六种石斑鱼核型特征比较和染色体进化研究
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摘要
采用常规核型分析、C-带、Ag-NORs、DAPI荧光染料染色以及几种重复序列(5SrDNA、18S rDNA和(TTAGGG)_n)的荧光原位杂交技术对斜带石斑鱼E.coioides、青石斑鱼E.awara、拟青石斑鱼E.fasciatomaculosus、赤点石斑鱼E.akaara、褐石斑鱼E.bruneus和镶点石斑鱼E.amblycephalus的核型特征进行描述比较和研究,结果如下:
1、6种石斑鱼二倍体染色体数目均为48,但其染色体组组成具有一定的差异,核型公式分别为:斜带石斑鱼2n=48,2sm+46t,NF=50;青石斑鱼2n=48,48t,NF=48;拟青石斑鱼2n=48,1st+47t,NF=48;赤点石斑鱼2n=48,2sm+8st+38t,NF=50;褐石斑鱼2n=48,2m+4sm+42t,NF=54;镶点石斑鱼2n=48,2m+46t,NF=50。
2、分析了C-带和DAPI荧光带在6种石斑鱼染色体上的分布,6种石斑鱼染色体C-带和DAPI荧光带的分布模式均不同。
根据染色体组中是否绝大多数染色体的着丝粒区域为C—带异染色质这一特征,可将6种石斑鱼分为2种类型:(1)染色体组中绝大多数染色体具有着丝粒带,此类型中包括斜带石斑鱼和褐石斑鱼;(2)染色体组中绝大多数染色体不具有着丝粒带,此类型又可根据是否具有恒定的异染色质带细分为:a、绝大多数染色体无着丝粒带,仅双臂染色体的整条短臂为恒定的异染色质带,如拟青石斑鱼和赤点石斑鱼,b、绝大多数染色体无着丝粒带,仅在部分染色体的着丝点位置具有变动带,此类型包括青石斑鱼和镶点石斑鱼。
DAPI荧光带的研究结果表明,除斜带石斑鱼外,其它5种石斑鱼均具有DAPI荧光亮带,具体模式如下:斜带石斑鱼各条染色体被均匀染色,无明显DAPI荧光带;青石斑鱼与褐石斑鱼在其着丝点位置具有荧光亮带,但青石斑鱼的荧光带信号非常微弱而褐石斑鱼则较强;拟青石斑鱼和赤点石斑鱼染色体组中双臂染色体的整个短臂被DAPI染成明亮的荧光带;镶点石斑鱼在一些染色体的着丝点位置和第24对染色体上靠近端粒位置显示为荧光亮带。
褐石斑鱼绝大多数染色体着丝粒位置,拟青石斑鱼和赤点石斑鱼所有双臂染色体的整个短臂既表现为C带阳性也表现为DAPI荧光亮带,说明这些位置染色体的结构和碱基组成相似,可能具有同源性。
3、不论是常规的Giemsa染色,还是C—带以及DAPI荧光染色体,拟青石斑鱼染色体组中第1对染色体为异形。该对染色体由1条端部着丝粒染色体和1条亚端部着丝粒染色体组成,C—带和DAPI荧光带均表明该对染色体中的亚端部着丝粒染色体,其整个短臂为染色体组中唯一1条可以检测到C带阳性以及DAPI荧光亮带的染色体,而同源染色体中的另外1条则既无异染色质带也无DAPI荧光带,可见该对同源染色体为1对异形染色体。这种同源染色体异形的现象在石斑鱼属中属首次发现。
4、银染结果表明,在6种石斑鱼种内和种间,NORs数目和分布模式具有多态性。青石斑鱼、拟青石斑鱼、赤点石斑鱼和镶点石斑鱼染色体组中仅有1对核仁组织区位于第24对染色体靠近着丝粒的位置上;斜带石斑鱼仅在第24对染色体的短臂上具有1对NORs;褐石斑鱼染色体组中核仁组织区数目2～5个,位于双臂染色体,既第2、9、24对染色体短臂上。
5、利用荧光原位杂交技术研究5S rDNA和18S rDNA在6种石斑鱼染色体上的分布模式。结果表明5S rDNA在6种石斑鱼染色体组中,均位于1对中等大小端部着丝粒染色体且靠近着丝点的位置上,种间和种内未观察到差异,在染色体上的分布模式基本一致。这表明5S rDNA在石斑鱼属鱼类染色体上的分布模式具有较高的保守性;6种石斑鱼染色体组中,具有5S rDNA的染色体具有同源性。
利用FISH研究18S rDNA在6种石斑鱼染色体上的分布模式,结果表明:褐石斑鱼在3对(第2,9和24对)染色体上具有明显杂交信号,并且在一些染色体的末端也有较小和微弱的信号,检测到的rDNA位点多于银染检测到的位点数;其它5种石斑鱼只能在第24对染色体上检测到唯一1对杂交信号,与银染结果一致。6种石斑鱼第24对染色体上均具有NORs说明第24对染色体在6种石斑鱼中具有同源性。
6、利用FISH技术检测端粒序列在6种石斑鱼染色体上的分布,端粒探针杂交信号仅局限在所有染色体的端粒位置,在染色体上的中间部位未检测到杂交信号。但在褐石斑鱼中,发现有10对染色体的端粒位置,杂交信号强度明显比其它位点的强度高,信号大小也比其它染色体上的大,说明这些位置上的端粒序列拷贝数明显高于其它位点。
7、为了探索石斑鱼属鱼类进化分歧过程中染色体进化趋势与途径,本文综合了6种石斑鱼核型特征信息建立了系统进化树。通过与前人利用分子标记构建的系统进化树进行对比,发现2种进化树之间具有较好的一一对应关系。此外,系统进化树还揭示了石斑鱼属染色体的进化规律:石斑鱼属系统进化树中不同分支中的石斑鱼染色体进化采用不同形式的染色体重组以实现进化分歧。
The karyotypic characteristics of 6 groupers Epinephelus coioides、E. awara、E. fasciatomaculosus、E. akaara、E. bruneus and E. amblycephalus were investigated in the present paper by using Giemsa-staining, C-band, Ag-NORs, DAPI staining and Fluorescence in situ hybridization (FISH) with repetitive sequencing (5S rDNA, 18S rDNA, and (TTAGGG)_n ). The results are as following:
