罗汉果二倍体及四倍体遗传变异研究
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摘要
罗汉果Siraitia grosvenorii (Swingle) C. Jeffrey隶属于葫芦科(Cucurbitaceae)罗汉果属(Siraitia),为多年生草质藤本植物,是我国特有的药用和甜料植物,具止咳祛痰、凉血舒胃、润肠通便等作用。其主要活性成分为罗汉果甜苷V,具强烈甜味,为蔗糖的300-500倍,具抗氧化、免疫调节及抗癌的作用。罗汉果提取物低热、无毒,是一种纯天然的甜味剂及理想的保健品,可为糖尿病和肥胖病患者使用。罗汉果种子数量多、重量大,占果实鲜重的44%,干重的70%,但几乎不含罗汉果皂苷。因此罗汉果无籽化是罗汉果生产中亟待解决的关键问题,而且无籽罗汉果也成为罗汉果育种研究的热点。
2005年本课题组在罗汉果伯林3号和ND的后代中发现一突变株,植株高大,茎粗叶厚,花大败育,后代无籽。本研究首次对罗汉果正常株及其突变株从细胞学、基因组及基因表达水平进行初步探讨,研究罗汉果正正常株与突变株的遗传变异情况及无籽罗汉果基因表达情况。主要研究结果如下：
1、染色体核型的研究结果显示罗汉果正常株为二倍体,染色体数目为2n=2x=28,而突变株为四倍体,染色体数目为2n=4x=56,其后代无籽罗汉果为三倍体,染色体数目为2n=3x=42条,同时验证了突变株为四倍体。这是首次对罗汉果多倍体方面进行的报道研究。
2、本实验采用SRAP分子标记方法对罗汉果二倍体及四倍体的基因组水平进行遗传变异分析。用196对引物组合对两者的基因组进行扩增,其中9对引物组合未能扩增出条带。筛选得到的189对引物组合共扩增出约4573条带,其中577条(12.6%)为差异条带,1998条(87.4%)为共有条带。大部分引物组合均能扩增出条带,获得的条带长度大多集中在100-800bp之间,平均每对引物组合扩增条带数为12.1。结果表明通过SRAP分子标记方法得出罗汉果二倍体及四倍体株系之间基因组DNA的SRAP多态性较低,遗传差异较小。
3、采用SRAP-cDNA方法对罗汉果二倍体及四倍体的基因表达水平进行差异分析。用196对SRAP引物组合对两者的基因表达水平cDNA进行多态性扩增,其中63对引物组合未能扩增出条带。筛选得到的133对引物组合共扩增出约2917条带,其中289条(9.9%)为差异条带,1313条(90.1%)为共有条带。对其中稳定出现的明显差异片段进行回收、克隆及测序,共获得92条差异表达的基因片段,其中77.2%与已知基因具高度同源性,9.8%为编码未知蛋白基因,13.0%为可能的编码新蛋白基因。对已知序列的结果分析发现大多数序列都与光合、呼吸及抗性相关基因具有很高的同源性,其中比较重要的蛋白质如：核铜糖-1,5-二磷酸羧化酶/加氧酶、磷酸烯醇丙酮酸羧基激酶、丙酮酸激酶、过氧化物酶膜转运蛋白、NBS-LRR抗性蛋白、蛋白磷酸酶等。序列的功能分析结果表明已知基因编码蛋白涉及到生物学的诸多方面,其编码的蛋白主要属于：离子通道、信号转导、代谢途径、转录因子、蛋白合成、生长发育、能量代谢等。这些蛋白质在植物生长发育中都起着重要的调节作用,如：锌指蛋白、分子伴侣、促有丝分裂素活化激酶、转录因子IWS1、信号转导蛋白、内膜蛋白、胞外孔道蛋白、纤维素合酶、细胞色素P450、糖基转移酶GT8、氧化还原酶等。以上实验结果从一定程度上表明罗汉果四倍体在抗逆性及光合能力方面更优于二倍体植株。这也为多倍体在表型及生物学方面优于二倍体的现象提供了分子证据。
4、利用Solexa高通量测序技术对罗汉果二倍体后代的正常种子及四倍体后代无籽罗汉果干瘪种子进行转录组和表达谱测序。转录组测序结果共得到64372条Unigene,与代谢途径相关的Unigene共17593条。表达谱测序得到87350条reference tags及41678条geneso通过分析发现占mRNA,总量76%-77%的是种类不到5.5%-5.8%的少数nRNA,而占种类58%-62%的mRNA全部加起来不到mRNA,总量的4.2%-4.5%。对差异基因进行比较发现有1748个基因上调,2037个基因下调。
本文首次对罗汉果多倍体及其后代无籽罗汉果进行报道研究,并从细胞学、基因组及基因表达水平进行初步探讨,研究罗汉果正正常株与突变株的遗传变异情况及无籽罗汉果基因表达情况。这为多倍体罗汉果及无籽罗汉果形成的遗传机理、种子形成及发育相关基因等方面的研究提供重要的基因资源,也为克隆基因、研究其表达和调控机制及功能提供了基础数据,同时为应用生物技术方法实现罗汉果无籽化,提高生产中罗汉果有效成分提取率提供了技术支持,为培育无籽罗汉果及新资源新品种的开发利用提供理论依据,提高育种的预见性和效率。
