不同MHC单倍型SPF鸡MHC I类和II类分子的差异表达与MD致病性的相关研究
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摘要
鸡主要组织相容性复合体（Major histocompatablity complex, MHC）与各种传染性疾病抗性或易感性有关，且与鸡恶性淋巴组织增生性疾病马立克氏病（Marek's disease,MD）关系尤为密切。不同MHC-B单倍型对MD抗性不同。前期根据4个微卫星位点的基因型，建立了6种纯合BW/G(n)系鸡群，本研究在此基础上，通过对后4代各5个个体中BF和BG1基因多态外显子进行直接测序，同时以B2、B19和B5/B21单倍型为免疫原，制备了3种同种免疫抗血清，用血清学方法对鸡群进行了验证，成功地获得了6种MHC-B单倍型鸡群，不存在个体差异，具有高度的遗传稳定性。
随后以建立的MHC-B单倍型鸡群为基础来研究MHC基因座中BF基因和BLB基因的表达。BF基因包括BF1和BF2两个基因，BLB基因也包括BLB1和BLB2两个基因，本研究建立了能鉴别检测B2和B19单倍型鸡中BF和BLB双拷贝基因表达的特异性TaqMan qRT-PCR方法，并发现在MDV感染后期（77dpi），不同单倍型鸡脾脏中BF和BLB双拷贝基因转录水平呈现出不同的变化，且BF2和BLB2基因分别为优势表达基因，这可能与双拷贝基因的启动子序列结构有关。
为了分析MD抗性较强的BW/G3系鸡群（MHC B2单倍型）和敏感性较强的BW/G7系鸡群（B19单倍型）接种马立克氏病毒（MDV）后鸡群脾脏中MHC I类蛋白表达量的差异，本研究利用原核表达系统体外表达了BF基因较为保守的序列，制备出一种能检测不同B单倍型MHC I类蛋白的通用抗体，利用此抗体对BW/G3和BW/G7系各3只健康鸡和MDV攻毒后11周各5只攻毒鸡的脾脏进行Western blot，利用ImageQuant5.2软件半定量分析其中MHC I类蛋白含量的变化。结果发现抗性鸡BW/G3和敏感鸡BW/G7脾脏中MHC I类蛋白的表达量都发生上调，且易感鸡中的表达量高于抗性鸡，为进一步揭示病毒性肿瘤病的致病机理提供了思路。
为了进一步研究脾脏中不同淋巴细胞和法氏囊中BF和BLB双拷贝基因的表达与MD致病性的关系，本研究选取BW/G3（B2单倍型）和BW/G7系（B19单倍型）鸡群，各分为正常组和攻毒组，每组18只，攻毒组腹腔接种MDV Md5毒，攻毒后利用流式细胞术检测外周血和脾脏中CD4+和CD8+T细胞比例以及B细胞含量的变化并对脾脏进行了细胞分选，根据real-time荧光定量PCR方法检测脾脏不同类型细胞和法氏囊中的病毒载量，同时结合临床症状、相对体增重、大体病变、脾脏和法氏囊的石蜡切片、HE染色观察和脾脏中病毒抗原免疫组化等方法分析两种鸡群的致病性差异。然后在病毒感染后的6个不同时间点采样，利用建立的TaqMan qRT-PCR方法检测脾脏B细胞、CD4+T细胞、CD8+T细胞和法氏囊中MHC BF和BLB双拷贝基因的表达差异，并用SAS统计学软件进行数理统计。结果表明，BW/G3系（B2单倍型）和BW/G7系（B19单倍型）在MDV感染后，病毒拷贝数无显著差异，但脾脏中BW/G7系中病毒抗原明显高于BW/G3系。而且在不同单倍型鸡脾脏中，MDV主要感染的细胞种类有差异，并且引起了不同类型细胞数量发生改变。MDV感染后，在脾脏3类细胞中，MDV感染后诱导两种单倍型鸡中BF1基因的表达同时上调，BF2基因在较抗性鸡（BW/G3系）中表达下调而在易感鸡（BW/G7系）中上调，对于BLB基因，主要在B细胞中表达，且MDV对其表达具有抑制的趋势。在法氏囊中，BF1基因仅在易感鸡（BW/G7系）中表达上调，而BF2基因在两系中的表达全部上调。因此MDV与MHC I类分子有密切的关系，而BLB虽为MD的候选抗病基因，但MDV对其表达具有较小的影响。
本研究首先为我国研究各种传染性疾病、致病机理以及防控措施提供了宝贵的实验材料，同时也为研究各种疾病引起的宿主免疫应答机制提供了一定的理论依据。
There exists a strong and reproducible association between chicken majorhistocompatablity complex (MHC) haplotypes and the resistance to all kinds of diseases. Themost accepted example is related to lymphoproliferative tumor Marek’s disease (MD). Inearlier stage, we had successfully established six SPF BW/G(n) lines chickens carryingdifferent MHC-B haplotypes based on genotypes at four MHC microsatellite DNA markers.In this study, the sequences of the polymorphic exons of duplicated BF and BG genes insubsequent4generations (5chickens of each haplotype) were determined by direct sequence.We prepared three kinds of anti-BF and anti-BG allo-antisera, and serologically confirmed theMHC-B haplotypes using serological typing method simultaneously. It was shown that therewas a highly genetic stability with no individual difference for the six haplotype lines.
