茄属种及其种间体细胞杂种的染色体特征分析
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摘要
通过细胞融合的方式将马铃薯野生种或其他茄属近缘种的优良性状引入马铃薯栽培种是马铃薯资源创制的重要途径之一。研究体细胞杂种基因组中来自双亲的染色体的组成和重排,并在染色体水平解析性状的改变与染色体导入的联系,对基于染色体水平的马铃薯育种和细胞遗传学研究具有重要意义。荧光原位杂交是联系DNA序列和染色体生物学研究的重要工具。因此,本研究建立并优化了马铃薯荧光原位杂交体系,在此基础上对可能用于马铃薯资源创制的茄属几个相关种的染色体特征进行了研究,同时对马铃薯S. tuberosum和S. melongena的几个体细胞杂种进行了染色体分析。主要研究结果如下：
1.基于BAC-FISH标记的马铃薯染色体核型
本研究首先采用5个来自S. tuberosum和7个来自S. bulbocastanum的染色体特异BAC,通过BAC-FISH识别了马铃薯野生种S. chacoense的全套12条染色体。然后据已报道的遗传图谱、BAC连锁群以及前人核型分析信息,按照染色体和对应的BAC-FISH杂交结果将染色体进行排列,建立了S. chacoense12条染色体标准图,为后续研究奠定了基础。
2.马铃薯、番茄、茄子染色体的ITR分布特征
以二倍体茄属代表种S. tuberosum (A genome)、S. chacoense (A genome)、S. bulbocastanum (B genome)、S. pinnatisectum (B genome)、S. paucissectum (P genome)、 S. lycopersicum和S. melongena为材料,开展了端粒探针FISH杂交。结果显示,内部端粒序列(ITR)在供试种染色体着丝粒区域大量扩增。Dot blot定量分析发现,各个种中端粒序列相对含量的增加与FISH检测到的端粒信号强弱基本一致。大量来源于ITR的FISH信号被定位于S. pinnatisectum (B genome)粗线期染色体的主缢痕中。另外,在马铃薯A genome种S. chacoense第12染色体长臂的亚着丝粒区域检测到一对杂合的ITR位点。这些结果暗示,ITR可能与功能着丝粒有关。
3.茄属代表种rDNA分布
通过PCR扩增分别得到S.tuberosum、S. chacoense和S. melongena的5S rRNA基因和45S rRNA基因间隔区(IGS)的5'ETS。序列分析显示,S. tuberosum和S.chacoense的5S rRNA基因间隔区高度同源,5'ETS序列也高度同源。而S. tuberosum和S. melongena的5S rRNA基因间隔区序列差异较大,5'ETS序列差异也较大。用S. tuberosum5S rDNA和25S rDNA作为FISH探针,与7个马铃薯种和1个番茄、1个茄子种染色体进行原位杂交,结果显示,在所杂交的材料中,5S rDNA只在1组同源染色体上有杂交信号,25S rDNA除了S. melongena在3对非同源染色体上均有信号外,其它种也只在1组同源染色体上有信号。配对分析显示,5S rDNA和25SrDNA位于不同的染色体。信号分析还表明,在S. tuberosum、S. berthaultii、S. bulbocastanum、S. acaule和S. melongena5个种中,同源染色体之间的5S rDNA或25S rDNA信号强弱存在明显差异。
4.寡核苷酸SSR在茄属几个种染色体上的分布
通过FISH分别研究了单、二和三核苷酸SSR,包括(A)n、(C) n、(CG) n、(AC) n、(TC)n、(AT) n、(AAC) n、(AAG)n、(AAT)n、(AGG) n、(CAC) n、(CAT)n、(CAG)n、(ACT) n、(ACG)n和(GCC)n在马铃薯栽培种S. tuberosum和野生种S. chacoense中期染色体上的物理分布。总体上,所有核苷酸SSR在染色体上呈散点状分布。(A)。在S. chacoense中信号主要位于染色体末端和部分着丝粒区域,而在S. tuberosum中信号较少且分布没有明显规律。(AAT)n、(ACT)n和(CAG)n在S. chacoense中几乎出现在所有染色体的末端,但在其他部位很少发现,而在S.tuberosum中除去少数染色体末端外其他多数信号呈点状分散分布于整个染色体上。进一步分别采用混合的单、二和三核苷酸SSR在马铃薯种S. chacoense、S. bulbocastanum和S. pinnatisectum以及番茄有丝分裂中期染色体上开展了FISH定位。结果显示,单和二核苷酸SSR在马铃薯种和番茄染色体上的分布模式类似。三核苷酸SSR在马铃薯种间的分布模式类似,但在马铃薯和番茄间表现出很大的差异,在番茄中几乎所有的信号都集中于染色体着丝粒附近并且信号强烈,而在马铃薯中信号主要位于大多数染色体末端并呈点状分布。在一定程度上,该研究对马铃薯种间以及马铃薯和番茄之间差异SSR的筛选提供了依据。
5. S. tuberosum与几个相关种染色体GISH分析
