猪戊型肝炎病毒V1蛋白单克隆抗体的制备及其抗原表位的鉴定
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摘要
戊型肝炎(Hepatitis E)是由戊型肝炎病毒(Hepatitis E virus,HEV)引起的急性自限性疾病。其临床特征为发热、全身疲乏、食欲不振、厌油腻、恶心、呕吐、尿色深黄如浓茶、眼睛和皮肤发黄,肝功能检查出现转氨酶迅速升高到几百甚至几千单位,黄疸较常见,并可持续1周。在亚洲、非洲和美洲等发展中国家人群中发生比较普遍,在一些地区可占到急性病毒性肝炎的50%,是导致发病和病死的重要因素,尤其对孕妇可造成有20%的病死率。由于在灵长类、啮齿类以及各种家禽家畜体内也检测到HEV抗体,提示其可能为一种人畜共患病。HEV过去曾被认为是一种经肠道传播的非甲-非乙型肝炎病毒。Balagan等用免疫电镜技术从一名志愿受试者粪便中观察到直径为27~30nm的病毒样颗粒,从而证实该病毒为新型的非甲非乙型肝炎病毒,1989年东京国际会议正式将这一类型的肝炎及其相关病毒分别命名为戊型肝炎和戊型肝炎病毒。HEV基因组为单股正链RNA,约7.5kb,具有3个开放读码框(ORF)。病毒基因组的5′端到3′端依次为5′端非编码区(5′-NCR)、ORF1、ORF3、ORF2、3′-NCR及polyA尾。
HEV全球分布,危害严重。虽然HEV诊断方法不断得到改进,但是安全有效疫苗的研制是控制HEV感染、预防HE发生的根本措施。明确HEV抗原表位是开发HEV诊断试剂和研制疫苗的基础。国内外曾就重组蛋白和合成肽对HEV ORF2编码结构蛋白的免疫原性做了大量研究,发现在其羧基端集中了具有较强免疫反应性的主要抗原表位。ORF2编码的衣壳蛋白的上抗原表位数量多,结构复杂,其富含疏水区的C端2/3部分存在多个免疫优势B细胞表位,包括能诱导中和抗体的抗原结合位点。因此本研究使用的免疫原是原核表达的含有swDQ株HEV ORF2 C端220个氨基酸残基的V1蛋白,制备了6株针对V1蛋白的单抗,而通过单克隆抗体技术对HEV抗原表位进行深入研究,不仅有助于了解HEV抗原结构与功能的关系,而且对该病的诊断以及设计安全有效的基因工程表位疫苗等具有重要的意义。本研究首先构建了含swDQ株HEV V1蛋白的原核表达载体pET32a-V1,并在大肠杆菌中进行了表达,利用HEV感染猪血清对表达产物进行Western blot分析,证实表达的融合蛋白都具有较好的反应活性。然后以表达的融合蛋白为抗原,免疫BAL B/c小鼠,最终共获得了6株针对V1蛋白的单抗。利用肽扫描技术对6株V1蛋白的单克隆抗体(αC11、αC12、γH1、γF8、BC4、CH8)的抗原表位进行了研究。
将V1蛋白基因分为两个相互重叠的基因片段M1和M2,对6株抗V1蛋白的单抗进行鉴定,结果显示有3株单抗既能特异性识别M1又能识别M2的表达产物,说明这三株单抗所针对的表位位于M1和M2的重叠区,即位于ORF2 492~500aa (HEV ORF2编码的病毒结构蛋白的492~500aa);其余3株单抗中有两株单抗仅能识别M1的表达产物,另外一株单抗仅能识别M2的表达产物。将片段M1(386-492aa)和M2(500-606aa)人工合成一套彼此重叠8个氨基酸长度为16个氨基酸的25个短肽,融合表达后与MAbs进行Western blot和ELISA反应性扫描,对3株单抗鉴定结果显示,一株单抗与融合多肽E4和E5反应而不与其他短肽反应,说明这株单抗所针对的表位位于E4与E5的重叠区ORF2 418~425aa(HEV ORF2编码的病毒结构蛋白的418~425aa);有一株单抗能特异性识别融合多肽E1而不识别融合多肽E2,说明这株单抗所针对的表位位于E1与E2的不重叠区ORF2 386~394aa(HEV ORF2编码的病毒结构蛋白的386~394aa);另外一株单抗与融合多肽E24和E25反应而不与其他短肽反应,说明这株单抗所针对的表位位于E24与E25的重叠区ORF2 588~595aa(HEV ORF2编码的病毒结构蛋白的588~595aa)。其中,表位418~425aa和表位492~500aa为首次发现。应用纯化的表位融合蛋白对小鼠进行免疫,结果表明4个融合蛋白均能诱导小鼠产生针对短肽的特异性抗体。将纯化的表位融合蛋白进行SDS-PAGE电泳,与抗swDQ株HEV猪血清进行Western blot分析,结果表明这四个表位融合蛋白具有较好的反应原性。
本研究对swDQ株HEV V1蛋白进行抗原表位的鉴定,将有助于进一步阐明HEV的结构与功能,将为建立以表位为基础的抗原抗体诊断方法和基于表位疫苗的设计研究提供重要的研究工作基础。
Hepatitis E is a self-limited acute illness which caused by Hepatitis E virus (HEV). The illness may be particularly severe among pregnant women, with mortality rates reaching as high as 20%. Human healthy will be severe harmed. The antibody against HEV is widely detected in the body of primates, rodents, domestic animals and poultry, so it is considered as a possible diseases contracted commonly by both human beings and livestockHepatitis E virus (HEV) was previously considered an enterically transmitted non-A, non-B hepatitis virus. Using an immunoelectronmicroscopic technique. Balagan et al observed a type of particles 27 to 30 min diameter in the stool sample of a volunteer subject. They further proved this virus was a new non-A, non-B type that represents a serious threat to human health. Reyes et al cloned the genome of this virus. The virus and the hepatitis caused by the virus were named as HEV and hepatitis E, respectively, at an international conference held in Tokyo Japan in 1989. The HEV genome is a single-stranded, positive-sense RNA molecule of approximately 7.5 kb. The viral genome contains three open reading frames (ORFs) flanked by two non-coding regions (NCRs) at the 5′- and 3′-ends. The order of the sequence is 5′-NCR, ORF1, ORF2, ORF3, 3′-NCR and a polyadenylated tail.
