串联不同分子佐剂的小鹅瘟病毒VP3基因疫苗的构建及其免疫原性研究
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摘要
小鹅瘟(Gosling Plague)是严重危害水禽养殖业的重要传染病之一,其防制措施主要依靠疫苗接种,传统疫苗在预防和控制小鹅瘟时,存在着散毒和免疫应答不完全等缺点。基因疫苗符合未来疫苗的发展方向,具有安全和诱导全面的免疫应答等诸多优点。本研究以小鹅瘟病毒(Goose Parvovirus,GPV)VP3为研究对象,构建了VP3单基因DNA疫苗,为了增强VP3基因疫苗的免疫原性,将鹅白介素-2(Goose Interleukin-2,gIL-2)和VP3基因进行串联融合表达,还嵌入了一段CpG免疫刺激序列,将构建的三种基因疫苗分别免疫28日龄雏鹅和种鹅,并对其诱导鹅产生的体液免疫和细胞免疫进行了比较研究,为研制高效小鹅瘟病毒VP3基因疫苗打下了基础。
1.为了探索GPV VP3基因构建基因疫苗的可行性,通过PCR扩增和基因重组技术克隆了GPV VP3基因并将其连接到pMD18T-Simple上,在对VP3基因分子特性、遗传特点和抗原性进行了分析之后,亚克隆到pcDNA3.1(+)CMV启动子下游的HindⅢ和BamHⅠ酶切位点之间,构建了单独表达GPVVP3基因的基因疫苗载体pcDNA-VP3,将其转染真核细胞COS-7后,利用间接免疫荧光抗体试验可以在转染后48h检测到VP3基因的高效表达。
2.通过重叠延伸PCR(Splicing by overlap extention PCR)扩增gIL2-VP3串联基因,包含缺失了TAA终止密码子的gIL-2基因和缺失了ATG起始密码子的VP3基因,两个基因之间以高亲水性氨基酸(Gly-Gly-Gly-Ser)编码核苷酸相连接;将gIL2-VP3融合基因通过基因重组技术正向插入到到pcDNA3.1(+)CMV启动子的下游HindⅢ和BamHⅠ酶切位点之间,构建了表达gIL2-VP3融合基因的pcDNA-gIL2-VP3基因疫苗载体,可以在COS-7细胞内高效表达,表达产物的IL-2生物活性可达31.57U/ml。pcDNA-gIL2-VP3基因疫苗载体,可以在COS-7细胞内高效表达,表达产物的IL-2生物活性可达31.57U/ml。
3.人工合成在禽体内具有免疫刺激作用的CpG序列,克隆到pMD18T-Simple载体上,然后亚克隆定向插入到pcDNA-gIL2-VP3目的基因终止密码子后的BamHⅠ和XbaⅠ酶切位点之间,构建成pcDNA-gIL2-VP3/CpG基因疫苗载体,对鹅外周淋巴细胞增殖具有较强的刺激活性(SI=3.06)。
4.将三种基因疫苗以50μg、100μg和200μg的不同剂量分别免疫28日龄鹅,分别于基因免疫后第3d、7d、14d、21d、28d、35d和49d颈静脉采集免疫鹅的抗凝血,分离外周血淋巴细胞后,利用MTT比色法检测ConA对鹅外周血淋巴细胞增殖的免疫刺激作用,发现鹅在免疫pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG后第28d体内细胞免疫最强,200μg pcDNA-gIL2-VP3/CpG免疫组外周血淋巴细胞的ConA刺激指数显著高于pcDNA-gIL2-VP3各组(P＜0.05);显著高于单基因基因疫苗pcDNA-VP3各个剂量组(P＜0.05)。含有佐剂的pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG两种基因疫苗细胞免疫的峰值比单基因VP3 DNA疫苗提前7d。
5.将三种基因疫苗以50μg、100μg和200μg不同剂量免疫28日龄鹅,分别于免疫后第3d、7d、14d、21d、35d、49d、63d、77d、105d、133d、161d、189d、217d,采用间接ELISA的方法检测鹅体内IgG的动态变化规律,结果显示,pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG所诱导的体液免疫水平明显高于pcDNA-VP3单基因组,这两种基因疫苗的各个剂量组均在第35d抗体水平达到最高,其中以200μg pcDNA-gIL2-VP3/CpG的ELISA水平最高,显著区别于pcDNA-VP3各剂量免疫组(P＜0.05)和极显著高于各个对照组(P＜0.01)。从pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG诱导鹅的体液免疫强度和维持时间来看,均优于pcDNA-VP3免疫鹅和GPV弱毒免疫鹅。