以马铃薯栽培种S. tuberosum基因组DNA为探针,与包括S. tuberosum (A genome)、S. chacoense (A genome)、S. pinnatisectum (B genome)、S. bulbocastanum (B genome)、S. paucissectum(P genome)、S. etuberosum (E genome)、S. lycopersicum (L genome)和S. melongena在内的茄属种进行GISH杂交,结果显示,S. tuberosum与S. chacoense、S. paucissectum的基因组DNA同源性最高,其次是S. etuberosum和S. pinnatisectum,再次是S. bulbocastanum。S. lycopersicum和S. melongena与S.tuberosum同源性最低。因此,如果GISH用于体细胞杂种染色体组分分析,能够有效鉴定S. tuberosum与S. bulbocastanum (B genome)、S. lycopersicum和S. melongena的体细胞杂种,但S. tuberosum与S. chacoense (A genome)和S. paucissectum (P genome)的体细胞杂种很难用GISH分析鉴定染色体组分。
6.体细胞杂种染色体组成鉴定
研究以S. chacoense基因组DNA为探针,分别采用不封阻、封阻两种方式对S. tuberosum和S. chacoense的体细胞杂种进行GISH分析,结果显示,由于双亲同源性较高,不能清楚地分析杂种的染色体组分。进一步分别采用地高辛和生物素标记S. tuberosum和S. chacoense基因组DNA,开展了双色GISH,对3C10-2(混倍体)、3C28-1(5x)和3C33-2(6x)三个体细胞杂种的双色GISH结果显示,在3个体细胞杂种染色体中,除能够观察到少数单色荧光信号外,大部分染色体均同时被S. chacoense的红色信号和S. tuberosum的绿色信号覆盖。表明采用GISH的方法不能直观地区分S. tuberosum和S. chacoense体细胞杂种中染色体的组成。该结果说明如果两个融合亲本亲缘关系较近,采用GISH技术很难鉴定杂种的染色体组分。
马铃薯和茄子体细胞杂种的双色GISH结果显示,在供试的3个体细胞杂种中均能够清晰的鉴定出染色体的亲本来源,包括整条染色体和重排的染色体。进一步结合rDNA探针、端粒探针,在马铃薯和茄子的体细胞杂种中检测到了多种染色体重排现象,包括末端对末端的染色体融合、含有rDNA位点染色体的易位重排等。同时,在体细胞杂种60-10中还发现了2条具有双着丝粒的重排染色体,分别由端粒对端粒的末端融合和易位重排产生。另外,在60-13中1条重排染色体上的重排位点发现了ITR信号。结果表明,在GISH基础上结合丰富的细胞遗传学标记的检测方法是研究马铃薯和茄子体细胞杂种染色体组成的有效工具。
Introgression of desired traits from wild potato species or other closely related Solanum species into potato cultivars by cell fusion is an applicable way to create new germplasms. Investigations into the genome compositions, chromosome rearrangement, and the association of trait variation with chromosome introgression of the somatic hybrids are of great significance for potato breeding and cytogenetic studies on the chromosomal level. Fluorescence in situ hybridization plays an important role in connecting DNA sequences and chromosome biology. Therefore, the present study established and optimized a fluorescence in situ hybridization system in potato, and further examined the characteristics of chromosomes of several related Solanum species that would probably being involved in potato genetic-resource creation. Additionally, chromosome analysis was performed with several somatic hybrids derived from S. tuberosum and S. melongena. The main results obtained are as following:
1. The karyotype of potato chromosomes based on BAC-FISH markers
First, a full set of12chromosomes of the wild potato species S. chacoense were identified through BAC-FISH with chromosome-specific BACs, including5from S. tuberosum and7from S. bulbocastanum. Then, the standard karyotype of12S. chacoense chromosomes was set up according to the genetic maps, the BAC linkage groups, as well as previously reported karyotype information, which would lay a foundation for follow-up study.
