影响绵羊高繁殖力和非季节性发情基因的研究
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摘要
本论文以褪黑激素受体1b(melatonin receptor 1b, MTNR1B)、抑制素βA亚基基因(inhibinβA, INHBA)、kisspeptin及其受体G蛋白偶联受体54(Gprotein-coupled receptor 54,GPR54)、绵羊骨形态发生蛋白6(bone morphogenetic protein 6,BMP6)基因为研究对象,用PCR单链构象多态性(PCR Single-Strand Conformation Polymorphism,PCR-SSCP)分析、RT-PCR、基因克隆等技术研究这些基因在常年发情的高繁殖力绵羊品种以及季节性发情的低繁殖力绵羊品种中的遗传变异及其对绵羊繁殖性能的影响,寻找影响小尾寒羊高繁殖力和常年发情的主效基因或有较大关联的基因型或基因位点,以探索小尾寒羊高繁殖力及常年发情的分子遗传机理,为绵羊高繁殖力的标记辅助选择提供科学依据。实验主要得出如下一些结果：
根据人褪黑激素受体lb基因外显子2和预测的牛褪黑激素受体mRNA序列设计5对引物,检测MTNR1B外显子2在常年发情绵羊品种(小尾寒羊、多赛特、湖羊)和季节性发情绵羊品种(特克塞尔、考力代)中的单核苷酸多态性。测序结果表明突变型与野生型序列相比存在33个碱基突变,其中14个引起氨基酸改变。分析表明这些突变均与绵羊季节性发情相关性不大,MTNR1B基因可能不是影响绵羊季节性发情的主要基因。比较发现获得的绵羊褪黑激素受体1b基因外显子2序列与牛、猪的系统进化关系较与人、小鼠的近,它与牛(95%)和猪(79%)的褪黑激素受体1b氨基酸同源性高于与人(76%)和小鼠(71%),而且与绵羊、人和小鼠的褪黑激素受体la的氨基酸序列也有较高的同源性(61%-63%)。结构分析发现绵羊褪黑激素受体lb具有35个特异变化的氨基酸,并在第3跨膜区后具有DRY和CYVCR序列,这与牛的褪黑激素受体1b一致,而不同于已知的其它褪黑激素受体家族中的NRY和CYICH结构,但和已知的其它褪黑激素受体家族一样,它在第7跨膜区内含有一个NAXXY序列。我们认为可能是由于绵羊褪黑激素受体lb这种结构上的差异导致了它与配体(主要是褪黑激素)结合功能的改变,或者导致其受体的特定结构域改变和磷酸化途径,并由于基因替代效应的影响,使它在绵羊体内的功能逐渐被弱化,在进化中其功能逐渐被其它基因取代(可能主要是褪黑激素受体1a),以致于其表达现在很难检测到。
根据牛抑制素βA亚基基因外显子1、外显子2和绵羊抑制素βA亚基mRNA序列设计5对引物,检测抑制素βA亚基基因的全部编码区和部分3’UTR序列在2个高繁殖力绵羊品种(小尾寒羊、湖羊)以及6个低繁殖力绵羊品种(多赛特羊、特克塞尔羊、德国肉用美利奴羊、南非肉用美利奴、考力代、中国美利奴绵羊)中的单核苷酸多态性,分析该基因对绵羊高繁殖力的影响。结果发现引物3、引物4与引物5扩增片段有多态性,其余2对引物的扩增片段不存在多态性。对于引物3,发现AA、BB、CC三种基因型型,中国美利奴绵羊都为CC型,湖羊含有AA和BB型,其余6个品种均为AA型。BB型与AA型相比有8个碱基突变,CC型与AA型相比有4个碱基突变,这些突变均没引起氨基酸改变。对于引物4,发现EE、EF、FF、GG和EG五种基因型型,其中小尾寒羊、湖羊、特克塞尔羊、南非肉用美利奴和考力代羊等5个品种都为EE型,多赛特羊和德国肉用美利奴羊均含有EE、GG和EG三种基因型,中国美利奴绵羊中含有EE、FF和EF三种基因型。FF型在对应EE序列的第114 bp处有一个G→A的单碱基沉默突变。GG型在对应EE序列的第143bp处有一个C→T的单碱基突变,即TCG→TTG,引起亲水的丝氨酸(Ser)→疏水的亮氨酸(Leu),该位点对应GenBank发表的绵羊抑制素βA亚基的第287位丝氨酸(Ser)残基。对于引物5,发现KK、LL、KL和MM四种基因型,其中MM基因型只在湖羊中检测到。LL型在KK型第218bp处有一个G→A的单碱基突变,MM型与KK型相比有9个的碱基突变,这些突变均为没引起氨基酸改变。小尾寒羊只在引物5中检测到多态,LL型小尾寒羊产羔数最小二乘均值分别比KL型和KK型小尾寒羊多0.53只(P<0.05)和0.63只(P<0.05)。KL基因型小尾寒羊的产羔数的最小二乘均值比KK基因型的分别多0.10 (P>0.05)。INHBA基因编码区在湖羊、多赛特羊和德国肉用美利奴羊中有较高的碱基突变率,以湖羊为最高,在小尾寒羊、特克塞尔羊、南非肉用美利奴、考力代和中国美利奴绵羊中的碱基突变率较低。
采用PCR-SSCP技术检测kisspeptin基因外显子1和GPR54基因外显子1、外显子2和部分外显子5序列在常年发情的高繁殖力绵羊品种(小尾寒羊、湖羊)和季节性发情的低繁殖力绵羊品种(多赛特、特克塞尔、考力代)中的单核苷酸多态性,并分析其对绵羊繁殖性能的影响。结果发现Kisspeptin基因的外显子1在常年发情的小尾寒羊(出现AA、BB和AB 3种基因型)、湖羊(出现AA朋和CC两种基因型)中均有多态性,在季节性发情的绵羊品种中均无多态性(只有AA型)：GPR54基因的外显子2在常年发情的湖羊中有多态性(出现DD和EE两种基因型),而在常年发情的小尾寒羊和季节性发情的绵羊品种中均无多态性(只有DD型)；GPR54基因的外显子1和部分外显子5序列在5个绵羊品种中均不存在多态性。测序发现BB与AA型相比有1个突变(G73A),导致Val25Met; CC型与AA型相比有7个碱基突变(C19T、C33T、T34C、C63A、T64C、C72G、C77T),其中4个突变引起氨基酸改变,即分别为Arg7Trp、Phe12Leu、Asn24Lys、Ala26Val。EE型与DD型相比有T269C、A320C两个突变,分别导致Met90Thr和Aspl07Ala。小尾寒羊AA、BB和AB基因型频率分别为0.62、0.05和0.33。BB型和AB型小尾寒羊平均产羔数分别比AA型多0.78只(P<0.05)和0.45只(P<0.05),BB型和AB型小尾寒羊平均产羔数差异不显著(P>0.05)。这些结果初步表明kisspeptin基因与绵羊的常年发情和高产羔数有一定的相关性。