HEV is found all around the world and causes great damage. Although methods of HEV diagnosis were always improved, the primary measure of control and prevention is to develop special vaccine. The basis of HEV diagnosis reagent and vaccine is making clear the HEV antigenic epitopes. The comprehensive studies have been conducted to further elaborate the antigenic composition of the ORF2-encoded protein with recombinant proteins and synthetic peptides, confirming the existence of a strong antigenic region located at the C terminus. The ORF2 protein existed many antigenic epitopes, complex structure, and was riched in B-cell antigenic epitopes in approximately two thirth c-terminal region of the ORF2 protein.In this research, the immunogenic antigen was ORF2-V1pritein ,which was approximately 220aa c-terminal region of the ORF2 protein. Six hybridoma cell lines designated steadily secreting monoclonal antibody (McAb) against ORF2-V1 protein of swine Hepatitis E virus were obtained by hybridomatechnology. Antigenic epitope information of the virus is not only useful for investigating the relationship between antigenic structure and function of the HEV but also useful for diagnosis of HEV infection and developing safe and effective multi-epitope vaccine against the disease.
In this study, recombinant plasmids harboring structural protein of HEV swDQ strain designated as pET32a-V1, were constructed and expressed in E.coli. The expressed fusion proteins were detected with sera of HEV infected pigs by Western-blotting. BALB/c mice were immunized with expressed fusion proteins and monoclonal antibodies (MAbs) were developed. 6 anti-ORF2-V1 MAbs were identified. This study identified the antigenic epitopes of 6 McAbs to HEV ORF2-V1(αC11、αC12、γH1、γF8、BC4、CH8) with pep scan.The ORF2-V1 gene were divided into two overlapping fragments and expressed in E. coli respectively. The reactivity of different fragments expressed in E. coli were probed with 6 anti-ORF2-V1 MAbs by Western-blot. Three out of 6 MAbs could react with ORF2-M1 and ORF2-M2 fusion Proteins. The results showed that both fragments could react with MAbsαC11、αC12andγF8 , suggesting that the epitope recognized by MAbsαC11、αC12andγF8 located in the overlaping region (ORF2 492-500aa) of the two fragments. The ORF2-M1 protein(386-492aa)and ORF2-M2 protein(500-606aa)was dissected into 25 overlapping fragments, expressed as fusion products in E. coli, and used for epitope mapping by pepscan analysis. Three antigenic epitopes were identified by ELISA and Western blot assays. MAb CH8 could react with E4 and E5 short synthetic peptides. The results showed that the epitope recognized by MAb CH8 located in the overlapingregion(ORF2 418~425aa)of the two fragments. MAbγH1 could only react with E1 short synthetic peptides,and could not react with other short synthetic peptides. The results showed that the epitope recognized by MAbγH1 located in the HEV ORF2-encoded protein(ORF2 386~394aa). MAb BC4 could react with E24 and E25 short synthetic peptides. The results showed that the epitope recognized by MAb BC4located in the overlapingregion(ORF2 588~595aa)of the two fragments. The epitope 418~425aa and the epitope492-500aa were identified first.Immunization of mice with each of the four antigenic epitope-fused proteins revealed that all four proteins could elicit short peptide specific antisera.The identified epitopes were synthesized as polypeptides. SDS-PAGE analysis showed that the four antigenic epitope-fused proteins was expressed in E.coli B121(DE3)and identified positive antisera of the swine by the Western-blot.