2. Distribution characteristics of ITRs in potato, tomato, and eggplant chromosomes
FISH hybridizations using telomeric probe were conducted in representative diploid Solanum species, including S. tuberosum (A genome), S. chacoense (A genome), S. bulbocastanum (B genome), S. pinnatisectum (B genome), S. paucissectum (P genome), S. lycopersicum, and S. melongena. The results showed that interstitial telomeric repeats (ITRs) had been undergone an intense amplification in centromeric regions of chromosomes in tested species. Dot blot analysis revealed basically the same trends between the increasing of the relative contents of telomeric sequences and the strengthening of FISH signals from telomeric probe in species assayed. Large amount of ITR FISH signals were located in primary constrictions of pachytene chromosomes in S. pinnatisectum (B genome). In addition, a pair of heterozygous ITR sites were detected in the pericentromeric regions on the long arms of chromosome12in potato A genome species S. chacoense. These results suggest that ITRs may be related to the functional centromeres.
3. Distribution of rDNA in representative Solanum species
The5S rRNA genes and the5'ETS of45S rRNA gene spacer regions (IGS) were obtained by the PCR amplification in S. tuberosum, S. chacoense and S. melongena, respectively. Sequence analysis showed that the spacer regions of5S rRNA genes and also the5'ETS sequences between S. tuberosum and S. chacoense were highly homologous. However, the sequences of5S rRNA gene spacer regions and that of the5'ETS between S. tuberosum and S. melongena were diverged. The FISH assays were performed on chromosomes from seven potato, one tomato, and one eggplant species using S. tuberosum derived5S rDNA and25S rDNA probes. The results showed that5S rDNA signals were detected in only one set of homologous chromosome, while25S rDNA signals were detected in another set of homologous chromosome except for S. melongena in which25S rDNA signals presented on three pairs of homologous chromosomes. The5S rDNA and25S rDNA were found on distinct chromosomes when paring the homologous chromosomes. The strength of the5S rDNA or25S rDNA signals varied significantly between homologous chromosomes in S. tuberosum, S. berthaultii, S. bulbocastanum, S. acaule and S. melongena.
4. Distribution of oligonucleotide SSRs on chromosomes in several Solanum species
The physical distributions of mono-, di-, and trinucleotide SSRs, including (A)n,(C)n,(CG)n,(AC)n,(TC)n,(AT)n,(AAC)n,(AAG)n,(AAT)n,(AGG)n,(CAC)n,(CAT)n,(CAG)n,(ACT)n,(ACG)n, and (GCC)n on metaphase chromosomes were investigated by FISH in potato cultivated species S. tuberosum and wild species S. chacoense. In general, all kinds of nucleotide SSRs were scattered on chromosomes. The signals of (A)n were mainly detected at the ends and parts of the centromeric regions of chromosomes in S. chacoense, but fewer signals were detected in S. tuberosum with random distributions. Signals from (AAT)n,(ACT)n, and (CAG)n were restricted at almost all the chromosomal ends in S. chacoense, while they were scattered throughout the chromosomes in S. tuberosum excluding a few located at chromosome ends. FISH mapping of the mixture of mono-, di-, and trinucleotide SSRs on mitotic metaphase chromosomes were conducted respectively in potato species S. chacoense, S. bulbocastanum, and&pinnatisectum, as well as in tomato. The distribution patterns of mono-and dinucleotide SSRs were similar between potato and tomato species. However, great discrepancy in distribution of trinucleotide SSRs was found between tomato and potato, almost all the trinucleotide SSRs signals were concentrated in or around the centromeric regions of tomato chromosomes with strong intensities while they were spotted principally at most chromosomal ends in potato species. These findings provide, to a certain extent, a basic information for screening differential SSRs among potato species and between potato and tomato.