Siraitia grosvenorii (Swingle) C. Jeffrey, belonging to the genus Siraitia Merr under the family Cucurbitaceae, is a precious and economically important species endemic to southern China and has been cultivated for several centuries. The fruits of S. grosvenorii, called Luohanguo, are used for food, beverages and traditional Chinese medicine. The fruit has evident effects on dry cough, sore throat, extreme thirst and constipation. The major components of Luohanguo are mogrosides, a group of terpene glycosides, estimated to be about 300 times as sweet as sucrose. Moreover, Luohanguo extracts have chemopreventive and antioxidant properties. Its non-caloric properties and effects on the cancer chemopreventive and antioxidant, are being used as sweetening agents for the patients with diabetes mellitus, adiposis, hypertension, heart disease and so on. Mogrosides only exist in flesh, occupying 15% of the dry weigh, whereas seeds constitute 70% of the weight but contain no mogrosides. Thus, seedless Luohanguo will no doubt increase the utilization and extraction rate of mogrosides.
Our group isolated a novel male mutant plant from hybrids of'Bolin' and 'ND'. The mutant exhibited growth vigor, larger body and increased organ size compared to wild type. We then used pollen to pollinate 'Nongyuan', and the following year fruits of female hybrids were seedless. In this study, we carried out preliminary studies on karyotype analysis and the molecular mechanism of mutant. The results will provide reference information and resources for molecular breeding of seedless Luohanguo.
1. The chromosome numbers of wild-type, mutant and hybrid were 2n=2x=28,2n =4x=56 and 2n=3x=42, respectively. The results showed that wild-type was a diploid strain. Mutant was a tetraploid, and hybrid was a triploid. To our knowledge, this is the first report on karyotype and molecular studied of polyploids in S. grosvenorii.
2. A total of 196 SRAP primer combinations were used for DNA amplifications,9 combinations of which failed to amplify banding patterns.189 SRAP primer combinations generated 4573 bands,577 (12.6%) of which were polymorphic while 1998 pairs (87.4%) were identical. Fragment bands obtained were in the size range of 100-800 bp, with a mean of 12.1. There were no apparent differences in the bands of M036 and M038 amplified by most primer pairs. The results generally indicated that the genetic diversity would be quite low between the diploid and tetraploid strains of S. grosvenorii.
