猪、鸡脂肪代谢相关基因的分子特征及表观遗传调控
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摘要
畜禽脂肪代谢受多种因素影响,包括遗传、营养、内分泌和环境等因素,它们在脂肪代谢中形成复杂的调控网络。脂类是动物和人体必需的营养成分,也是生物体重要的组成成分,但猪、鸡体内过多的脂肪沉积给人类带来一系列的问题。基于此,以猪、鸡为研究对象,本试验对影响脂肪代谢相关基因的分子特征和表观遗传调控进行了研究。利用分子生物学和生物信息学相结合的方法对猪BMAL1(pBMAL1)基因的分子特征进行分析,以便寻找影响猪脂肪代谢的主效基因,为猪的分子育种实践提供理论支持。分析不同剂量的甜菜碱对鸡部分生产性状和屠宰性状的作用,进而探讨甜菜碱对脂肪代谢相关基因表达和甲基化水平的影响,为进一步深入研究甜菜碱调控基因表达和甲基化的分子机理奠定基础,对于提高优质高产鸡的选育具有重要意义。
第一部分:以人和小鼠BMAL1基因序列设计简并引物成功克隆了猪BMAL1 cDNA和非编码区部分序列,分析了pBMAL1基因在不同部位脂肪组织和其它组织中的表达,并对其进行了精细定位。结果表明,pBMAL1基因序列长为2073bp,开放阅读框为1878bp,编码626个氨基酸,该氨基酸包括bHLH、PAS-A和PAS-B结构域。编码区核酸序列与人、小鼠和大鼠的同源性分别为94.36%, 89.85%和89.79%;氨基酸序列与人、小鼠和大鼠的同源性分别为99.20%, 98.24%和97.92%。在长白猪和莱芜猪背膘中发现了两个剪接变异体。pBMAL1 mRNA在长白猪5种脂肪组织均有表达,在肠系膜脂肪表达量最高,在花油表达量最低,但差异不显著(P>0.05);pBMAL1 mRNA在成年莱芜猪14种组织均有表达,在背膘中表达最高,在眼肌中表达最低。用辐射杂种板把pBMAL1基因定位在SSC 2p11-q21。
第二部分:通过在肉鸡和蛋鸡饲料中添加不同剂量的甜菜碱,研究甜菜碱对鸡部分生产性状和屠宰性状的影响;采用半定量RT-PCR方法分析了甜菜碱对脂肪代谢相关基因LPL、PPARγ、A-FABP、FAS、Adiponectin、BMAL1和CLOCK mRNA表达的影响,并通过亚硫酸盐测序的方法对LPL和PPARγ基因的启动子区甲基化进行了分析。
结果表明,在饲料中添加0.1%的甜菜碱对肉鸡平均日增重、腹脂率和肝重率作用不显著(P>0.05)。在饲料中添加0.08%的甜菜碱使165日龄蛋鸡的肝重率比对照组降低15.66%(P<0.05),使180日龄的日均产蛋率提高26.18%(P<0.05)。不同剂量的甜菜碱对其余性状在两个试验阶段各组之间影响均不显著( P>0.05 )。
饲料中添加0.1%的甜菜碱分别使56日龄肉鸡脂肪中A-FABP mRNA和66日龄肉鸡脂肪中LPL mRNA表达丰度显著降低(P<0.05);使56和66日龄肉鸡脂肪中FAS mRNA的表达丰度极显著降低(P<0.01)。但是,同样剂量的甜菜碱对56日龄肉鸡脂肪中LPL,66日龄肉鸡脂肪中A-FABP,56和66日龄肉鸡脂肪中PPARγ、Adiponectin、BMAL1和CLOCK mRNA的表达丰度影响不明显(P>0.05)。
对于蛋鸡半定量RT-PCR分析表明,LPL mRNA的表达量在180日龄时甜菜碱Ⅱ组(添加0.06%的甜菜碱)极显著低于对照组和甜菜碱Ⅲ组(添加0.08%的甜菜碱)(P<0.01);FAS mRNA的表达量在180日龄时甜菜碱Ⅲ组均极显著大于其余3组;Adiponectin mRNA的表达量在165日龄时甜菜碱Ⅲ组显著大于对照组(P<0.05);BMAL1 mRNA的表达量在180日龄时甜菜碱Ⅲ组显著大于对照组(P<0.05)。其余基因的表达量在不同试验组和不同试验阶段差异均不显著(P>0.05)。
亚硫酸盐测序结果显示,66日龄肉鸡脂肪中LPL基因启动子区处于低甲基化状态;对照组和甜菜碱组总体甲基化比率分别为4.55%和2.77%。56日龄肉鸡脂肪中PPARγ基因5′上游调控区处于高甲基化状态,对照组和甜菜碱组总体甲基化比率分别为76.7%和71.7%。甜菜碱对肉鸡脂肪LPL基因和PPARγ基因甲基化调控作用不明显(P>0.05)。
与肉鸡相似,180日龄蛋鸡脂肪中LPL基因启动子区处于低甲基化状态,对照组、甜菜碱Ⅰ组(添加0.04%的甜菜碱)、甜菜碱Ⅱ组和甜菜碱Ⅲ组的总体甲基化比率分别为3.20%、4.31%、1.83%和3.65%,各组之间差异不显著( P>0.05 )。PPARγ基因5′上游调控区处于高甲基化状态,对照组、甜菜碱Ⅰ组、甜菜碱Ⅱ组、甜菜碱Ⅲ组的总体甲基化比率分别为89.6%、85.7%、85.7%和71.2%,对照组与甜菜碱Ⅲ组相比差异显著(P<0.05)。
Fat metabolism is affected by many factors in livestock, including heredity, nutrition, endocrine secretion, environment and so on, which can inform complicated regulation network. Lipid is essential for the development of human and animals, and is also the very important composition in organism. However, too much fat deposition brings about many problems in swine and chicken. On account of this, the