5. GISH analysis of chromosomes between S. tuberosum and several related species
Using the genomic DNA of cultivated potato S. tuberosum as a probe, GISH hybridizations were performed in species including S. tuberosum (A genome), S. chacoense (A genome), S. pinnatisectum (B genome), S. bulbocastanum (B genome), S. paucissectum (P genome), S. etuberosum (E genome), S. lycopersicum (L genome) and S. melongena. The results showed that S. tuberosum shared the highest homology of genomic DNA with S. chacoense and S. paucissectum, lower with S. etuberosum and S. pinnatisectum, lowest with S. bulbocastanum, S. lycopersicum and S. melongen. Therefore, if GISH is employed in discriminating the chromosomal compositions of somatic hybrids, it would be effective to discriminate S. tuberosum from S. bulbocastanum, S. lycopersicum and S. melongena, but it could be impractical when S. tuberosum is fused with S. chacoense and S. paucissectum.
6. Identification of chromosome composition in the somatic hybrids
GISH analysis with and without blocking were carried out separately with somatic hybrids derived from S. tuberosum and S. chacoense using S. chacoense genomic DNA as probe. The results demonstrated an infeasibility in identifying the chromosomal compositions of the hybrids owing to a high homology between the parents. For further tests, double-color GISH employing digoxin and biotin labeled genomic DNA probes from S. tuberosum and S. chacoense were conducted in somatic hybrids3C10-2(mixoploid),3C28-1(5x), and3C33-2(6x). The results showed that apart from a small number of chromosomal segments covered by apparent single color signals, most of the chromosomes were simultaneously painted on red or green signals from S. chacoense or S. tuberosum. It was illustrated impossible to discriminate the chromosomal compositions of the somatic hybrids between S. tuberosum and S. chacoense through GISH. The results indicate that GISH method is incapable of distinguishing chromosomal components in somatic hybrids derived from closely related species.
Results from double-color GISH investigation of three somatic hybrids between potato and eggplant showed that the parental origins of the chromosomes, including whole chromosomes and rearranged chromosomes, were clearly identified. In combination with the rDNA and the telomere probe in further study, a variety of chromosomal rearrangements, involving end-to-end chromosome fusion and chromosome translocation containing the rDNA locus, were detected in the somatic hybrids. Two rearranged double-centromeric chromosomes, which might be resulted from telomere-to-telomere fusion and translocation, respectively, were found in somatic hybrid60-10. Additionally, ITR signal was found at the site where the rearrangement had occurred on a rearranged chromosome of60-13. The results demonstrated that the GISH detection method that assists with abundant cytogenetic markers serves as an effective tool in diagnosing the chromosome constituents of somatic hybrids.
1.基于BAC-FISH标记的马铃薯染色体核型
本研究首先采用5个来自S. tuberosum和7个来自S. bulbocastanum的染色体特异BAC,通过BAC-FISH识别了马铃薯野生种S. chacoense的全套12条染色体。然后据已报道的遗传图谱、BAC连锁群以及前人核型分析信息,按照染色体和对应的BAC-FISH杂交结果将染色体进行排列,建立了S. chacoense12条染色体标准图,为后续研究奠定了基础。
2.马铃薯、番茄、茄子染色体的ITR分布特征