1. All six species presented uniform diploid numbers (2n =48), yet differed in their karyotypic formulae: E. coioides is 2n=48, 2sm+46t, NF=50; E. awara is 2n=48, 48t, NF=48;E.fasciatomaculosus is 2n=48, 1st+47t, NF=48; E. akaara is 2n=48, 2sm+8st +38t, NF=50; E. bruneus is 2n=48, 2m+4sm+42t, NF=54; E. amblycephalus is 2n= 48,2m+46t,NF = 50.
2. All six species showed species-specific C-band and DAPI-band patterns.
Approximately two groups of C-band patterns could be obtained: (1) the constitutiveheterochromatin was observed at the centromeric region on most chromosomes, as in E. coioides and E. bruneus; 2) heterochromatin was absent at the centromeric region of most chromosomes, as in the other four species. And in the second group, still two sub-types could be observed: a) C-banding heterochromatin constantly occupied the total short arm of the bi-armed chromosomes, as in E. fasciatomaculosus and E. akaara; b) C-banding heterochromatin was occasionally observed at the centromeric region on some chromosomes, as in E. awara and E. amblycephalus.
DAPI fluorochrome analyses showed that all six species of Epinephelus possess distinct DAPI fluorescent bands except E. coioides. Fluorescent band signals were observed at the centromeric regions of chromosomes in E. awara and E. bruneus, fainter in E. awara and stronger in E. bruneus; the whole short arms of biarmed chromosomes showed bright DAPI fluorescent bands in E. fasciatomaculosus and E. akaara; and fluorescent bands were observed at the centromeric regions of some chromosomes and the subtelomeric region of pair No.24 chromosome in E. amblycephalus.
The concurrence of C-positive bands and DAPI fluorescent bands at the pericentromeric regions of most chromosomes in E. bruneus and on the whole short arm of bi-armed chromosomes in E. fasciatomaculosus and E. akaara indicated that at these particular locations, the chromosome structure and bases were similar and therefore could be homeologous.
3. Chromosome pair No.1 of E. fasciatomaculosus was heteromorphic. Not only Giemsa-staining results showed that it is consisted of a subtelocentric chromosome and a telocentric chromosome, C-bands and DAPI fluorescent bands also occurred only on one of the homeologous chromosomes-the subtelocentric chromosome. No C-banding or DAPI fluorescent signals were detected on the telocentric chromosome. In the genus of Epinephelus, no such pair of heteromorphic homeologous chromosomes has ever been reported.
