山羊DQA1、DQB1、DQA2基因外显子2多态性及与线虫抗性、免疫性状的相关研究
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摘要
主要组织相容性复合物(MHC)是一群紧密连锁高度多态的基因,编码主要组织相容性抗原,在脊椎动物机体的免疫系统中具有重要作用,因此MHC常作为家畜抗病育种的候选基因。本研究采用PCR-SSCP、PCR-RFLP技术对莱芜黑山羊、波尔山羊、鲁波杂交山羊3个山羊种群共175只个体的DQA1、DQA2、DQB1基因外显子2进行了遗传多态性分析,并对DQB1、DQA2基因多态性与线虫抗性、免疫性状的相关性进行了研究,其结果如下:
1.采用PCR-SSCP技术对DQA1基因外显子2多态性进行了分析,共检测到8种基因型,并对其纯合子进行了测序,检测到23处碱基突变。
2.对人类、山羊、绵羊、牛DQA1基因外显子2核苷酸序列进行同源性比较,结果发现同源性很高。
3.采用PCR-RFLP技术对莱芜黑山羊、鲁波杂交山羊和波尔山羊DQA2、DQB1基因外显子2进行多态性分析,共检测到4种DQA2基因外显子2基因型,7种DQB1基因外显子2基因型。通过酶切图谱和测序分析表明,DQA2基因外显子2的第77、79、80和169位的碱基表现出多态性,DQB1基因外显子2的第24、151位的碱基表现出多态性。
4.应用SAS 8.0软件GLM程序对DQA2、DQB1基因不同基因型与线虫抗性进行了相关分析,在DQA2基因中,基因型与线虫EPG有一定的相关性,但没有达到显著水平(P>0.05)。在DQB1基因中,BC、DD基因型的线虫EPG显著低于BB基因型(P<0.05),在种群内的分析中发现,鲁波杂交山羊中,BC基因型的线虫EPG显著低于AC基因型(P<0.05)。其它基因型与线虫EPG也有一定的相关性,但没有达到显著水平(P>0.05)。
5.应用SAS 8.0软件GLM程序对DQA2、DQB1基因不同基因型与免疫性状进行了相关分析,在DQA2基因中,莱芜黑山羊中AB基因型的RBC低于BB、BC基因型(P>0.05),而MCH高于BB、BC基因型,但没有达到显著水平(P>0.05);BC基因型的W-LCR高于AB、BB基因型(P>0.05),而W-SCC低于AB、BB基因型,但没有达到显著水平(P>0.05)。鲁波杂交山羊中AB基因型的RBC显著高于BB基因型(P<0.05),高于BC基因型(P>0.05);AB基因型的WBC显著高于BC基因型(P<0.05),高于BB基因型(P>0.05);AB基因型的HCT显著高于BB、BC基因型(P<0.05);BC基因型的W-LCR显著高于BB基因型(P<0.05),高于AB基因型(P>0.05)。在DQB1基因中,莱芜黑山羊中BC基因型的W-SCR显著高于AC、CC基因型(P<0.05),W-LCR显著低于CC基因型(P<0.05),W-LCC低于AC、CC基因型,但差异不显著(P>0.05);波尔山羊中BC基因型的W-LCC低于AA、AB基因型,但差异不显著(P>0.05);鲁波杂交山羊中BC、AC基因型的W-LCC显著低于AA基因型(P<0.05)。
MHC is a chromosomal region consisting of a group of closely linked loci which are highly polymorphic and play a central role in the immune system.So it is a candidate gene in disease resistance breeding. PCR-SSCP/PCR-RFLP were used to analyze the polymorphisms of exon 2 of DQA1,DQA2,DQB1 gene in Laiwu black goats, Lubo goats and Boer goats.The effects ofgenotypes of exon 2 of DQB1,DQA2 gene on nematode resistance ,immune traits were estimated.The results were as followed:
1. Polymorphisms of exon 2 of DQA1 was analyzed by PCR-SSCP , Eight genotypes were detected.and Its homozygous were sequenced, 23 polymorphic sites were detected .
