副猪嗜血杆菌重要免疫原性相关蛋白发掘及GAPDH免疫调节机理研究
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摘要
副猪嗜血杆菌能够引起猪的多发性浆膜炎,关节炎和脑膜炎,已成为引起猪细菌性疾病最重要病原之一。仔猪感染副猪嗜血杆菌之后几天就可以发病,并经常引起死亡。从急性期发病康复的仔猪经常会出现慢性关节炎症状。抗生素治疗副猪嗜血杆菌有一定效果。但长期使用一方面导致细菌耐药情况严重,另一方面抗生素的滥用导致药物在动物食品的药物残留。因此,疫苗仍然是控制该病原菌最佳的手段。
到目前为止,副猪嗜血杆菌共鉴定出15种血清型,每种血清型菌株的毒力又有差异。此外,还有很多菌株没有被定型。各种血清型之间的交叉免疫保护十分有限。当前使用的商业化灭活疫苗能够对同源血清型菌株感染提供一定的免疫保护,但不能提供异源血清型菌株感染的交叉保护。因此,急需研制开发出一种能够提供持久免疫保护和针对多种血清型菌株感染具有交叉免疫保护的新型疫苗。
目前,关于副猪嗜血杆菌毒力相关因子很少被报道。外膜蛋白是细菌的表面重要组成部分,能够和宿主细胞发生互作,并诱导宿主产生保护性抗体和引起细胞免疫反应。因此针对靶向外膜蛋白发展疫苗非常有价值。本课题在实验室前期完成的副猪嗜血杆菌流行优势血清5型SH0165菌株全基因组序列基础上,基于生物信息学及SignalP2.0软件,预测了副猪嗜血杆菌可能的重要外膜蛋白。进一步将编码外膜蛋白基因克隆、表达纯化后利用动物感染模型进行了蛋白免疫原性研究。同时,开展了外膜蛋白的免疫生物学活性研究,并从中筛选到了一个能够促进B细胞和T细胞增值的蛋白3-磷酸甘油醛脱氢酶(GAPDH),可能是一个重要免疫调节蛋白,深入探讨了该免疫调节蛋白免疫调节的机理。构建了编码GAPDH蛋白的DNA疫苗,并开展了相关免疫原性及免疫保护研究。主要的研究内容包括如下：
1.副猪嗜血杆菌SH0165菌株可能外膜蛋白预测、蛋白免疫原性和保护效力研究
本研究首先利用SignalP2.0软件,通过同源相似性比对预测了cdtB, GAPDH, HPS-2037, OmpP1, HPS-0220, HPS-0675, HPS-1406,HPS-1834, OapA, IdgA等10个可能的外膜蛋白,克隆表达并纯化了10个外膜蛋白。将10种重组外膜蛋白免疫小鼠后28天进行了5×LD50剂量的攻击。结果表明, rGAPDH, rOapA和rHPS-0675三种重组蛋白具有很好的免疫保护效果,分别为75%,87.5%和75%。进一步将这三种蛋白单独或混合形式以猪为感染模型进行了免疫反应和保护效力研究。结果表明,三种蛋白都能够引起很好的体液免疫反应并分别提供70%,50%和70%的免疫保护,而混合蛋白免疫提供的免疫保护力最高,为80%。而3种蛋白产生的抗体和混合蛋白的抗体在全血杀菌试验中都能够抑制细菌的生长。研究结果进一步提示rGAPDH, rOapA和rHPS-0675重组蛋白能够介导针对副猪嗜血杆菌SH0165菌株感染的免疫保护,是重要的疫苗候选抗原,能够研制开发成一种多组分通用型亚单位疫苗。
2.副猪嗜血杆菌3-磷酸甘油醛脱氢酶DNA疫苗的构建及免疫效力研究
研究结果表明编码3-磷酸甘油醛脱氢酶的gapA基因广泛存在于15种标准血清型菌株中而且基因序列比较保守,前期研究也显示其是一种很好的疫苗候选抗原。因此我们进一步构建了GAPDH蛋白的DNA疫苗pCgap,并在小鼠感染模型中评价了该DNA疫苗免疫反应和针对血清4型菌株MD0322和血清5型菌株SH0165的攻击所提供的免疫效力。结果表明,DNA疫苗通过肌内免疫小鼠之后能够产生显著的抗体反应。而且产生的抗体具有杀菌作用。DNA疫苗免疫小鼠7天之后能够在肌肉、肝脏、脾脏和肾脏中检测到转录产物。IgG亚类分析表明DNA疫苗可以同时诱导Thl和Th2型免疫反应,但是IgGl型免疫反应强于IgG2a型免疫反应。此外,DNA疫苗能够提供分别针对血清4型菌株MD0322攻击的保护率83.3%和血清5型菌株SH0165攻击的保护率50%。免疫组免疫保护力显著高于阴性对照组和空白对照组。以上结果提示DNA疫苗pCgap为控制副猪嗜血杆菌的感染提供一个新的策略。
3.副猪嗜血杆菌3-磷酸甘油醛脱氢酶(GAPDH)免疫调节机理研究
