家禽繁殖性状相关基因的遗传效应及组织表达规律研究
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摘要
本实验选取我国地方遗传资源中具有特殊繁殖性状的文昌鸡和高邮鸭作为实验素材,研究影响禽类繁殖性能的外观性状标记和分子遗传标记,为文昌鸡和高邮鸭选种和繁育提供依据。实验分析了不同羽色和羽速文昌鸡生长和繁殖时期的发育规律以及繁殖性状差异；同时,对文昌鸡的促卵泡素受体(Follicle-stimulating hormone receptor, FSHR)基因,以及高邮鸭的FSHR基因、神经肽Y(NeuropeptideY,NPY)基因、促甲状腺激素β (Thyroid-stimulating hormone beta, TSH-β)基因、钙结合蛋白-D28K (Calbindin-D28k, CaBP-28K)基因序列进行了克隆、单核苷酸多态以及表达水平分析,研究了文昌鸡、高邮鸭FSHR基因序列的SNPs、基因多态与繁殖性状相关关系,根据获得的鸭NPY基因序列与其它物种进行进化地位分析,结合实时荧光定量方法对NPY、TSH-β、CaBP-28K在中枢神经组织和周围组织器官的表达情况进行了相关研究,从分子水平上探讨研究以上繁殖相关基因的生理学效应；对FSHR基因mRNA在不同组织部位存在的选择性剪切进行初步验证和总结。
研究结果如下：
1.采用Gompertz (?)非线性模型拟合黑羽、珍珠羽、黄麻色羽慢羽系、黄麻色羽快羽系四个群体文昌鸡生长曲线模型,对其生长发育规律进行比较,分析不同羽色和不同羽速文昌鸡生产性能差异。结果表明：较之于珍珠羽系、黑羽系、黄麻慢羽系,在产蛋期之前的生长发育期,黄麻快羽系文昌鸡群体体重增长突出,差异显著(P<0.05)；进入产蛋期后黄麻慢羽系产蛋性状突出,开产日龄早,300日龄产蛋数较多,与其他群体差异显著(P<0.05)。由此揭示了文昌鸡不同羽系的生产性能差异。采用Gompertz模型拟合文昌鸡生长曲线,通过比较各拟合参数,拟合参数符合其种质特性。
2.以促卵泡素受体(FSHR)基因作为影响鸡繁殖性状的候选基因,采用PCR-SSCP技术结合测序对FSHR进行单核苷酸多态性分析。结果发现,在第二外显子(exon2)、第四外显子(exon4)、第六外显子(exon6)、第八外显子(exon8)区域存在SNPs位点,分别为在exon2片段中编码区5’端-49bp处的C→T突变；exon4片段中编码区43bp处T→C突变,但没有引起氨基酸的改变；exon6片段中编码区3’端+12bp处的A→G突变；距exon8编码区3’端+38bp处的G→T突变。经适合性检验,各基因的基因频率在群体内的分布均处于Hardy-Weinberg平衡状态p>0.05)。
3.对文昌鸡FSHR基因4个区域基因多态与繁殖性状相关性进行分析表明,FSHR基因第四外显子呈现三种基因型(CC/CD/DD),其多态性对文昌鸡300日龄产蛋数及平均联产天数显著相关,CD杂合基因型与文昌鸡个体的300日龄产蛋数及平均联产天数存在显著相关(P<0.05)。文昌鸡FSHR基因所检测的其它突变位点的多态性与文昌鸡各繁殖性状相关性不显著(P>0.05)。采用鸡FSHR基因exon2、exon4、exon6、exon8相同的扩增引物对高邮鸭FSHR基因进行PCR-SSCP分析,发现exon4存在多态,基因多态性与繁殖性状相关性分析表明,高邮鸭FSHR基因第四外显子基因多态(基因型ZZ/ZW/WW)对高邮鸭380日龄产蛋数存在显著相关(P<0.05)。
4.通过实时荧光定量(real-time quantitative PCR)方法对繁殖期高邮鸭繁殖相关基因NPY、TSH-β、CaBP-28K克隆,并且分析其mRNA在下丘脑、垂体、心脏、肝脏、脾脏、胰腺、肾脏、子宫、卵巢、小肠、胸肌、腿肌等组织器官基因相对表达量。结果显示：进化保守基因NPY序列的系统进化分析构建出的系统进化树,符合鸭在动物进化中的定位；NPY基因在第三外显子至3’末端之间不存在选择性拼接调控机制；NPY基因在下丘脑中表达丰富,在垂体和小肠中表达较少,在卵巢、心脏、肝脏、脾脏、肾脏、胰腺、子宫、胸肌、腿肌等组织部位痕量表达；TSH-β基因在垂体中表达量丰富,在卵巢和子宫中表达量次之,在心脏、肝脏、脾脏、肾脏、胰腺、胸肌、腿肌等部位呈痕量表达；CaBP-28K在卵巢、肾脏和小肠中表达量较多,子宫、垂体中表达量次之,在下丘脑、心脏、肝脏、脾脏、胰腺、胸肌、腿肌等组织部位呈痕量表达。
5采用荧光定量PCR分析方法对高邮鸭卵巢、子宫的组织的FSHR基因mRNA是否存在剪切进行了验证性试验。结果表明：鸭卵巢、子宫中存在FSHR基因第二外显子选择性剪切,其nRNA剪切体在子宫中的含量与FSHR基因mRNA总量比为0.425：1；在卵巢中含量占FSHR基因表达总量比为0.394：1,在其他组织中未发现选择性剪切。
This study was carried out to discuss the markers of apparent characters and molecular genetic markers which affect the fowl productive capability, and provide reference for the fowl breeding. Wenchang chicken and Gaoyou duck were selected as the materials. In this experiment, the rules of growth and development and laying traits of four lines of Wenchang chicken were analyzed comparatively and the reproductive performance was compared among four lines of Wenchang chicken and Gaoyou duck respectively. The polymorphisms of FSHR gene in