2. The exon 2 of DQA1gene were amplified by PCR respectively. DNA sequences are compared. Comparative analysis showed that the homologies of DQA1 gene among Human, Bovine, sheep, goats, Laiwu black goats are very high.
3. PCR-RFLP was used to analyze the polymorphisms of exon 2 of DQA2 ,DQB1 gene in Laiwu black goats,Lubo goats and Boer goats.it was showed four DQA2 gene exon 2 genotypes and seven DQB1 gene exon 2 genotypes by analyzing restriction map and sequencing. Polymorphic sites were detected at base position 77, 79,80,169 of exon 2 of DQA2 gene. 24,151 of exon 2 of DQB1 gene.
4. The association of polymorphisms of exon 2 of DQA2, DQB1 gene with nematode resistance in three goat breeds was analyzed by SAS 8.0 software GLM program, In DQB1 gene, the nematode eggs count of genotype BC, DD was significantly lower than that with genotype BB (P <0.05). by analysising in the breeds , we found that in Lubo goats, the nematode eggs count with genotype BC was significantly lower than that with genotype AC (P <0.05). other genotypes and nematode eggs count had some relevance, but did not reach significant levels (P> 0.05).
5. The association of polymorphisms of exon 2 of DQA2 , DQB1 gene with
immune traits in three goats was analyzed by SAS 8.0 software GLM program. The results suggested that for DQA2 gene, RBC in Laiwu black goats with genotype AB was lower than that with genotype BB,BC(P>0.05), MCH in Laiwu black goats with genotype AB was higher than that with genotype BB, BC(P>0.05);W-LCR in Laiwu black goats with genotype BC was higher than that with genotype AB,BB(P>0.05);W-SCC in Laiwu black goats with genotype BC was lower than that with genotype AB, BB(P>0.05). RBC in Lubo goats with genotype AB was significantly higher than that with genotype BB (P<0.05), RBC in Lubo goats with genotype AB was higher than that with genotype BC(P>0.05). WBC in Lubo goats with genotype AB was significantly higher than that with genotype AC (P<0.05). WBC in Lubo goats with genotype AB was higher than that with genotype BB(P>0.05).HCT in Lubo goats with genotype AB was significantly higher than that with genotype BB ,BC (P<0.05).W-LCR in Lubo goats with genotype BC was significantly higher than that with genotype BB (P<0.05). W-LCR in Lubo goats with genotype BC was higher than that with genotype AC(P>0.05). for DQB1 gene,W-SCR in Laiwu black goats with genotype BC was significantly higher than that with genotype AC and CC (P<0.05), W-LCR in Laiwu black goats with genotype BC was significantly lower than that with genotype CC (P<0.05); W-LCC in Laiwu black goats with genotype BC was lower than that with genotype AC, CC (P>0.05). W-LCC in Boer goats with genotype BC was lower than that with genotype AA , AB (P>0.05) ; W-LCC in Lubo goats with genotype BC and AC was significantly lower than that with genotype AA (P<0.05).
These results may be applied to marker assisted selection in disease resistance breeding of goats.
