担子菌毛头鬼伞的TMV抗性蛋白y3基因的分离、鉴定和表达
摘要
许多具有生物活性的蛋白质具有潜在的应用价值,因而吸引着人们的研究兴趣。目前已被纯化和鉴定的真菌蛋白质有：核糖体失活蛋白、抗真菌蛋白、类-泛素蛋白、免疫调节蛋白、漆酶、凝集素、抗病毒蛋白,以及其他酶类：纤维素酶木聚糖酶、转化酶、核糖核酸酶等。对毛头鬼伞(Coprinus comatus (Mull.:Fr.) S. F. Gray)生物活性蛋白研究较深入的当属毛头鬼伞抗性蛋白Y3。该蛋白具有抗植物病毒TMV的活性、红细胞凝集活性,也与核糖体失活蛋白(RIP)有所相似。抗性蛋白Y3作为一种新的蛋白资源,有希望开发成转基因的抗病毒工程产品用于生物农药生产。为了达到此目的,首先需要获得抗蛋白基因y3的全长序列,并鉴定该基因抗病毒特性。
鉴于毛头鬼伞的抗性蛋白Y3具有潜在的应用价值和开发前景,本文分离了抗性蛋白y3基因,并分析了它的结构与功能。取得的主要结果如下：
(1)用YADE法从毛头鬼伞菌丝体中分离了抗性蛋白基因y3的全长DNA序列,发现其由1289 bp核苷酸组成(GenBank Accession No. HM204931),包括5L侧翼和3L侧翼区。
(2)用5'-RACE从毛头鬼伞菌丝体中克隆出y3基因的全长cDNA序列,该全长序列由534个bp组成,含1个ORF,编码130个氨基酸残基。GenBank和EMBL中索引号(Accession No.)分别为GQ859168和FN546262。
(3)y3基因的DNA和cDNA比对表明, y3基因含5个外显子和4个内含子。本研究获得的抗性蛋白基因y3的全长序列与前人报道的研究结果之间,有101个氨基酸残基共享序列。全长序列推导的氨基酸序列比原先增加了29个氨基酸残基。依据本研究的实验数据,使用在线网络启动子预测系统,发现了抗性蛋白基因y3的转录起始位点A和TATA盒,为进一步研究该基因的生物学功能奠定了基础。
(4)本研究得到的cDNA全长序列和由此推导的氨基酸序列与前人发表的基因y3的部分对应片断都表现出了高度的相似性,相似率达94%。利用y3基因全长序列信息,克隆了y3基因cDNA序列,并用它与pBI121和pCAMBIA1301构建了植物表达载体pCAMBIA1301-y3。
(5)用pCAMBIA1301-y3转化烟草,获得了转基因植株。经PCR, RT-PCR和Northern blot分析证实,y3基因已经整合到烟草基因组中,并在其中得到表达。对TMV的抗性鉴定也表明,y3基因导入使转基因烟草植株表现出一定抗病毒活性。y3基因全长序列的获得和对烟草的成功转化为y3基因的进一步研究奠定了基础。
(6)对前人关于真菌核酸的快速提取方法做了改进,用于真菌和烟草DNA和RNA的提取。同时,通过低温保藏试验评价核酸提取液和提取产物的稳定性和有效性；利用凝胶电泳、PCR和RT-PCR等方法比较分析核酸质量；并将提取效果与传统方法或试剂盒的提取效果进行比较。结果表明,改进后的提取方法是一种简单、方便和有效的实验方法,不仅可用于真菌也可用于植物DNA和RNA提取,用这种方法提取得到的DNA和RNA在低温保存条件下比较稳定,能较长时间保持其原有活性。
(7)在从毛头鬼伞中克隆抗性蛋白基因y3时意外获得的一条非目的条带,长度为906 bp (GenBank Accession No. GU568178)。将此序列通过NCBI的BLAST搜索,以及与其同源序列进行Clustal w和MEGA4.1聚类分析,证实该序列是28S rRNA。同时还发现毛头鬼伞的系统进化关系比较离散。此外,在这一新28S rRNA与TMV的抗性蛋白基因y3之间发现有两个同源区段有可能是PCR扩增y3基因时出现非目的条带的原因。在这两个同源区段中,其一与克隆y3基因时所用的PCR引物之一有较高的相似性,另一区段也是一般PCR引物的类似物。本研究中新28S rRNA序列的获得是PCR扩增中出现非目的条带的新例,这一新序列的发现及聚类分析的结果有助于该真菌基因组学及真菌生物分子分类系统的建立。
A variety of fungus proteins with biological activity possess potentially applicable value. They include ribosome inactivating proteins, antifungal proteins, ribonucleases, ubiquitin-like proteins, lectins, cellulases, laccases, invertases etc. Among them, there is an anti-TMV protein, designated as Y3 protein and encoded by y3 gene. Y3 protein was isolated from the fungus shaggy mane (Coprinus comatus (Mull.:Fr.) S. F. Gray), and showed inhibition on multiplication of tobacco mosaic virus (TMV). Because y3 gene might be a potential gene resource for antiviral genetic engineering, further studies, including full-length cDNA sequencing and identification of antiviral activity, should be completed.