根据GenBank发表的绵羊骨形态发生蛋白6基因部分序列所包含的外显子5、6、7和小鼠BMP6基因外显子5、6、7序列设计3对引物,检测了BMP6基因外显子5、外显子6和外显子7在高繁殖力绵羊品种(小尾寒羊、湖羊)和低繁殖力绵羊品种(多赛特、特克塞尔、考力代)中的单核苷酸多态性,同时研究该基因对小尾寒羊高繁殖力的影响。结果发现该3对引物在所检测的5个品种中均无多态性,说明所检测的BMP6外显子5、6、7序列比较保守,该区域可能不是影响绵羊高繁殖力的功能结构域。同时克隆了绵羊(695bp)和山羊(699bp)部分BMP6 mRNA和绵羊exon3-4的内含子序列和两端部分外显子序列(785bp)。发现绵羊和山羊的碱基和氨基酸序列差异很少。两者的碱基序列同源性高达99.28%,氨基酸序列同源性为98.1%,除山羊中插入一个丙氨酸外,.只有4个氨基酸的差异。绵羊与牛、人、小鼠和大鼠的碱基同源性分别为97.04%、81.82%、84.68%和85.06%；氨基酸序列同源性分别为99.05%、91.43%、90.48%和92.38%,均大于90%,说明BMP6在各物种碱基序列虽然差异较大,但氨基酸序列却非常保守。
The melatonin receptor 1b(MTNR1B), inhibinβA(INHBA),kisspeptin, G protein-coupled receptor 54(GPR54),bone morphogenetic protein 6 (BMP6) were studied as candidate genes for the prolificacy and year-round estrus in sheep in the present study by techniques of PCR single-strand conformational polymorphism (PCR-SSCP),RT-PCR, gene cloning. The results were as follows:
To analyze the structure and function of the melatonin 1b receptor (MT2) in sheep, single nucleotide polymorphisms were detected in exon 2 of sheep MT2 gene using genomic DNA from five sheep breeds by five primers. Polymorphisms were found.33 nucleotide mutations were revealed by comparing the mutant types with the wild types. Among them,14 give rise to deduced amino acid changes.However, none is likely to be associated with non-seasonal or seasonal estrus in sheep breeds tested. Sequence of exon 2 of MT2 of Small Tail Han sheep shows much closer phylogenetic relationship with predicted bovine and porcine MT2 than with human and mouse. The deduced amino acid sequence shows higher identity with the MT2 of cattle (95%) and pig (79%) than with human (76%) and mouse (71%).A rather high identity (61-63%) with the MT1 of sheep, human and mouse was also found. Compared with the other known MT2,35 unique altered amino acids were revealed. Albeit it also contains a NAXXY motif in transmembrane 7, both a DRY motif and a CYVCR motif were detected just downstream from its third transmembrane domain rather than NRY and CYICH found in other melatonin receptor groups.We presumed that it is possible that the structural changes make its binding function to the ligands attenuated or disrupted, and other genes (most probably MT1)were substituted in the progress of evolution, which ultimately resulted in no detectable expression in current breeds of sheep.