Identification of HEV envelope protein antigenic epitopes may provide the basis for the development of immunity-based prophylactic, therapeutic, and diagnostic clinical techniques for HEV, and further structural and function analysis of envelope protein.
HEV全球分布,危害严重。虽然HEV诊断方法不断得到改进,但是安全有效疫苗的研制是控制HEV感染、预防HE发生的根本措施。明确HEV抗原表位是开发HEV诊断试剂和研制疫苗的基础。国内外曾就重组蛋白和合成肽对HEV ORF2编码结构蛋白的免疫原性做了大量研究,发现在其羧基端集中了具有较强免疫反应性的主要抗原表位。ORF2编码的衣壳蛋白的上抗原表位数量多,结构复杂,其富含疏水区的C端2/3部分存在多个免疫优势B细胞表位,包括能诱导中和抗体的抗原结合位点。因此本研究使用的免疫原是原核表达的含有swDQ株HEV ORF2 C端220个氨基酸残基的V1蛋白,制备了6株针对V1蛋白的单抗,而通过单克隆抗体技术对HEV抗原表位进行深入研究,不仅有助于了解HEV抗原结构与功能的关系,而且对该病的诊断以及设计安全有效的基因工程表位疫苗等具有重要的意义。本研究首先构建了含swDQ株HEV V1蛋白的原核表达载体pET32a-V1,并在大肠杆菌中进行了表达,利用HEV感染猪血清对表达产物进行Western blot分析,证实表达的融合蛋白都具有较好的反应活性。然后以表达的融合蛋白为抗原,免疫BAL B/c小鼠,最终共获得了6株针对V1蛋白的单抗。利用肽扫描技术对6株V1蛋白的单克隆抗体(αC11、αC12、γH1、γF8、BC4、CH8)的抗原表位进行了研究。
将V1蛋白基因分为两个相互重叠的基因片段M1和M2,对6株抗V1蛋白的单抗进行鉴定,结果显示有3株单抗既能特异性识别M1又能识别M2的表达产物,说明这三株单抗所针对的表位位于M1和M2的重叠区,即位于ORF2 492~500aa (HEV ORF2编码的病毒结构蛋白的492~500aa);其余3株单抗中有两株单抗仅能识别M1的表达产物,另外一株单抗仅能识别M2的表达产物。将片段M1(386-492aa)和M2(500-606aa)人工合成一套彼此重叠8个氨基酸长度为16个氨基酸的25个短肽,融合表达后与MAbs进行Western blot和ELISA反应性扫描,对3株单抗鉴定结果显示,一株单抗与融合多肽E4和E5反应而不与其他短肽反应,说明这株单抗所针对的表位位于E4与E5的重叠区ORF2 418~425aa(HEV ORF2编码的病毒结构蛋白的418~425aa);有一株单抗能特异性识别融合多肽E1而不识别融合多肽E2,说明这株单抗所针对的表位位于E1与E2的不重叠区ORF2 386~394aa(HEV ORF2编码的病毒结构蛋白的386~394aa);另外一株单抗与融合多肽E24和E25反应而不与其他短肽反应,说明这株单抗所针对的表位位于E24与E25的重叠区ORF2 588~595aa(HEV ORF2编码的病毒结构蛋白的588~595aa)。其中,表位418~425aa和表位492~500aa为首次发现。应用纯化的表位融合蛋白对小鼠进行免疫,结果表明4个融合蛋白均能诱导小鼠产生针对短肽的特异性抗体。将纯化的表位融合蛋白进行SDS-PAGE电泳,与抗swDQ株HEV猪血清进行Western blot分析,结果表明这四个表位融合蛋白具有较好的反应原性。
本研究对swDQ株HEV V1蛋白进行抗原表位的鉴定,将有助于进一步阐明HEV的结构与功能,将为建立以表位为基础的抗原抗体诊断方法和基于表位疫苗的设计研究提供重要的研究工作基础。
Hepatitis E is a self-limited acute illness which caused by Hepatitis E virus (HEV). The illness may be particularly severe among pregnant women, with mortality rates reaching as high as 20%. Human healthy will be severe harmed. The antibody against HEV is widely detected in the body of primates, rodents, domestic animals and poultry, so it is considered as a possible diseases contracted commonly by both human beings and livestockHepatitis E virus (HEV) was previously considered an enterically transmitted non-A, non-B hepatitis virus. Using an immunoelectronmicroscopic technique. Balagan et al observed a type of particles 27 to 30 min diameter in the stool sample of a volunteer subject. They further proved this virus was a new non-A, non-B type that represents a serious threat to human health. Reyes et al cloned the genome of this virus. The virus and the hepatitis caused by the virus were named as HEV and hepatitis E, respectively, at an international conference held in Tokyo Japan in 1989. The HEV genome is a single-stranded, positive-sense RNA molecule of approximately 7.5 kb. The viral genome contains three open reading frames (ORFs) flanked by two non-coding regions (NCRs) at the 5′- and 3′-ends. The order of the sequence is 5′-NCR, ORF1, ORF2, ORF3, 3′-NCR and a polyadenylated tail.