In this work gene y3 was isolated from fungus C. comatus. The sequencing of this gene was carried out. Plant expression vector harboring gene Y3 was constructed and transformed into tobacco. The summary is as follows:
(1) Using Y-shaped adapter dependant extension (YADE), y3 gene of full-length DNA sequence was isolated from C. comatus mycelia and characterized. The putative full DNA sequence of y3 gene consisted of 1289 bps (GenBank Accession No. HM204931), which included an encoding region and two flanking regions at the 5'-and 3'-terminal.
(2) Cloning and characterization of the full-length cDNA y3 from of C. comatus mycelia was carried out with 5'-RACE. The putative full sequence of cDNA of y3 gene was composed of 534 bps with an ORF. The ORF encoded a polypeptide of 130 amino acid residues. Accession No. in GenBank is GQ859168和Accession No. in EMBL is FN546262。Comparison of the polypeptide sequence encoded by full-length y3 gene with the partial amino acid sequence reported before showed that there were 101 amino acids shared with the published Y3 sequence, and there were additional 29 amino acids newly derived from the full-length cDNA sequence.
(3) Through alignment of the DNA sequence of y3 gene with its cDNA, it was revealed that y3 gene was composed of 5 exons and 4 introns. Utilizing the software of promoter prediction on line and based on our experimental data, the transcription start site of y3 gene was predicted at A (No.321 bp) that followed TaTa box in DNA sequence.
(4) The full length of y3 gene we obtained exhibited high similarity (up to 94%) to partial known fragment of gene y3 sequence. According to the full-length cDNA sequence, gene y3 had been cloned and used to construct plant expression plasmid.
(5) The plasmid was constructed via inserting gene y3 sequence, at the MCS of pCAMBIA1301 along with CaMV 35S promoter and NOS terminator. The expression plasmids were transferred into Nicotiana tabacumvia via Agrobacterium mediated method, resulting in transgenic plantlets. The transcription of y3 gene was characterized by Northern blot analysis. Inoculation tests on transgenic plantlets with TMV demonstrated inhibitory activity against TMV. This illustrated that y3 gene had been transcripted and expressed in transgenic tobacco plants and y3 gene was proved to have certain inhibitory function against TMV in vivo.
(6) In order to extract DNA or RNA simply and effectively, a published protocol for extracting DNA and RNA was improved and applied to isolate DNA and RNA of fungi and tobacco. Compared with commonly used method and commercial kits, the simplified protocol could be used to extract DNA and RNA not only from fungi but also from plants. The isolated DNA and RNA were stable and their activities could be kept when stored under low temperatures.
