奶牛精液品质和抗病及抗运输应激的分子机制研究
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摘要
种公牛的精液品质、疾病和应激是制约奶牛生产和经济效益的重要因素,这些因素的遗传力比较低,通过直接选择的方式很难取得较大的遗传进展。为了选育抗病力强的个体,本文研究了Toll样受体家族相关基因(TLR1、TLR2、TLR4、TLR6和TLR9)多态性与奶牛乳腺炎、结核和布病易感性之间的相关,并分析了BMP15基因多态性与奶牛精液品质性状之间的相关,以期利用分子育种手段进行抗性选择,提高奶牛的繁殖力和抗病力。运输作为动物生产中经常发生的、不可避免的生产活动,对动物造成很大的应激,如免疫功能的改变、发病率的显著增加、产奶量的降低和肉品质的改变。然而目前却鲜有关于长途运输应激对动物生殖生理影响的报道。因此本试验以小鼠为模式动物,一方面研究了运输前后雄性动物睾丸组织中主要激素受体(LHR和AR)及应激蛋白Hsp70mRNA和蛋白表达的变化,另一方面采用Agilent小鼠表达谱芯片筛选了不同孕期小鼠垂体、子宫和胎盘中的差异表达基因,为研究运输应激对牛生殖生理的影响及抗运输应激新产品的开发提供参考和依据。
1.BMP15基因多态性与中国荷斯坦奶牛精液品质性状的关系
采用候选基因法分析了BMP15基因多态性与中国荷斯坦奶牛精液品质性状的关系。在所设计的引物中只检测到一个突变位点C5697T,年龄效应和该突变位点的基因型效应对鲜精活力有显著的影响,然而二者的互作却对精液品质性状无显著影响。进一步分析了该突变位点不同基因型对鲜精活力的影响,CT基因型的鲜精活力显著低于CC和TT基因型。结果表明,BMP15基因可能是控制牛精液品质性状的候选基因。
2.牛抗病候选基因的鉴定
采用PCR-SSCP、PCR-RFLP和测序技术相结合,以TLRs基因家族中的TLR1、 TLR2、TLR4、TLR6和TLR9为候选基因,研究了其在中国荷斯坦牛群体中的遗传多态性,并分析了多态位点不同基因型与牛乳腺炎、结核和布病的相关性。主要研究结果如下：
(1)TLR1基因检测到了5个多态位点,分别是C632T、G1409A、A1475C、 G1550A和G1596A,其中C632T、G1409A和G1550A多态位点不同基因型在牛结核自然感染组和对照组间差异不显著,而G1596A位点不同基因型间有显著差异,GH基因型牛患结核的危险程度是GG基因型的2.43倍(95%CI1.47-4.03),A1475C位点不同基因型有显著差异的趋势(P=0.056)。
(2)TLR2基因检测到了3个突变位点T385G.G398A和C1828T,分别被EcoRV、 HaeⅢ和DraI限制性内切酶识别,T385G位点不同基因型与SCS及布病均存在显著性差异,GG基因型的SCS显著高于GT和TT基因型,其布病发生的危险程度是TT基因型牛的2.084倍(95%CI1.116-3.893),GG基因型可能是疾病易感基因型；G398A和C1828T位点不同基因型与SCS和布病间差异不显著。
(3)TLR4基因多态位点不同基因型在牛结核病感染组和对照组间没有显著差异,提示TLR4基因可能不是控制牛结核病性状的候选基因。
(4)TLR6基因检测到了4个突变位点,分别是G1793A、C1859A、G1934A和A1980G。其中C1859A和A1980G位点不同基因型在牛结核自然感染组和对照组间有显著差异,C1859A位点AA和CA基因型牛患结核的危险程度分别是CC基因型牛的2.73倍(95%CI0.869-8.573)和1.547倍(95%CI0.803-2.982),CC基因型可能是牛结核抗性基因型；A1980G位点GG和GA基因型牛患结核的危险程度分别是AA基因型牛的2.986倍(95%CI1.245-7.165)和1.582倍(95%C10.734-3.409),AA基因型可能是牛结核抗性基因型。
(5)TLR9基因多态位点不同基因型与牛结核和布病感染均没有显著相关,提示TLR9可能不是牛结核和布病的易感基因。
3.运输应激对动物生殖的影响
为了研究长途运输应激对动物生殖生理的影响,试验以健康成年昆明小鼠为模型,进行约15h(1000km)的长途运输,研究运输应激对雄性生殖和妊娠的影响。
(1)比较了运输前后小鼠附睾尾精子相对计数、体重和附睾指数的变化。结果表明,长途运输显著降低了模式动物小鼠的体重、附睾指数和附睾尾精子相对计数。为了分析其可能机制,进一步采用QPCR和Western blot方法分析了下丘脑-垂体-性腺轴主要激素受体和应激相关基因(LHR、AR和Hsp70) mRNA和蛋白表达的变化。AR的表达水平在经历长途运输后显著的降低,而LHR和Hsp70稍有升高,但差异不显著。
(2)与对照组相比,孕晚期长途运输对活胎率、死胎率和吸收胎率没有显著影响,但造成1只孕鼠早产；而孕中期长途运输则导致活胎率的显著下降和死胎率的明显增加。
(3)E10垂体、E10子宫、E15垂体和E15胎盘中分别筛选出133条、586条、146条和562条差异表达基因,这些差异基因的Go分类主要集中在细胞过程、生物调节、代谢过程、生物过程的调节等。
(4)以HPG轴中对生殖起主要调控作用的垂体和性腺组织为研究对象,筛选出了运输过程中可能对生殖起主要调控作用的信号通路和基因。