1.采用PCR-SSCP技术对DQA1基因外显子2多态性进行了分析,共检测到8种基因型,并对其纯合子进行了测序,检测到23处碱基突变。
2.对人类、山羊、绵羊、牛DQA1基因外显子2核苷酸序列进行同源性比较,结果发现同源性很高。
3.采用PCR-RFLP技术对莱芜黑山羊、鲁波杂交山羊和波尔山羊DQA2、DQB1基因外显子2进行多态性分析,共检测到4种DQA2基因外显子2基因型,7种DQB1基因外显子2基因型。通过酶切图谱和测序分析表明,DQA2基因外显子2的第77、79、80和169位的碱基表现出多态性,DQB1基因外显子2的第24、151位的碱基表现出多态性。
4.应用SAS 8.0软件GLM程序对DQA2、DQB1基因不同基因型与线虫抗性进行了相关分析,在DQA2基因中,基因型与线虫EPG有一定的相关性,但没有达到显著水平(P>0.05)。在DQB1基因中,BC、DD基因型的线虫EPG显著低于BB基因型(P<0.05),在种群内的分析中发现,鲁波杂交山羊中,BC基因型的线虫EPG显著低于AC基因型(P<0.05)。其它基因型与线虫EPG也有一定的相关性,但没有达到显著水平(P>0.05)。
5.应用SAS 8.0软件GLM程序对DQA2、DQB1基因不同基因型与免疫性状进行了相关分析,在DQA2基因中,莱芜黑山羊中AB基因型的RBC低于BB、BC基因型(P>0.05),而MCH高于BB、BC基因型,但没有达到显著水平(P>0.05);BC基因型的W-LCR高于AB、BB基因型(P>0.05),而W-SCC低于AB、BB基因型,但没有达到显著水平(P>0.05)。鲁波杂交山羊中AB基因型的RBC显著高于BB基因型(P<0.05),高于BC基因型(P>0.05);AB基因型的WBC显著高于BC基因型(P<0.05),高于BB基因型(P>0.05);AB基因型的HCT显著高于BB、BC基因型(P<0.05);BC基因型的W-LCR显著高于BB基因型(P<0.05),高于AB基因型(P>0.05)。在DQB1基因中,莱芜黑山羊中BC基因型的W-SCR显著高于AC、CC基因型(P<0.05),W-LCR显著低于CC基因型(P<0.05),W-LCC低于AC、CC基因型,但差异不显著(P>0.05);波尔山羊中BC基因型的W-LCC低于AA、AB基因型,但差异不显著(P>0.05);鲁波杂交山羊中BC、AC基因型的W-LCC显著低于AA基因型(P<0.05)。
MHC is a chromosomal region consisting of a group of closely linked loci which are highly polymorphic and play a central role in the immune system.So it is a candidate gene in disease resistance breeding. PCR-SSCP/PCR-RFLP were used to analyze the polymorphisms of exon 2 of DQA1,DQA2,DQB1 gene in Laiwu black goats, Lubo goats and Boer goats.The effects ofgenotypes of exon 2 of DQB1,DQA2 gene on nematode resistance ,immune traits were estimated.The results were as followed:
1. Polymorphisms of exon 2 of DQA1 was analyzed by PCR-SSCP , Eight genotypes were detected.and Its homozygous were sequenced, 23 polymorphic sites were detected .
2. The exon 2 of DQA1gene were amplified by PCR respectively. DNA sequences are compared. Comparative analysis showed that the homologies of DQA1 gene among Human, Bovine, sheep, goats, Laiwu black goats are very high.
3. PCR-RFLP was used to analyze the polymorphisms of exon 2 of DQA2 ,DQB1 gene in Laiwu black goats,Lubo goats and Boer goats.it was showed four DQA2 gene exon 2 genotypes and seven DQB1 gene exon 2 genotypes by analyzing restriction map and sequencing. Polymorphic sites were detected at base position 77, 79,80,169 of exon 2 of DQA2 gene. 24,151 of exon 2 of DQB1 gene.
4. The association of polymorphisms of exon 2 of DQA2, DQB1 gene with nematode resistance in three goat breeds was analyzed by SAS 8.0 software GLM program, In DQB1 gene, the nematode eggs count of genotype BC, DD was significantly lower than that with genotype BB (P <0.05). by analysising in the breeds , we found that in Lubo goats, the nematode eggs count with genotype BC was significantly lower than that with genotype AC (P <0.05). other genotypes and nematode eggs count had some relevance, but did not reach significant levels (P> 0.05).