HEV is found all around the world and causes great damage. Although methods of HEV diagnosis were always improved, the primary measure of control and prevention is to develop special vaccine. The basis of HEV diagnosis reagent and vaccine is making clear the HEV antigenic epitopes. The comprehensive studies have been conducted to further elaborate the antigenic composition of the ORF2-encoded protein with recombinant proteins and synthetic peptides, confirming the existence of a strong antigenic region located at the C terminus. The ORF2 protein existed many antigenic epitopes, complex structure, and was riched in B-cell antigenic epitopes in approximately two thirth c-terminal region of the ORF2 protein.In this research, the immunogenic antigen was ORF2-V1pritein ,which was approximately 220aa c-terminal region of the ORF2 protein. Six hybridoma cell lines designated steadily secreting monoclonal antibody (McAb) against ORF2-V1 protein of swine Hepatitis E virus were obtained by hybridomatechnology. Antigenic epitope information of the virus is not only useful for investigating the relationship between antigenic structure and function of the HEV but also useful for diagnosis of HEV infection and developing safe and effective multi-epitope vaccine against the disease.
In this study, recombinant plasmids harboring structural protein of HEV swDQ strain designated as pET32a-V1, were constructed and expressed in E.coli. The expressed fusion proteins were detected with sera of HEV infected pigs by Western-blotting. BALB/c mice were immunized with expressed fusion proteins and monoclonal antibodies (MAbs) were developed. 6 anti-ORF2-V1 MAbs were identified. This study identified the antigenic epitopes of 6 McAbs to HEV ORF2-V1(αC11、αC12、γH1、γF8、BC4、CH8) with pep scan.The ORF2-V1 gene were divided into two overlapping fragments and expressed in E. coli respectively. The reactivity of different fragments expressed in E. coli were probed with 6 anti-ORF2-V1 MAbs by Western-blot. Three out of 6 MAbs could react with ORF2-M1 and ORF2-M2 fusion Proteins. The results showed that both fragments could react with MAbsαC11、αC12andγF8 , suggesting that the epitope recognized by MAbsαC11、αC12andγF8 located in the overlaping region (ORF2 492-500aa) of the two fragments. The ORF2-M1 protein(386-492aa)and ORF2-M2 protein(500-606aa)was dissected into 25 overlapping fragments, expressed as fusion products in E. coli, and used for epitope mapping by pepscan analysis. Three antigenic epitopes were identified by ELISA and Western blot assays. MAb CH8 could react with E4 and E5 short synthetic peptides. The results showed that the epitope recognized by MAb CH8 located in the overlapingregion(ORF2 418~425aa)of the two fragments. MAbγH1 could only react with E1 short synthetic peptides,and could not react with other short synthetic peptides. The results showed that the epitope recognized by MAbγH1 located in the HEV ORF2-encoded protein(ORF2 386~394aa). MAb BC4 could react with E24 and E25 short synthetic peptides. The results showed that the epitope recognized by MAb BC4located in the overlapingregion(ORF2 588~595aa)of the two fragments. The epitope 418~425aa and the epitope492-500aa were identified first.Immunization of mice with each of the four antigenic epitope-fused proteins revealed that all four proteins could elicit short peptide specific antisera.The identified epitopes were synthesized as polypeptides. SDS-PAGE analysis showed that the four antigenic epitope-fused proteins was expressed in E.coli B121(DE3)and identified positive antisera of the swine by the Western-blot.
Identification of HEV envelope protein antigenic epitopes may provide the basis for the development of immunity-based prophylactic, therapeutic, and diagnostic clinical techniques for HEV, and further structural and function analysis of envelope protein.
引文
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