牛MBL1基因多态性及其与产奶性能和乳腺炎相关性研究
摘要
本研究运用分子生物学方法分析牛甘露糖结合凝集素(Mannan-Binding Lectinl, MBL1)基因单核苷酸多态性及其多态性与产奶性能、血清中MBL-A和补体活性(CH50 andACH50)的相关性,检验MBL-A抗菌活性和MBL1基因在组织中表达情况,为中国荷斯坦奶牛的抗病高产育种工作提供一定的理论依据和基础资料。主要内容如下：
1.采用PCR-SSCP、CRS-PCR、PCR-RFLP、巢式PCR以及直接测序的方法检测469头中国荷斯坦奶牛、44头鲁西黄牛及24头渤海黑牛的MBL1基因启动子区和第一外显子的单核苷酸多态性,比较三个品种牛MBL1基因5个SNPs的差异。结果显示：多态位点g.-2194 A>C、g.-1446 T>C、g.-1330 G>A和g.70 G>A在三个品种牛上的优势等位基因相同,分别是A、C、G和G,频率为A(CH0.9 909/LYC0.7273/BBC0.9375)、C(CH0.6 098/LYC0.7 841/BBC0.6 250)、G(CH0.7 111/LYC0.8 523/BBC0.8 958)和G(CH0.6130/LYC0.7 614/0.7 500)。在中国荷斯坦牛中,只有g.-2 194 G>A满足Hardy-Weinberg平衡(P>0.05)。在鲁西黄牛中,只有g.105 T>C未满足Hardy-Weinberg平衡(P<0.05)。多态位点g.-2194G>A、g.-1446 T>C、g.-1 330 G>A、g.70 G>A和g.105T>C都在渤海黑牛中满足Hardy-Weinberg平衡(P>0.05)。
2.运用SHEsis软件对469头中国荷斯坦奶牛MBL1基因5个SNPs构建单倍型,利用SAS8.0软件的线性回归模型程序比较奶牛产奶性能(305天产奶量、乳脂率、乳蛋白率、体细胞评分)、CH50、ACH50和血清中MBL-A含量在不同基因型和单倍型组合之间的差异。结果如下：多态位点g.-1330 G>A显著影响CH50(P<0.05),g.70G>A与血清中MBL-A水平和CH50显著相关(P<0.05),g.-2194 A>C、g.-1446 T>C和g.105 T>C的不同基因型与产奶性能、CH50、ACH50和血清中MBL-A含量之间的差异不显著(P>0.05)；单倍型组合为H16H17个体的SCS显著低于单倍型组合H8H21个体(P<0.05),单倍型组合为H14H1个体的305天产奶量显著高于H16H1、H16H16、H16H17、H16H21、H16H3、H16H5、H16H6、H16H8、H8H1、H8H22和H8H5个体(P<0.05),单倍型组合H16H7个体的乳脂率显著低于H14H1、H16H16、H6H1和H8H1(P<0.05)；单倍型组合H16H5个体显著低于H14H1和H8H1个体(P<0.05),单倍型组合为H16H21(P<0.05)和H5H1(P<0.05)个体的乳蛋白率分别显著高于单倍型组合H14H1、H14H5、H15H1、H16H17、H16H22和H8H17个体,单倍型组合为H7H3的个体血清中MBL-A含量显著低于H14H5、H15H1、H16H1、H16H13、H16H15、H16H16、H16H17、H16H21、H16H3、H16H5、H16H6、H6H1、H8H1、H8H17和H8H5(P<0.05),单倍型组合H7H3个体CH50显著低于H14H5、H15H1、H16H1、H16H21、H16H22、H16H3、H16H5、H16H7、H8H1、H8H21、H8H22和H8H5(P<0.05),单倍型组合为H16H16个体的ACH50显著高于H16H21、H16H3、H16H5、H5H1和H8H1(P<0.05)。
3.选取奶牛乳腺炎主要致病菌金黄色葡萄球菌和大肠杆菌,运用平板计数方法研究牛血清中MBL-A的抗菌活性。结果显示：抗病性强试验组(H16H17单倍型组合个体血清)的抗金黄色葡萄球菌活性和抗大肠杆菌活性都显著高于(P<0.05)易受感染试验组(H8H21单倍型组合个体血清)、D-甘露糖抑制试验组和N-乙酰基-D-葡萄糖胺试验组个体血清。
4.利用实时荧光定量PCR技术检测牛MBLl基因在健康和患乳腺炎中国荷斯坦奶牛的心、肝、脾、肺、肾、肌肉和乳腺等7种组织中表达量差异,结果发现MBL1基因在中国荷斯坦牛不同组织中的表达存在差异,患乳腺炎个体的肝组织中MBL1基因的表达量显著高于健康个体(P<0.05),在肺、肾、肌肉和乳腺等组织中表达量低,脾中不表达。
This study analysis single nucleotide polymorphisms of mannose-combining lectin gene in bovine using molecular biology, and associations with milk performance traits, MBL-A level of serum and complement activity (CH50 and ACH50), inspection MBL-A antibacterial activity and MBL1 gene expression in the organization. For Chinese holstein cow with high-yielding breeding and resistance to provide certain theoretical basis and material. The results were as followed:
1. In this study, three single nucleotide polymorphisms (SNPs) in promoter and two SNPs in exonl of MBL1 gene were detected by PCR-SSCP, PCR-RFLP, CRS-PCR, Nested-PCR and DNA sequencing methods in 537 individuals from three Chinese cattle breeds. The dominant alleles in 3 three breeds were A、C、G and G, and the allelic frequencies are A (CH0.9909 /LYC0.7 273/BBC0.9375)、C (CH0.6 098/LYC0.7 841/BBC0.6250)、G (CH0.7111 /LYC0.8523/BBC0.8958)和G (CH0.6130/LYC0.7614/0.7500). Theχ2 test for the Bohai Black Cattle of all sites achieved balance about the Hardy-Weinberg equilibrium (P> 0.05); Holstein cow in the mutation g.-2194 G>A has achieved Hardy-Weinberg equilibrium (P> 0.05); Luxi cattle only in the g.105 T>C sites has not achieved Hardy-Weinberg equilibrium (P＜0.05)