4. There are inter-species variations in the number and distribution pattern of NORs in the six species of Epinephelus. In E. awara、E. fasciatomaculosus、E. akaara and E. amblycephalus, a single pair of NORs was detected at the paracentromeric region of chromosome pair No.24. In E. coioides, a single pair of NORs was also detected on chromosome pair No.24, but it was located on the telomere of the short arms of chromosome pair No.24. In E. bruneus, the number of NORs varies from 2 to 6, and they always appeared on the telomere of the short arm of biarmed chromosomes (No.2, 9 and 24) .
5. Fluorescence in situ hybridization (FISH) technique was used to study the distribution patterns of 5S rDNA and 18S rDNA on chromosmoes of six species of Epinephelus. No inter- or intra- species differences were detected in the distribution pattern of 5S rDNA. In all 6 species, 5S rDNA was located at the paracentromeric region of a pair of middle-sized telocentric chromosome. This indicated that 5S rDNA distribution patterns were conservative and the 5S rDNA bearing chromosomes were homeologous in these 6 species. In five of the six species, a single pair of 18S rDNA hybridization signals could be detected only on chromosome pair No.24, corresponding to silver staining results. The exception was E. bruneus, in which strong hybridization signals were observed on three pairs of chromosomes (No.2, 9 and 24) as well as weak signals on some other chromosomes. 18S rDNA distribution patterns revealed that chromosome pair No.24 were homeologous in these 6 species.
6. The localization of the telomeric sequence (TTAGGG)_n on chromosomes of the six species was also analyzed. In all species, the telomeric sequences, detected by FISH, were restricted to telomeres, and no interstitial sites were observed. However, in E. bruneus, significant increase of hybridization signal intensity at one end of chromosome was observed in 10 pairs of chromosomes. This suggested that at these specific locations, telomeric sequences were repeated many more times than at the other locations.
7. To explore the evolutionary trend and process of chromosomes in the Epinephelus during diversification, this study summarized the karyotypic characteristics of six species of groupers and built a phylogenetic tree based on the Giemsa-staining, C-band, Ag-NORs, DAPI and FISH with repetitive sequencing (5S rDNA, 18S rDNA, and (TTAGGG)_n ) experimental results. The groupings of this tree corresponded well with many other former studies in which phylogenetic trees were built based on molecular marker information. Moreover, this karyotypic phylogenetic tree further revealed that species of different clade tend to evolve through different approaches.
