巴音布鲁克羊、德国美利奴羊、多浪羊BMPR-IB、BMP15、GDF9、MC4R、CAST基因的研究
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摘要
绵羊育种主要是改善其生长和繁殖性状。迄今为止已发现了一些控制这些生长和繁殖性状的基因,比如,BMPR-IB、BMP15和GDF9已被确认为绵羊的高繁殖力主效基因；黑素皮质素受体-4(melanocortin-4receptor,MC4R)是一类G-蛋白偶联受体,主要在下丘脑表达,在体重、能量平稳和采食量的调控中发挥重要作用。钙蛋白酶抑制蛋白(calpastatin,CAST)是钙蛋白酶系统的主要成员之一,它是钙激活蛋白酶的抑制因子,可以通过激活钙蛋白酶或降低钙蛋白酶抑制蛋白的活性来提高肌肉的嫩度,改善肉质。
本研究利用连接酶检测反应(ligase detection reaction,LDR)技术及生物信息学技术,检测了绵羊繁殖候选基因BMPR-IB、BMP15和GDF9在巴音布鲁克羊(含低产和高产两个群体)、德国美利奴羊、多浪羊中的多态性,并分析了其多态性与产羔性状的关系；同时检测了生长性状候选基因MC4R和CAST在德国美利奴羊中的多态性并分析了其与部分产肉性状的关系,为绵羊的标记辅助选择提供了参考依据。本研究主要包括以下内容：
Ⅰ以巴音布鲁克羊普通群(100只,B群)和多羔小群(20只,A群)、内蒙古有完整产羔记录的德国美利奴羊、巴州的多浪羊为研究对象,采用LDR技术,对BMPR-IB, BMP15和GDF9三个高繁殖力基因的10个位点：BMPR-IB基因FecB突变位点,BMP15基因上FecXI、FecXB、FecXL、FecXH、FecXG和FecXR突变位点以及GDF9基因上G1、G8和FecTT突变位点,进行单核苷酸多态性(single nucleotide polymorphism,SNP)检测,以比较BMPR-IB基因、BMP15基因和GDF9基因在巴音布鲁克羊普通群和多羔小群、德国美利奴羊及多浪羊中的多态性,并与产羔性能进行关联分析。
Ⅱ以内蒙古有完整生长、产肉性能(初生重、背膘厚度、眼肌面积、日均增重)测定记录的德国美利奴羔羊为研究对象,采用LDR技术对MC4R基因上4个突变位点、CAST基因第二外显子上7个突变位点进行SNP检测,并与生长、产肉性能进行关联分析。
主要结果是：
1BMPR-IB基因在巴音布鲁克羊、德国美利奴羊、多浪羊中的多态性及其与产羔数的相关性
在巴音布鲁克羊群中发现了BMPR-IB/FecB基因突变。其中,A、B两群的野生型(Fee++)、杂合型(FecB+)和突变型(FecBB)基因型频率分别为0.15、0.7、0.15和1.0、0、0,对应的野生型和突变型基因频率分别为0.5、0.5和1.0、O. A群与B群独立性检验结果表明,FecB多态位点对巴音布鲁克羊群体的产羔数具有极显著影响。据此推测：BMPR-IB/FecB可确认为影响巴音布鲁克羊多羔群产羔数的主效基因,并可以此遗传标记对该群体的繁殖力性状实施标记辅助选择。在德国美利奴羊和多浪羊中未发现BMPR-IB/FecB突变,由此推断,BMPR-IB/FecB突变与德国美利奴羊和多浪羊的繁殖性能无关联。
2BMP15基因在巴音布鲁克羊、德国美利奴羊、多浪羊中的多态性及其与产羔数的相关性
在三个品种中均未检测到BMP15基因上的FecX1、 FecXB、FeCXL、FecXH、FeCXG和FecXT等6个位点发生突变,据此推测：6个关联高繁殖力的BMP15基因的突变与巴音布鲁克羊、德国美利奴羊、多浪羊的繁殖力无关联。
3GDF9基因在巴音布鲁克羊、德国美利奴羊、多浪羊中的多态性及其与产羔数的相关性
在巴音布鲁克羊、德国美利奴羊GDF9基因上检测到了G1突变,但未检测到FecGH(G8)、FecTT突变,G1突变在德国美利奴羊达到中度多态,在所检测群体中均处于Hardy-Weinberg平衡状态,但其多态与产羔数的相关性不显著；在多浪羊中未检测到GDF9基因的三个突变位点。据此推测：GDF9基因的上述位点不是影响巴音布鲁克羊、德国美利奴羊、多浪羊产羔数的主效基因。
4MC4R基因在德国美利奴羊中的多态性及其与部分产肉性状的相关性
在德国美利奴羊MC4R基因上检测到了G309A、G508A、G1232A和T1456C4个突变。各突变位点在所检测群体中处于Hardy-Weinberg平衡状态,其中G1232A突变位点达到中度多态,其余三个位点为低度多态。4个突变位点中,G309A突变AA型背膘厚极显著高于GG型(P<0.01)；AG型背膘厚显著高于GG型(P<0.05),眼肌面积、出生重AA型显著高于AG型、GG型(P<0.05)；G1232A突变背膘厚AA型、AG型显著高于GG型(P<0.05)；日增重AA型显著高于GG型(P<0.05)。据此推测：MC4R基因是影响德国美利奴羊生产性状的QTL或与控制这些性状的QTL有连锁,因此有关标记将来可能用于这些性状的标记辅助选择。
5CAST基因在德国美利奴羊中的多态性及其与部分肉用性状的相关性
德国美利奴羊在CAST基因上存在CAST3-1、CAST3-2、CAST3-3、CAST3-4、CAST7-1、CAST7-2和CAST7-3等7个突变位点,统计结果：CAST3-2、CAST3-3和CAST7-1突变位点达到中度多态,其余4个突变位点为低度多态。但这7个突变位点与出生重、日均增重、背膘厚、眼肌面积等生产性能均未达到显著相关,这些突变位点是否影响其他生产或肉品质性状还有待于进一步研究。
Sheep breeding involves in improving growth and reproductive characteristics. To date many genes have been found to control these traits. BMPR-IB, BMP15, and GDF9, have been found to be the major genes that influence sheep reproductive capacity. Melanocortin-4receptor (MC4R) is a G-protein coupled receptor expressed in the brain, predominantly in the hypothalamus where it controls food intake, energy homeostasis and body weight. Calpastatin (CAST), one member of calpains system, is a specific inhibitor of calpains and is believed to be involved in the degradation of myofibrillar proteins in post-mortem proteolysis through regulating calpain activity, and hence it is implicated in meat tenderization.
