationship. One pair of telomeric NORs reveal that Verasper sp. are the more primitive groups.
4. The G-banding was carried out by the trypsin method. The study showed that V. moseri and V. variegates have more difference in G-bands of chromosomes,but they have the same total numbers of G-bands which are 59. V. moseri have 37 deeply stained bands and 22 shallow stained bands,while V. variegates have 38 deeply stained bands and 21 shallow stained bands,but the region of G-bands in Verasper sp. chromosomes are different. G-banding is susceptible to interference by variations such as the degree of chromosome contraction,so the results in this paper are not satisfactory. It also may be due to the larger number and the tiny shape of fish chromosome and their DNA composition of chromosomes.
5. Calculated the karyotype resemblance near coefficients (λ) and the distance of karyotype evolution ( De ) of Verasper sp.,the resultλ=0.98, De =0.02 indicate that the difference between V. moseri and V. variegates are very little,they have very close relationship.
引文
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