研究表明,一些外膜蛋白通过修改宿主的免疫系统来影响细菌在体内定植能力并且可以刺激宿主产生IL-2或者IL-10的能力。因此,本研究探讨副猪嗜血杆菌外膜蛋白是否在参与调节宿主的免疫反应方面起着重要作用。通过将12种外膜蛋白(cdtB, GAPDH, HPS.2037, OmpP1, HPS-0220, HPS-0675, HPS.1406, HPS.1834, OapA,6PGD, HPS.1889, IdgA)体外刺激小鼠脾淋巴细胞,从中筛选到了外膜蛋白GAPDH能够显著上调脾脏细胞表面CD69分子的表达。通过MTT实验结果也表明GAPDH蛋白能够促进脾脏淋巴细胞的增值和B细胞的分化。进一步构建了GAPDH过表达菌株GAO,开展了过表达菌株小鼠体内定植、诱导宿主产生细胞因子、中性粒细胞所介导的杀伤和对宿主免疫相关基因表达的影响等实验。结果表明：GAPDH能够促进机体对副猪嗜血杆菌的清除和诱导机体产生IL-2。与野生菌株相比,过表达菌株GAO更容易被中性粒细胞和小鼠的巨噬细胞所杀伤。而且,机体感染过表达菌株GAO之后能够促进所检测的除了MHC Ⅱα分子以外IFN-γ, MHC Ⅰα, TCR1和TLR3分子免疫相关基因上调表达。总之,GAPDH蛋白参与了宿主免疫系统的调节,增强了宿主的免疫反应和对外源病原菌的清除能力。
Haemophilus parasuis (H. parasuis), the causative agent of swine polyserositis, polyarthritis, and meningitis, is one of the most important bacterial diseases of pigs worldwide. In naive pigs disease onset occurs few days after H. parasuis exposure and is usually lethal. These animals recovering from the acute phase of the disease can develop chronic arthritis.
To date,15serovars of H. parasuis, which typically have a range of different virulence potentials, have been described. In addition, a high percentage of strains are non-serotypeable. To date, only a few virulence related factors have been reported in relation to the pathogenicity of Glasser's disease. Limited cross-protection among strains has complicated the control of Glasser's disease. Maternally derived antibodies also have an important role in protection, allowing the colonization of piglets and the development of a protective immune response without the induction of disease. Animals protected against H. parasuis infection after immunization with a bacterin may develop antibodies against outer membrane proteins (OMPs). The currently commercially available inactivated vaccine confers protection against homologous challenge, but further research is needed to develop a new generation of vaccine that can stimulate long-term immunity and provide cross-protection against infections by several H. parasuis serovars.