Here we used ligase detection reaction (LDR) and bioinformatics techniques to investigate the polymorphisms of BMPR-IB, BMP15, and GDF9genes in Bayanbulak (including low and high yielding groups), German Mutton Merino, and Duolang sheep, and correlated the polymorphisms with litter size. The polymorphisms of MC4R and CAST gene in German Mutton Merino were also determined, hoping to find clues to the marker assisted selection (MAS) in sheep production. This study included two experiments:
I One hundred ordinary Bayanbulak sheep having single lamb (Flock B),20special Bayanbulak sheep having more than one lamb (Flock A), German Mutton Merino with complete lambing record in Inner Mongolia, and Bazhou Duolang sheep were studied using LDR to detect10SNPs in BMPR-IB, BMP15, and GDF9. The loci included FecB in BMPR-IB, FecXI, FecXB, FecXL, FecXH, FecXG, and FecXR in BMP15, and G1, G8, and FecTT in GDF9. The polymorphisms of BMPR-IB, BMP15, and GDF9were then compared among ordinary Bayanbulak group, special Bayanbulak group, German Mutton Merino, and Duolang Sheep. An association of the SNPs with litter size was subsequently conducted.
II Lambs of German Mutton Merino in Inner Mongolia with complete record of production performance (including birth weight, back-fat, eye muscle area and average daily gain) were studied using LDR to detect SNPs in MC4R gene and the second exon of the CAST gene. An association of the SNPs with growth trait and meat performance. The results were as follows:
1Polymorphisms of BMPR-IB gene in Bayanbulak, German Mutton Merino, and Duolang sheep, and its association with litter size
Polymorphisms of BMPR-1B/FecB gene were found in the Bayanbulak sheep. The genotypic frequency of the wild type (Fec++), heterozygous type (FecB+), and mutant (FecBB) were0.15,0.7, and0.15in flock A, and1.0,0, and0in flock B, respectively. The corresponding genotype frequency of wild type and mutation type were0.5,0.5in flock A and1.0,0.0in flock B, respectively. The independence test results of flock A and B suggested that the FecB locus significantly affects the litter size in the Bayanbulak sheep. BMPR-IB/FecB can be considered as the major gene that determines litter size in Bayanbulak sheep. Accordingly, this genetic marker may be used to improve litter size through MAS. No polymorphism of BMPR-IB/FecB mutation was found in German Mutton Merino and Duolang sheep. Therefore, the BMPR-IB/FecB mutation may not be correlated with the reproductive performance in the German Mutton Merino and Duolang sheep.