(7) A new 28S rRNA sequence with 906 bps was isolated from mycelia of C. comatus (GenBank Accession No. GU568178). The sequence was obtained from an untargeted band when we cloned antiviral protein gene y3. This sequence was used to search homologous sequences using BLAST algorithm in NCBI, and to perform cluster with its homologues using the Clustal w and MEGA methods. The results confirmed that this sequence was a new 28S rRNA, and the phylogenic relationship of C. comatus was found to be dispersed from each other. Moreover, two homologous counterparts between the sequences of new 28S rRNA and the y3 gene were the possible cause of formation of the untargeted band in PCR amplification of y3 gene. Within these two homologous counterparts, one counterpart had the sequence similar to a PCR primer for y3 gene, and the other one was a primer-like fragment. In this research, obtaining of the new sequence of 28S rRNA was a new example of the formation of untargeted bands during PCR amplification. This finding and clustering analysis results of 28S rRNA would be helpful for researches in Fungus Genomics and the establishment of their molecular classification system.
鉴于毛头鬼伞的抗性蛋白Y3具有潜在的应用价值和开发前景,本文分离了抗性蛋白y3基因,并分析了它的结构与功能。取得的主要结果如下：
(1)用YADE法从毛头鬼伞菌丝体中分离了抗性蛋白基因y3的全长DNA序列,发现其由1289 bp核苷酸组成(GenBank Accession No. HM204931),包括5L侧翼和3L侧翼区。
(2)用5'-RACE从毛头鬼伞菌丝体中克隆出y3基因的全长cDNA序列,该全长序列由534个bp组成,含1个ORF,编码130个氨基酸残基。GenBank和EMBL中索引号(Accession No.)分别为GQ859168和FN546262。
(3)y3基因的DNA和cDNA比对表明, y3基因含5个外显子和4个内含子。本研究获得的抗性蛋白基因y3的全长序列与前人报道的研究结果之间,有101个氨基酸残基共享序列。全长序列推导的氨基酸序列比原先增加了29个氨基酸残基。依据本研究的实验数据,使用在线网络启动子预测系统,发现了抗性蛋白基因y3的转录起始位点A和TATA盒,为进一步研究该基因的生物学功能奠定了基础。
(4)本研究得到的cDNA全长序列和由此推导的氨基酸序列与前人发表的基因y3的部分对应片断都表现出了高度的相似性,相似率达94%。利用y3基因全长序列信息,克隆了y3基因cDNA序列,并用它与pBI121和pCAMBIA1301构建了植物表达载体pCAMBIA1301-y3。
(5)用pCAMBIA1301-y3转化烟草,获得了转基因植株。经PCR, RT-PCR和Northern blot分析证实,y3基因已经整合到烟草基因组中,并在其中得到表达。对TMV的抗性鉴定也表明,y3基因导入使转基因烟草植株表现出一定抗病毒活性。y3基因全长序列的获得和对烟草的成功转化为y3基因的进一步研究奠定了基础。
(6)对前人关于真菌核酸的快速提取方法做了改进,用于真菌和烟草DNA和RNA的提取。同时,通过低温保藏试验评价核酸提取液和提取产物的稳定性和有效性；利用凝胶电泳、PCR和RT-PCR等方法比较分析核酸质量；并将提取效果与传统方法或试剂盒的提取效果进行比较。结果表明,改进后的提取方法是一种简单、方便和有效的实验方法,不仅可用于真菌也可用于植物DNA和RNA提取,用这种方法提取得到的DNA和RNA在低温保存条件下比较稳定,能较长时间保持其原有活性。
(7)在从毛头鬼伞中克隆抗性蛋白基因y3时意外获得的一条非目的条带,长度为906 bp (GenBank Accession No. GU568178)。将此序列通过NCBI的BLAST搜索,以及与其同源序列进行Clustal w和MEGA4.1聚类分析,证实该序列是28S rRNA。同时还发现毛头鬼伞的系统进化关系比较离散。此外,在这一新28S rRNA与TMV的抗性蛋白基因y3之间发现有两个同源区段有可能是PCR扩增y3基因时出现非目的条带的原因。在这两个同源区段中,其一与克隆y3基因时所用的PCR引物之一有较高的相似性,另一区段也是一般PCR引物的类似物。本研究中新28S rRNA序列的获得是PCR扩增中出现非目的条带的新例,这一新序列的发现及聚类分析的结果有助于该真菌基因组学及真菌生物分子分类系统的建立。
A variety of fungus proteins with biological activity possess potentially applicable value. They include ribosome inactivating proteins, antifungal proteins, ribonucleases, ubiquitin-like proteins, lectins, cellulases, laccases, invertases etc. Among them, there is an anti-TMV protein, designated as Y3 protein and encoded by y3 gene. Y3 protein was isolated from the fungus shaggy mane (Coprinus comatus (Mull.:Fr.) S. F. Gray), and showed inhibition on multiplication of tobacco mosaic virus (TMV). Because y3 gene might be a potential gene resource for antiviral genetic engineering, further studies, including full-length cDNA sequencing and identification of antiviral activity, should be completed.