molecular characterization and epigenetic regulation of genes, which are involved in fat metabolism, were studied in swine and chicken here. And pig BMAL1 (pBMAL1) was analyzed with both molecular biology and bioinformatics methods in order to search major genes affecting fat metabolism, which can contribute to the molecular breeding of swine. In addition , the effects of betaine supplementation to diet on partial growth performance and carcass traits were analyzed in chicken, and thus to approach the effects of dietary betaine supplementation on expression and methylation of fat metabolism genes, which laid the foundation for further studying gene expression regulated by betaine and the molecular mechanism of methylation. It is of important significance for the breeding of high quality and high yield chicken.
PartⅠ:In present study, we cloned and sequenced the cDNA and partial non-coding regions of pBMAL1 with degenerate primers designed according to the sequences of human and mouse BMAL1 gene, analyzed its expression in adipose tissues and other tissues and further mapped its position on porcine chromosome. The results indicated that pBMAL cDNA had 2073 bp, including 1878 bp open reading frame (ORF), which coded 626 amino acids including bHLH, PAS-A and PAS-B domains. They shared 94.36%, 89.85%, 89.79% identity in coding region nucleotide and 99.20%, 98.24%, 97.92% in amino acid with the sequences of human, mouse and rat BMAL1, respectively. Two splice-variants were found in Landrace pig and Laiwu Black pig backfat. And the expression of pBMAL1 was detected in five adipose tissues of Landrace pig,with the highest expression level in mesentery fat tissue and the lowest in caul fat tissue, but the expression levels were not significantly different among them (P>0.05). And also the expression of pBMAL1 was detected in 14 tissues of adult Laiwu pigs, with the highest in backfat and the lowest in eye muscle. Furthermore, the pBMAL1 gene was mapped SSC 2p11-q21 with a whole genome porcine radiation hybrid (RH) panel (IMpRH).
PartⅡ:This experiment was conducted to determine the effect of dietary betaine supplementation on partial growth performance and carcass traits in broiler and laying hen; the expressions of LPL, PPARγ, A-FABP, FAS, Adiponectin, BMAL1 and CLOCK were analyzed by semi-quantitative RT-PCR method. And the methylation status of LPL and PPARγpromoters were examined by bisulfite sequencing.