The expression of duplicated MHC-B BF and BLB genes were studied based onestablished MHC-B haplotypes. BF loci contain BF1and BF2genes, and BLB loci containBLB1and BLB2genes. In this study, we developed specific TaqMan probed real-timequantitative reverse transcription PCR (TaqMan qRT-PCR) methods based on the diagnosticnucleotide polymorphisms present in duplicated BF or BLB genes of B2and B19haplotypes.Spleen mRNA samples of MD-infected and control chickens of B2and B19haplotypes wereused to validate these TaqMan qRT-PCR methods. We observed that there was differenttranscribed change for duplicated BF and BLB genes of spleens in both lines during the latestage of the MDV infection (77dpi); furthermore, BF2and BLB2gene was dominantlytranscribed. Our findings verified the impact of diversified promoter sequences on thefunction of duplicated BF or BLB genes.
To semi-quantitatively analyze differential expressions of MHC class I proteins ofMDV-infected and control chickens of MD-resistent BW/G3and MD-susceptible BW/G7lines, we cloned and expressed class I α chain protein coded by conserved BF sequences usingprokaryotic expression system and prepared the polyclonal antibody of the rabbit serum todetect MHC class I protein of different MHC-B haplotypes. The differential expression ofMHC class I protein of MDV-infected (5chickens of each line) and control chickens (3chickens of each line) of BW/G3and BW/G7lines were semi-quantitatively analyzed using the ImageQuant5.2software. As showed by Western blotting, the expression of MHC class Igenes of the resistant and susceptible chickens were up-regulated during the late stage of theinfection, and the expression of MHC class I genes of susceptible chickens was higher thanresistant chickens.
In order to further study the association of the expression of duplicated BF or BLB genesin different kinds of splenocytes and bursa of Fabricius with the pathogenicity of MD, oneday old BW/G3(B2haplotype) and BW/G7(B19haplotype) chickens were divided intocontrol and infection group (18chickens of each group), The infection groups were infectedwith MDV Md5strain. After infection, the ration of CD4/CD8lymphocytes of PBL andspleens and B cells composition of spleens were detected using flow cytometry. In addition,three kinds of spleen lymphocytes were sorted using flow cytometry. MDV meq gene mRNAlevels of sorting cells and bursa of Fabricius were assessed by real-time fluorescencequantitative PCR. The disease resistance/susceptibility of BW/G3and BW/G7lines wereanalysed by clinical symptom, relative weight gain rate, the gross lesion, histological and pathologicalchanges (HE staining) of spleens and bursa of Fabricius and immunohistochemistry of viralantigens in spleens. The differential expression of duplicated BF or BLB genes of splenocytesB cell, CD4+and CD8+T cell and bursa of Fabricius were deceted using TaqMan qRT-PCRmethods per sampling time point and data were analysed using the SAS System. All of theresults above indicated that there was no significant difference of virus copy numbers betweenBW/G3line (B2haplotype) and BW/G7line (B19haplotype) after MDV infection, but theantigen amounts in BW/G7line spleens were dramatically higher than BW/G3line. The kindsof cells of spleens infected MDV were different between the lines, and the change of amountof different kinds of cells was induced correspondingly. The significantly up-regulatedtranscriptions of BF1gene in three kinds of spleen lymphocytes were detected in both lines ofchickens after MDV infection. The mRNA level of BF2gene was reduced in the MD mediumresistant chickens (BW/G3line), but increased in the susceptible chickens (BW/G7line). Inaddition, BLB genes mainly expressed in B cells, and the expression can be inhibited due toMDV infection. For bursa of Fabricius, the expression of BF1was up-regulated in thesusceptible chickens (BW/G7lines); meanwhile, the expression of BF2gene was increased inboth lines. So there was the close relationship between MHC I molecular and MDV, while the expression of BLB loci as the candidate gene for MD resistance was less affected by MDV.