6.将三种基因疫苗以200μg的剂量免疫28日龄鹅,分别于免疫后第15d、30d、45d、60d和75d,采用微量血清中和试验检测基因免疫所诱导的中和抗体水平,结果显示,免疫后第15d就可以检测到中和抗体,鹅血清中和抗体随时间的推移稳定中有所缓慢上升,pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG免疫后第60d的鹅血清中和抗体平均效价分别达到峰值1:178.5和1:198.2,二者之间差异不显著,但是显著高于pcDNA-VP3和GPV弱毒免疫鹅(P＜0.05)。
7.将三种GPV VP3基因疫苗以200μg的剂量肌肉注射免疫开产前母鹅,分别在免疫后第7d、14d、28d、42d、56d和第70d收集免疫母鹅所产鹅蛋用作卵黄抗体的检测,另外在免疫后第14d、28d、42d取一批鹅蛋入孵用以孵化小鹅用作雏鹅攻毒保护试验。结果显示不同时间卵黄抗体和中和抗体水平与雏鹅攻毒保护率呈现一定的正相关性,三种基因疫苗以pcDNA-gIL2-VP3/CpG效果最好,具有类似于GPV弱毒苗的免疫保护效果。
Goose parvovirus(GPV)infection is still a great threat for waterfowl industry in waterfowl breeding countries,its main prevention depends on the vaccination,but common vaccines usually have some defficincies,such as back to virulence or incomplete immune responses.Gene vaccine meets the future of vaccine development and could induce safe,low cost and complete immune responses in animals.This study took GPV VP3 gene as target gene for DNA vaccine construction,taken goose interleukin-2(gIL-2)and CpG motifs as adjuvant to improve the immunogenicity of GPV VP3 gene vaccines.
1.In order to study the feasibility of VP3 gene vaccine,we firstly cloned VP3 gene using PCR,and then cloned the VP3 gene into pMD 18T-Simple T vector.VP3 gene was analylized its molecular character and antigenicity after sequencing,then subcloned into pcDNA3.1(+)for construction of pcDNA-VP3,it coube be expressed in COS-7 at 48h post transfection using indirect immunofluorescence antibody test.
2.Using splicing by overlap extention PCR(SOE-PCR),gIL2-VP3 fusion gene was acquired,which lacked TAA stop codon of gIL2 gene and ATG iniation codon of VP3 gene,the two genes were linked by high hydrophilia amino acid(Gly-Gly-Gly-Ser) encoding nucleotides,gIL2-VP3 gene was then inserted into pcDNA3.1(+)under the control of CMV,pcDNA-gIL2-VP3 was constructed,which could be expressed in COS-7 cells 48h post transfection and its expression protein had bioactivity of IL-2(31.57U/ml).
3.CpG ODN had been systhesized and ligated into pMD18T-Simple,then subcloned to pcDNA-gIL2-VP3,thus pcDNA-gIL2-VP3/CpG was constructed,which could stimulate the proliferation of PBMC in goose,the stimulation index was 3.06.