The sequencing of H. parasuis SH0165has been completed by our group. The current knowledge of the genomic sequence of pathogenic bacteria has facilitated recent vaccine development. Because OMPs are the primary bacterial components that interact with host cells, targeting OMPs for the development of recombinant vaccines is of value. Based on this, we focused on putative OMPs of H. parasuis which were predicted by SignalP2.0software. They were cloned, expressed, and purified as HIS fusion proteins and screened for protective efficacy in a piglet model of infection. Furthermore, we have studied the immune biological activity of the OMPs and constructed a DNA vaccine to prevent and control the infection of H. parasuis. Below is detailed information on this study:
1. Immunogenicity and protective efficacy of recombinant Haemophilus parasuis SH0165putative outer membrane proteins as vaccine candidates
H. parasuis putative outer membrane proteins (OMPs) are potentially essential components of more effective vaccines. Recently, the genomic sequence of H. parasuis serovar5strain SH0165was completed in our laboratory, which allow us to target OMPs for the development of recombinant vaccines. In this study, we focused on10putative OMPs and all the putative OMPs were cloned, expressed and purified as HIS fusion proteins. Primary screening for immunoprotective potential was performed in mice challenged with an LD50challenge. Out of these10OMPs three fusion proteins rGAPDH, rOapA, and rHPS-0675were found to be protective in a mouse model of H. parasuis infection. We further evaluated the immune responses and protective efficacy of rGAPDH, rOapA, and rHPS-0675in pig models. All three proteins elicited humoral antibody responses and conferred different levels of protection against challenge with a lethal dose of H. parasuis SH0165in pig models. In addition, the antisera against the three individual proteins and the the synergistic protein efficiently inhibited bacterial growth in a whole blood assay. The data demonstrated that the three proteins showed high value individually and the combination of rGAPDH, rOapA, and rHPS-0675offered the best protection. Our results indicate that rGAPDH, rOapA, and rHPS-0675induced protection against H. parasuis SH0165infection, which may facilitate the development of a multi-component vaccine.
2. Construction and immune effect of Haemophilus parasuis DNA vaccine encoding glyceraldehyde-3-phosphate dehydrogenase (GAPDH) in mice
In this study the gap A gene that encodes glyceraldehyde-3-phosphate dehydrogenase (GAPDH) was shown to be present and highly conserved in various serotypes of H. parasuis and we constructed a novel DNA vaccine encoding GAPDH (pCgap) to evaluate the immune response and protective efficacy against infection with H. parasuis MD0322serovar4or SH0165serovar5in mice. A significant antibody response against GAPDH was generated following pCgap intramuscular immunization; moreover, antibodies to the pCgap DNA vaccine were bactericidal, suggesting that it was expressed in vivo. The gapA transcript was detected in muscle, liver, spleen, and kidney of the mice seven days post-vaccination. The IgG subclass (IgGl and IgG2a) analysis indicated that the DNA vaccine induced both Thl and Th2immune responses, but the IgG1response was greater than the IgG2a response. Moreover, the groups vaccinated with the pCgap vaccine exhibited83.3%and50%protective efficacy against the H. parasuis MD0322serovar4or SH0165serovar5challenge, respectively. The pCgap DNA vaccine provided significantly greater protective efficacy compared to the negative control groups or blank control groups (P<0.05for both). Taken together, these findings indicate that the pCgap DNA vaccine provides a novel strategy against infection of H. parasuis and offer insight concerning the underlying immune mechanisms of a bacterial DNA vaccine.
3. Multifunctional Glyceraldehyde-3-phosphate Dehydrogenase of Haemophilus parasuis enhances host immune response
Some research showed that some OMPs of the diverse pathogenic microbes impact bacteria colonization by modulation of the host immune system. And these OMPs also elicit the production of IL-2and or IL-10in the host cells. So we studied whether the outer membrane proteins play an important role in regulating the host's immune response. In this study, the GAPDH protein was screened from12OMPs which had a significant B cell stimulatoty effect regulated CD69expression observed on lymphocytes. In the MTT assay the GAPDH protein induces lymphocyte proliferation and B cell differentiation. A H. parasuis strain GAO overexpressing GAPDH protein demonstrated that it can promote the clearance of body against H. parasuis and induce the host IL-2production. Compared with the WT strain, the GAO strain was less able to resist killing by PMN cells and the RAW cells. Furthermore, infection with the GAO strain significantly upregulated the expression of all the examined immune-related genes except that encoding MHC II a. Taken together, GAPDH protein involved in regulating the host immune defense system which promote the immune response and bacterial clearance in vivo.