5. The association of polymorphisms of exon 2 of DQA2 , DQB1 gene with
immune traits in three goats was analyzed by SAS 8.0 software GLM program. The results suggested that for DQA2 gene, RBC in Laiwu black goats with genotype AB was lower than that with genotype BB,BC(P>0.05), MCH in Laiwu black goats with genotype AB was higher than that with genotype BB, BC(P>0.05);W-LCR in Laiwu black goats with genotype BC was higher than that with genotype AB,BB(P>0.05);W-SCC in Laiwu black goats with genotype BC was lower than that with genotype AB, BB(P>0.05). RBC in Lubo goats with genotype AB was significantly higher than that with genotype BB (P<0.05), RBC in Lubo goats with genotype AB was higher than that with genotype BC(P>0.05). WBC in Lubo goats with genotype AB was significantly higher than that with genotype AC (P<0.05). WBC in Lubo goats with genotype AB was higher than that with genotype BB(P>0.05).HCT in Lubo goats with genotype AB was significantly higher than that with genotype BB ,BC (P<0.05).W-LCR in Lubo goats with genotype BC was significantly higher than that with genotype BB (P<0.05). W-LCR in Lubo goats with genotype BC was higher than that with genotype AC(P>0.05). for DQB1 gene,W-SCR in Laiwu black goats with genotype BC was significantly higher than that with genotype AC and CC (P<0.05), W-LCR in Laiwu black goats with genotype BC was significantly lower than that with genotype CC (P<0.05); W-LCC in Laiwu black goats with genotype BC was lower than that with genotype AC, CC (P>0.05). W-LCC in Boer goats with genotype BC was lower than that with genotype AA , AB (P>0.05) ; W-LCC in Lubo goats with genotype BC and AC was significantly lower than that with genotype AA (P<0.05).
These results may be applied to marker assisted selection in disease resistance breeding of goats.
引文
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66. M.Amill et.A PCR-RFLP trping method for the caprine MHC classII DRB gene .Veterinary immunology and Immunopathology, 1996,55:255-260.
67. Meng-Hua et Allelic variation in exon 2 of capine MHC classII DRB3 gene in Chineseindigenous goats .Small Ruminant Research,2006 66:236-243. Immunology,1993,12:6977-6985.
68. Meng-Hua et al,Allelic variation in exon 2 of capine MHC classII DRB3 gene in Chineseindigenous goats .Small Ruminant Research ,2006,66: 236-243.
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70. M. Amills, O. Francino and A. Sánchez Nested PCR allows the characterization of TaqI and PstI RFLPs in the second exon of the caprine MHC class II DRB gene .Vet Immmunol Immunopathol 48:313-321,1995
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72. Ostergard H.K.,Kristensen B.,Adersen S.,Investigations informanmals of associations between the MHC system and disease resistence and fertility, Livestock Production Science.1989,22:49-67.
73. OTT J.. Strategies for characterizing highly polymorphic markers in human gene mapping. Am. J. Hum. Genet. 1992, 51: 283-290.
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75. R.T.Stone etal,Nucletide sequence f the second exon of a BoLA-DRB gene,Animal Genetics,1992,23:273-274.
76. Reche, P.A., Reinherz, E.L., Sequence variability analysis of human class I and class II MHC molecules: functional and structural correlates of amino acid polymorphisms. J. Mol. Biol. 2003,331:623–641.
77. S.A.Fabb etal, Isolation, characterization and evoluation of ovine MHC classⅡDRA and DQA genes,AnimalGenetics,1993,24:249-255.
78. Sayers G,Good B, et al,Major histocompatibility complex DRB1gene :its role in nematode resistance in Suffolk and Texe sheep breeds.Parasitology, 2005 ,131:403-409.
79. T. Takada, Y. Kikkawa, H. Yonekawa and T. Amano, Analysis of goat MHC class II DRA and DRB genes: identification of the expressed gene and new DRB alleles. Immunogenetics .1998,.48: 408–412.
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82. Saruhan-Direskeneli, G., Uyar, F.A., Bas, F., Gunoz, H., Bundak, R.,Saka, N., Darendeliler, F.,. HLA-DR and–DQ associations within sulin-depend diabetes mellitus in a population of Turkey. Hum.Immunol.2006,61:296- 302.
83. Saruhan-Direskeneli, G., Uyar, F.A., Bas, F., Gunoz, H., Bundak, R.,Saka, N., Darendeliler, F.,. HLA-DR and–DQ associations within sulin-depend diabetes mellitus in population of Turkey. Hum.Immunol.2000,61:296– 302.