Reproduction, stress and disease are important factors that restricting dairy production and economic benefits, the estimated heritability values of these traits tend to be relatively low, and it is difficult to make great genetic progress through direct selection. In order to breed disease resistance individuals, we studied the polymorphisms of toll like receptor family and analyzed its association with mastitis, bovine tuberculosis and bovine brucellosis; we further analyzed the variants of BMP15with sperm quality traits, expecting to conduct resistance selection by molecular breeding and improve fertility and disease resistance of animals. Transport, as an frequent and inevitably behavior in animal production, would cause big stress to animals, for example, significant increase in disease incidence, however, up to now, there is little literature about the impact of transport stress on animal reproduction. Therefore, in this experiment, we use mice as model, investigated the mRNA and protein levels of LHR, AR and Hsp70in mice testis after long road transport; Second, screened differentially expressed genes in pituitary, uterus and placenta of different pregnancy phase by use of Agilent mouse microarray, and further functional annotation to some of the differentially expressed genes. To provide reference or basis for the study of transport stress on cattle reproductive physiology and product development of anti-transport stress.
1. BMP15gene polymorphisms and its association with sperm quality traits in Chinese Holstein cattle
We analyzed the association between polymorphism of BMP15gene and sperm quality traits. One SNP in intron (C5697T) was detected. Age and genotype had significant effect on sperm quality trait of fresh sperm motility, however, the interaction of age and genotype had no effect on any sperm quality traits. We further analyzed the effect of different genotype on fresh sperm motility and found that fresh sperm motility of CT genotype was significantly lower than that of CC or TT genotype. These results indicate that BMP15may be candidate genes for sperm quality traits.
2. Identification of candidate genes associated with bovine disease resistance
In this study, we first chose TLR1、TLR2、TLR、TLR6and TLR9as candidate genes to study their variants with mastitis, bovine tuberculosis and bovine brucellosis by PCR-SSCP, PCR-RFLP combined with sequencing. The main results are as follows:
(1) Five SNPs C632T, G1409A, A1475C, G1550A and G1596A were detected in TLR1gene. The allele and genotype frequencies of C632T, G1409A and G1550A did not vary significantly between bovine tuberculosis (bTB) infected and non-infected cattle. In G1596A site, genotype frequencies differed significantly between bTB-infected and non-infected cattle, and relative risk of bovine tuberculosis incidence of GH is2.43times than that of GG genotype; In A1475C site, there was a trend toward an increased relative risk of tuberculosis incidence in CD genotype (P=0.056).