4.28-day-old geese were immunized with three kinds of VP3 gene vaccine with different doses of 50μg,100μg and 200μg,anticoagulated blood was prepared and used for lymphocytes proliferation assay at 3d,7d,14d,21d,28d,35d and 49d post immunizaiton,the results indicated that,at 28d post immunization,pcDNA-gIL2-VP3 and pcDNA-gIL2-VP3/CpG had the highest values,stimulation index(SI)of pcDNA-gIL2-VP3/CpG immunized with 200μg doses was significantly different with pcDNA-gIL2-VP3 and pcDNA-VP3(P＜0.05).VP3 gene vaccine with adjuvant showed earlier peak of T lymphocytes proliferation abilities than pcDNA-VP3.
5.Serum samples were collected from immunized geese at 3,7,14,21,28,35,49,63, 77,105,133,161,189,217 days post immunization,and indirect ELISA was used to detect the kinetics of IgG,the results indicated that,pcDNA-gIL2-VP3 and pcDNA-gIL2-VP3/CpG induced superior humoral immunity than pcDNA-VP3,200μg of pcDNA-gIL2-VP3/CpG induced the highest level of IgG,which was significantly different from pcDNA-VP3(P＜0.05)and other control groups(P＜0.01),VP3 gene vaccines possessed adjuvant induced better and longer immune responses than pcDNA-VP3 and GPV live attenuated vaccine group.
6.28-day-old geese were immunized with 200μg each kind of gene vaccine,sera were collected for virus neutralization test at 15d,30d,45d,60d and 75d post immunization, the results indicated that,low levels of neutralization antibodies could be detected at 15d post immunization,then the neutralizaiotn in each gene vaccine group climbed slowly, pcDNA-gIL2-VP3 and pcDNA-gIL2-VP3/CpG peaked at 60d post immunization with titers of 1:178.5 and 1:198.2,there was no significant difference between them,but they were significantly different with pcDNA-VP3 group and GPV live attenuated vaccine group(P＜0.05).
7.Three kinds of GPV VP3 gene vaccine were delivered into maternao geese with doses of 200μg each vaccine,yolk antibody and yolk neutralizing antibody were tested at 7d,14d,28d,42d,56d and 70d post immunization,geese eggs collected at 14d,28d and 42d post immunization were hatched for goslings used for virus protection assay.The results indicated that,yolk antibodies and yolk neutralizing antibodies at different times post immunization showed some correlations with the gosling virus protection test, pcDNA-gIL2-VP3/CpG was superior in the three gene vaccines and had the similar results with GPV-live attenuated vaccine.
