绒山羊绒毛生长相关基因的筛选、鉴定和多态性分析
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摘要
绒山羊是重要的绒用经济动物,其绒毛生长具有较强的季节性,但不同品种之间有很大的差异,大多数绒山羊在非长绒季节(5-8月份)不长绒,而辽宁绒山羊一些品系可以常年长绒(6月开始长绒)。埋植褪黑激素可以促进绒山羊在非长绒季节长绒,从而达到四季长绒的目的。本研究以内蒙古白绒山羊为主要研究对象,采用分子标记和基因芯片技术,从候选基因和差异表达基因筛选入手,研究绒山羊产绒经济性状的遗传多态性和非长绒季节埋植褪黑激素绒山羊皮肤组织差异表达基因,初步探讨促进绒山羊绒毛生长的因素及其分子调控机理。
1、绒山羊内分泌激素类基因SNP分析
应用PCR-SSCP和DNA测序技术,以内蒙古阿尔巴斯型、阿拉善型、二郎山型以及罕山白绒山羊4个品种(系)共456个样本为实验材料,分析PRL、PRLR、 MLTR1a、MLTR1b、IGF-I和TG6个候选基因共16个位点的遗传多态性,结果表明：
(1)在绒山羊PRL基因Exon5区域P2引物位点存在X76049:g.576bp处C-A突变,该突变导致第176个密码子CCC变成ACC,从而使脯氨酸变成苏氨酸(Pro-Thr),该位点SNP对绒山羊绒厚度存在显著影响(P＜0.01),同时还对阿尔巴斯绒山羊产羔数存在边际显著影响(P=0.055)；在PRLR基因Exon10区域P3引物位点存在HQ266606:g.1076bp处T-C突变,该突变导致第359个密码子CTT变成CCT,从而使亮氨酸变成脯氨酸(Lue-Pro),该位点SNP对阿尔巴斯绒山羊羊绒细度存在显著影响(P＜0.05)；
(2)在绒山羊MLTR1b基因Exon2区域P8引物位点存在NM_001206907:g.648bp处C-G突变,该突变没有导致氨基酸的改变,还存在NM_001206907:g.665bp处G-A,该突变导致第223个密码子CGC变成CAC,从而使精氨酸变成组氨酸(Arg-His),该位点SNP对绒山羊产绒量有显著影响(P＜0.05)。同时,该多态位点基因型与年龄互作效应对绒山羊绒厚度有显著影响(P＜0.05)；MLTR1b基因Exon2区域P9引物位点存在NM_001206907:g.1015bp处G-A突变,该突变导致第339个密码子GGC变成AGC,从而使甘氨酸变成丝氨酸(Gly-Ser),该位点SNP与年龄互作效应对绒山羊绒细度存在边际显著影响(P=0.056)；这两个位点多态对绒山羊产羔数、羔羊初生重等繁殖性状均无显著影响(P>0.05)
(3)在绒山羊TG基因Exon9区域P14引物位点存在NM_173883:g.1895bp处C-T突变,该突变导致第632个密码子GCG变成GTG,从而使丙氨酸变成缬氨酸(Ala-Val);在Exon11区域P16引物位点存在NM_173883:g.2849bp处G-A突变,该突变导致第950个密码子CGC变成CAC,从而使精氨酸变成组氨酸(Arg-His)。这两个位点SNP对绒山羊体重、产羔数和产绒性状均无显著影响(P>0.05)。
2、绒山羊皮肤差异表达基因的筛选、鉴定和特征分析
(1)通过基因芯片技术筛选绒山羊在非长绒期埋植MLT促进羊绒生长的相关基因。筛选出差异表达的基因95个,其中61个表达上调、34个下调。这些异常表达的基因按照功能分类,参与组成细胞成分的基因占18.33%,参与分子功能的基因占47.78%,参与生物学过程的基因占33.89%。
(2) Pathway信号通路分析表明：差异基因共参与了55条信号通路,其中CTNNB1基因参与的Wnt信号通路,CYP2B基因参与的维生素A代谢和外源药物代谢等信号通路可能与埋植MLT促进绒山羊非长绒期绒毛生长和毛囊发育有关。
(3)采用Real time定量PCR检测技术,选择芯片试验结果中表达差异显著上调的PAX6基因和显著下调的CSN2基因,进一步扩大样本对芯片结果准确性做验证。结果表明,PAX6基因在埋植MLT试验组羊皮肤中的表达量是对照组羊皮肤中表达量的2.399倍,CSN2基因在试验组羊皮肤中的表达量是对照组羊皮肤中表达量的0.044倍,与基因芯片结果完全相符,说明本研究基因芯片数据准确、可靠。
本研究揭示了CTNNB1、CYP2B、PAX6、CSN2等基因可能参与了埋植MLT之后促进绒山羊长绒的过程,有必要对这些基因功能作进一步研究,以期探明绒山羊绒毛生长和毛囊发育的分子机制。
Cashmere goat is a kind of economic animal for production cashmere,which cashmere growth have strong seasonal and great difference between different breeds,which can't produce cashmere for most of cahsmere goats in the telogen period(May-August), while some strains of Liaoning cashmere goats can product cahsmere in the whole year(beginning from June). Embedding melatonin can promote cashmere growth in the telogen period so as to production cashmere in the whole year.In this paper,Inner Mongolian Ardrs white cashmere goats were used as main experimental samples, selection of candidate and differentially expressed genes was engaged in studying differentially expressed gene between genetic polymorphism of production cashmere economic traits and skin organism embedding melatonin in the telogen period by PCR-SSCP and gene chip technology, and preliminary discuss factors and molecular regulation mechanism of promoting cashmere growth.