到目前为止,副猪嗜血杆菌共鉴定出15种血清型,每种血清型菌株的毒力又有差异。此外,还有很多菌株没有被定型。各种血清型之间的交叉免疫保护十分有限。当前使用的商业化灭活疫苗能够对同源血清型菌株感染提供一定的免疫保护,但不能提供异源血清型菌株感染的交叉保护。因此,急需研制开发出一种能够提供持久免疫保护和针对多种血清型菌株感染具有交叉免疫保护的新型疫苗。
目前,关于副猪嗜血杆菌毒力相关因子很少被报道。外膜蛋白是细菌的表面重要组成部分,能够和宿主细胞发生互作,并诱导宿主产生保护性抗体和引起细胞免疫反应。因此针对靶向外膜蛋白发展疫苗非常有价值。本课题在实验室前期完成的副猪嗜血杆菌流行优势血清5型SH0165菌株全基因组序列基础上,基于生物信息学及SignalP2.0软件,预测了副猪嗜血杆菌可能的重要外膜蛋白。进一步将编码外膜蛋白基因克隆、表达纯化后利用动物感染模型进行了蛋白免疫原性研究。同时,开展了外膜蛋白的免疫生物学活性研究,并从中筛选到了一个能够促进B细胞和T细胞增值的蛋白3-磷酸甘油醛脱氢酶(GAPDH),可能是一个重要免疫调节蛋白,深入探讨了该免疫调节蛋白免疫调节的机理。构建了编码GAPDH蛋白的DNA疫苗,并开展了相关免疫原性及免疫保护研究。主要的研究内容包括如下：
1.副猪嗜血杆菌SH0165菌株可能外膜蛋白预测、蛋白免疫原性和保护效力研究
本研究首先利用SignalP2.0软件,通过同源相似性比对预测了cdtB, GAPDH, HPS-2037, OmpP1, HPS-0220, HPS-0675, HPS-1406,HPS-1834, OapA, IdgA等10个可能的外膜蛋白,克隆表达并纯化了10个外膜蛋白。将10种重组外膜蛋白免疫小鼠后28天进行了5×LD50剂量的攻击。结果表明, rGAPDH, rOapA和rHPS-0675三种重组蛋白具有很好的免疫保护效果,分别为75%,87.5%和75%。进一步将这三种蛋白单独或混合形式以猪为感染模型进行了免疫反应和保护效力研究。结果表明,三种蛋白都能够引起很好的体液免疫反应并分别提供70%,50%和70%的免疫保护,而混合蛋白免疫提供的免疫保护力最高,为80%。而3种蛋白产生的抗体和混合蛋白的抗体在全血杀菌试验中都能够抑制细菌的生长。研究结果进一步提示rGAPDH, rOapA和rHPS-0675重组蛋白能够介导针对副猪嗜血杆菌SH0165菌株感染的免疫保护,是重要的疫苗候选抗原,能够研制开发成一种多组分通用型亚单位疫苗。
2.副猪嗜血杆菌3-磷酸甘油醛脱氢酶DNA疫苗的构建及免疫效力研究
研究结果表明编码3-磷酸甘油醛脱氢酶的gapA基因广泛存在于15种标准血清型菌株中而且基因序列比较保守,前期研究也显示其是一种很好的疫苗候选抗原。因此我们进一步构建了GAPDH蛋白的DNA疫苗pCgap,并在小鼠感染模型中评价了该DNA疫苗免疫反应和针对血清4型菌株MD0322和血清5型菌株SH0165的攻击所提供的免疫效力。结果表明,DNA疫苗通过肌内免疫小鼠之后能够产生显著的抗体反应。而且产生的抗体具有杀菌作用。DNA疫苗免疫小鼠7天之后能够在肌肉、肝脏、脾脏和肾脏中检测到转录产物。IgG亚类分析表明DNA疫苗可以同时诱导Thl和Th2型免疫反应,但是IgGl型免疫反应强于IgG2a型免疫反应。此外,DNA疫苗能够提供分别针对血清4型菌株MD0322攻击的保护率83.3%和血清5型菌株SH0165攻击的保护率50%。免疫组免疫保护力显著高于阴性对照组和空白对照组。以上结果提示DNA疫苗pCgap为控制副猪嗜血杆菌的感染提供一个新的策略。
3.副猪嗜血杆菌3-磷酸甘油醛脱氢酶(GAPDH)免疫调节机理研究