Wenchang chicken were detected by PCR-SSCP and sequencing, and analyzed with the reproductive traits, and also was in Gaoyou duck. The relative expression of the FSHR, NPY, TSH-β, and Calbindin-D28k mRNA in central nervous and visceral tissues were detected by real-time qPCR respectively, on which the phylogenetic analysis of Gaoyou duck and the physiology effects of those reproductive genes were carried out on the molecular level. The alternative splicing of FSHR RNA was observed and analyzed. The follows were the results of this study.
1. The rules of growth and development and laying traits of four lines of Wenchang chicken were analyzed comparatively by nonlinear Gompertz model, which were black spotted (zz), fast yellow spotted (hk), black (h) and slow yellow spotted (hm). The capability of product was compared among the four lines of Wenchang chicken. The result indicated that the body weight of fast yellow spotted (hk) line went up higher than other lines before the laying period (P<0.05). During the laying period, the slow yellow spotted(hm) line had higher laying production, including earlier first egg(AFG) and more number of egg to300days old(NE-300d)(P<0.05). This conclusion proved objective references for the research on the difference of product performance of Wenchang among different feathering genotype and the production practice. According to the statistical test, the Gompertz models fitted the experimental data quite well, and the parameters were according with the characteristic of Wenchang chicken.
2. The objective of the present study was to detect the polymorphisms of the follicle stimulating hormone receptor (FSHR) gene as the candidate gene of chicken's reproductive trait, and provide a scientific basis for marker assisted selection for high prolificacy in Wenchang chicken. In this study, the regions of FSHR which encoded the entire extracellular domain were analyzed by PCR-SSCP and sequencing. The result showed that4SNPs sites were found at the region of exon2, exon4, exon6and exon8respectively. They are C-49→T-49mutation before the CDS'5-flanking terminal of the exon2region, T43→C43mutation in the CDS of the exon4region, A12→G12mutation after the CDS'3-flanking terminal of the exon6region, G38→T38mutation after the CDS'3-flanking terminal of the exon8region. The fitness test indicated the gene frequencies were all in Hardy-Weinberg balanced state (P>0.05). In conclusions, the polymorphisms of theses sites lay a foundation for the further research on relationship between the polymorphisms of FSHR gene and the reproductive performance of Wenchang chicken.