Inhibins participate in the regulation of pituitary follicle-stimulating hormone synthesis and secretion, follicular maturation and steroidogenesis in the female. Inhibin (3a gene (INHBA)was studied as a candidate gene for the prolificacy of sheep. Single nucleotide polymorphisms of entire coding region and partial 3' untranslated region of INHBA were detected by PCR-SSCP in both two high fecundity breeds (Small Tail Han and Hu sheep) and six low fecundity breeds (Dorset, Texel, German Mutton Merino, South African Mutton Merino, Chinese Merino and Corriedale sheep).Only the PCR products amplified by primers 3,4 and 5 displayed polymorphisms.For primer 3,genotype CC was only detected in Chinese Merino sheep, genotype AA was detected in other seven sheep breeds. Genotype BB was only detected in Hu sheep. Only Hu sheep displayed polymorphism. Eight or four nucleotide mutations were revealed between BB or CC and AA, respectively, and these mutations did not result in any amino acid change. For primer 4, genotypes EE, EG and GG were detected in Dorset and German Mutton Merino sheep, genotypes EE, EF and FF were detected in Chinese Merino sheep, only genotype EE was detected in other five sheep breeds. Only Dorset, German Mutton Merino and Chinese Merino sheep displayed polymorphism. Sequencing revealed one nucleotide mutation (114G→A) of exon 2 of INHBA gene between genotype FF and genotype EE, and this mutation did not cause any amino acid change. Another nucleotide change(143C→T) was identified between genotype GG and genotype EE, and this mutation resulted in an amino acid change of serine→leucine.For primer 5,genotypes KK and KL were detected in German Mutton Merino and Corriedale sheep, genotypes KK, LL and KL were detected in other six sheep breeds. Genotype MM was only detected in Hu sheep.All of these eight sheep breeds displayed polymorphism. Sequencing revealed one nucleotide mutation (218A→G) of exon 2 of INHBA gene between genotype LL and genotype KK, and nine nucleotide mutations between genotype MM and genotype KK. These mutations did not alter amino acid sequence.The partial sequence (395 bp for exon 1 and 933 bp for exon 2) of INHBA gene in Small Tail Han sheep (with genotype KK for primer 5)was submitted into GenBank (accession number EF192431).Small Tail Han sheep displayed polymorphisms only in fragment amplified by primer 5.The Small Tail Han ewes with genotype LL had 0.53 (p<0.05)or 0.63 (p<0.05)lambs more than those with genotype KL or KK, respectively. The Small Tail Han ewes with genotype KL had 0.10 (p>0.05) lambs more than those with genotype KK.