2. Furthermore, analyzing the presence of the genotypes and haplotypes, we show the polymorphisms and their possible implications, while attempting to define the association of discovered polymorphisms with the serum MBL-A levels, complement activity (CH50 and ACH50) and the milk production traits. Homozygous genotype AA had significant lower than those with genotypes GA and AA(P<0.05) for CH50 in g.-1330 G>A. At g.70 G>A, the homozygous genotype GG had significant lower than those with genotypes GA and AA(P<0.05) for concentration of serum; genotype AA hah significant higher CH50 than genotype GG(P<0.05). However, no other significant associations were observed g.-2194 A>C, g.-1 446 and g.105 T>C polymorphisms and milk production traits and concentration of serum,CH50 and ACH50 in the 469 Chinese Holstein population (P>0.05).Statistical analyses revealed that cows with the combined genotypes of H16H17 had significantly lower SCS than combined genotypes H8H21The individuals with haplotype combination H14H1(P<0.05) presented a higher 305d milk yield than the ones with haplotype combination H16H1, H16H16, H16H17, H16H21, H16H3, H16H5, H16H6, H16H8, H8H1, H8H22 and H8H5, respectively;For fat percentage, the individuals with haplotype combination H16H7 (P＜0.05) showed significantly lower contrast to the haplotype combination H14H1,H16H16, H6H1 and H8H1; In protein percentage, the individuals with haplotype combination H16H21 (P＜0.05) and H5H1 (P＜0.05) presented higher than the ones with haplotype combination H14H1, H14H5,H15H1, H16H17, H16H22 and H8H17; For concentration of serum, the individuals with haplotype combination H7H3(P＜0.05) showed significantly lower contrast to the haplotype combination H14H5, H15H1, H16H1, H16H13, H16H15, H16H16, H16H17, H16H21, H16H3, H16H5, H16H6, H6H1, H8H1, H8H17 and H8H5; In the CH50, the individuals with haplotype combination H7H3 (P＜0.05) showed significantly lower than the haplotype combination H14H5, H15H1, H16H1, H16H21, H16H22, H16H3, H16H5, H16H7, H8H1, H8H21, H8H22 and H8H5; The individuals with haplotype combination H16H16 (P＜0.05) showed significantly higher ACH50 in contrast to the haplotype combination H16H21, H16H3, H16H5, H5H1 and H8H1.
3. To probe the biological activity of MBL-A influence Staphylococcus Aureus and Escherichia coli, we counted the CFU. The results shown that serum from H16H17(resistant) animals displayed a significantly(P<0.05) higher antibacterial activity(Staphylococcus Aureus and E.coli infection) compared with serum from H8H21 (susceptible) animals, serum samples pre-incubated with mannose and serum samples pre-incubated with N-acetyl-D-glucosamine. The evidence that genetic analysis and functional (serum antibacterial activity) analysis yield fully concordant results lends biological significance to the protective role of the H16H17 genotype against Staphylococcus Aureus and E.coli infection and the predisposing role of the H8H21 genotype.