3. In this study, the correlative analyses between the polymorphisms in Wenchang chicken of the FSHR exon2, exon4, exon6and exon8and the reproductive trait were carried out by Least square analysis. The result showed that the polymorphisms(gene types CC/CD/DD) of exon4was distinct correlative with the egg number of300days and the average days of continual laying. The CD chicken had more egg number of300days and longer average days of continual laying (P<0.05).
The polymorphisms in Gaoyou duck of the FSHR exon2,exon4,exon6and exon8were detected by PCR-SSCP, and the correlative analyses between the polymorphisms and the reproductive trait were carried out by Least square analysis. The result showed that only exon4was polymorphic (3gene types ZZ/ZW/WW), and the WW duck had more egg number of380days than others. The polymorphisms of FSHR exon4was higher correlative with egg number of380days in Gaoyou duck (P<0.05).
4. The expression level of NPY (Neuropeptide Y),TSH-β,CaBP-28K of Gaoyou Duck in reproduction period were analyzed in the central nervous tissue and surrounding tissues(hypothalamus, pituitary, heart, liver, spleen, pancreas, kidney, uterus, ovary, small intestine, and muscle of chest). By real-time fluorescent quantitative PCR (RT-PCR), we cloned the core alignment of NPY gene, and tested the relative expression level and the fragment size of NPY mRNA in the central nervous tissue and surrounding tissues. The result showed that the phylogenetic tree constructed from the cloned core alignment of NPY by phylogenetic analysis correspond to duck's position in zoogenesis. NPY gene of duck was relatively conservative in evolution, and there was no alternative splicing between the third exon and untranslated3'terminal. The result of real-time fluorescent quantitative analysis showed NPY gene expressed abundantly in hypothalamus, and less in pituitary and small intestine, trace in ovary, heart, liver, spleen, kidney, pancreas, uterus, pectorales and crureus. NPY also expressed in surrounding tissue, which declared that NPY may regulate the physiological function by autocrine and/or paracrine. TSH-β gene expressed abundantly in pituitary, less in ovary and uterus, and least in other tissues. The expression of Calbindin-D28k was higher in ovary, kidney and small intestine, less in pituitary and uterus, and trace in other tissues.
5. The Identification of transcripts from the FSHR mRNA in ovary and uterus tissues of Gaoyou duck was carried out by real-time fluorescent quantitative PCR. This test proved that there were alternative splicings at the exon2of FSHR gene. The relative expression ratio of the exceptional transcript was42.5%in uterus, close to that in ovary (39.4%).There was no alternative splicing found in other tissues.