2Polymorphisms of BMP15gene in Bayanbulak, German Mutton Merino, and Duolang sheep, and its association with litter size
No polymorphism was detected at FecX-I、FecXB、FecXL、FecXH、FecX and FecXR of BMP15gene in the above three breeds. Therefore, the six loci of the BMP15gene cannot be used as a genetic marker for litter size in the three sheep breeds.
3Polymorphisms of GDF9gene in Bayanbulak, German Mutton Merino, and Duolang sheep, and its relation to litter size
The G1mutation was detected in the GDF9gene of Bayanbulak and German Mutton Merino sheep. However, FecG (G8) and FecTT mutations were not detected in the two sheep breeds. The polymorphism of G1mutation was moderate, and the locus was within the Hardy-Weinberg equilibrium in the two groups. No association of GDF9polymorphisms with litter size was found. There was no SNP of GDF9gene in Duolang sheep. These results indicated that the three site of GDF9gene does not significantly influence litter size of Bayanbulak, German Mutton Merino, and Duolang sheep.
4Polymorphisms of MC4R gene in German Mutton Merino and its association with meat performance
Four SNPs of MC4R gene were detected in German Mutton Merino sheep, namely, G309A, G508A, G1232A, and T1456C. All SNPs were within Hardy-Weinberg equilibrium. The SNP G1232A was moderate polymorphic, whereas the other three were slightly polymorphic. The sheep with AA type at G309A SNP had higher back fat than those with GG type (P<0.01). The back fat of sheep with AG type was higher than those with GG type (P<0.05). The eye muscle area and birth weight of AA sheep were higher than those of AG and GG sheep (P<0.05). Sheep with AA and AG genotype at the SNP G1232A had higher back fat than those with GG type (P<0.05). Average daily gain of AA genotype sheep was higher than that of GG genotypes (P<0.05). These findings indicated that the MC4R gene might be the QTL that influences production traits or it is linkage to the QTL that controls the traits. The markers may be applied in the MAS.
5Polymorphisms of CAST gene in German Mutton Merino and its association with meat performance
Seven SNPs of CAST gene were found in German Mutton Merino, namely, CAST3-1, CAST3-2, CAST3-3, CAST3-4, CAST7-1, CAST7-2, and CAST7-3. The statistical results showed that CAST3-2, CAST3-3, and CAST7-1were moderately polymorphic, whereas the other four SNPs were slightly polymorphic. The seven SNPs, however, were not significantly correlated to birth weight, average daily gain, back fat, eye muscle area. Further studies should be focused on find association of these SNPs with other production characteristcs.