In this work gene y3 was isolated from fungus C. comatus. The sequencing of this gene was carried out. Plant expression vector harboring gene Y3 was constructed and transformed into tobacco. The summary is as follows:
(1) Using Y-shaped adapter dependant extension (YADE), y3 gene of full-length DNA sequence was isolated from C. comatus mycelia and characterized. The putative full DNA sequence of y3 gene consisted of 1289 bps (GenBank Accession No. HM204931), which included an encoding region and two flanking regions at the 5'-and 3'-terminal.
(2) Cloning and characterization of the full-length cDNA y3 from of C. comatus mycelia was carried out with 5'-RACE. The putative full sequence of cDNA of y3 gene was composed of 534 bps with an ORF. The ORF encoded a polypeptide of 130 amino acid residues. Accession No. in GenBank is GQ859168和Accession No. in EMBL is FN546262。Comparison of the polypeptide sequence encoded by full-length y3 gene with the partial amino acid sequence reported before showed that there were 101 amino acids shared with the published Y3 sequence, and there were additional 29 amino acids newly derived from the full-length cDNA sequence.
(3) Through alignment of the DNA sequence of y3 gene with its cDNA, it was revealed that y3 gene was composed of 5 exons and 4 introns. Utilizing the software of promoter prediction on line and based on our experimental data, the transcription start site of y3 gene was predicted at A (No.321 bp) that followed TaTa box in DNA sequence.
(4) The full length of y3 gene we obtained exhibited high similarity (up to 94%) to partial known fragment of gene y3 sequence. According to the full-length cDNA sequence, gene y3 had been cloned and used to construct plant expression plasmid.
(5) The plasmid was constructed via inserting gene y3 sequence, at the MCS of pCAMBIA1301 along with CaMV 35S promoter and NOS terminator. The expression plasmids were transferred into Nicotiana tabacumvia via Agrobacterium mediated method, resulting in transgenic plantlets. The transcription of y3 gene was characterized by Northern blot analysis. Inoculation tests on transgenic plantlets with TMV demonstrated inhibitory activity against TMV. This illustrated that y3 gene had been transcripted and expressed in transgenic tobacco plants and y3 gene was proved to have certain inhibitory function against TMV in vivo.
(6) In order to extract DNA or RNA simply and effectively, a published protocol for extracting DNA and RNA was improved and applied to isolate DNA and RNA of fungi and tobacco. Compared with commonly used method and commercial kits, the simplified protocol could be used to extract DNA and RNA not only from fungi but also from plants. The isolated DNA and RNA were stable and their activities could be kept when stored under low temperatures.
(7) A new 28S rRNA sequence with 906 bps was isolated from mycelia of C. comatus (GenBank Accession No. GU568178). The sequence was obtained from an untargeted band when we cloned antiviral protein gene y3. This sequence was used to search homologous sequences using BLAST algorithm in NCBI, and to perform cluster with its homologues using the Clustal w and MEGA methods. The results confirmed that this sequence was a new 28S rRNA, and the phylogenic relationship of C. comatus was found to be dispersed from each other. Moreover, two homologous counterparts between the sequences of new 28S rRNA and the y3 gene were the possible cause of formation of the untargeted band in PCR amplification of y3 gene. Within these two homologous counterparts, one counterpart had the sequence similar to a PCR primer for y3 gene, and the other one was a primer-like fragment. In this research, obtaining of the new sequence of 28S rRNA was a new example of the formation of untargeted bands during PCR amplification. This finding and clustering analysis results of 28S rRNA would be helpful for researches in Fungus Genomics and the establishment of their molecular classification system.
引文
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