Our results showed that the 0.1% of betaine addition to the diet showed no significant effect on average daily gain, percentage of abdominal fat and the ratio of liver to body weight in broiler (P>0.05). However, the addition 0.08% betaine to the diet decreased the ratio of liver to body weight by 15.66% compared with the control diet (P<0.05) in 165 d laying hen. Similarly, the daily average laying rates were increased by 26.18% (P<0.05) in 180 d laying hen by 0.08% betaine supplementation. Moreover, there were no significant differences on other traits by the addition different dosage of betaine to the diet during two test stages (P>0.05).
For broiler, the addition of 0.1% betaine to the diet decreased significantly A-FABP mRNA expression level in 56 d and LPL mRNA expression level in 66 d of subcutaneous adipose tissue (P<0.05), respectively. Also, the FAS mRNA expression level of 0.1% betaine treatment groups were extremely significantly decreased compared with that of control groups in 56 d and in 66 d (P<0.01), respectively. Nevertheless, no significant differences were found in LPL mRNA expression in 56 d and A-FABP mRNA expression in 66 d by the same betaine supplementation (P>0.05), and neither 56 d broiler nor 66 d broiler of PPARγ, Adiponectin, BMAL1 and CLOCK mRNA expression levels were significant differences by the same betaine supplementation (P>0.05).
For laying hen, semi-quantitative RT-PCR results indicated that LPL mRNA expression level in 180 d for betaineⅡgroup (0.06% betaine addition) was extremely significantly lower than those of the control group and betaineⅢgroup (0.08% betaine addition) (P<0.01); and FAS mRNA expression level in 180 d for betaineⅢgroup was extremely significantly higher than those of the other groups (P<0.01); and also both Adiponectin mRNA in 165 d and BMAL1 mRNA in 180 d for betaineⅢgroups were significantly higher than those for controls (P<0.05). The expression levels of other genes did not reach significance among groups in different test stages (P>0.05).
Bisulfite sequencing analysis results revealed that LPL promoter region was hypomethylated in subcutaneous adipose tissue of 66 d broiler; percentages of global DNA methylation in control and betaine groups were 4.55% and 2.77%, respectively. However, PPARγ5′upstream regulatory region was in hypermethylation status in subcutaneous adipose tissue of 56 d broiler. And the global DNA methylation levels of PPARγdisplayed 76.7% and 71.7% in control and betaine groups, respectively. The DNA methylation level of LPL and PPARγwere not affected by betaine supplement in broiler subcutaneous adipose tissue (P>0.05).
Similar observations were made for 180 d laying hen, in which the LPL promoters was hypomethylated. The global DNA methylation levels were 3.20%, 4.31%, 1.83% and 3.65% among control and betaineⅠ(0.04% betaine addition),ⅡandⅢgroups, respectively, and no significant differences were found among groups (P>0.05). And, PPARγ5′upstream regulatory region was also hypermethylated in subcutaneous adipose tissue of 180 d laying hen. The global DNA methylation status indicated 89.6%, 85.7%, 85.7% and 71.2%, respectively. And the difference of methylation levels was significant between control and betaineⅢgroup (P<0.05).