(2) Three SNPs T385G, G398A and C1828T were detected in TLR2gene, they can be recognized by restriction enzyme EcoRV, HaeⅢ and Dral respectively. In T385G site, GG genotype had significantly higher SCS than GT and TT genotype, and its relative risk of bovine brucellosis incidence is2.084times than that of TT genotype and GG may be disease susceptibility genotype; No significant differences were observed between G398A and C1828T site in SCS and bovine brucellosis incidence.
(3) There were no significant differences between different genotypes of TLR4and bovine tuberculosis, and this may indicate TLR4is not a candidate gene in controlling bovine tuberculosis.
(4) Four SNPs G1793A, C1859A、G1934A and A1980G were detected in TLR6gene. There were significant differences between different genotypes of C1859A and A1980G and bovine tuberculosis. In C1859A, cattle with AA or CA genotype had greater relative risk of disease incidence (OR=2.73,95%CI0.869-8.573; OR=1.547,95%CI0.803-2.982) than those with CC genotype, and CC maybe bovine tuberculosis resistance genotype; In A1980G, cattle with GG or GA genotype had greater relative risk of disease incidence (OR=2.986,95%CI1.245-7.165; OR=1.582,95%CI0.734-3.409) than those of AA genotype, and AA maybe bovine tuberculosis resistance genotype.
(5) There were no significant differences between different genotypes of TLR9and bovine tuberculosis or bovine brucellosis, and this may indicate TLR9is not a candidate gene in controlling bovine tuberculosis or bovine brucellosis.
3. Effect of transport stress on animal reproduction
In order to study the effect of transport stress on animal reproduction, we chose Kunming mice as model to study its role in male and pregnant for about15h (1000km) long road transport.
(1) Long road transport significantly reduced body weight, epididymis index and relative sperm count in epididymis. In order to analyze the possible mechanism, the mRNA and protein expression of LHR, AR and Hsp70were conducted. Results showed that expression of AR was significantly lower than before transport, LHR and Hsp70expression were somewhat higher after transport, however, not significant.
(2) Compared with control group, there are no significant differences in living birth rate, dead birth rate and absorbed fetus rate on long road transport during the last trimester of pregnancy, while declined living birth rate and increased dead birth rate after transport during the second trimester of pregnancy.
(3)133genes,586genes,146genes and562genes were scanned out in E10pituitary, E10uterus, E15pituitary and E15placenta respectively. These differential expression genes, according to biological process of Go analysis, mainly focused on the cellular process, biological regulation, metabolic process and regulation of biological process.
(4) We use pituitary and gonadal tissue, which play main roles in reproduction, as our taget, screeded several pathways and genes that may play major role in reproduction during transportation.
1.BMP15基因多态性与中国荷斯坦奶牛精液品质性状的关系