3. A total of 196 SRAP primer combinations were used for cDNA amplifications,63 of which failed to amplify any banding patterns. A total of 133 SRAP primer combinations generated 2917 bands,289 (9.9%) of which were polymorphic while 1313 pairs (90.1%) were identical. Fragment bands obtained were in the size range of 100-800 bp. Stable and clearly differentiated fragments were sorted out, cloned and sequenced. Ninety-two differentially expressed fragments were successfully sequenced,77.2% of them were highly homologous to known genes,9.8% were hypothetical genes and 13.0% were possible new genes as they had no significant similarity to known genes. Sequence analysis revealed that most fragments had significant homologous nucleotide sequence to photosynthesis, respiration and stress response genes. These genes included ribulose-1,5-bisphosphate carboxylase/oxygenase, phosphoenolpyruvate carboxykinase, pyruvate kinase, peroxisomal membrane transporter, NBS-LRR type resistance protein, protein phosphatase and others. Functional analysis showed that the gene-encoded proteins were involved in a broad range of biological pathways, including transporters, signal transduction, metabolism, transcription, protein synthesis, development, energy, etc. There proteins play important roles in regulating palnt growth and development and include zinc finger protein, molecular chaperone, mitogen-activated kinase, transcription factor IWS1, transducin, endomembrane protein, porin, cellulose synthase, cytochrome P450, glycosyltransferase, oxidoreductase and other. The results, to some extent, indicated that the tetraploid of S. grosvenorii has more resistance to stress and photosynthesis ability than its diploid relatives. It provided molecular evidences for the phenomenon of polyploids exceeding their diploid relatives based on the phenotypic and biological features.
4. With Solexa high-throughput sequencing technology for transcriptome assembly of seeds from F049 and F050,43,891 Unigenes are obtained. There are 17593 Unigenes involving in metabolic pathway. In the DGE analysis of seeds from F049 and F050, 87350 reference tags and 41678 genes are obtained. The analysis shows that mRNA, which is less than 5.5%-5.8%of the types of mRNA, accounts for nearly 76%-77%of the total ammount, but all which accounts for 58%-62% of types, is less than 4.2%-4.5% of total amount mRNA. Statistics on the differentially expressed genes show that, in the F050/F049 genes, there are 1748 genes up-regulated, while 2037 genes are down-regulated.
In summary, in this study we carried out preliminary studies on karyotype analysis and the molecular mechanism of mutant strain. The interesting tetraploid mutant contains changes in genome structure and gene expression compared to the diploid strain. As this is the first study on tetraploid of S. grosvenorii, which will provide solid basis for future studies on tetraploid and hybrid triploid seedless Luohanguo. The results will provide reference information and resources for molecular breeding of seedless Luohanguo.