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85. S.Sharif,B.A.Mallard et al. Association of the bovine MHC DRB3 alleles with occurrence of disease and mild somatic cell in Canadian dairy cattle, Animal Genetics,1998,29:185-193.
86. SolbuH.,Spooner R.L.Lie O.A.,Aposible influence of the bovine major histocompatibility complex (BoLA) on mastitis.Proceedings of 2nd Word Congresson Genetics Applided to Livestock Production Madrid,2003,7: 368-371.
87. S.Konnai et al.DNA Typing for Ovine MHC DRB1 Using Polymerase Chain Reaction-Restriction ragment Length Polymorphism (PCR-RFLP),J. Dairy Sci,2003,86:3362–3365.
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64. M.Amills et . Structure characterization of the caprine major histocompatibility Complex classII DQB1gene,Molecular Immunology, 2004, 418:43-846.
65. M.Amill et. Nucleotide sequence and polymorphism of the caprine major histocompatibility complex class II DQA(Cahi-DQA1)gene . Molecular Immunology. 2005, 42:375-379.
66. M.Amill et.A PCR-RFLP trping method for the caprine MHC classII DRB gene .Veterinary immunology and Immunopathology, 1996,55:255-260.
67. Meng-Hua et Allelic variation in exon 2 of capine MHC classII DRB3 gene in Chineseindigenous goats .Small Ruminant Research,2006 66:236-243. Immunology,1993,12:6977-6985.
68. Meng-Hua et al,Allelic variation in exon 2 of capine MHC classII DRB3 gene in Chineseindigenous goats .Small Ruminant Research ,2006,66: 236-243.
69. M. Amills, O. Francino and A. Sànchez, A PCR-RFLP typing method for the caprine Mhc classII DRB gene, Vet. Immunol. Immunop, 1996, 55:255–260.
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71. M.Amills et al.Structure characterization of the caprine major histocompatibility classII DQB1gene,Molecular Immunology 2004,418:43-846.
72. Ostergard H.K.,Kristensen B.,Adersen S.,Investigations informanmals of associations between the MHC system and disease resistence and fertility, Livestock Production Science.1989,22:49-67.
73. OTT J.. Strategies for characterizing highly polymorphic markers in human gene mapping. Am. J. Hum. Genet. 1992, 51: 283-290.
74. R.A.Hvan et al Characterization and evolution of ovine MHC classⅡDQB sequence polymorphism, AnialGenetcs,1994,25:417-424.
75. R.T.Stone etal,Nucletide sequence f the second exon of a BoLA-DRB gene,Animal Genetics,1992,23:273-274.
76. Reche, P.A., Reinherz, E.L., Sequence variability analysis of human class I and class II MHC molecules: functional and structural correlates of amino acid polymorphisms. J. Mol. Biol. 2003,331:623–641.
77. S.A.Fabb etal, Isolation, characterization and evoluation of ovine MHC classⅡDRA and DQA genes,AnimalGenetics,1993,24:249-255.
78. Sayers G,Good B, et al,Major histocompatibility complex DRB1gene :its role in nematode resistance in Suffolk and Texe sheep breeds.Parasitology, 2005 ,131:403-409.
79. T. Takada, Y. Kikkawa, H. Yonekawa and T. Amano, Analysis of goat MHC class II DRA and DRB genes: identification of the expressed gene and new DRB alleles. Immunogenetics .1998,.48: 408–412.
80. Scott et al., 1991. P.C. Scott, J.F. Maddox, K.J. Gogolin-Ewens and M.R. Brandon, The nucleotide sequence and evolution of ovine MHC class II B genes: DQB and DRB. Immunogenetics 34 (1991), pp. 80–87.
81. Scott P.C.and Maddox J.F., The nucleotide sequence and evoluation of ovine MHC classⅡgenes:DQB andDRB .Immunogenetics,1991,34:80-87
82. Saruhan-Direskeneli, G., Uyar, F.A., Bas, F., Gunoz, H., Bundak, R.,Saka, N., Darendeliler, F.,. HLA-DR and–DQ associations within sulin-depend diabetes mellitus in a population of Turkey. Hum.Immunol.2006,61:296- 302.