采用候选基因法分析了BMP15基因多态性与中国荷斯坦奶牛精液品质性状的关系。在所设计的引物中只检测到一个突变位点C5697T,年龄效应和该突变位点的基因型效应对鲜精活力有显著的影响,然而二者的互作却对精液品质性状无显著影响。进一步分析了该突变位点不同基因型对鲜精活力的影响,CT基因型的鲜精活力显著低于CC和TT基因型。结果表明,BMP15基因可能是控制牛精液品质性状的候选基因。
2.牛抗病候选基因的鉴定
采用PCR-SSCP、PCR-RFLP和测序技术相结合,以TLRs基因家族中的TLR1、 TLR2、TLR4、TLR6和TLR9为候选基因,研究了其在中国荷斯坦牛群体中的遗传多态性,并分析了多态位点不同基因型与牛乳腺炎、结核和布病的相关性。主要研究结果如下：
(1)TLR1基因检测到了5个多态位点,分别是C632T、G1409A、A1475C、 G1550A和G1596A,其中C632T、G1409A和G1550A多态位点不同基因型在牛结核自然感染组和对照组间差异不显著,而G1596A位点不同基因型间有显著差异,GH基因型牛患结核的危险程度是GG基因型的2.43倍(95%CI1.47-4.03),A1475C位点不同基因型有显著差异的趋势(P=0.056)。
(2)TLR2基因检测到了3个突变位点T385G.G398A和C1828T,分别被EcoRV、 HaeⅢ和DraI限制性内切酶识别,T385G位点不同基因型与SCS及布病均存在显著性差异,GG基因型的SCS显著高于GT和TT基因型,其布病发生的危险程度是TT基因型牛的2.084倍(95%CI1.116-3.893),GG基因型可能是疾病易感基因型；G398A和C1828T位点不同基因型与SCS和布病间差异不显著。
(3)TLR4基因多态位点不同基因型在牛结核病感染组和对照组间没有显著差异,提示TLR4基因可能不是控制牛结核病性状的候选基因。
(4)TLR6基因检测到了4个突变位点,分别是G1793A、C1859A、G1934A和A1980G。其中C1859A和A1980G位点不同基因型在牛结核自然感染组和对照组间有显著差异,C1859A位点AA和CA基因型牛患结核的危险程度分别是CC基因型牛的2.73倍(95%CI0.869-8.573)和1.547倍(95%CI0.803-2.982),CC基因型可能是牛结核抗性基因型；A1980G位点GG和GA基因型牛患结核的危险程度分别是AA基因型牛的2.986倍(95%CI1.245-7.165)和1.582倍(95%C10.734-3.409),AA基因型可能是牛结核抗性基因型。
(5)TLR9基因多态位点不同基因型与牛结核和布病感染均没有显著相关,提示TLR9可能不是牛结核和布病的易感基因。
3.运输应激对动物生殖的影响
为了研究长途运输应激对动物生殖生理的影响,试验以健康成年昆明小鼠为模型,进行约15h(1000km)的长途运输,研究运输应激对雄性生殖和妊娠的影响。
(1)比较了运输前后小鼠附睾尾精子相对计数、体重和附睾指数的变化。结果表明,长途运输显著降低了模式动物小鼠的体重、附睾指数和附睾尾精子相对计数。为了分析其可能机制,进一步采用QPCR和Western blot方法分析了下丘脑-垂体-性腺轴主要激素受体和应激相关基因(LHR、AR和Hsp70) mRNA和蛋白表达的变化。AR的表达水平在经历长途运输后显著的降低,而LHR和Hsp70稍有升高,但差异不显著。
(2)与对照组相比,孕晚期长途运输对活胎率、死胎率和吸收胎率没有显著影响,但造成1只孕鼠早产；而孕中期长途运输则导致活胎率的显著下降和死胎率的明显增加。
(3)E10垂体、E10子宫、E15垂体和E15胎盘中分别筛选出133条、586条、146条和562条差异表达基因,这些差异基因的Go分类主要集中在细胞过程、生物调节、代谢过程、生物过程的调节等。
(4)以HPG轴中对生殖起主要调控作用的垂体和性腺组织为研究对象,筛选出了运输过程中可能对生殖起主要调控作用的信号通路和基因。
Reproduction, stress and disease are important factors that restricting dairy production and economic benefits, the estimated heritability values of these traits tend to be relatively low, and it is difficult to make great genetic progress through direct selection. In order to breed disease resistance individuals, we studied the polymorphisms of toll like receptor family and analyzed its association with mastitis, bovine tuberculosis and bovine brucellosis; we further analyzed the variants of BMP15with sperm quality traits, expecting to conduct resistance selection by molecular breeding and improve fertility and disease resistance of animals. Transport, as an frequent and inevitably behavior in animal production, would cause big stress to animals, for example, significant increase in disease incidence, however, up to now, there is little literature about the impact of transport stress on animal reproduction. Therefore, in this experiment, we use mice as model, investigated the mRNA and protein levels of LHR, AR and Hsp70in mice testis after long road transport; Second, screened differentially expressed genes in pituitary, uterus and placenta of different pregnancy phase by use of Agilent mouse microarray, and further functional annotation to some of the differentially expressed genes. To provide reference or basis for the study of transport stress on cattle reproductive physiology and product development of anti-transport stress.