以二倍体茄属代表种S. tuberosum (A genome)、S. chacoense (A genome)、S. bulbocastanum (B genome)、S. pinnatisectum (B genome)、S. paucissectum (P genome)、 S. lycopersicum和S. melongena为材料,开展了端粒探针FISH杂交。结果显示,内部端粒序列(ITR)在供试种染色体着丝粒区域大量扩增。Dot blot定量分析发现,各个种中端粒序列相对含量的增加与FISH检测到的端粒信号强弱基本一致。大量来源于ITR的FISH信号被定位于S. pinnatisectum (B genome)粗线期染色体的主缢痕中。另外,在马铃薯A genome种S. chacoense第12染色体长臂的亚着丝粒区域检测到一对杂合的ITR位点。这些结果暗示,ITR可能与功能着丝粒有关。
3.茄属代表种rDNA分布
通过PCR扩增分别得到S.tuberosum、S. chacoense和S. melongena的5S rRNA基因和45S rRNA基因间隔区(IGS)的5'ETS。序列分析显示,S. tuberosum和S.chacoense的5S rRNA基因间隔区高度同源,5'ETS序列也高度同源。而S. tuberosum和S. melongena的5S rRNA基因间隔区序列差异较大,5'ETS序列差异也较大。用S. tuberosum5S rDNA和25S rDNA作为FISH探针,与7个马铃薯种和1个番茄、1个茄子种染色体进行原位杂交,结果显示,在所杂交的材料中,5S rDNA只在1组同源染色体上有杂交信号,25S rDNA除了S. melongena在3对非同源染色体上均有信号外,其它种也只在1组同源染色体上有信号。配对分析显示,5S rDNA和25SrDNA位于不同的染色体。信号分析还表明,在S. tuberosum、S. berthaultii、S. bulbocastanum、S. acaule和S. melongena5个种中,同源染色体之间的5S rDNA或25S rDNA信号强弱存在明显差异。
4.寡核苷酸SSR在茄属几个种染色体上的分布
通过FISH分别研究了单、二和三核苷酸SSR,包括(A)n、(C) n、(CG) n、(AC) n、(TC)n、(AT) n、(AAC) n、(AAG)n、(AAT)n、(AGG) n、(CAC) n、(CAT)n、(CAG)n、(ACT) n、(ACG)n和(GCC)n在马铃薯栽培种S. tuberosum和野生种S. chacoense中期染色体上的物理分布。总体上,所有核苷酸SSR在染色体上呈散点状分布。(A)。在S. chacoense中信号主要位于染色体末端和部分着丝粒区域,而在S. tuberosum中信号较少且分布没有明显规律。(AAT)n、(ACT)n和(CAG)n在S. chacoense中几乎出现在所有染色体的末端,但在其他部位很少发现,而在S.tuberosum中除去少数染色体末端外其他多数信号呈点状分散分布于整个染色体上。进一步分别采用混合的单、二和三核苷酸SSR在马铃薯种S. chacoense、S. bulbocastanum和S. pinnatisectum以及番茄有丝分裂中期染色体上开展了FISH定位。结果显示,单和二核苷酸SSR在马铃薯种和番茄染色体上的分布模式类似。三核苷酸SSR在马铃薯种间的分布模式类似,但在马铃薯和番茄间表现出很大的差异,在番茄中几乎所有的信号都集中于染色体着丝粒附近并且信号强烈,而在马铃薯中信号主要位于大多数染色体末端并呈点状分布。在一定程度上,该研究对马铃薯种间以及马铃薯和番茄之间差异SSR的筛选提供了依据。
5. S. tuberosum与几个相关种染色体GISH分析
以马铃薯栽培种S. tuberosum基因组DNA为探针,与包括S. tuberosum (A genome)、S. chacoense (A genome)、S. pinnatisectum (B genome)、S. bulbocastanum (B genome)、S. paucissectum(P genome)、S. etuberosum (E genome)、S. lycopersicum (L genome)和S. melongena在内的茄属种进行GISH杂交,结果显示,S. tuberosum与S. chacoense、S. paucissectum的基因组DNA同源性最高,其次是S. etuberosum和S. pinnatisectum,再次是S. bulbocastanum。S. lycopersicum和S. melongena与S.tuberosum同源性最低。因此,如果GISH用于体细胞杂种染色体组分分析,能够有效鉴定S. tuberosum与S. bulbocastanum (B genome)、S. lycopersicum和S. melongena的体细胞杂种,但S. tuberosum与S. chacoense (A genome)和S. paucissectum (P genome)的体细胞杂种很难用GISH分析鉴定染色体组分。
6.体细胞杂种染色体组成鉴定
研究以S. chacoense基因组DNA为探针,分别采用不封阻、封阻两种方式对S. tuberosum和S. chacoense的体细胞杂种进行GISH分析,结果显示,由于双亲同源性较高,不能清楚地分析杂种的染色体组分。进一步分别采用地高辛和生物素标记S. tuberosum和S. chacoense基因组DNA,开展了双色GISH,对3C10-2(混倍体)、3C28-1(5x)和3C33-2(6x)三个体细胞杂种的双色GISH结果显示,在3个体细胞杂种染色体中,除能够观察到少数单色荧光信号外,大部分染色体均同时被S. chacoense的红色信号和S. tuberosum的绿色信号覆盖。表明采用GISH的方法不能直观地区分S. tuberosum和S. chacoense体细胞杂种中染色体的组成。该结果说明如果两个融合亲本亲缘关系较近,采用GISH技术很难鉴定杂种的染色体组分。
马铃薯和茄子体细胞杂种的双色GISH结果显示,在供试的3个体细胞杂种中均能够清晰的鉴定出染色体的亲本来源,包括整条染色体和重排的染色体。进一步结合rDNA探针、端粒探针,在马铃薯和茄子的体细胞杂种中检测到了多种染色体重排现象,包括末端对末端的染色体融合、含有rDNA位点染色体的易位重排等。同时,在体细胞杂种60-10中还发现了2条具有双着丝粒的重排染色体,分别由端粒对端粒的末端融合和易位重排产生。另外,在60-13中1条重排染色体上的重排位点发现了ITR信号。结果表明,在GISH基础上结合丰富的细胞遗传学标记的检测方法是研究马铃薯和茄子体细胞杂种染色体组成的有效工具。