1、SNP analysis of endocrine hormone gene on cashmere goats
456samples were used as experimental materials including four types:Inner Mongolian Ardrs, Alashan, Erlangshan and Hanshan white cashmere goats by PCR-SSCP and DNA sequencing technology to analyze16loci genetic polymorphism of6candidate genes including PRL、PRLR、MLTR1a、MLTR1b、IGF-I and TG.
Results were as follows:
(1)There were SSCP at P2primer of PRL gene and P3primer of PRLR gene. P2primer of exon5of PRL gene existed in C-A mutation at X76049:g.576bp, which resulted in CCC-ACC of the176th codon from Pro into Thr. The results showed that the its SNP was the most significant correlation with cashmere thickness(P<0.01), and the marginal significant correlation(P=0.055)with kidding number of Ardrs cashmere goat; P3primer was in T-C mutation at HQ266606:g of exon10of PRLR gene, which lead to CTT-CCT of the359th codon from Lue into Pro, of which SNP was significant difference (P<0.05) to cashmere fineness.
(2) There were SSCP in the exon2of MLTRlb gene and three of SNP were detected. P8primer existed in C-G mutation at NM_001206907:g.648bp which didn't result in change of amino acid, and also be G-A mutation at NM_001206907:g.665bp which led to CGC-CAC of the223th codon from Arg into His. The results showed that the SNP of P8locus had an significant effect on cashmere yield(P<0.05), meanwhile, interaction effect between the SNP genotype and age was a significant correlation with cashmere thickness(P <0.05); P9primer existed in G-A mutation at NM_001206907:g.1015bp, which led to GGC-AGC of the339th codon from Gly into Ser. The analysis indicated that interaction effect of the SNP and age was the marginal significant correlation(P=0.056) with cashmere fineness.
(3) There were SSCP at P14primer and P16primer of TG gene. P14primer of the exon9existed in C-T mutation at NM_173883:g.1895bp, which resulted in GCG-GTG of the632th codon from Ala into Val. P16primer of exon11existed in G-A mutation at NM_173883:g.2849bp, which led to CGC-CAC of the950th codon from Arg into His, which both were no significant effect on weight, kidding number and production cashmere traits (p>0.05). P8and P9primes were not significance(P>0.05) with kidding number and birth weight.
2^Selection, identification and characteristic analysis of skin differentially expressed gene on cashmere Goat
(1) To select correlation genes which promoted cashmere growth embedding melatonin(MLT) in the telogen period by gene chip technology.To have selected95of differentially expressed genes,61of which were up-regulated expression, while34genes were down-regulated expression.Those abnormal expression genes were assorted by function,18.33%,47.78%,33.89%of genes involved in cellular component, molecular function, biological process, respectively.
(2) Analyses of signaling pathway showed that differentially expressed genes totally involved in55pieces of signaling pathway. CTNNB1gene participated in WNT signaling pathway, and CYP2B gene related to metabolism of retinol and xenobiotics by cytochrome P450, which probably promoted cashmere growth and follicle development related with embedding MLT in the telogen period.
(3) Significant Up-regulated PAX6gene and down-regulated CSN2gene of differentially expressed gene were further identified by real time quantitative so as to validate result of gene chip. Resulted showed that expression level of skin in PAX6gene was2.399times that of control group, and also expression level of skin in CNS2gene was0.044times compared to control group. The results were the same with gene chip. It indicated that gene chip had character of data precision and reliability.