第一部分:以人和小鼠BMAL1基因序列设计简并引物成功克隆了猪BMAL1 cDNA和非编码区部分序列,分析了pBMAL1基因在不同部位脂肪组织和其它组织中的表达,并对其进行了精细定位。结果表明,pBMAL1基因序列长为2073bp,开放阅读框为1878bp,编码626个氨基酸,该氨基酸包括bHLH、PAS-A和PAS-B结构域。编码区核酸序列与人、小鼠和大鼠的同源性分别为94.36%, 89.85%和89.79%;氨基酸序列与人、小鼠和大鼠的同源性分别为99.20%, 98.24%和97.92%。在长白猪和莱芜猪背膘中发现了两个剪接变异体。pBMAL1 mRNA在长白猪5种脂肪组织均有表达,在肠系膜脂肪表达量最高,在花油表达量最低,但差异不显著(P>0.05);pBMAL1 mRNA在成年莱芜猪14种组织均有表达,在背膘中表达最高,在眼肌中表达最低。用辐射杂种板把pBMAL1基因定位在SSC 2p11-q21。
第二部分:通过在肉鸡和蛋鸡饲料中添加不同剂量的甜菜碱,研究甜菜碱对鸡部分生产性状和屠宰性状的影响;采用半定量RT-PCR方法分析了甜菜碱对脂肪代谢相关基因LPL、PPARγ、A-FABP、FAS、Adiponectin、BMAL1和CLOCK mRNA表达的影响,并通过亚硫酸盐测序的方法对LPL和PPARγ基因的启动子区甲基化进行了分析。
结果表明,在饲料中添加0.1%的甜菜碱对肉鸡平均日增重、腹脂率和肝重率作用不显著(P>0.05)。在饲料中添加0.08%的甜菜碱使165日龄蛋鸡的肝重率比对照组降低15.66%(P<0.05),使180日龄的日均产蛋率提高26.18%(P<0.05)。不同剂量的甜菜碱对其余性状在两个试验阶段各组之间影响均不显著( P>0.05 )。
饲料中添加0.1%的甜菜碱分别使56日龄肉鸡脂肪中A-FABP mRNA和66日龄肉鸡脂肪中LPL mRNA表达丰度显著降低(P<0.05);使56和66日龄肉鸡脂肪中FAS mRNA的表达丰度极显著降低(P<0.01)。但是,同样剂量的甜菜碱对56日龄肉鸡脂肪中LPL,66日龄肉鸡脂肪中A-FABP,56和66日龄肉鸡脂肪中PPARγ、Adiponectin、BMAL1和CLOCK mRNA的表达丰度影响不明显(P>0.05)。
对于蛋鸡半定量RT-PCR分析表明,LPL mRNA的表达量在180日龄时甜菜碱Ⅱ组(添加0.06%的甜菜碱)极显著低于对照组和甜菜碱Ⅲ组(添加0.08%的甜菜碱)(P<0.01);FAS mRNA的表达量在180日龄时甜菜碱Ⅲ组均极显著大于其余3组;Adiponectin mRNA的表达量在165日龄时甜菜碱Ⅲ组显著大于对照组(P<0.05);BMAL1 mRNA的表达量在180日龄时甜菜碱Ⅲ组显著大于对照组(P<0.05)。其余基因的表达量在不同试验组和不同试验阶段差异均不显著(P>0.05)。
亚硫酸盐测序结果显示,66日龄肉鸡脂肪中LPL基因启动子区处于低甲基化状态;对照组和甜菜碱组总体甲基化比率分别为4.55%和2.77%。56日龄肉鸡脂肪中PPARγ基因5′上游调控区处于高甲基化状态,对照组和甜菜碱组总体甲基化比率分别为76.7%和71.7%。甜菜碱对肉鸡脂肪LPL基因和PPARγ基因甲基化调控作用不明显(P>0.05)。
与肉鸡相似,180日龄蛋鸡脂肪中LPL基因启动子区处于低甲基化状态,对照组、甜菜碱Ⅰ组(添加0.04%的甜菜碱)、甜菜碱Ⅱ组和甜菜碱Ⅲ组的总体甲基化比率分别为3.20%、4.31%、1.83%和3.65%,各组之间差异不显著( P>0.05 )。PPARγ基因5′上游调控区处于高甲基化状态,对照组、甜菜碱Ⅰ组、甜菜碱Ⅱ组、甜菜碱Ⅲ组的总体甲基化比率分别为89.6%、85.7%、85.7%和71.2%,对照组与甜菜碱Ⅲ组相比差异显著(P<0.05)。
Fat metabolism is affected by many factors in livestock, including heredity, nutrition, endocrine secretion, environment and so on, which can inform complicated regulation network. Lipid is essential for the development of human and animals, and is also the very important composition in organism. However, too much fat deposition brings about many problems in swine and chicken. On account of this, the molecular characterization and epigenetic regulation of genes, which are involved in fat metabolism, were studied in swine and chicken here. And pig BMAL1 (pBMAL1) was analyzed with both molecular biology and bioinformatics methods in order to search major genes affecting fat metabolism, which can contribute to the molecular breeding of swine. In addition , the effects of betaine supplementation to diet on partial growth performance and carcass traits were analyzed in chicken, and thus to approach the effects of dietary betaine supplementation on expression and methylation of fat metabolism genes, which laid the foundation for further studying gene expression regulated by betaine and the molecular mechanism of methylation. It is of important significance for the breeding of high quality and high yield chicken.