Introgression of desired traits from wild potato species or other closely related Solanum species into potato cultivars by cell fusion is an applicable way to create new germplasms. Investigations into the genome compositions, chromosome rearrangement, and the association of trait variation with chromosome introgression of the somatic hybrids are of great significance for potato breeding and cytogenetic studies on the chromosomal level. Fluorescence in situ hybridization plays an important role in connecting DNA sequences and chromosome biology. Therefore, the present study established and optimized a fluorescence in situ hybridization system in potato, and further examined the characteristics of chromosomes of several related Solanum species that would probably being involved in potato genetic-resource creation. Additionally, chromosome analysis was performed with several somatic hybrids derived from S. tuberosum and S. melongena. The main results obtained are as following:
1. The karyotype of potato chromosomes based on BAC-FISH markers
First, a full set of12chromosomes of the wild potato species S. chacoense were identified through BAC-FISH with chromosome-specific BACs, including5from S. tuberosum and7from S. bulbocastanum. Then, the standard karyotype of12S. chacoense chromosomes was set up according to the genetic maps, the BAC linkage groups, as well as previously reported karyotype information, which would lay a foundation for follow-up study.
2. Distribution characteristics of ITRs in potato, tomato, and eggplant chromosomes
FISH hybridizations using telomeric probe were conducted in representative diploid Solanum species, including S. tuberosum (A genome), S. chacoense (A genome), S. bulbocastanum (B genome), S. pinnatisectum (B genome), S. paucissectum (P genome), S. lycopersicum, and S. melongena. The results showed that interstitial telomeric repeats (ITRs) had been undergone an intense amplification in centromeric regions of chromosomes in tested species. Dot blot analysis revealed basically the same trends between the increasing of the relative contents of telomeric sequences and the strengthening of FISH signals from telomeric probe in species assayed. Large amount of ITR FISH signals were located in primary constrictions of pachytene chromosomes in S. pinnatisectum (B genome). In addition, a pair of heterozygous ITR sites were detected in the pericentromeric regions on the long arms of chromosome12in potato A genome species S. chacoense. These results suggest that ITRs may be related to the functional centromeres.
3. Distribution of rDNA in representative Solanum species
The5S rRNA genes and the5'ETS of45S rRNA gene spacer regions (IGS) were obtained by the PCR amplification in S. tuberosum, S. chacoense and S. melongena, respectively. Sequence analysis showed that the spacer regions of5S rRNA genes and also the5'ETS sequences between S. tuberosum and S. chacoense were highly homologous. However, the sequences of5S rRNA gene spacer regions and that of the5'ETS between S. tuberosum and S. melongena were diverged. The FISH assays were performed on chromosomes from seven potato, one tomato, and one eggplant species using S. tuberosum derived5S rDNA and25S rDNA probes. The results showed that5S rDNA signals were detected in only one set of homologous chromosome, while25S rDNA signals were detected in another set of homologous chromosome except for S. melongena in which25S rDNA signals presented on three pairs of homologous chromosomes. The5S rDNA and25S rDNA were found on distinct chromosomes when paring the homologous chromosomes. The strength of the5S rDNA or25S rDNA signals varied significantly between homologous chromosomes in S. tuberosum, S. berthaultii, S. bulbocastanum, S. acaule and S. melongena.