According to the sequence of the predicted Bos taurus KiSS-1 gene and its receptor GPR54 gene, four pairs of primers were designed to detect single nucleotide polymorphisms of exon 1 of KiSS-1 gene and exon 1,exon 2 and partial exon 5 of GPR54 gene in high fecundity and year-round estrous breeds(Small Tail Han and Hu sheep) and low fecundity and seasonal estrous breeds (Dorset, Texel and Corriedale sheep) by PCR-SSCP. Polymorphisms in exon 1 of KiSS-1 gene were found in year-round estrous Small Tail Han sheep (AA,BB and AB genotypes) and Hu sheep (AA and CC genotypes), no polymorphism was found in seasonal estrous sheep breeds (only AA genotype).Polymorphisms in exon 2 of GPR54 gene were found in year-round estrous Hu sheep (DD and EE genotypes), no polymorphism was found in year-round estrous Small Tail Han sheep and seasonal estrous sheep breeds (only DD genotype).No polymorphism was detected in exon 1 and partial exon 5 of GPR54 gene in five sheep breeds. The polymorphic genotypes were sequenced. While compared the BB genotype with the AA genotype, one nucleotide mutation (G73A) was detected, which resulted in amino acid change, Val25Met. Seven nucleotide mutations were detected from AA to CC genotype (C19T, C33T, T34C,C63A, T64C,C72G, C77T),and among them 4 make amino acid changes, that is, Arg7Trp,Phel2Leu, Asn24Lys, Ala26Val.While compared the EE genotype with the DD genotype, two nucleotide mutations (T269C, A320C) were detected, which give rise to amino acid changes, Met90Thr and Asp107Ala, respectively. Genotype frequencies of AA,BB and AB were 0.62,0.05 and 0.33,respectively. The Small Tail Han ewes with genotypes BB and AB had 0.78 (P<0.05) or 0.45 (P<0.05) lambs more than those with genotype AA;the Small Tail Han ewes with genotype BB had 0.33 (P>0.05)lambs more than those with genotype AB. These results preliminarily indicate that the KiSS-1 gene may have some association with prolificacy and year-round estrus in sheep.
The bone morphogenetic protein 6 gene was studied as a candidate gene for the prolificacy of Small Tail Han sheep.Single nucleotide polymorphisms(SNPs) of exons 5,6,7 of BMP6 gene were detected in high fecundity breeds (Small Tail Han and Hu sheep) and low fecundity breeds (Dorset, Texel and Corriedale sheep) with three,pairs of primers by polymerase chain reaction (PCR)-single strand conformation polymorphism (SSCP).No polymorphisms were detected in the amplified regions of the three pairs of primers in the five sheep breeds tested, which may means that the sequences of exons 5,6,7 of BMP6 gene were rather conserved, and these regions may not be the functional domains that affect the prolific performance of sheep.On the other hand, partial sequences of BMP6 mRNA were cloned from sheep and goat,695bp and 699bp respectively, and the sequences of intron between exon 3 and exon 4 with some of the two flanking exons were also sequenced. The two BMP6 sequences show high identical with 99.28% in nucleotide and 98.1% in amino acid sequence, and besides an insert of alanine in goat, only 4 alterations of amino acids were found between them (211 in total).Compared the ovine nucleotide and amino acid sequence of BMP6 with that of cattle, human, mouse and rat, identity of 97.04%、81.82%,84.68%,85.06% and identity of 99.05%, 91.43%,90.48%,92.38% were revealed respectively.The results suggested that although the change of nucleotides is a little big, but the sequence of amino acid of BMP6 is very conservative among species.