研究表明,一些外膜蛋白通过修改宿主的免疫系统来影响细菌在体内定植能力并且可以刺激宿主产生IL-2或者IL-10的能力。因此,本研究探讨副猪嗜血杆菌外膜蛋白是否在参与调节宿主的免疫反应方面起着重要作用。通过将12种外膜蛋白(cdtB, GAPDH, HPS.2037, OmpP1, HPS-0220, HPS-0675, HPS.1406, HPS.1834, OapA,6PGD, HPS.1889, IdgA)体外刺激小鼠脾淋巴细胞,从中筛选到了外膜蛋白GAPDH能够显著上调脾脏细胞表面CD69分子的表达。通过MTT实验结果也表明GAPDH蛋白能够促进脾脏淋巴细胞的增值和B细胞的分化。进一步构建了GAPDH过表达菌株GAO,开展了过表达菌株小鼠体内定植、诱导宿主产生细胞因子、中性粒细胞所介导的杀伤和对宿主免疫相关基因表达的影响等实验。结果表明：GAPDH能够促进机体对副猪嗜血杆菌的清除和诱导机体产生IL-2。与野生菌株相比,过表达菌株GAO更容易被中性粒细胞和小鼠的巨噬细胞所杀伤。而且,机体感染过表达菌株GAO之后能够促进所检测的除了MHC Ⅱα分子以外IFN-γ, MHC Ⅰα, TCR1和TLR3分子免疫相关基因上调表达。总之,GAPDH蛋白参与了宿主免疫系统的调节,增强了宿主的免疫反应和对外源病原菌的清除能力。
Haemophilus parasuis (H. parasuis), the causative agent of swine polyserositis, polyarthritis, and meningitis, is one of the most important bacterial diseases of pigs worldwide. In naive pigs disease onset occurs few days after H. parasuis exposure and is usually lethal. These animals recovering from the acute phase of the disease can develop chronic arthritis.
To date,15serovars of H. parasuis, which typically have a range of different virulence potentials, have been described. In addition, a high percentage of strains are non-serotypeable. To date, only a few virulence related factors have been reported in relation to the pathogenicity of Glasser's disease. Limited cross-protection among strains has complicated the control of Glasser's disease. Maternally derived antibodies also have an important role in protection, allowing the colonization of piglets and the development of a protective immune response without the induction of disease. Animals protected against H. parasuis infection after immunization with a bacterin may develop antibodies against outer membrane proteins (OMPs). The currently commercially available inactivated vaccine confers protection against homologous challenge, but further research is needed to develop a new generation of vaccine that can stimulate long-term immunity and provide cross-protection against infections by several H. parasuis serovars.