1.为了探索GPV VP3基因构建基因疫苗的可行性,通过PCR扩增和基因重组技术克隆了GPV VP3基因并将其连接到pMD18T-Simple上,在对VP3基因分子特性、遗传特点和抗原性进行了分析之后,亚克隆到pcDNA3.1(+)CMV启动子下游的HindⅢ和BamHⅠ酶切位点之间,构建了单独表达GPVVP3基因的基因疫苗载体pcDNA-VP3,将其转染真核细胞COS-7后,利用间接免疫荧光抗体试验可以在转染后48h检测到VP3基因的高效表达。
2.通过重叠延伸PCR(Splicing by overlap extention PCR)扩增gIL2-VP3串联基因,包含缺失了TAA终止密码子的gIL-2基因和缺失了ATG起始密码子的VP3基因,两个基因之间以高亲水性氨基酸(Gly-Gly-Gly-Ser)编码核苷酸相连接;将gIL2-VP3融合基因通过基因重组技术正向插入到到pcDNA3.1(+)CMV启动子的下游HindⅢ和BamHⅠ酶切位点之间,构建了表达gIL2-VP3融合基因的pcDNA-gIL2-VP3基因疫苗载体,可以在COS-7细胞内高效表达,表达产物的IL-2生物活性可达31.57U/ml。pcDNA-gIL2-VP3基因疫苗载体,可以在COS-7细胞内高效表达,表达产物的IL-2生物活性可达31.57U/ml。
3.人工合成在禽体内具有免疫刺激作用的CpG序列,克隆到pMD18T-Simple载体上,然后亚克隆定向插入到pcDNA-gIL2-VP3目的基因终止密码子后的BamHⅠ和XbaⅠ酶切位点之间,构建成pcDNA-gIL2-VP3/CpG基因疫苗载体,对鹅外周淋巴细胞增殖具有较强的刺激活性(SI=3.06)。
4.将三种基因疫苗以50μg、100μg和200μg的不同剂量分别免疫28日龄鹅,分别于基因免疫后第3d、7d、14d、21d、28d、35d和49d颈静脉采集免疫鹅的抗凝血,分离外周血淋巴细胞后,利用MTT比色法检测ConA对鹅外周血淋巴细胞增殖的免疫刺激作用,发现鹅在免疫pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG后第28d体内细胞免疫最强,200μg pcDNA-gIL2-VP3/CpG免疫组外周血淋巴细胞的ConA刺激指数显著高于pcDNA-gIL2-VP3各组(P＜0.05);显著高于单基因基因疫苗pcDNA-VP3各个剂量组(P＜0.05)。含有佐剂的pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG两种基因疫苗细胞免疫的峰值比单基因VP3 DNA疫苗提前7d。
5.将三种基因疫苗以50μg、100μg和200μg不同剂量免疫28日龄鹅,分别于免疫后第3d、7d、14d、21d、35d、49d、63d、77d、105d、133d、161d、189d、217d,采用间接ELISA的方法检测鹅体内IgG的动态变化规律,结果显示,pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG所诱导的体液免疫水平明显高于pcDNA-VP3单基因组,这两种基因疫苗的各个剂量组均在第35d抗体水平达到最高,其中以200μg pcDNA-gIL2-VP3/CpG的ELISA水平最高,显著区别于pcDNA-VP3各剂量免疫组(P＜0.05)和极显著高于各个对照组(P＜0.01)。从pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG诱导鹅的体液免疫强度和维持时间来看,均优于pcDNA-VP3免疫鹅和GPV弱毒免疫鹅。
6.将三种基因疫苗以200μg的剂量免疫28日龄鹅,分别于免疫后第15d、30d、45d、60d和75d,采用微量血清中和试验检测基因免疫所诱导的中和抗体水平,结果显示,免疫后第15d就可以检测到中和抗体,鹅血清中和抗体随时间的推移稳定中有所缓慢上升,pcDNA-gIL2-VP3和pcDNA-gIL2-VP3/CpG免疫后第60d的鹅血清中和抗体平均效价分别达到峰值1:178.5和1:198.2,二者之间差异不显著,但是显著高于pcDNA-VP3和GPV弱毒免疫鹅(P＜0.05)。
7.将三种GPV VP3基因疫苗以200μg的剂量肌肉注射免疫开产前母鹅,分别在免疫后第7d、14d、28d、42d、56d和第70d收集免疫母鹅所产鹅蛋用作卵黄抗体的检测,另外在免疫后第14d、28d、42d取一批鹅蛋入孵用以孵化小鹅用作雏鹅攻毒保护试验。结果显示不同时间卵黄抗体和中和抗体水平与雏鹅攻毒保护率呈现一定的正相关性,三种基因疫苗以pcDNA-gIL2-VP3/CpG效果最好,具有类似于GPV弱毒苗的免疫保护效果。
Goose parvovirus(GPV)infection is still a great threat for waterfowl industry in waterfowl breeding countries,its main prevention depends on the vaccination,but common vaccines usually have some defficincies,such as back to virulence or incomplete immune responses.Gene vaccine meets the future of vaccine development and could induce safe,low cost and complete immune responses in animals.This study took GPV VP3 gene as target gene for DNA vaccine construction,taken goose interleukin-2(gIL-2)and CpG motifs as adjuvant to improve the immunogenicity of GPV VP3 gene vaccines.