研究结果如下：
1.采用Gompertz (?)非线性模型拟合黑羽、珍珠羽、黄麻色羽慢羽系、黄麻色羽快羽系四个群体文昌鸡生长曲线模型,对其生长发育规律进行比较,分析不同羽色和不同羽速文昌鸡生产性能差异。结果表明：较之于珍珠羽系、黑羽系、黄麻慢羽系,在产蛋期之前的生长发育期,黄麻快羽系文昌鸡群体体重增长突出,差异显著(P<0.05)；进入产蛋期后黄麻慢羽系产蛋性状突出,开产日龄早,300日龄产蛋数较多,与其他群体差异显著(P<0.05)。由此揭示了文昌鸡不同羽系的生产性能差异。采用Gompertz模型拟合文昌鸡生长曲线,通过比较各拟合参数,拟合参数符合其种质特性。
2.以促卵泡素受体(FSHR)基因作为影响鸡繁殖性状的候选基因,采用PCR-SSCP技术结合测序对FSHR进行单核苷酸多态性分析。结果发现,在第二外显子(exon2)、第四外显子(exon4)、第六外显子(exon6)、第八外显子(exon8)区域存在SNPs位点,分别为在exon2片段中编码区5’端-49bp处的C→T突变；exon4片段中编码区43bp处T→C突变,但没有引起氨基酸的改变；exon6片段中编码区3’端+12bp处的A→G突变；距exon8编码区3’端+38bp处的G→T突变。经适合性检验,各基因的基因频率在群体内的分布均处于Hardy-Weinberg平衡状态p>0.05)。
3.对文昌鸡FSHR基因4个区域基因多态与繁殖性状相关性进行分析表明,FSHR基因第四外显子呈现三种基因型(CC/CD/DD),其多态性对文昌鸡300日龄产蛋数及平均联产天数显著相关,CD杂合基因型与文昌鸡个体的300日龄产蛋数及平均联产天数存在显著相关(P<0.05)。文昌鸡FSHR基因所检测的其它突变位点的多态性与文昌鸡各繁殖性状相关性不显著(P>0.05)。采用鸡FSHR基因exon2、exon4、exon6、exon8相同的扩增引物对高邮鸭FSHR基因进行PCR-SSCP分析,发现exon4存在多态,基因多态性与繁殖性状相关性分析表明,高邮鸭FSHR基因第四外显子基因多态(基因型ZZ/ZW/WW)对高邮鸭380日龄产蛋数存在显著相关(P<0.05)。
4.通过实时荧光定量(real-time quantitative PCR)方法对繁殖期高邮鸭繁殖相关基因NPY、TSH-β、CaBP-28K克隆,并且分析其mRNA在下丘脑、垂体、心脏、肝脏、脾脏、胰腺、肾脏、子宫、卵巢、小肠、胸肌、腿肌等组织器官基因相对表达量。结果显示：进化保守基因NPY序列的系统进化分析构建出的系统进化树,符合鸭在动物进化中的定位；NPY基因在第三外显子至3’末端之间不存在选择性拼接调控机制；NPY基因在下丘脑中表达丰富,在垂体和小肠中表达较少,在卵巢、心脏、肝脏、脾脏、肾脏、胰腺、子宫、胸肌、腿肌等组织部位痕量表达；TSH-β基因在垂体中表达量丰富,在卵巢和子宫中表达量次之,在心脏、肝脏、脾脏、肾脏、胰腺、胸肌、腿肌等部位呈痕量表达；CaBP-28K在卵巢、肾脏和小肠中表达量较多,子宫、垂体中表达量次之,在下丘脑、心脏、肝脏、脾脏、胰腺、胸肌、腿肌等组织部位呈痕量表达。
5采用荧光定量PCR分析方法对高邮鸭卵巢、子宫的组织的FSHR基因mRNA是否存在剪切进行了验证性试验。结果表明：鸭卵巢、子宫中存在FSHR基因第二外显子选择性剪切,其nRNA剪切体在子宫中的含量与FSHR基因mRNA总量比为0.425：1；在卵巢中含量占FSHR基因表达总量比为0.394：1,在其他组织中未发现选择性剪切。
This study was carried out to discuss the markers of apparent characters and molecular genetic markers which affect the fowl productive capability, and provide reference for the fowl breeding. Wenchang chicken and Gaoyou duck were selected as the materials. In this experiment, the rules of growth and development and laying traits of four lines of Wenchang chicken were analyzed comparatively and the reproductive performance was compared among four lines of Wenchang chicken and Gaoyou duck respectively. The polymorphisms of FSHR gene in Wenchang chicken were detected by PCR-SSCP and sequencing, and analyzed with the reproductive traits, and also was in Gaoyou duck. The relative expression of the FSHR, NPY, TSH-β, and Calbindin-D28k mRNA in central nervous and visceral tissues were detected by real-time qPCR respectively, on which the phylogenetic analysis of Gaoyou duck and the physiology effects of those reproductive genes were carried out on the molecular level. The alternative splicing of FSHR RNA was observed and analyzed. The follows were the results of this study.