The sequencing of H. parasuis SH0165has been completed by our group. The current knowledge of the genomic sequence of pathogenic bacteria has facilitated recent vaccine development. Because OMPs are the primary bacterial components that interact with host cells, targeting OMPs for the development of recombinant vaccines is of value. Based on this, we focused on putative OMPs of H. parasuis which were predicted by SignalP2.0software. They were cloned, expressed, and purified as HIS fusion proteins and screened for protective efficacy in a piglet model of infection. Furthermore, we have studied the immune biological activity of the OMPs and constructed a DNA vaccine to prevent and control the infection of H. parasuis. Below is detailed information on this study:
1. Immunogenicity and protective efficacy of recombinant Haemophilus parasuis SH0165putative outer membrane proteins as vaccine candidates
H. parasuis putative outer membrane proteins (OMPs) are potentially essential components of more effective vaccines. Recently, the genomic sequence of H. parasuis serovar5strain SH0165was completed in our laboratory, which allow us to target OMPs for the development of recombinant vaccines. In this study, we focused on10putative OMPs and all the putative OMPs were cloned, expressed and purified as HIS fusion proteins. Primary screening for immunoprotective potential was performed in mice challenged with an LD50challenge. Out of these10OMPs three fusion proteins rGAPDH, rOapA, and rHPS-0675were found to be protective in a mouse model of H. parasuis infection. We further evaluated the immune responses and protective efficacy of rGAPDH, rOapA, and rHPS-0675in pig models. All three proteins elicited humoral antibody responses and conferred different levels of protection against challenge with a lethal dose of H. parasuis SH0165in pig models. In addition, the antisera against the three individual proteins and the the synergistic protein efficiently inhibited bacterial growth in a whole blood assay. The data demonstrated that the three proteins showed high value individually and the combination of rGAPDH, rOapA, and rHPS-0675offered the best protection. Our results indicate that rGAPDH, rOapA, and rHPS-0675induced protection against H. parasuis SH0165infection, which may facilitate the development of a multi-component vaccine.
2. Construction and immune effect of Haemophilus parasuis DNA vaccine encoding glyceraldehyde-3-phosphate dehydrogenase (GAPDH) in mice
In this study the gap A gene that encodes glyceraldehyde-3-phosphate dehydrogenase (GAPDH) was shown to be present and highly conserved in various serotypes of H. parasuis and we constructed a novel DNA vaccine encoding GAPDH (pCgap) to evaluate the immune response and protective efficacy against infection with H. parasuis MD0322serovar4or SH0165serovar5in mice. A significant antibody response against GAPDH was generated following pCgap intramuscular immunization; moreover, antibodies to the pCgap DNA vaccine were bactericidal, suggesting that it was expressed in vivo. The gapA transcript was detected in muscle, liver, spleen, and kidney of the mice seven days post-vaccination. The IgG subclass (IgGl and IgG2a) analysis indicated that the DNA vaccine induced both Thl and Th2immune responses, but the IgG1response was greater than the IgG2a response. Moreover, the groups vaccinated with the pCgap vaccine exhibited83.3%and50%protective efficacy against the H. parasuis MD0322serovar4or SH0165serovar5challenge, respectively. The pCgap DNA vaccine provided significantly greater protective efficacy compared to the negative control groups or blank control groups (P<0.05for both). Taken together, these findings indicate that the pCgap DNA vaccine provides a novel strategy against infection of H. parasuis and offer insight concerning the underlying immune mechanisms of a bacterial DNA vaccine.
3. Multifunctional Glyceraldehyde-3-phosphate Dehydrogenase of Haemophilus parasuis enhances host immune response
Some research showed that some OMPs of the diverse pathogenic microbes impact bacteria colonization by modulation of the host immune system. And these OMPs also elicit the production of IL-2and or IL-10in the host cells. So we studied whether the outer membrane proteins play an important role in regulating the host's immune response. In this study, the GAPDH protein was screened from12OMPs which had a significant B cell stimulatoty effect regulated CD69expression observed on lymphocytes. In the MTT assay the GAPDH protein induces lymphocyte proliferation and B cell differentiation. A H. parasuis strain GAO overexpressing GAPDH protein demonstrated that it can promote the clearance of body against H. parasuis and induce the host IL-2production. Compared with the WT strain, the GAO strain was less able to resist killing by PMN cells and the RAW cells. Furthermore, infection with the GAO strain significantly upregulated the expression of all the examined immune-related genes except that encoding MHC II a. Taken together, GAPDH protein involved in regulating the host immune defense system which promote the immune response and bacterial clearance in vivo.
引文
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