犬冠状病毒基因疫苗与重组腺病毒的构建、表达及免疫原性研究
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摘要
犬冠状病毒(Canine coronavirus, CCV)是引起幼犬发生胃肠炎的重要病原之一,不但对我国养犬业危害较大,同时也对一些野生动物构成较大的威胁。血清学调查发现,该病呈世界分布,在我国很多地区犬群中也时有流行,导致较高的发病率和死亡率。免疫接种是预防此病的有效途径之一,因此为控制该病的流行须研制安全有效的疫苗。本研究首次对犬冠状病毒大熊猫株(CCV DXMV)纤突蛋白(S)、膜蛋白(M) 和核蛋白(N) 三种主要结构蛋白基因进行了克隆、测序及遗传进化分析,并用DNAstar软件对DXMV 株与疫苗株Insavc-1 S 蛋白和M 蛋白疏水性、Jameson-Wolf 抗原表位和表面极性进行了分析。结果表明,DXMV 野毒株的基因型为CCV 基因II 型;与疫苗株Insavc-1 相比,两毒株S 蛋白在糖基化位点、疏水性、Jameson-Wolf 抗原表位上存在比较明显的差异;DXMV 株S 蛋白和M 蛋白Jameson-Wolf 抗原表位指数要显著高于Insavc-1 疫苗弱毒,从分子水平上揭示了DXMV 株免疫原性要好于Insavc-1 疫苗株的原因。以pVAX1 为载体成功构建了三种CCV 基因疫苗表达载体pVAXS1 、pVAXM 和pVAXN。将三种真核表达载体转染MDCK 细胞后,间接ELISA 和间接荧光染色试验检测结果表明,三种目的蛋白均能在MDCK 细胞中表达。基因疫苗载体在小鼠上免疫试验结果表明,pVAXS1、pVAXM 和pVAXN 均可诱导小鼠产生特异性免疫应答;其中pVAXS1+pVAXM+pVAXN 三个基因疫苗混合免疫小鼠可诱导产生较强的体液免疫和细胞免疫应答。以感染性犬2 型腺病毒全基因组为载体,首次构建了的含有CCV S1 基因的重组CAV-2-S1 病毒。Western-blotting 检测结果证实,该重组病毒能表达具有免疫学活性的重组CCV S 1 蛋白。体外传代试验表明,该重组病毒具有良好的遗传稳定性。根据基因疫苗在小鼠上的免疫试验结果,将基因疫苗pVAXS1+pVAXM+pVAXN 和重组病毒CAV-2-S1 在犬上进行了免疫试验。结果表明,两者均可诱导犬产生特异性体液免疫和细胞免疫。其中pVAXS1+pVAXM+pVAXN 与重组病毒联合免疫能诱导机体产生较高水平的体液和细胞免疫应答。以上试验研究为CCV 基因疫苗和重组腺病毒疫苗的研制奠定了良好的基础。
In order to clarify the novel molecular biological characteristics of canine coronavirus strain DXMV (CCV DXMV) isolated from the liver of a died giant panda, three pairs of specific primers were designed for amplification of spike protein gene, membrane protein gene and nucleoprotein gene by RT-PCR, respectively, according to the genomic sequence of CCV strain Insavc-1 published in GenBank. The PCR products were cloned into pGEM-T and sequenced. Then the nucleotide sequences of the three structural protein genes were analysed by DNAMAN and DNAstar software, respectively. The full length of spike protein gene of strain DXMV was 4362bp,which encoded 1453 amino acids. The initiative 18 amino acids were signal peptide. The membrane protein gene was 789bp, encoding of polypeptide of 263 amino acids which had a 17 amino acids constituting signal peptide. The overall length of nucleoprotein gene was 1146bp, which encoded 382 amino acids. The homologies of nucleic acid and deduced amino acid sequence of spike protein, membrane protein and nucleoprotein among the different isolates of CCV strain accessed by GenBank were 22.7-99.7% and 38.3-99.7%, 77.7-99.5% and 81.7-98.9%, 91.4-99.8% and 92.1-99.7%, respectively. The variation parts of spike gene mainly existed in front half of nucleotide sequence, especially the region 350-370nt, 439-478nt and1718-1818nt, while the region 1060-1700 showed highly conservative bases. Phylogenetic analysis based on the complete spike and membrane gene, indicated that there were two different genotypes of CCV, genotype I and genotype II. The strain DXMV, together with most classical isolates such as K378, C54, V6, V1 and Insavc-1 belonged to genotype II, whereas E1mo02 and 2303 were included into genotype I. There were high homologies among the strains of genotype II, while the low identities between the genotype I and genotype II. There are 34 potential N-glycosylation sites in deduced S protein for strain DXMV and V54, but 33 for strain Insavc-1. The N-glycosylation site 566-568 was possessed for most virulent strains but not for vaccinal strain Insavc-1. Moreover, analysis of hydrophobicity,antigenic index
and surface probability plot of spike and membrane proteins, indicated that there were some differences between strain DXMV and Insavc-1. Jameson-Wolf antigenic index of spike and membrane proteins of strain were significantly higher than that of the vaccinal strain Insavc-1, which revealed the possibly reasons that strain DXMV had better immunogenicity than that of Insavc-1 in molecular aspects. Three eukaryotic expression vectors, pVAXS1, pVAXM and pVAXN, used as gene vaccines of canine coronavirus were successfully constructed, and expressed in MDCK cell line. The S1 gene fragment of strain DXMV, which encoded major antigenic region A, B, C and D of S protein, was amplified from pTS by PCR with a pair of specific primers and cloned into pVAX1 vector. The recombinant pTM and pTN were digested by KpnI/NotI and interesting gene fragments were recovered and further subcloned into pVAX1, respectively. The three constructed expression vectors, namely pVAXS1, pVAXM and pVAXN were used for transfection in MDCK cell. Transcriptions of interesting genes were detected by RT-PCR. The interesting proteins were detected by indirect immunofluorescent technique and indirect ELISA method. The results showed that transcription of interesting genes could be detected at 36 h post-transformation, and expressed proteins could be detected by the two methods mentioned-above after 48-96 hours of transformation. In order to construct a recombinant canine adenovirus type 2 (CAV-2) to express the spike glycoprotein of canine coronavirus (CCV), complete expression cassette harboring S1 gene of CCV was subcloned into the shuttle vector pVAXE3, then further cloned into the backbone vector pPoly2-CAV2 containing complete genome of CAV-2. To gain the recombinant canine adenovirus, the recombinant plasmid pCAV-2-CCV-S1 was linearized by ClaI/AscI to release recombinant genome, and then transfected into MDCK cell. The recombinant virus CAV-2-S1 was gained through 4 times of passaging in MDCK, and the classical cytopathic effects (CPE) of CAV-2 could be observed. The recombinant S1 protein expressed by recombinant virus, which was identified by western blot analysis, could be specifically recognized by polyclonal serum against CCV. Eight groups of mice were immunized by intramuscular injection with the different combinations as following: sole expression plasmid, two expression plasmids or three expression plasmids. Indirect ELISA, serum neutralization test and lymphocyte