1、6种石斑鱼二倍体染色体数目均为48,但其染色体组组成具有一定的差异,核型公式分别为:斜带石斑鱼2n=48,2sm+46t,NF=50;青石斑鱼2n=48,48t,NF=48;拟青石斑鱼2n=48,1st+47t,NF=48;赤点石斑鱼2n=48,2sm+8st+38t,NF=50;褐石斑鱼2n=48,2m+4sm+42t,NF=54;镶点石斑鱼2n=48,2m+46t,NF=50。
2、分析了C-带和DAPI荧光带在6种石斑鱼染色体上的分布,6种石斑鱼染色体C-带和DAPI荧光带的分布模式均不同。
根据染色体组中是否绝大多数染色体的着丝粒区域为C—带异染色质这一特征,可将6种石斑鱼分为2种类型:(1)染色体组中绝大多数染色体具有着丝粒带,此类型中包括斜带石斑鱼和褐石斑鱼;(2)染色体组中绝大多数染色体不具有着丝粒带,此类型又可根据是否具有恒定的异染色质带细分为:a、绝大多数染色体无着丝粒带,仅双臂染色体的整条短臂为恒定的异染色质带,如拟青石斑鱼和赤点石斑鱼,b、绝大多数染色体无着丝粒带,仅在部分染色体的着丝点位置具有变动带,此类型包括青石斑鱼和镶点石斑鱼。
DAPI荧光带的研究结果表明,除斜带石斑鱼外,其它5种石斑鱼均具有DAPI荧光亮带,具体模式如下:斜带石斑鱼各条染色体被均匀染色,无明显DAPI荧光带;青石斑鱼与褐石斑鱼在其着丝点位置具有荧光亮带,但青石斑鱼的荧光带信号非常微弱而褐石斑鱼则较强;拟青石斑鱼和赤点石斑鱼染色体组中双臂染色体的整个短臂被DAPI染成明亮的荧光带;镶点石斑鱼在一些染色体的着丝点位置和第24对染色体上靠近端粒位置显示为荧光亮带。
褐石斑鱼绝大多数染色体着丝粒位置,拟青石斑鱼和赤点石斑鱼所有双臂染色体的整个短臂既表现为C带阳性也表现为DAPI荧光亮带,说明这些位置染色体的结构和碱基组成相似,可能具有同源性。
3、不论是常规的Giemsa染色,还是C—带以及DAPI荧光染色体,拟青石斑鱼染色体组中第1对染色体为异形。该对染色体由1条端部着丝粒染色体和1条亚端部着丝粒染色体组成,C—带和DAPI荧光带均表明该对染色体中的亚端部着丝粒染色体,其整个短臂为染色体组中唯一1条可以检测到C带阳性以及DAPI荧光亮带的染色体,而同源染色体中的另外1条则既无异染色质带也无DAPI荧光带,可见该对同源染色体为1对异形染色体。这种同源染色体异形的现象在石斑鱼属中属首次发现。
4、银染结果表明,在6种石斑鱼种内和种间,NORs数目和分布模式具有多态性。青石斑鱼、拟青石斑鱼、赤点石斑鱼和镶点石斑鱼染色体组中仅有1对核仁组织区位于第24对染色体靠近着丝粒的位置上;斜带石斑鱼仅在第24对染色体的短臂上具有1对NORs;褐石斑鱼染色体组中核仁组织区数目2～5个,位于双臂染色体,既第2、9、24对染色体短臂上。
5、利用荧光原位杂交技术研究5S rDNA和18S rDNA在6种石斑鱼染色体上的分布模式。结果表明5S rDNA在6种石斑鱼染色体组中,均位于1对中等大小端部着丝粒染色体且靠近着丝点的位置上,种间和种内未观察到差异,在染色体上的分布模式基本一致。这表明5S rDNA在石斑鱼属鱼类染色体上的分布模式具有较高的保守性;6种石斑鱼染色体组中,具有5S rDNA的染色体具有同源性。
利用FISH研究18S rDNA在6种石斑鱼染色体上的分布模式,结果表明:褐石斑鱼在3对(第2,9和24对)染色体上具有明显杂交信号,并且在一些染色体的末端也有较小和微弱的信号,检测到的rDNA位点多于银染检测到的位点数;其它5种石斑鱼只能在第24对染色体上检测到唯一1对杂交信号,与银染结果一致。6种石斑鱼第24对染色体上均具有NORs说明第24对染色体在6种石斑鱼中具有同源性。
6、利用FISH技术检测端粒序列在6种石斑鱼染色体上的分布,端粒探针杂交信号仅局限在所有染色体的端粒位置,在染色体上的中间部位未检测到杂交信号。但在褐石斑鱼中,发现有10对染色体的端粒位置,杂交信号强度明显比其它位点的强度高,信号大小也比其它染色体上的大,说明这些位置上的端粒序列拷贝数明显高于其它位点。
7、为了探索石斑鱼属鱼类进化分歧过程中染色体进化趋势与途径,本文综合了6种石斑鱼核型特征信息建立了系统进化树。通过与前人利用分子标记构建的系统进化树进行对比,发现2种进化树之间具有较好的一一对应关系。此外,系统进化树还揭示了石斑鱼属染色体的进化规律:石斑鱼属系统进化树中不同分支中的石斑鱼染色体进化采用不同形式的染色体重组以实现进化分歧。
The karyotypic characteristics of 6 groupers Epinephelus coioides、E. awara、E. fasciatomaculosus、E. akaara、E. bruneus and E. amblycephalus were investigated in the present paper by using Giemsa-staining, C-band, Ag-NORs, DAPI staining and Fluorescence in situ hybridization (FISH) with repetitive sequencing (5S rDNA, 18S rDNA, and (TTAGGG)_n ). The results are as following:
1. All six species presented uniform diploid numbers (2n =48), yet differed in their karyotypic formulae: E. coioides is 2n=48, 2sm+46t, NF=50; E. awara is 2n=48, 48t, NF=48;E.fasciatomaculosus is 2n=48, 1st+47t, NF=48; E. akaara is 2n=48, 2sm+8st +38t, NF=50; E. bruneus is 2n=48, 2m+4sm+42t, NF=54; E. amblycephalus is 2n= 48,2m+46t,NF = 50.