本研究利用连接酶检测反应(ligase detection reaction,LDR)技术及生物信息学技术,检测了绵羊繁殖候选基因BMPR-IB、BMP15和GDF9在巴音布鲁克羊(含低产和高产两个群体)、德国美利奴羊、多浪羊中的多态性,并分析了其多态性与产羔性状的关系；同时检测了生长性状候选基因MC4R和CAST在德国美利奴羊中的多态性并分析了其与部分产肉性状的关系,为绵羊的标记辅助选择提供了参考依据。本研究主要包括以下内容：
Ⅰ以巴音布鲁克羊普通群(100只,B群)和多羔小群(20只,A群)、内蒙古有完整产羔记录的德国美利奴羊、巴州的多浪羊为研究对象,采用LDR技术,对BMPR-IB, BMP15和GDF9三个高繁殖力基因的10个位点：BMPR-IB基因FecB突变位点,BMP15基因上FecXI、FecXB、FecXL、FecXH、FecXG和FecXR突变位点以及GDF9基因上G1、G8和FecTT突变位点,进行单核苷酸多态性(single nucleotide polymorphism,SNP)检测,以比较BMPR-IB基因、BMP15基因和GDF9基因在巴音布鲁克羊普通群和多羔小群、德国美利奴羊及多浪羊中的多态性,并与产羔性能进行关联分析。
Ⅱ以内蒙古有完整生长、产肉性能(初生重、背膘厚度、眼肌面积、日均增重)测定记录的德国美利奴羔羊为研究对象,采用LDR技术对MC4R基因上4个突变位点、CAST基因第二外显子上7个突变位点进行SNP检测,并与生长、产肉性能进行关联分析。
主要结果是：
1BMPR-IB基因在巴音布鲁克羊、德国美利奴羊、多浪羊中的多态性及其与产羔数的相关性
在巴音布鲁克羊群中发现了BMPR-IB/FecB基因突变。其中,A、B两群的野生型(Fee++)、杂合型(FecB+)和突变型(FecBB)基因型频率分别为0.15、0.7、0.15和1.0、0、0,对应的野生型和突变型基因频率分别为0.5、0.5和1.0、O. A群与B群独立性检验结果表明,FecB多态位点对巴音布鲁克羊群体的产羔数具有极显著影响。据此推测：BMPR-IB/FecB可确认为影响巴音布鲁克羊多羔群产羔数的主效基因,并可以此遗传标记对该群体的繁殖力性状实施标记辅助选择。在德国美利奴羊和多浪羊中未发现BMPR-IB/FecB突变,由此推断,BMPR-IB/FecB突变与德国美利奴羊和多浪羊的繁殖性能无关联。
2BMP15基因在巴音布鲁克羊、德国美利奴羊、多浪羊中的多态性及其与产羔数的相关性
在三个品种中均未检测到BMP15基因上的FecX1、 FecXB、FeCXL、FecXH、FeCXG和FecXT等6个位点发生突变,据此推测：6个关联高繁殖力的BMP15基因的突变与巴音布鲁克羊、德国美利奴羊、多浪羊的繁殖力无关联。
3GDF9基因在巴音布鲁克羊、德国美利奴羊、多浪羊中的多态性及其与产羔数的相关性
在巴音布鲁克羊、德国美利奴羊GDF9基因上检测到了G1突变,但未检测到FecGH(G8)、FecTT突变,G1突变在德国美利奴羊达到中度多态,在所检测群体中均处于Hardy-Weinberg平衡状态,但其多态与产羔数的相关性不显著；在多浪羊中未检测到GDF9基因的三个突变位点。据此推测：GDF9基因的上述位点不是影响巴音布鲁克羊、德国美利奴羊、多浪羊产羔数的主效基因。
4MC4R基因在德国美利奴羊中的多态性及其与部分产肉性状的相关性
在德国美利奴羊MC4R基因上检测到了G309A、G508A、G1232A和T1456C4个突变。各突变位点在所检测群体中处于Hardy-Weinberg平衡状态,其中G1232A突变位点达到中度多态,其余三个位点为低度多态。4个突变位点中,G309A突变AA型背膘厚极显著高于GG型(P<0.01)；AG型背膘厚显著高于GG型(P<0.05),眼肌面积、出生重AA型显著高于AG型、GG型(P<0.05)；G1232A突变背膘厚AA型、AG型显著高于GG型(P<0.05)；日增重AA型显著高于GG型(P<0.05)。据此推测：MC4R基因是影响德国美利奴羊生产性状的QTL或与控制这些性状的QTL有连锁,因此有关标记将来可能用于这些性状的标记辅助选择。
5CAST基因在德国美利奴羊中的多态性及其与部分肉用性状的相关性
德国美利奴羊在CAST基因上存在CAST3-1、CAST3-2、CAST3-3、CAST3-4、CAST7-1、CAST7-2和CAST7-3等7个突变位点,统计结果：CAST3-2、CAST3-3和CAST7-1突变位点达到中度多态,其余4个突变位点为低度多态。但这7个突变位点与出生重、日均增重、背膘厚、眼肌面积等生产性能均未达到显著相关,这些突变位点是否影响其他生产或肉品质性状还有待于进一步研究。
Sheep breeding involves in improving growth and reproductive characteristics. To date many genes have been found to control these traits. BMPR-IB, BMP15, and GDF9, have been found to be the major genes that influence sheep reproductive capacity. Melanocortin-4receptor (MC4R) is a G-protein coupled receptor expressed in the brain, predominantly in the hypothalamus where it controls food intake, energy homeostasis and body weight. Calpastatin (CAST), one member of calpains system, is a specific inhibitor of calpains and is believed to be involved in the degradation of myofibrillar proteins in post-mortem proteolysis through regulating calpain activity, and hence it is implicated in meat tenderization.