根据人褪黑激素受体lb基因外显子2和预测的牛褪黑激素受体mRNA序列设计5对引物,检测MTNR1B外显子2在常年发情绵羊品种(小尾寒羊、多赛特、湖羊)和季节性发情绵羊品种(特克塞尔、考力代)中的单核苷酸多态性。测序结果表明突变型与野生型序列相比存在33个碱基突变,其中14个引起氨基酸改变。分析表明这些突变均与绵羊季节性发情相关性不大,MTNR1B基因可能不是影响绵羊季节性发情的主要基因。比较发现获得的绵羊褪黑激素受体1b基因外显子2序列与牛、猪的系统进化关系较与人、小鼠的近,它与牛(95%)和猪(79%)的褪黑激素受体1b氨基酸同源性高于与人(76%)和小鼠(71%),而且与绵羊、人和小鼠的褪黑激素受体la的氨基酸序列也有较高的同源性(61%-63%)。结构分析发现绵羊褪黑激素受体lb具有35个特异变化的氨基酸,并在第3跨膜区后具有DRY和CYVCR序列,这与牛的褪黑激素受体1b一致,而不同于已知的其它褪黑激素受体家族中的NRY和CYICH结构,但和已知的其它褪黑激素受体家族一样,它在第7跨膜区内含有一个NAXXY序列。我们认为可能是由于绵羊褪黑激素受体lb这种结构上的差异导致了它与配体(主要是褪黑激素)结合功能的改变,或者导致其受体的特定结构域改变和磷酸化途径,并由于基因替代效应的影响,使它在绵羊体内的功能逐渐被弱化,在进化中其功能逐渐被其它基因取代(可能主要是褪黑激素受体1a),以致于其表达现在很难检测到。
根据牛抑制素βA亚基基因外显子1、外显子2和绵羊抑制素βA亚基mRNA序列设计5对引物,检测抑制素βA亚基基因的全部编码区和部分3’UTR序列在2个高繁殖力绵羊品种(小尾寒羊、湖羊)以及6个低繁殖力绵羊品种(多赛特羊、特克塞尔羊、德国肉用美利奴羊、南非肉用美利奴、考力代、中国美利奴绵羊)中的单核苷酸多态性,分析该基因对绵羊高繁殖力的影响。结果发现引物3、引物4与引物5扩增片段有多态性,其余2对引物的扩增片段不存在多态性。对于引物3,发现AA、BB、CC三种基因型型,中国美利奴绵羊都为CC型,湖羊含有AA和BB型,其余6个品种均为AA型。BB型与AA型相比有8个碱基突变,CC型与AA型相比有4个碱基突变,这些突变均没引起氨基酸改变。对于引物4,发现EE、EF、FF、GG和EG五种基因型型,其中小尾寒羊、湖羊、特克塞尔羊、南非肉用美利奴和考力代羊等5个品种都为EE型,多赛特羊和德国肉用美利奴羊均含有EE、GG和EG三种基因型,中国美利奴绵羊中含有EE、FF和EF三种基因型。FF型在对应EE序列的第114 bp处有一个G→A的单碱基沉默突变。GG型在对应EE序列的第143bp处有一个C→T的单碱基突变,即TCG→TTG,引起亲水的丝氨酸(Ser)→疏水的亮氨酸(Leu),该位点对应GenBank发表的绵羊抑制素βA亚基的第287位丝氨酸(Ser)残基。对于引物5,发现KK、LL、KL和MM四种基因型,其中MM基因型只在湖羊中检测到。LL型在KK型第218bp处有一个G→A的单碱基突变,MM型与KK型相比有9个的碱基突变,这些突变均为没引起氨基酸改变。小尾寒羊只在引物5中检测到多态,LL型小尾寒羊产羔数最小二乘均值分别比KL型和KK型小尾寒羊多0.53只(P<0.05)和0.63只(P<0.05)。KL基因型小尾寒羊的产羔数的最小二乘均值比KK基因型的分别多0.10 (P>0.05)。INHBA基因编码区在湖羊、多赛特羊和德国肉用美利奴羊中有较高的碱基突变率,以湖羊为最高,在小尾寒羊、特克塞尔羊、南非肉用美利奴、考力代和中国美利奴绵羊中的碱基突变率较低。
采用PCR-SSCP技术检测kisspeptin基因外显子1和GPR54基因外显子1、外显子2和部分外显子5序列在常年发情的高繁殖力绵羊品种(小尾寒羊、湖羊)和季节性发情的低繁殖力绵羊品种(多赛特、特克塞尔、考力代)中的单核苷酸多态性,并分析其对绵羊繁殖性能的影响。结果发现Kisspeptin基因的外显子1在常年发情的小尾寒羊(出现AA、BB和AB 3种基因型)、湖羊(出现AA朋和CC两种基因型)中均有多态性,在季节性发情的绵羊品种中均无多态性(只有AA型)：GPR54基因的外显子2在常年发情的湖羊中有多态性(出现DD和EE两种基因型),而在常年发情的小尾寒羊和季节性发情的绵羊品种中均无多态性(只有DD型)；GPR54基因的外显子1和部分外显子5序列在5个绵羊品种中均不存在多态性。测序发现BB与AA型相比有1个突变(G73A),导致Val25Met; CC型与AA型相比有7个碱基突变(C19T、C33T、T34C、C63A、T64C、C72G、C77T),其中4个突变引起氨基酸改变,即分别为Arg7Trp、Phe12Leu、Asn24Lys、Ala26Val。EE型与DD型相比有T269C、A320C两个突变,分别导致Met90Thr和Aspl07Ala。小尾寒羊AA、BB和AB基因型频率分别为0.62、0.05和0.33。BB型和AB型小尾寒羊平均产羔数分别比AA型多0.78只(P<0.05)和0.45只(P<0.05),BB型和AB型小尾寒羊平均产羔数差异不显著(P>0.05)。这些结果初步表明kisspeptin基因与绵羊的常年发情和高产羔数有一定的相关性。