1. The rules of growth and development and laying traits of four lines of Wenchang chicken were analyzed comparatively by nonlinear Gompertz model, which were black spotted (zz), fast yellow spotted (hk), black (h) and slow yellow spotted (hm). The capability of product was compared among the four lines of Wenchang chicken. The result indicated that the body weight of fast yellow spotted (hk) line went up higher than other lines before the laying period (P<0.05). During the laying period, the slow yellow spotted(hm) line had higher laying production, including earlier first egg(AFG) and more number of egg to300days old(NE-300d)(P<0.05). This conclusion proved objective references for the research on the difference of product performance of Wenchang among different feathering genotype and the production practice. According to the statistical test, the Gompertz models fitted the experimental data quite well, and the parameters were according with the characteristic of Wenchang chicken.
2. The objective of the present study was to detect the polymorphisms of the follicle stimulating hormone receptor (FSHR) gene as the candidate gene of chicken's reproductive trait, and provide a scientific basis for marker assisted selection for high prolificacy in Wenchang chicken. In this study, the regions of FSHR which encoded the entire extracellular domain were analyzed by PCR-SSCP and sequencing. The result showed that4SNPs sites were found at the region of exon2, exon4, exon6and exon8respectively. They are C-49→T-49mutation before the CDS'5-flanking terminal of the exon2region, T43→C43mutation in the CDS of the exon4region, A12→G12mutation after the CDS'3-flanking terminal of the exon6region, G38→T38mutation after the CDS'3-flanking terminal of the exon8region. The fitness test indicated the gene frequencies were all in Hardy-Weinberg balanced state (P>0.05). In conclusions, the polymorphisms of theses sites lay a foundation for the further research on relationship between the polymorphisms of FSHR gene and the reproductive performance of Wenchang chicken.
3. In this study, the correlative analyses between the polymorphisms in Wenchang chicken of the FSHR exon2, exon4, exon6and exon8and the reproductive trait were carried out by Least square analysis. The result showed that the polymorphisms(gene types CC/CD/DD) of exon4was distinct correlative with the egg number of300days and the average days of continual laying. The CD chicken had more egg number of300days and longer average days of continual laying (P<0.05).
The polymorphisms in Gaoyou duck of the FSHR exon2,exon4,exon6and exon8were detected by PCR-SSCP, and the correlative analyses between the polymorphisms and the reproductive trait were carried out by Least square analysis. The result showed that only exon4was polymorphic (3gene types ZZ/ZW/WW), and the WW duck had more egg number of380days than others. The polymorphisms of FSHR exon4was higher correlative with egg number of380days in Gaoyou duck (P<0.05).
4. The expression level of NPY (Neuropeptide Y),TSH-β,CaBP-28K of Gaoyou Duck in reproduction period were analyzed in the central nervous tissue and surrounding tissues(hypothalamus, pituitary, heart, liver, spleen, pancreas, kidney, uterus, ovary, small intestine, and muscle of chest). By real-time fluorescent quantitative PCR (RT-PCR), we cloned the core alignment of NPY gene, and tested the relative expression level and the fragment size of NPY mRNA in the central nervous tissue and surrounding tissues. The result showed that the phylogenetic tree constructed from the cloned core alignment of NPY by phylogenetic analysis correspond to duck's position in zoogenesis. NPY gene of duck was relatively conservative in evolution, and there was no alternative splicing between the third exon and untranslated3'terminal. The result of real-time fluorescent quantitative analysis showed NPY gene expressed abundantly in hypothalamus, and less in pituitary and small intestine, trace in ovary, heart, liver, spleen, kidney, pancreas, uterus, pectorales and crureus. NPY also expressed in surrounding tissue, which declared that NPY may regulate the physiological function by autocrine and/or paracrine. TSH-β gene expressed abundantly in pituitary, less in ovary and uterus, and least in other tissues. The expression of Calbindin-D28k was higher in ovary, kidney and small intestine, less in pituitary and uterus, and trace in other tissues.
5. The Identification of transcripts from the FSHR mRNA in ovary and uterus tissues of Gaoyou duck was carried out by real-time fluorescent quantitative PCR. This test proved that there were alternative splicings at the exon2of FSHR gene. The relative expression ratio of the exceptional transcript was42.5%in uterus, close to that in ovary (39.4%).There was no alternative splicing found in other tissues.
引文
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