In order to clarify the novel molecular biological characteristics of canine coronavirus strain DXMV (CCV DXMV) isolated from the liver of a died giant panda, three pairs of specific primers were designed for amplification of spike protein gene, membrane protein gene and nucleoprotein gene by RT-PCR, respectively, according to the genomic sequence of CCV strain Insavc-1 published in GenBank. The PCR products were cloned into pGEM-T and sequenced. Then the nucleotide sequences of the three structural protein genes were analysed by DNAMAN and DNAstar software, respectively. The full length of spike protein gene of strain DXMV was 4362bp,which encoded 1453 amino acids. The initiative 18 amino acids were signal peptide. The membrane protein gene was 789bp, encoding of polypeptide of 263 amino acids which had a 17 amino acids constituting signal peptide. The overall length of nucleoprotein gene was 1146bp, which encoded 382 amino acids. The homologies of nucleic acid and deduced amino acid sequence of spike protein, membrane protein and nucleoprotein among the different isolates of CCV strain accessed by GenBank were 22.7-99.7% and 38.3-99.7%, 77.7-99.5% and 81.7-98.9%, 91.4-99.8% and 92.1-99.7%, respectively. The variation parts of spike gene mainly existed in front half of nucleotide sequence, especially the region 350-370nt, 439-478nt and1718-1818nt, while the region 1060-1700 showed highly conservative bases. Phylogenetic analysis based on the complete spike and membrane gene, indicated that there were two different genotypes of CCV, genotype I and genotype II. The strain DXMV, together with most classical isolates such as K378, C54, V6, V1 and Insavc-1 belonged to genotype II, whereas E1mo02 and 2303 were included into genotype I. There were high homologies among the strains of genotype II, while the low identities between the genotype I and genotype II. There are 34 potential N-glycosylation sites in deduced S protein for strain DXMV and V54, but 33 for strain Insavc-1. The N-glycosylation site 566-568 was possessed for most virulent strains but not for vaccinal strain Insavc-1. Moreover, analysis of hydrophobicity,antigenic index
and surface probability plot of spike and membrane proteins, indicated that there were some differences between strain DXMV and Insavc-1. Jameson-Wolf antigenic index of spike and membrane proteins of strain were significantly higher than that of the vaccinal strain Insavc-1, which revealed the possibly reasons that strain DXMV had better immunogenicity than that of Insavc-1 in molecular aspects. Three eukaryotic expression vectors, pVAXS1, pVAXM and pVAXN, used as gene vaccines of canine coronavirus were successfully constructed, and expressed in MDCK cell line. The S1 gene fragment of strain DXMV, which encoded major antigenic region A, B, C and D of S protein, was amplified from pTS by PCR with a pair of specific primers and cloned into pVAX1 vector. The recombinant pTM and pTN were digested by KpnI/NotI and interesting gene fragments were recovered and further subcloned into pVAX1, respectively. The three constructed expression vectors, namely pVAXS1, pVAXM and pVAXN were used for transfection in MDCK cell. Transcriptions of interesting genes were detected by RT-PCR. The interesting proteins were detected by indirect immunofluorescent technique and indirect ELISA method. The results showed that transcription of interesting genes could be detected at 36 h post-transformation, and expressed proteins could be detected by the two methods mentioned-above after 48-96 hours of transformation. In order to construct a recombinant canine adenovirus type 2 (CAV-2) to express the spike glycoprotein of canine coronavirus (CCV), complete expression cassette harboring S1 gene of CCV was subcloned into the shuttle vector pVAXE3, then further cloned into the backbone vector pPoly2-CAV2 containing complete genome of CAV-2. To gain the recombinant canine adenovirus, the recombinant plasmid pCAV-2-CCV-S1 was linearized by ClaI/AscI to release recombinant genome, and then transfected into MDCK cell. The recombinant virus CAV-2-S1 was gained through 4 times of passaging in MDCK, and the classical cytopathic effects (CPE) of CAV-2 could be observed. The recombinant S1 protein expressed by recombinant virus, which was identified by western blot analysis, could be specifically recognized by polyclonal serum against CCV. Eight groups of mice were immunized by intramuscular injection with the different combinations as following: sole expression plasmid, two expression plasmids or three expression plasmids. Indirect ELISA, serum neutralization test and lymphocyte
引文
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