1.In order to study the feasibility of VP3 gene vaccine,we firstly cloned VP3 gene using PCR,and then cloned the VP3 gene into pMD 18T-Simple T vector.VP3 gene was analylized its molecular character and antigenicity after sequencing,then subcloned into pcDNA3.1(+)for construction of pcDNA-VP3,it coube be expressed in COS-7 at 48h post transfection using indirect immunofluorescence antibody test.
2.Using splicing by overlap extention PCR(SOE-PCR),gIL2-VP3 fusion gene was acquired,which lacked TAA stop codon of gIL2 gene and ATG iniation codon of VP3 gene,the two genes were linked by high hydrophilia amino acid(Gly-Gly-Gly-Ser) encoding nucleotides,gIL2-VP3 gene was then inserted into pcDNA3.1(+)under the control of CMV,pcDNA-gIL2-VP3 was constructed,which could be expressed in COS-7 cells 48h post transfection and its expression protein had bioactivity of IL-2(31.57U/ml).
3.CpG ODN had been systhesized and ligated into pMD18T-Simple,then subcloned to pcDNA-gIL2-VP3,thus pcDNA-gIL2-VP3/CpG was constructed,which could stimulate the proliferation of PBMC in goose,the stimulation index was 3.06.
4.28-day-old geese were immunized with three kinds of VP3 gene vaccine with different doses of 50μg,100μg and 200μg,anticoagulated blood was prepared and used for lymphocytes proliferation assay at 3d,7d,14d,21d,28d,35d and 49d post immunizaiton,the results indicated that,at 28d post immunization,pcDNA-gIL2-VP3 and pcDNA-gIL2-VP3/CpG had the highest values,stimulation index(SI)of pcDNA-gIL2-VP3/CpG immunized with 200μg doses was significantly different with pcDNA-gIL2-VP3 and pcDNA-VP3(P＜0.05).VP3 gene vaccine with adjuvant showed earlier peak of T lymphocytes proliferation abilities than pcDNA-VP3.
5.Serum samples were collected from immunized geese at 3,7,14,21,28,35,49,63, 77,105,133,161,189,217 days post immunization,and indirect ELISA was used to detect the kinetics of IgG,the results indicated that,pcDNA-gIL2-VP3 and pcDNA-gIL2-VP3/CpG induced superior humoral immunity than pcDNA-VP3,200μg of pcDNA-gIL2-VP3/CpG induced the highest level of IgG,which was significantly different from pcDNA-VP3(P＜0.05)and other control groups(P＜0.01),VP3 gene vaccines possessed adjuvant induced better and longer immune responses than pcDNA-VP3 and GPV live attenuated vaccine group.
6.28-day-old geese were immunized with 200μg each kind of gene vaccine,sera were collected for virus neutralization test at 15d,30d,45d,60d and 75d post immunization, the results indicated that,low levels of neutralization antibodies could be detected at 15d post immunization,then the neutralizaiotn in each gene vaccine group climbed slowly, pcDNA-gIL2-VP3 and pcDNA-gIL2-VP3/CpG peaked at 60d post immunization with titers of 1:178.5 and 1:198.2,there was no significant difference between them,but they were significantly different with pcDNA-VP3 group and GPV live attenuated vaccine group(P＜0.05).
7.Three kinds of GPV VP3 gene vaccine were delivered into maternao geese with doses of 200μg each vaccine,yolk antibody and yolk neutralizing antibody were tested at 7d,14d,28d,42d,56d and 70d post immunization,geese eggs collected at 14d,28d and 42d post immunization were hatched for goslings used for virus protection assay.The results indicated that,yolk antibodies and yolk neutralizing antibodies at different times post immunization showed some correlations with the gosling virus protection test, pcDNA-gIL2-VP3/CpG was superior in the three gene vaccines and had the similar results with GPV-live attenuated vaccine.
引文
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