根据GenBank发表的绵羊骨形态发生蛋白6基因部分序列所包含的外显子5、6、7和小鼠BMP6基因外显子5、6、7序列设计3对引物,检测了BMP6基因外显子5、外显子6和外显子7在高繁殖力绵羊品种(小尾寒羊、湖羊)和低繁殖力绵羊品种(多赛特、特克塞尔、考力代)中的单核苷酸多态性,同时研究该基因对小尾寒羊高繁殖力的影响。结果发现该3对引物在所检测的5个品种中均无多态性,说明所检测的BMP6外显子5、6、7序列比较保守,该区域可能不是影响绵羊高繁殖力的功能结构域。同时克隆了绵羊(695bp)和山羊(699bp)部分BMP6 mRNA和绵羊exon3-4的内含子序列和两端部分外显子序列(785bp)。发现绵羊和山羊的碱基和氨基酸序列差异很少。两者的碱基序列同源性高达99.28%,氨基酸序列同源性为98.1%,除山羊中插入一个丙氨酸外,.只有4个氨基酸的差异。绵羊与牛、人、小鼠和大鼠的碱基同源性分别为97.04%、81.82%、84.68%和85.06%；氨基酸序列同源性分别为99.05%、91.43%、90.48%和92.38%,均大于90%,说明BMP6在各物种碱基序列虽然差异较大,但氨基酸序列却非常保守。
The melatonin receptor 1b(MTNR1B), inhibinβA(INHBA),kisspeptin, G protein-coupled receptor 54(GPR54),bone morphogenetic protein 6 (BMP6) were studied as candidate genes for the prolificacy and year-round estrus in sheep in the present study by techniques of PCR single-strand conformational polymorphism (PCR-SSCP),RT-PCR, gene cloning. The results were as follows:
To analyze the structure and function of the melatonin 1b receptor (MT2) in sheep, single nucleotide polymorphisms were detected in exon 2 of sheep MT2 gene using genomic DNA from five sheep breeds by five primers. Polymorphisms were found.33 nucleotide mutations were revealed by comparing the mutant types with the wild types. Among them,14 give rise to deduced amino acid changes.However, none is likely to be associated with non-seasonal or seasonal estrus in sheep breeds tested. Sequence of exon 2 of MT2 of Small Tail Han sheep shows much closer phylogenetic relationship with predicted bovine and porcine MT2 than with human and mouse. The deduced amino acid sequence shows higher identity with the MT2 of cattle (95%) and pig (79%) than with human (76%) and mouse (71%).A rather high identity (61-63%) with the MT1 of sheep, human and mouse was also found. Compared with the other known MT2,35 unique altered amino acids were revealed. Albeit it also contains a NAXXY motif in transmembrane 7, both a DRY motif and a CYVCR motif were detected just downstream from its third transmembrane domain rather than NRY and CYICH found in other melatonin receptor groups.We presumed that it is possible that the structural changes make its binding function to the ligands attenuated or disrupted, and other genes (most probably MT1)were substituted in the progress of evolution, which ultimately resulted in no detectable expression in current breeds of sheep.