2. All six species showed species-specific C-band and DAPI-band patterns.
Approximately two groups of C-band patterns could be obtained: (1) the constitutiveheterochromatin was observed at the centromeric region on most chromosomes, as in E. coioides and E. bruneus; 2) heterochromatin was absent at the centromeric region of most chromosomes, as in the other four species. And in the second group, still two sub-types could be observed: a) C-banding heterochromatin constantly occupied the total short arm of the bi-armed chromosomes, as in E. fasciatomaculosus and E. akaara; b) C-banding heterochromatin was occasionally observed at the centromeric region on some chromosomes, as in E. awara and E. amblycephalus.
DAPI fluorochrome analyses showed that all six species of Epinephelus possess distinct DAPI fluorescent bands except E. coioides. Fluorescent band signals were observed at the centromeric regions of chromosomes in E. awara and E. bruneus, fainter in E. awara and stronger in E. bruneus; the whole short arms of biarmed chromosomes showed bright DAPI fluorescent bands in E. fasciatomaculosus and E. akaara; and fluorescent bands were observed at the centromeric regions of some chromosomes and the subtelomeric region of pair No.24 chromosome in E. amblycephalus.
The concurrence of C-positive bands and DAPI fluorescent bands at the pericentromeric regions of most chromosomes in E. bruneus and on the whole short arm of bi-armed chromosomes in E. fasciatomaculosus and E. akaara indicated that at these particular locations, the chromosome structure and bases were similar and therefore could be homeologous.
3. Chromosome pair No.1 of E. fasciatomaculosus was heteromorphic. Not only Giemsa-staining results showed that it is consisted of a subtelocentric chromosome and a telocentric chromosome, C-bands and DAPI fluorescent bands also occurred only on one of the homeologous chromosomes-the subtelocentric chromosome. No C-banding or DAPI fluorescent signals were detected on the telocentric chromosome. In the genus of Epinephelus, no such pair of heteromorphic homeologous chromosomes has ever been reported.
4. There are inter-species variations in the number and distribution pattern of NORs in the six species of Epinephelus. In E. awara、E. fasciatomaculosus、E. akaara and E. amblycephalus, a single pair of NORs was detected at the paracentromeric region of chromosome pair No.24. In E. coioides, a single pair of NORs was also detected on chromosome pair No.24, but it was located on the telomere of the short arms of chromosome pair No.24. In E. bruneus, the number of NORs varies from 2 to 6, and they always appeared on the telomere of the short arm of biarmed chromosomes (No.2, 9 and 24) .
5. Fluorescence in situ hybridization (FISH) technique was used to study the distribution patterns of 5S rDNA and 18S rDNA on chromosmoes of six species of Epinephelus. No inter- or intra- species differences were detected in the distribution pattern of 5S rDNA. In all 6 species, 5S rDNA was located at the paracentromeric region of a pair of middle-sized telocentric chromosome. This indicated that 5S rDNA distribution patterns were conservative and the 5S rDNA bearing chromosomes were homeologous in these 6 species. In five of the six species, a single pair of 18S rDNA hybridization signals could be detected only on chromosome pair No.24, corresponding to silver staining results. The exception was E. bruneus, in which strong hybridization signals were observed on three pairs of chromosomes (No.2, 9 and 24) as well as weak signals on some other chromosomes. 18S rDNA distribution patterns revealed that chromosome pair No.24 were homeologous in these 6 species.
6. The localization of the telomeric sequence (TTAGGG)_n on chromosomes of the six species was also analyzed. In all species, the telomeric sequences, detected by FISH, were restricted to telomeres, and no interstitial sites were observed. However, in E. bruneus, significant increase of hybridization signal intensity at one end of chromosome was observed in 10 pairs of chromosomes. This suggested that at these specific locations, telomeric sequences were repeated many more times than at the other locations.
7. To explore the evolutionary trend and process of chromosomes in the Epinephelus during diversification, this study summarized the karyotypic characteristics of six species of groupers and built a phylogenetic tree based on the Giemsa-staining, C-band, Ag-NORs, DAPI and FISH with repetitive sequencing (5S rDNA, 18S rDNA, and (TTAGGG)_n ) experimental results. The groupings of this tree corresponded well with many other former studies in which phylogenetic trees were built based on molecular marker information. Moreover, this karyotypic phylogenetic tree further revealed that species of different clade tend to evolve through different approaches.
引文
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