4. Form the fluorescence quantitative real-time PCR (qPCR) analyses, the MBL1 gene was observed to be differentially expressed in bovine tissues (Fig.4). The liver of diseased cows had higher MBL1 expression than the ones with healthy (P＜0.05). However, no other significant associations were observed heart, spleen, lung, kidney, muscle and mammary tissue between the healthy samples and the diseased samples (P>0.05). It was low expression of MBL1 in spleen, lung, kidney, muscle and mammary and the less expression of MBL1 in spleen.
1.采用PCR-SSCP、CRS-PCR、PCR-RFLP、巢式PCR以及直接测序的方法检测469头中国荷斯坦奶牛、44头鲁西黄牛及24头渤海黑牛的MBL1基因启动子区和第一外显子的单核苷酸多态性,比较三个品种牛MBL1基因5个SNPs的差异。结果显示：多态位点g.-2194 A>C、g.-1446 T>C、g.-1330 G>A和g.70 G>A在三个品种牛上的优势等位基因相同,分别是A、C、G和G,频率为A(CH0.9 909/LYC0.7273/BBC0.9375)、C(CH0.6 098/LYC0.7 841/BBC0.6 250)、G(CH0.7 111/LYC0.8 523/BBC0.8 958)和G(CH0.6130/LYC0.7 614/0.7 500)。在中国荷斯坦牛中,只有g.-2 194 G>A满足Hardy-Weinberg平衡(P>0.05)。在鲁西黄牛中,只有g.105 T>C未满足Hardy-Weinberg平衡(P<0.05)。多态位点g.-2194G>A、g.-1446 T>C、g.-1 330 G>A、g.70 G>A和g.105T>C都在渤海黑牛中满足Hardy-Weinberg平衡(P>0.05)。
2.运用SHEsis软件对469头中国荷斯坦奶牛MBL1基因5个SNPs构建单倍型,利用SAS8.0软件的线性回归模型程序比较奶牛产奶性能(305天产奶量、乳脂率、乳蛋白率、体细胞评分)、CH50、ACH50和血清中MBL-A含量在不同基因型和单倍型组合之间的差异。结果如下：多态位点g.-1330 G>A显著影响CH50(P<0.05),g.70G>A与血清中MBL-A水平和CH50显著相关(P<0.05),g.-2194 A>C、g.-1446 T>C和g.105 T>C的不同基因型与产奶性能、CH50、ACH50和血清中MBL-A含量之间的差异不显著(P>0.05)；单倍型组合为H16H17个体的SCS显著低于单倍型组合H8H21个体(P<0.05),单倍型组合为H14H1个体的305天产奶量显著高于H16H1、H16H16、H16H17、H16H21、H16H3、H16H5、H16H6、H16H8、H8H1、H8H22和H8H5个体(P<0.05),单倍型组合H16H7个体的乳脂率显著低于H14H1、H16H16、H6H1和H8H1(P<0.05)；单倍型组合H16H5个体显著低于H14H1和H8H1个体(P<0.05),单倍型组合为H16H21(P<0.05)和H5H1(P<0.05)个体的乳蛋白率分别显著高于单倍型组合H14H1、H14H5、H15H1、H16H17、H16H22和H8H17个体,单倍型组合为H7H3的个体血清中MBL-A含量显著低于H14H5、H15H1、H16H1、H16H13、H16H15、H16H16、H16H17、H16H21、H16H3、H16H5、H16H6、H6H1、H8H1、H8H17和H8H5(P<0.05),单倍型组合H7H3个体CH50显著低于H14H5、H15H1、H16H1、H16H21、H16H22、H16H3、H16H5、H16H7、H8H1、H8H21、H8H22和H8H5(P<0.05),单倍型组合为H16H16个体的ACH50显著高于H16H21、H16H3、H16H5、H5H1和H8H1(P<0.05)。
3.选取奶牛乳腺炎主要致病菌金黄色葡萄球菌和大肠杆菌,运用平板计数方法研究牛血清中MBL-A的抗菌活性。结果显示：抗病性强试验组(H16H17单倍型组合个体血清)的抗金黄色葡萄球菌活性和抗大肠杆菌活性都显著高于(P<0.05)易受感染试验组(H8H21单倍型组合个体血清)、D-甘露糖抑制试验组和N-乙酰基-D-葡萄糖胺试验组个体血清。
4.利用实时荧光定量PCR技术检测牛MBLl基因在健康和患乳腺炎中国荷斯坦奶牛的心、肝、脾、肺、肾、肌肉和乳腺等7种组织中表达量差异,结果发现MBL1基因在中国荷斯坦牛不同组织中的表达存在差异,患乳腺炎个体的肝组织中MBL1基因的表达量显著高于健康个体(P<0.05),在肺、肾、肌肉和乳腺等组织中表达量低,脾中不表达。
This study analysis single nucleotide polymorphisms of mannose-combining lectin gene in bovine using molecular biology, and associations with milk performance traits, MBL-A level of serum and complement activity (CH50 and ACH50), inspection MBL-A antibacterial activity and MBL1 gene expression in the organization. For Chinese holstein cow with high-yielding breeding and resistance to provide certain theoretical basis and material. The results were as followed:
1. In this study, three single nucleotide polymorphisms (SNPs) in promoter and two SNPs in exonl of MBL1 gene were detected by PCR-SSCP, PCR-RFLP, CRS-PCR, Nested-PCR and DNA sequencing methods in 537 individuals from three Chinese cattle breeds. The dominant alleles in 3 three breeds were A、C、G and G, and the allelic frequencies are A (CH0.9909 /LYC0.7 273/BBC0.9375)、C (CH0.6 098/LYC0.7 841/BBC0.6250)、G (CH0.7111 /LYC0.8523/BBC0.8958)和G (CH0.6130/LYC0.7614/0.7500). Theχ2 test for the Bohai Black Cattle of all sites achieved balance about the Hardy-Weinberg equilibrium (P> 0.05); Holstein cow in the mutation g.-2194 G>A has achieved Hardy-Weinberg equilibrium (P> 0.05); Luxi cattle only in the g.105 T>C sites has not achieved Hardy-Weinberg equilibrium (P＜0.05)