1. BMP15gene polymorphisms and its association with sperm quality traits in Chinese Holstein cattle
We analyzed the association between polymorphism of BMP15gene and sperm quality traits. One SNP in intron (C5697T) was detected. Age and genotype had significant effect on sperm quality trait of fresh sperm motility, however, the interaction of age and genotype had no effect on any sperm quality traits. We further analyzed the effect of different genotype on fresh sperm motility and found that fresh sperm motility of CT genotype was significantly lower than that of CC or TT genotype. These results indicate that BMP15may be candidate genes for sperm quality traits.
2. Identification of candidate genes associated with bovine disease resistance
In this study, we first chose TLR1、TLR2、TLR、TLR6and TLR9as candidate genes to study their variants with mastitis, bovine tuberculosis and bovine brucellosis by PCR-SSCP, PCR-RFLP combined with sequencing. The main results are as follows:
(1) Five SNPs C632T, G1409A, A1475C, G1550A and G1596A were detected in TLR1gene. The allele and genotype frequencies of C632T, G1409A and G1550A did not vary significantly between bovine tuberculosis (bTB) infected and non-infected cattle. In G1596A site, genotype frequencies differed significantly between bTB-infected and non-infected cattle, and relative risk of bovine tuberculosis incidence of GH is2.43times than that of GG genotype; In A1475C site, there was a trend toward an increased relative risk of tuberculosis incidence in CD genotype (P=0.056).
(2) Three SNPs T385G, G398A and C1828T were detected in TLR2gene, they can be recognized by restriction enzyme EcoRV, HaeⅢ and Dral respectively. In T385G site, GG genotype had significantly higher SCS than GT and TT genotype, and its relative risk of bovine brucellosis incidence is2.084times than that of TT genotype and GG may be disease susceptibility genotype; No significant differences were observed between G398A and C1828T site in SCS and bovine brucellosis incidence.
(3) There were no significant differences between different genotypes of TLR4and bovine tuberculosis, and this may indicate TLR4is not a candidate gene in controlling bovine tuberculosis.
(4) Four SNPs G1793A, C1859A、G1934A and A1980G were detected in TLR6gene. There were significant differences between different genotypes of C1859A and A1980G and bovine tuberculosis. In C1859A, cattle with AA or CA genotype had greater relative risk of disease incidence (OR=2.73,95%CI0.869-8.573; OR=1.547,95%CI0.803-2.982) than those with CC genotype, and CC maybe bovine tuberculosis resistance genotype; In A1980G, cattle with GG or GA genotype had greater relative risk of disease incidence (OR=2.986,95%CI1.245-7.165; OR=1.582,95%CI0.734-3.409) than those of AA genotype, and AA maybe bovine tuberculosis resistance genotype.
(5) There were no significant differences between different genotypes of TLR9and bovine tuberculosis or bovine brucellosis, and this may indicate TLR9is not a candidate gene in controlling bovine tuberculosis or bovine brucellosis.
3. Effect of transport stress on animal reproduction
In order to study the effect of transport stress on animal reproduction, we chose Kunming mice as model to study its role in male and pregnant for about15h (1000km) long road transport.
(1) Long road transport significantly reduced body weight, epididymis index and relative sperm count in epididymis. In order to analyze the possible mechanism, the mRNA and protein expression of LHR, AR and Hsp70were conducted. Results showed that expression of AR was significantly lower than before transport, LHR and Hsp70expression were somewhat higher after transport, however, not significant.
(2) Compared with control group, there are no significant differences in living birth rate, dead birth rate and absorbed fetus rate on long road transport during the last trimester of pregnancy, while declined living birth rate and increased dead birth rate after transport during the second trimester of pregnancy.
(3)133genes,586genes,146genes and562genes were scanned out in E10pituitary, E10uterus, E15pituitary and E15placenta respectively. These differential expression genes, according to biological process of Go analysis, mainly focused on the cellular process, biological regulation, metabolic process and regulation of biological process.
(4) We use pituitary and gonadal tissue, which play main roles in reproduction, as our taget, screeded several pathways and genes that may play major role in reproduction during transportation.
引文
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