83. Saruhan-Direskeneli, G., Uyar, F.A., Bas, F., Gunoz, H., Bundak, R.,Saka, N., Darendeliler, F.,. HLA-DR and–DQ associations within sulin-depend diabetes mellitus in population of Turkey. Hum.Immunol.2000,61:296– 302.
84. Scott P.C.and Maddox J,F, The nucleotide sequence polymorphisms and evoluation of ovine MHC classⅡgenes:DQA gene,Immunogenetics,1991 34:69-79.
85. S.Sharif,B.A.Mallard et al. Association of the bovine MHC DRB3 alleles with occurrence of disease and mild somatic cell in Canadian dairy cattle, Animal Genetics,1998,29:185-193.
86. SolbuH.,Spooner R.L.Lie O.A.,Aposible influence of the bovine major histocompatibility complex (BoLA) on mastitis.Proceedings of 2nd Word Congresson Genetics Applided to Livestock Production Madrid,2003,7: 368-371.
87. S.Konnai et al.DNA Typing for Ovine MHC DRB1 Using Polymerase Chain Reaction-Restriction ragment Length Polymorphism (PCR-RFLP),J. Dairy Sci,2003,86:3362–3365.
88. Sharif S, Mallard BA et al.Associations of the bovine major histocompatibility complex DRB3 (BoLA-DRB3) with production traits in Canadian dairy cattle, Anim Genet, 1999 ,Apr;30(2):157-60.
89. S.Konnai et al.DNA Typing for Ovine MHC DRB1 Using Polymerase Chain Reaction-Restriction ragment Length Polymorphism (PCR-RFLP), J. Dairy Sci,2003,86:3362–3365.
90. Sally L. Debenhama et al. Genomic sequence of the class II region of the canine MHC:comparison with the MHC of other mammalian species,Genomics,2005,85:48–59.
91. Sally L. Debenhama et al. Genomic sequence of the class II region of the canine MHC:comparison with the MHC of other mammalian species, Genomics,2005,85:48–59.
92. Sun DX, Zhang Y. Polymorphism of the second exon of MHC-DRB gene in Chinese local sheep and goat [J].Biochemical Genetics,2004,42:385-390
93. Stear M J,T ierney,T J,et al,BoLA antigens are association with increased frequency of persistent lymphocytosis in bovine leukemia virusin fected cattle and with increased incidence of antibodies to bovine leukemia virus.Animal Genetics, 1988, 19: 151 -158.
94. Toyoyuki Takada et al, Analysis of goat MHCclassII DRA and DRB gene identification of the expressed gene and new DRB alleles. Immunogenetics 1998,48:408-412.
95. Van Oorschot R.A.H,Maddox J.F.,Adams., et al, Characterization and evolution of ovine MHC classII DQB sequence polymorphism [J].Animal genetics,1994,25:417-424.
96. Wright, H., Ballingall, K.T.,. Mapping and characterization of theDQ subregion of the ovine MHC. Anim. Genet. 25, 243–249.
97. Xu A L. Polymorphism in BoLA-DRB3 exon2 correlated with resistance to persistent lymphocytosis caused by Bovine leukemia virus.Journal of Immunology,1 993,15(2):6977 -6 985.
98. XU Ru-Hai,et al Sequences and Their polymorphisms of swine MHC-DQB、DRB proximal promoter region. Acta Genetica Sinica, 2005, 32(3): 282-288.
99. YoshikoNagaoka,Hidenori Kabeya ,Misao Onuma,Noriyuki Kasai,Kosuke Okada,and Yoko Aida.Ovine MHC Class IIDRB1 Alleles Associated with Resistance or Susceptibility to Development of Bovine Leukernia Virus-induced Ovine Lymphorna[J].Cancer research 59,975-981.
100.Zanoti M,Poll G.,et al ,Association of BoLA classI I haplotype with sunclinical progression of bovine leuka emia virus infection in Holstein-Friesian. Animal Genetics,1996,27:337-341.