2. Furthermore, analyzing the presence of the genotypes and haplotypes, we show the polymorphisms and their possible implications, while attempting to define the association of discovered polymorphisms with the serum MBL-A levels, complement activity (CH50 and ACH50) and the milk production traits. Homozygous genotype AA had significant lower than those with genotypes GA and AA(P<0.05) for CH50 in g.-1330 G>A. At g.70 G>A, the homozygous genotype GG had significant lower than those with genotypes GA and AA(P<0.05) for concentration of serum; genotype AA hah significant higher CH50 than genotype GG(P<0.05). However, no other significant associations were observed g.-2194 A>C, g.-1 446 and g.105 T>C polymorphisms and milk production traits and concentration of serum,CH50 and ACH50 in the 469 Chinese Holstein population (P>0.05).Statistical analyses revealed that cows with the combined genotypes of H16H17 had significantly lower SCS than combined genotypes H8H21The individuals with haplotype combination H14H1(P<0.05) presented a higher 305d milk yield than the ones with haplotype combination H16H1, H16H16, H16H17, H16H21, H16H3, H16H5, H16H6, H16H8, H8H1, H8H22 and H8H5, respectively;For fat percentage, the individuals with haplotype combination H16H7 (P＜0.05) showed significantly lower contrast to the haplotype combination H14H1,H16H16, H6H1 and H8H1; In protein percentage, the individuals with haplotype combination H16H21 (P＜0.05) and H5H1 (P＜0.05) presented higher than the ones with haplotype combination H14H1, H14H5,H15H1, H16H17, H16H22 and H8H17; For concentration of serum, the individuals with haplotype combination H7H3(P＜0.05) showed significantly lower contrast to the haplotype combination H14H5, H15H1, H16H1, H16H13, H16H15, H16H16, H16H17, H16H21, H16H3, H16H5, H16H6, H6H1, H8H1, H8H17 and H8H5; In the CH50, the individuals with haplotype combination H7H3 (P＜0.05) showed significantly lower than the haplotype combination H14H5, H15H1, H16H1, H16H21, H16H22, H16H3, H16H5, H16H7, H8H1, H8H21, H8H22 and H8H5; The individuals with haplotype combination H16H16 (P＜0.05) showed significantly higher ACH50 in contrast to the haplotype combination H16H21, H16H3, H16H5, H5H1 and H8H1.
3. To probe the biological activity of MBL-A influence Staphylococcus Aureus and Escherichia coli, we counted the CFU. The results shown that serum from H16H17(resistant) animals displayed a significantly(P<0.05) higher antibacterial activity(Staphylococcus Aureus and E.coli infection) compared with serum from H8H21 (susceptible) animals, serum samples pre-incubated with mannose and serum samples pre-incubated with N-acetyl-D-glucosamine. The evidence that genetic analysis and functional (serum antibacterial activity) analysis yield fully concordant results lends biological significance to the protective role of the H16H17 genotype against Staphylococcus Aureus and E.coli infection and the predisposing role of the H8H21 genotype.
4. Form the fluorescence quantitative real-time PCR (qPCR) analyses, the MBL1 gene was observed to be differentially expressed in bovine tissues (Fig.4). The liver of diseased cows had higher MBL1 expression than the ones with healthy (P＜0.05). However, no other significant associations were observed heart, spleen, lung, kidney, muscle and mammary tissue between the healthy samples and the diseased samples (P>0.05). It was low expression of MBL1 in spleen, lung, kidney, muscle and mammary and the less expression of MBL1 in spleen.
引文
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