4. Distribution of oligonucleotide SSRs on chromosomes in several Solanum species
The physical distributions of mono-, di-, and trinucleotide SSRs, including (A)n,(C)n,(CG)n,(AC)n,(TC)n,(AT)n,(AAC)n,(AAG)n,(AAT)n,(AGG)n,(CAC)n,(CAT)n,(CAG)n,(ACT)n,(ACG)n, and (GCC)n on metaphase chromosomes were investigated by FISH in potato cultivated species S. tuberosum and wild species S. chacoense. In general, all kinds of nucleotide SSRs were scattered on chromosomes. The signals of (A)n were mainly detected at the ends and parts of the centromeric regions of chromosomes in S. chacoense, but fewer signals were detected in S. tuberosum with random distributions. Signals from (AAT)n,(ACT)n, and (CAG)n were restricted at almost all the chromosomal ends in S. chacoense, while they were scattered throughout the chromosomes in S. tuberosum excluding a few located at chromosome ends. FISH mapping of the mixture of mono-, di-, and trinucleotide SSRs on mitotic metaphase chromosomes were conducted respectively in potato species S. chacoense, S. bulbocastanum, and&pinnatisectum, as well as in tomato. The distribution patterns of mono-and dinucleotide SSRs were similar between potato and tomato species. However, great discrepancy in distribution of trinucleotide SSRs was found between tomato and potato, almost all the trinucleotide SSRs signals were concentrated in or around the centromeric regions of tomato chromosomes with strong intensities while they were spotted principally at most chromosomal ends in potato species. These findings provide, to a certain extent, a basic information for screening differential SSRs among potato species and between potato and tomato.
5. GISH analysis of chromosomes between S. tuberosum and several related species
Using the genomic DNA of cultivated potato S. tuberosum as a probe, GISH hybridizations were performed in species including S. tuberosum (A genome), S. chacoense (A genome), S. pinnatisectum (B genome), S. bulbocastanum (B genome), S. paucissectum (P genome), S. etuberosum (E genome), S. lycopersicum (L genome) and S. melongena. The results showed that S. tuberosum shared the highest homology of genomic DNA with S. chacoense and S. paucissectum, lower with S. etuberosum and S. pinnatisectum, lowest with S. bulbocastanum, S. lycopersicum and S. melongen. Therefore, if GISH is employed in discriminating the chromosomal compositions of somatic hybrids, it would be effective to discriminate S. tuberosum from S. bulbocastanum, S. lycopersicum and S. melongena, but it could be impractical when S. tuberosum is fused with S. chacoense and S. paucissectum.
6. Identification of chromosome composition in the somatic hybrids
GISH analysis with and without blocking were carried out separately with somatic hybrids derived from S. tuberosum and S. chacoense using S. chacoense genomic DNA as probe. The results demonstrated an infeasibility in identifying the chromosomal compositions of the hybrids owing to a high homology between the parents. For further tests, double-color GISH employing digoxin and biotin labeled genomic DNA probes from S. tuberosum and S. chacoense were conducted in somatic hybrids3C10-2(mixoploid),3C28-1(5x), and3C33-2(6x). The results showed that apart from a small number of chromosomal segments covered by apparent single color signals, most of the chromosomes were simultaneously painted on red or green signals from S. chacoense or S. tuberosum. It was illustrated impossible to discriminate the chromosomal compositions of the somatic hybrids between S. tuberosum and S. chacoense through GISH. The results indicate that GISH method is incapable of distinguishing chromosomal components in somatic hybrids derived from closely related species.
Results from double-color GISH investigation of three somatic hybrids between potato and eggplant showed that the parental origins of the chromosomes, including whole chromosomes and rearranged chromosomes, were clearly identified. In combination with the rDNA and the telomere probe in further study, a variety of chromosomal rearrangements, involving end-to-end chromosome fusion and chromosome translocation containing the rDNA locus, were detected in the somatic hybrids. Two rearranged double-centromeric chromosomes, which might be resulted from telomere-to-telomere fusion and translocation, respectively, were found in somatic hybrid60-10. Additionally, ITR signal was found at the site where the rearrangement had occurred on a rearranged chromosome of60-13. The results demonstrated that the GISH detection method that assists with abundant cytogenetic markers serves as an effective tool in diagnosing the chromosome constituents of somatic hybrids.
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