The paper disclosed that CTNNB1gene, CYP2B gene, PAX6gene and CSN2gene etc. probably participated in promoting cashmere growth process after embedding MLT, which was necessary to further study their gene function, so as to find out molecular mechanism of cashmere growth and follicle development on cashmere goats.
1、绒山羊内分泌激素类基因SNP分析
应用PCR-SSCP和DNA测序技术,以内蒙古阿尔巴斯型、阿拉善型、二郎山型以及罕山白绒山羊4个品种(系)共456个样本为实验材料,分析PRL、PRLR、 MLTR1a、MLTR1b、IGF-I和TG6个候选基因共16个位点的遗传多态性,结果表明：
(1)在绒山羊PRL基因Exon5区域P2引物位点存在X76049:g.576bp处C-A突变,该突变导致第176个密码子CCC变成ACC,从而使脯氨酸变成苏氨酸(Pro-Thr),该位点SNP对绒山羊绒厚度存在显著影响(P＜0.01),同时还对阿尔巴斯绒山羊产羔数存在边际显著影响(P=0.055)；在PRLR基因Exon10区域P3引物位点存在HQ266606:g.1076bp处T-C突变,该突变导致第359个密码子CTT变成CCT,从而使亮氨酸变成脯氨酸(Lue-Pro),该位点SNP对阿尔巴斯绒山羊羊绒细度存在显著影响(P＜0.05)；
(2)在绒山羊MLTR1b基因Exon2区域P8引物位点存在NM_001206907:g.648bp处C-G突变,该突变没有导致氨基酸的改变,还存在NM_001206907:g.665bp处G-A,该突变导致第223个密码子CGC变成CAC,从而使精氨酸变成组氨酸(Arg-His),该位点SNP对绒山羊产绒量有显著影响(P＜0.05)。同时,该多态位点基因型与年龄互作效应对绒山羊绒厚度有显著影响(P＜0.05)；MLTR1b基因Exon2区域P9引物位点存在NM_001206907:g.1015bp处G-A突变,该突变导致第339个密码子GGC变成AGC,从而使甘氨酸变成丝氨酸(Gly-Ser),该位点SNP与年龄互作效应对绒山羊绒细度存在边际显著影响(P=0.056)；这两个位点多态对绒山羊产羔数、羔羊初生重等繁殖性状均无显著影响(P>0.05)
(3)在绒山羊TG基因Exon9区域P14引物位点存在NM_173883:g.1895bp处C-T突变,该突变导致第632个密码子GCG变成GTG,从而使丙氨酸变成缬氨酸(Ala-Val);在Exon11区域P16引物位点存在NM_173883:g.2849bp处G-A突变,该突变导致第950个密码子CGC变成CAC,从而使精氨酸变成组氨酸(Arg-His)。这两个位点SNP对绒山羊体重、产羔数和产绒性状均无显著影响(P>0.05)。
2、绒山羊皮肤差异表达基因的筛选、鉴定和特征分析
(1)通过基因芯片技术筛选绒山羊在非长绒期埋植MLT促进羊绒生长的相关基因。筛选出差异表达的基因95个,其中61个表达上调、34个下调。这些异常表达的基因按照功能分类,参与组成细胞成分的基因占18.33%,参与分子功能的基因占47.78%,参与生物学过程的基因占33.89%。
(2) Pathway信号通路分析表明：差异基因共参与了55条信号通路,其中CTNNB1基因参与的Wnt信号通路,CYP2B基因参与的维生素A代谢和外源药物代谢等信号通路可能与埋植MLT促进绒山羊非长绒期绒毛生长和毛囊发育有关。
(3)采用Real time定量PCR检测技术,选择芯片试验结果中表达差异显著上调的PAX6基因和显著下调的CSN2基因,进一步扩大样本对芯片结果准确性做验证。结果表明,PAX6基因在埋植MLT试验组羊皮肤中的表达量是对照组羊皮肤中表达量的2.399倍,CSN2基因在试验组羊皮肤中的表达量是对照组羊皮肤中表达量的0.044倍,与基因芯片结果完全相符,说明本研究基因芯片数据准确、可靠。
本研究揭示了CTNNB1、CYP2B、PAX6、CSN2等基因可能参与了埋植MLT之后促进绒山羊长绒的过程,有必要对这些基因功能作进一步研究,以期探明绒山羊绒毛生长和毛囊发育的分子机制。
Cashmere goat is a kind of economic animal for production cashmere,which cashmere growth have strong seasonal and great difference between different breeds,which can't produce cashmere for most of cahsmere goats in the telogen period(May-August), while some strains of Liaoning cashmere goats can product cahsmere in the whole year(beginning from June). Embedding melatonin can promote cashmere growth in the telogen period so as to production cashmere in the whole year.In this paper,Inner Mongolian Ardrs white cashmere goats were used as main experimental samples, selection of candidate and differentially expressed genes was engaged in studying differentially expressed gene between genetic polymorphism of production cashmere economic traits and skin organism embedding melatonin in the telogen period by PCR-SSCP and gene chip technology, and preliminary discuss factors and molecular regulation mechanism of promoting cashmere growth.