In conclusion, this study provides the basic information to explore the possibility ofMHC haplotypes to be used for the challenge experiments to investigate particularly thepathogenic mechanisms of and control measures to disease. In addition, it also helpsunderstanding the theoretical basis to study the host immune response mechanism after virusinfection.
随后以建立的MHC-B单倍型鸡群为基础来研究MHC基因座中BF基因和BLB基因的表达。BF基因包括BF1和BF2两个基因，BLB基因也包括BLB1和BLB2两个基因，本研究建立了能鉴别检测B2和B19单倍型鸡中BF和BLB双拷贝基因表达的特异性TaqMan qRT-PCR方法，并发现在MDV感染后期（77dpi），不同单倍型鸡脾脏中BF和BLB双拷贝基因转录水平呈现出不同的变化，且BF2和BLB2基因分别为优势表达基因，这可能与双拷贝基因的启动子序列结构有关。
为了分析MD抗性较强的BW/G3系鸡群（MHC B2单倍型）和敏感性较强的BW/G7系鸡群（B19单倍型）接种马立克氏病毒（MDV）后鸡群脾脏中MHC I类蛋白表达量的差异，本研究利用原核表达系统体外表达了BF基因较为保守的序列，制备出一种能检测不同B单倍型MHC I类蛋白的通用抗体，利用此抗体对BW/G3和BW/G7系各3只健康鸡和MDV攻毒后11周各5只攻毒鸡的脾脏进行Western blot，利用ImageQuant5.2软件半定量分析其中MHC I类蛋白含量的变化。结果发现抗性鸡BW/G3和敏感鸡BW/G7脾脏中MHC I类蛋白的表达量都发生上调，且易感鸡中的表达量高于抗性鸡，为进一步揭示病毒性肿瘤病的致病机理提供了思路。
为了进一步研究脾脏中不同淋巴细胞和法氏囊中BF和BLB双拷贝基因的表达与MD致病性的关系，本研究选取BW/G3（B2单倍型）和BW/G7系（B19单倍型）鸡群，各分为正常组和攻毒组，每组18只，攻毒组腹腔接种MDV Md5毒，攻毒后利用流式细胞术检测外周血和脾脏中CD4+和CD8+T细胞比例以及B细胞含量的变化并对脾脏进行了细胞分选，根据real-time荧光定量PCR方法检测脾脏不同类型细胞和法氏囊中的病毒载量，同时结合临床症状、相对体增重、大体病变、脾脏和法氏囊的石蜡切片、HE染色观察和脾脏中病毒抗原免疫组化等方法分析两种鸡群的致病性差异。然后在病毒感染后的6个不同时间点采样，利用建立的TaqMan qRT-PCR方法检测脾脏B细胞、CD4+T细胞、CD8+T细胞和法氏囊中MHC BF和BLB双拷贝基因的表达差异，并用SAS统计学软件进行数理统计。结果表明，BW/G3系（B2单倍型）和BW/G7系（B19单倍型）在MDV感染后，病毒拷贝数无显著差异，但脾脏中BW/G7系中病毒抗原明显高于BW/G3系。而且在不同单倍型鸡脾脏中，MDV主要感染的细胞种类有差异，并且引起了不同类型细胞数量发生改变。MDV感染后，在脾脏3类细胞中，MDV感染后诱导两种单倍型鸡中BF1基因的表达同时上调，BF2基因在较抗性鸡（BW/G3系）中表达下调而在易感鸡（BW/G7系）中上调，对于BLB基因，主要在B细胞中表达，且MDV对其表达具有抑制的趋势。在法氏囊中，BF1基因仅在易感鸡（BW/G7系）中表达上调，而BF2基因在两系中的表达全部上调。因此MDV与MHC I类分子有密切的关系，而BLB虽为MD的候选抗病基因，但MDV对其表达具有较小的影响。
本研究首先为我国研究各种传染性疾病、致病机理以及防控措施提供了宝贵的实验材料，同时也为研究各种疾病引起的宿主免疫应答机制提供了一定的理论依据。
There exists a strong and reproducible association between chicken majorhistocompatablity complex (MHC) haplotypes and the resistance to all kinds of diseases. Themost accepted example is related to lymphoproliferative tumor Marek’s disease (MD). Inearlier stage, we had successfully established six SPF BW/G(n) lines chickens carryingdifferent MHC-B haplotypes based on genotypes at four MHC microsatellite DNA markers.In this study, the sequences of the polymorphic exons of duplicated BF and BG genes insubsequent4generations (5chickens of each haplotype) were determined by direct sequence.We prepared three kinds of anti-BF and anti-BG allo-antisera, and serologically confirmed theMHC-B haplotypes using serological typing method simultaneously. It was shown that therewas a highly genetic stability with no individual difference for the six haplotype lines.