2005年本课题组在罗汉果伯林3号和ND的后代中发现一突变株,植株高大,茎粗叶厚,花大败育,后代无籽。本研究首次对罗汉果正常株及其突变株从细胞学、基因组及基因表达水平进行初步探讨,研究罗汉果正正常株与突变株的遗传变异情况及无籽罗汉果基因表达情况。主要研究结果如下：
1、染色体核型的研究结果显示罗汉果正常株为二倍体,染色体数目为2n=2x=28,而突变株为四倍体,染色体数目为2n=4x=56,其后代无籽罗汉果为三倍体,染色体数目为2n=3x=42条,同时验证了突变株为四倍体。这是首次对罗汉果多倍体方面进行的报道研究。
2、本实验采用SRAP分子标记方法对罗汉果二倍体及四倍体的基因组水平进行遗传变异分析。用196对引物组合对两者的基因组进行扩增,其中9对引物组合未能扩增出条带。筛选得到的189对引物组合共扩增出约4573条带,其中577条(12.6%)为差异条带,1998条(87.4%)为共有条带。大部分引物组合均能扩增出条带,获得的条带长度大多集中在100-800bp之间,平均每对引物组合扩增条带数为12.1。结果表明通过SRAP分子标记方法得出罗汉果二倍体及四倍体株系之间基因组DNA的SRAP多态性较低,遗传差异较小。
3、采用SRAP-cDNA方法对罗汉果二倍体及四倍体的基因表达水平进行差异分析。用196对SRAP引物组合对两者的基因表达水平cDNA进行多态性扩增,其中63对引物组合未能扩增出条带。筛选得到的133对引物组合共扩增出约2917条带,其中289条(9.9%)为差异条带,1313条(90.1%)为共有条带。对其中稳定出现的明显差异片段进行回收、克隆及测序,共获得92条差异表达的基因片段,其中77.2%与已知基因具高度同源性,9.8%为编码未知蛋白基因,13.0%为可能的编码新蛋白基因。对已知序列的结果分析发现大多数序列都与光合、呼吸及抗性相关基因具有很高的同源性,其中比较重要的蛋白质如：核铜糖-1,5-二磷酸羧化酶/加氧酶、磷酸烯醇丙酮酸羧基激酶、丙酮酸激酶、过氧化物酶膜转运蛋白、NBS-LRR抗性蛋白、蛋白磷酸酶等。序列的功能分析结果表明已知基因编码蛋白涉及到生物学的诸多方面,其编码的蛋白主要属于：离子通道、信号转导、代谢途径、转录因子、蛋白合成、生长发育、能量代谢等。这些蛋白质在植物生长发育中都起着重要的调节作用,如：锌指蛋白、分子伴侣、促有丝分裂素活化激酶、转录因子IWS1、信号转导蛋白、内膜蛋白、胞外孔道蛋白、纤维素合酶、细胞色素P450、糖基转移酶GT8、氧化还原酶等。以上实验结果从一定程度上表明罗汉果四倍体在抗逆性及光合能力方面更优于二倍体植株。这也为多倍体在表型及生物学方面优于二倍体的现象提供了分子证据。
4、利用Solexa高通量测序技术对罗汉果二倍体后代的正常种子及四倍体后代无籽罗汉果干瘪种子进行转录组和表达谱测序。转录组测序结果共得到64372条Unigene,与代谢途径相关的Unigene共17593条。表达谱测序得到87350条reference tags及41678条geneso通过分析发现占mRNA,总量76%-77%的是种类不到5.5%-5.8%的少数nRNA,而占种类58%-62%的mRNA全部加起来不到mRNA,总量的4.2%-4.5%。对差异基因进行比较发现有1748个基因上调,2037个基因下调。
本文首次对罗汉果多倍体及其后代无籽罗汉果进行报道研究,并从细胞学、基因组及基因表达水平进行初步探讨,研究罗汉果正正常株与突变株的遗传变异情况及无籽罗汉果基因表达情况。这为多倍体罗汉果及无籽罗汉果形成的遗传机理、种子形成及发育相关基因等方面的研究提供重要的基因资源,也为克隆基因、研究其表达和调控机制及功能提供了基础数据,同时为应用生物技术方法实现罗汉果无籽化,提高生产中罗汉果有效成分提取率提供了技术支持,为培育无籽罗汉果及新资源新品种的开发利用提供理论依据,提高育种的预见性和效率。
Siraitia grosvenorii (Swingle) C. Jeffrey, belonging to the genus Siraitia Merr under the family Cucurbitaceae, is a precious and economically important species endemic to southern China and has been cultivated for several centuries. The fruits of S. grosvenorii, called Luohanguo, are used for food, beverages and traditional Chinese medicine. The fruit has evident effects on dry cough, sore throat, extreme thirst and constipation. The major components of Luohanguo are mogrosides, a group of terpene glycosides, estimated to be about 300 times as sweet as sucrose. Moreover, Luohanguo extracts have chemopreventive and antioxidant properties. Its non-caloric properties and effects on the cancer chemopreventive and antioxidant, are being used as sweetening agents for the patients with diabetes mellitus, adiposis, hypertension, heart disease and so on. Mogrosides only exist in flesh, occupying 15% of the dry weigh, whereas seeds constitute 70% of the weight but contain no mogrosides. Thus, seedless Luohanguo will no doubt increase the utilization and extraction rate of mogrosides.