Inhibins participate in the regulation of pituitary follicle-stimulating hormone synthesis and secretion, follicular maturation and steroidogenesis in the female. Inhibin (3a gene (INHBA)was studied as a candidate gene for the prolificacy of sheep. Single nucleotide polymorphisms of entire coding region and partial 3' untranslated region of INHBA were detected by PCR-SSCP in both two high fecundity breeds (Small Tail Han and Hu sheep) and six low fecundity breeds (Dorset, Texel, German Mutton Merino, South African Mutton Merino, Chinese Merino and Corriedale sheep).Only the PCR products amplified by primers 3,4 and 5 displayed polymorphisms.For primer 3,genotype CC was only detected in Chinese Merino sheep, genotype AA was detected in other seven sheep breeds. Genotype BB was only detected in Hu sheep. Only Hu sheep displayed polymorphism. Eight or four nucleotide mutations were revealed between BB or CC and AA, respectively, and these mutations did not result in any amino acid change. For primer 4, genotypes EE, EG and GG were detected in Dorset and German Mutton Merino sheep, genotypes EE, EF and FF were detected in Chinese Merino sheep, only genotype EE was detected in other five sheep breeds. Only Dorset, German Mutton Merino and Chinese Merino sheep displayed polymorphism. Sequencing revealed one nucleotide mutation (114G→A) of exon 2 of INHBA gene between genotype FF and genotype EE, and this mutation did not cause any amino acid change. Another nucleotide change(143C→T) was identified between genotype GG and genotype EE, and this mutation resulted in an amino acid change of serine→leucine.For primer 5,genotypes KK and KL were detected in German Mutton Merino and Corriedale sheep, genotypes KK, LL and KL were detected in other six sheep breeds. Genotype MM was only detected in Hu sheep.All of these eight sheep breeds displayed polymorphism. Sequencing revealed one nucleotide mutation (218A→G) of exon 2 of INHBA gene between genotype LL and genotype KK, and nine nucleotide mutations between genotype MM and genotype KK. These mutations did not alter amino acid sequence.The partial sequence (395 bp for exon 1 and 933 bp for exon 2) of INHBA gene in Small Tail Han sheep (with genotype KK for primer 5)was submitted into GenBank (accession number EF192431).Small Tail Han sheep displayed polymorphisms only in fragment amplified by primer 5.The Small Tail Han ewes with genotype LL had 0.53 (p<0.05)or 0.63 (p<0.05)lambs more than those with genotype KL or KK, respectively. The Small Tail Han ewes with genotype KL had 0.10 (p>0.05) lambs more than those with genotype KK.