1、SNP analysis of endocrine hormone gene on cashmere goats
456samples were used as experimental materials including four types:Inner Mongolian Ardrs, Alashan, Erlangshan and Hanshan white cashmere goats by PCR-SSCP and DNA sequencing technology to analyze16loci genetic polymorphism of6candidate genes including PRL、PRLR、MLTR1a、MLTR1b、IGF-I and TG.
Results were as follows:
(1)There were SSCP at P2primer of PRL gene and P3primer of PRLR gene. P2primer of exon5of PRL gene existed in C-A mutation at X76049:g.576bp, which resulted in CCC-ACC of the176th codon from Pro into Thr. The results showed that the its SNP was the most significant correlation with cashmere thickness(P<0.01), and the marginal significant correlation(P=0.055)with kidding number of Ardrs cashmere goat; P3primer was in T-C mutation at HQ266606:g of exon10of PRLR gene, which lead to CTT-CCT of the359th codon from Lue into Pro, of which SNP was significant difference (P<0.05) to cashmere fineness.
(2) There were SSCP in the exon2of MLTRlb gene and three of SNP were detected. P8primer existed in C-G mutation at NM_001206907:g.648bp which didn't result in change of amino acid, and also be G-A mutation at NM_001206907:g.665bp which led to CGC-CAC of the223th codon from Arg into His. The results showed that the SNP of P8locus had an significant effect on cashmere yield(P<0.05), meanwhile, interaction effect between the SNP genotype and age was a significant correlation with cashmere thickness(P <0.05); P9primer existed in G-A mutation at NM_001206907:g.1015bp, which led to GGC-AGC of the339th codon from Gly into Ser. The analysis indicated that interaction effect of the SNP and age was the marginal significant correlation(P=0.056) with cashmere fineness.
(3) There were SSCP at P14primer and P16primer of TG gene. P14primer of the exon9existed in C-T mutation at NM_173883:g.1895bp, which resulted in GCG-GTG of the632th codon from Ala into Val. P16primer of exon11existed in G-A mutation at NM_173883:g.2849bp, which led to CGC-CAC of the950th codon from Arg into His, which both were no significant effect on weight, kidding number and production cashmere traits (p>0.05). P8and P9primes were not significance(P>0.05) with kidding number and birth weight.
2^Selection, identification and characteristic analysis of skin differentially expressed gene on cashmere Goat
(1) To select correlation genes which promoted cashmere growth embedding melatonin(MLT) in the telogen period by gene chip technology.To have selected95of differentially expressed genes,61of which were up-regulated expression, while34genes were down-regulated expression.Those abnormal expression genes were assorted by function,18.33%,47.78%,33.89%of genes involved in cellular component, molecular function, biological process, respectively.
(2) Analyses of signaling pathway showed that differentially expressed genes totally involved in55pieces of signaling pathway. CTNNB1gene participated in WNT signaling pathway, and CYP2B gene related to metabolism of retinol and xenobiotics by cytochrome P450, which probably promoted cashmere growth and follicle development related with embedding MLT in the telogen period.
(3) Significant Up-regulated PAX6gene and down-regulated CSN2gene of differentially expressed gene were further identified by real time quantitative so as to validate result of gene chip. Resulted showed that expression level of skin in PAX6gene was2.399times that of control group, and also expression level of skin in CNS2gene was0.044times compared to control group. The results were the same with gene chip. It indicated that gene chip had character of data precision and reliability.
The paper disclosed that CTNNB1gene, CYP2B gene, PAX6gene and CSN2gene etc. probably participated in promoting cashmere growth process after embedding MLT, which was necessary to further study their gene function, so as to find out molecular mechanism of cashmere growth and follicle development on cashmere goats.
引文
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