Our group isolated a novel male mutant plant from hybrids of'Bolin' and 'ND'. The mutant exhibited growth vigor, larger body and increased organ size compared to wild type. We then used pollen to pollinate 'Nongyuan', and the following year fruits of female hybrids were seedless. In this study, we carried out preliminary studies on karyotype analysis and the molecular mechanism of mutant. The results will provide reference information and resources for molecular breeding of seedless Luohanguo.
1. The chromosome numbers of wild-type, mutant and hybrid were 2n=2x=28,2n =4x=56 and 2n=3x=42, respectively. The results showed that wild-type was a diploid strain. Mutant was a tetraploid, and hybrid was a triploid. To our knowledge, this is the first report on karyotype and molecular studied of polyploids in S. grosvenorii.
2. A total of 196 SRAP primer combinations were used for DNA amplifications,9 combinations of which failed to amplify banding patterns.189 SRAP primer combinations generated 4573 bands,577 (12.6%) of which were polymorphic while 1998 pairs (87.4%) were identical. Fragment bands obtained were in the size range of 100-800 bp, with a mean of 12.1. There were no apparent differences in the bands of M036 and M038 amplified by most primer pairs. The results generally indicated that the genetic diversity would be quite low between the diploid and tetraploid strains of S. grosvenorii.
3. A total of 196 SRAP primer combinations were used for cDNA amplifications,63 of which failed to amplify any banding patterns. A total of 133 SRAP primer combinations generated 2917 bands,289 (9.9%) of which were polymorphic while 1313 pairs (90.1%) were identical. Fragment bands obtained were in the size range of 100-800 bp. Stable and clearly differentiated fragments were sorted out, cloned and sequenced. Ninety-two differentially expressed fragments were successfully sequenced,77.2% of them were highly homologous to known genes,9.8% were hypothetical genes and 13.0% were possible new genes as they had no significant similarity to known genes. Sequence analysis revealed that most fragments had significant homologous nucleotide sequence to photosynthesis, respiration and stress response genes. These genes included ribulose-1,5-bisphosphate carboxylase/oxygenase, phosphoenolpyruvate carboxykinase, pyruvate kinase, peroxisomal membrane transporter, NBS-LRR type resistance protein, protein phosphatase and others. Functional analysis showed that the gene-encoded proteins were involved in a broad range of biological pathways, including transporters, signal transduction, metabolism, transcription, protein synthesis, development, energy, etc. There proteins play important roles in regulating palnt growth and development and include zinc finger protein, molecular chaperone, mitogen-activated kinase, transcription factor IWS1, transducin, endomembrane protein, porin, cellulose synthase, cytochrome P450, glycosyltransferase, oxidoreductase and other. The results, to some extent, indicated that the tetraploid of S. grosvenorii has more resistance to stress and photosynthesis ability than its diploid relatives. It provided molecular evidences for the phenomenon of polyploids exceeding their diploid relatives based on the phenotypic and biological features.
4. With Solexa high-throughput sequencing technology for transcriptome assembly of seeds from F049 and F050,43,891 Unigenes are obtained. There are 17593 Unigenes involving in metabolic pathway. In the DGE analysis of seeds from F049 and F050, 87350 reference tags and 41678 genes are obtained. The analysis shows that mRNA, which is less than 5.5%-5.8%of the types of mRNA, accounts for nearly 76%-77%of the total ammount, but all which accounts for 58%-62% of types, is less than 4.2%-4.5% of total amount mRNA. Statistics on the differentially expressed genes show that, in the F050/F049 genes, there are 1748 genes up-regulated, while 2037 genes are down-regulated.
In summary, in this study we carried out preliminary studies on karyotype analysis and the molecular mechanism of mutant strain. The interesting tetraploid mutant contains changes in genome structure and gene expression compared to the diploid strain. As this is the first study on tetraploid of S. grosvenorii, which will provide solid basis for future studies on tetraploid and hybrid triploid seedless Luohanguo. The results will provide reference information and resources for molecular breeding of seedless Luohanguo.
引文
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