Here we used ligase detection reaction (LDR) and bioinformatics techniques to investigate the polymorphisms of BMPR-IB, BMP15, and GDF9genes in Bayanbulak (including low and high yielding groups), German Mutton Merino, and Duolang sheep, and correlated the polymorphisms with litter size. The polymorphisms of MC4R and CAST gene in German Mutton Merino were also determined, hoping to find clues to the marker assisted selection (MAS) in sheep production. This study included two experiments:
I One hundred ordinary Bayanbulak sheep having single lamb (Flock B),20special Bayanbulak sheep having more than one lamb (Flock A), German Mutton Merino with complete lambing record in Inner Mongolia, and Bazhou Duolang sheep were studied using LDR to detect10SNPs in BMPR-IB, BMP15, and GDF9. The loci included FecB in BMPR-IB, FecXI, FecXB, FecXL, FecXH, FecXG, and FecXR in BMP15, and G1, G8, and FecTT in GDF9. The polymorphisms of BMPR-IB, BMP15, and GDF9were then compared among ordinary Bayanbulak group, special Bayanbulak group, German Mutton Merino, and Duolang Sheep. An association of the SNPs with litter size was subsequently conducted.
II Lambs of German Mutton Merino in Inner Mongolia with complete record of production performance (including birth weight, back-fat, eye muscle area and average daily gain) were studied using LDR to detect SNPs in MC4R gene and the second exon of the CAST gene. An association of the SNPs with growth trait and meat performance. The results were as follows:
1Polymorphisms of BMPR-IB gene in Bayanbulak, German Mutton Merino, and Duolang sheep, and its association with litter size
Polymorphisms of BMPR-1B/FecB gene were found in the Bayanbulak sheep. The genotypic frequency of the wild type (Fec++), heterozygous type (FecB+), and mutant (FecBB) were0.15,0.7, and0.15in flock A, and1.0,0, and0in flock B, respectively. The corresponding genotype frequency of wild type and mutation type were0.5,0.5in flock A and1.0,0.0in flock B, respectively. The independence test results of flock A and B suggested that the FecB locus significantly affects the litter size in the Bayanbulak sheep. BMPR-IB/FecB can be considered as the major gene that determines litter size in Bayanbulak sheep. Accordingly, this genetic marker may be used to improve litter size through MAS. No polymorphism of BMPR-IB/FecB mutation was found in German Mutton Merino and Duolang sheep. Therefore, the BMPR-IB/FecB mutation may not be correlated with the reproductive performance in the German Mutton Merino and Duolang sheep.
2Polymorphisms of BMP15gene in Bayanbulak, German Mutton Merino, and Duolang sheep, and its association with litter size
No polymorphism was detected at FecX-I、FecXB、FecXL、FecXH、FecX and FecXR of BMP15gene in the above three breeds. Therefore, the six loci of the BMP15gene cannot be used as a genetic marker for litter size in the three sheep breeds.
3Polymorphisms of GDF9gene in Bayanbulak, German Mutton Merino, and Duolang sheep, and its relation to litter size
The G1mutation was detected in the GDF9gene of Bayanbulak and German Mutton Merino sheep. However, FecG (G8) and FecTT mutations were not detected in the two sheep breeds. The polymorphism of G1mutation was moderate, and the locus was within the Hardy-Weinberg equilibrium in the two groups. No association of GDF9polymorphisms with litter size was found. There was no SNP of GDF9gene in Duolang sheep. These results indicated that the three site of GDF9gene does not significantly influence litter size of Bayanbulak, German Mutton Merino, and Duolang sheep.
4Polymorphisms of MC4R gene in German Mutton Merino and its association with meat performance
Four SNPs of MC4R gene were detected in German Mutton Merino sheep, namely, G309A, G508A, G1232A, and T1456C. All SNPs were within Hardy-Weinberg equilibrium. The SNP G1232A was moderate polymorphic, whereas the other three were slightly polymorphic. The sheep with AA type at G309A SNP had higher back fat than those with GG type (P<0.01). The back fat of sheep with AG type was higher than those with GG type (P<0.05). The eye muscle area and birth weight of AA sheep were higher than those of AG and GG sheep (P<0.05). Sheep with AA and AG genotype at the SNP G1232A had higher back fat than those with GG type (P<0.05). Average daily gain of AA genotype sheep was higher than that of GG genotypes (P<0.05). These findings indicated that the MC4R gene might be the QTL that influences production traits or it is linkage to the QTL that controls the traits. The markers may be applied in the MAS.
5Polymorphisms of CAST gene in German Mutton Merino and its association with meat performance
Seven SNPs of CAST gene were found in German Mutton Merino, namely, CAST3-1, CAST3-2, CAST3-3, CAST3-4, CAST7-1, CAST7-2, and CAST7-3. The statistical results showed that CAST3-2, CAST3-3, and CAST7-1were moderately polymorphic, whereas the other four SNPs were slightly polymorphic. The seven SNPs, however, were not significantly correlated to birth weight, average daily gain, back fat, eye muscle area. Further studies should be focused on find association of these SNPs with other production characteristcs.
引文
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