The expression of duplicated MHC-B BF and BLB genes were studied based onestablished MHC-B haplotypes. BF loci contain BF1and BF2genes, and BLB loci containBLB1and BLB2genes. In this study, we developed specific TaqMan probed real-timequantitative reverse transcription PCR (TaqMan qRT-PCR) methods based on the diagnosticnucleotide polymorphisms present in duplicated BF or BLB genes of B2and B19haplotypes.Spleen mRNA samples of MD-infected and control chickens of B2and B19haplotypes wereused to validate these TaqMan qRT-PCR methods. We observed that there was differenttranscribed change for duplicated BF and BLB genes of spleens in both lines during the latestage of the MDV infection (77dpi); furthermore, BF2and BLB2gene was dominantlytranscribed. Our findings verified the impact of diversified promoter sequences on thefunction of duplicated BF or BLB genes.
To semi-quantitatively analyze differential expressions of MHC class I proteins ofMDV-infected and control chickens of MD-resistent BW/G3and MD-susceptible BW/G7lines, we cloned and expressed class I α chain protein coded by conserved BF sequences usingprokaryotic expression system and prepared the polyclonal antibody of the rabbit serum todetect MHC class I protein of different MHC-B haplotypes. The differential expression ofMHC class I protein of MDV-infected (5chickens of each line) and control chickens (3chickens of each line) of BW/G3and BW/G7lines were semi-quantitatively analyzed using the ImageQuant5.2software. As showed by Western blotting, the expression of MHC class Igenes of the resistant and susceptible chickens were up-regulated during the late stage of theinfection, and the expression of MHC class I genes of susceptible chickens was higher thanresistant chickens.
In order to further study the association of the expression of duplicated BF or BLB genesin different kinds of splenocytes and bursa of Fabricius with the pathogenicity of MD, oneday old BW/G3(B2haplotype) and BW/G7(B19haplotype) chickens were divided intocontrol and infection group (18chickens of each group), The infection groups were infectedwith MDV Md5strain. After infection, the ration of CD4/CD8lymphocytes of PBL andspleens and B cells composition of spleens were detected using flow cytometry. In addition,three kinds of spleen lymphocytes were sorted using flow cytometry. MDV meq gene mRNAlevels of sorting cells and bursa of Fabricius were assessed by real-time fluorescencequantitative PCR. The disease resistance/susceptibility of BW/G3and BW/G7lines wereanalysed by clinical symptom, relative weight gain rate, the gross lesion, histological and pathologicalchanges (HE staining) of spleens and bursa of Fabricius and immunohistochemistry of viralantigens in spleens. The differential expression of duplicated BF or BLB genes of splenocytesB cell, CD4+and CD8+T cell and bursa of Fabricius were deceted using TaqMan qRT-PCRmethods per sampling time point and data were analysed using the SAS System. All of theresults above indicated that there was no significant difference of virus copy numbers betweenBW/G3line (B2haplotype) and BW/G7line (B19haplotype) after MDV infection, but theantigen amounts in BW/G7line spleens were dramatically higher than BW/G3line. The kindsof cells of spleens infected MDV were different between the lines, and the change of amountof different kinds of cells was induced correspondingly. The significantly up-regulatedtranscriptions of BF1gene in three kinds of spleen lymphocytes were detected in both lines ofchickens after MDV infection. The mRNA level of BF2gene was reduced in the MD mediumresistant chickens (BW/G3line), but increased in the susceptible chickens (BW/G7line). Inaddition, BLB genes mainly expressed in B cells, and the expression can be inhibited due toMDV infection. For bursa of Fabricius, the expression of BF1was up-regulated in thesusceptible chickens (BW/G7lines); meanwhile, the expression of BF2gene was increased inboth lines. So there was the close relationship between MHC I molecular and MDV, while the expression of BLB loci as the candidate gene for MD resistance was less affected by MDV.
In conclusion, this study provides the basic information to explore the possibility ofMHC haplotypes to be used for the challenge experiments to investigate particularly thepathogenic mechanisms of and control measures to disease. In addition, it also helpsunderstanding the theoretical basis to study the host immune response mechanism after virusinfection.
引文
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