According to the sequence of the predicted Bos taurus KiSS-1 gene and its receptor GPR54 gene, four pairs of primers were designed to detect single nucleotide polymorphisms of exon 1 of KiSS-1 gene and exon 1,exon 2 and partial exon 5 of GPR54 gene in high fecundity and year-round estrous breeds(Small Tail Han and Hu sheep) and low fecundity and seasonal estrous breeds (Dorset, Texel and Corriedale sheep) by PCR-SSCP. Polymorphisms in exon 1 of KiSS-1 gene were found in year-round estrous Small Tail Han sheep (AA,BB and AB genotypes) and Hu sheep (AA and CC genotypes), no polymorphism was found in seasonal estrous sheep breeds (only AA genotype).Polymorphisms in exon 2 of GPR54 gene were found in year-round estrous Hu sheep (DD and EE genotypes), no polymorphism was found in year-round estrous Small Tail Han sheep and seasonal estrous sheep breeds (only DD genotype).No polymorphism was detected in exon 1 and partial exon 5 of GPR54 gene in five sheep breeds. The polymorphic genotypes were sequenced. While compared the BB genotype with the AA genotype, one nucleotide mutation (G73A) was detected, which resulted in amino acid change, Val25Met. Seven nucleotide mutations were detected from AA to CC genotype (C19T, C33T, T34C,C63A, T64C,C72G, C77T),and among them 4 make amino acid changes, that is, Arg7Trp,Phel2Leu, Asn24Lys, Ala26Val.While compared the EE genotype with the DD genotype, two nucleotide mutations (T269C, A320C) were detected, which give rise to amino acid changes, Met90Thr and Asp107Ala, respectively. Genotype frequencies of AA,BB and AB were 0.62,0.05 and 0.33,respectively. The Small Tail Han ewes with genotypes BB and AB had 0.78 (P<0.05) or 0.45 (P<0.05) lambs more than those with genotype AA;the Small Tail Han ewes with genotype BB had 0.33 (P>0.05)lambs more than those with genotype AB. These results preliminarily indicate that the KiSS-1 gene may have some association with prolificacy and year-round estrus in sheep.
The bone morphogenetic protein 6 gene was studied as a candidate gene for the prolificacy of Small Tail Han sheep.Single nucleotide polymorphisms(SNPs) of exons 5,6,7 of BMP6 gene were detected in high fecundity breeds (Small Tail Han and Hu sheep) and low fecundity breeds (Dorset, Texel and Corriedale sheep) with three,pairs of primers by polymerase chain reaction (PCR)-single strand conformation polymorphism (SSCP).No polymorphisms were detected in the amplified regions of the three pairs of primers in the five sheep breeds tested, which may means that the sequences of exons 5,6,7 of BMP6 gene were rather conserved, and these regions may not be the functional domains that affect the prolific performance of sheep.On the other hand, partial sequences of BMP6 mRNA were cloned from sheep and goat,695bp and 699bp respectively, and the sequences of intron between exon 3 and exon 4 with some of the two flanking exons were also sequenced. The two BMP6 sequences show high identical with 99.28% in nucleotide and 98.1% in amino acid sequence, and besides an insert of alanine in goat, only 4 alterations of amino acids were found between them (211 in total).Compared the ovine nucleotide and amino acid sequence of BMP6 with that of cattle, human, mouse and rat, identity of 97.04%、81.82%,84.68%,85.06% and identity of 99.05%, 91.43%,90.48%,92.38% were revealed respectively.The results suggested that although the change of nucleotides is a little big, but the sequence of amino acid of BMP6 is very conservative among species.
引文
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