PartⅠ:In present study, we cloned and sequenced the cDNA and partial non-coding regions of pBMAL1 with degenerate primers designed according to the sequences of human and mouse BMAL1 gene, analyzed its expression in adipose tissues and other tissues and further mapped its position on porcine chromosome. The results indicated that pBMAL cDNA had 2073 bp, including 1878 bp open reading frame (ORF), which coded 626 amino acids including bHLH, PAS-A and PAS-B domains. They shared 94.36%, 89.85%, 89.79% identity in coding region nucleotide and 99.20%, 98.24%, 97.92% in amino acid with the sequences of human, mouse and rat BMAL1, respectively. Two splice-variants were found in Landrace pig and Laiwu Black pig backfat. And the expression of pBMAL1 was detected in five adipose tissues of Landrace pig,with the highest expression level in mesentery fat tissue and the lowest in caul fat tissue, but the expression levels were not significantly different among them (P>0.05). And also the expression of pBMAL1 was detected in 14 tissues of adult Laiwu pigs, with the highest in backfat and the lowest in eye muscle. Furthermore, the pBMAL1 gene was mapped SSC 2p11-q21 with a whole genome porcine radiation hybrid (RH) panel (IMpRH).
PartⅡ:This experiment was conducted to determine the effect of dietary betaine supplementation on partial growth performance and carcass traits in broiler and laying hen; the expressions of LPL, PPARγ, A-FABP, FAS, Adiponectin, BMAL1 and CLOCK were analyzed by semi-quantitative RT-PCR method. And the methylation status of LPL and PPARγpromoters were examined by bisulfite sequencing.
Our results showed that the 0.1% of betaine addition to the diet showed no significant effect on average daily gain, percentage of abdominal fat and the ratio of liver to body weight in broiler (P>0.05). However, the addition 0.08% betaine to the diet decreased the ratio of liver to body weight by 15.66% compared with the control diet (P<0.05) in 165 d laying hen. Similarly, the daily average laying rates were increased by 26.18% (P<0.05) in 180 d laying hen by 0.08% betaine supplementation. Moreover, there were no significant differences on other traits by the addition different dosage of betaine to the diet during two test stages (P>0.05).
For broiler, the addition of 0.1% betaine to the diet decreased significantly A-FABP mRNA expression level in 56 d and LPL mRNA expression level in 66 d of subcutaneous adipose tissue (P<0.05), respectively. Also, the FAS mRNA expression level of 0.1% betaine treatment groups were extremely significantly decreased compared with that of control groups in 56 d and in 66 d (P<0.01), respectively. Nevertheless, no significant differences were found in LPL mRNA expression in 56 d and A-FABP mRNA expression in 66 d by the same betaine supplementation (P>0.05), and neither 56 d broiler nor 66 d broiler of PPARγ, Adiponectin, BMAL1 and CLOCK mRNA expression levels were significant differences by the same betaine supplementation (P>0.05).
For laying hen, semi-quantitative RT-PCR results indicated that LPL mRNA expression level in 180 d for betaineⅡgroup (0.06% betaine addition) was extremely significantly lower than those of the control group and betaineⅢgroup (0.08% betaine addition) (P<0.01); and FAS mRNA expression level in 180 d for betaineⅢgroup was extremely significantly higher than those of the other groups (P<0.01); and also both Adiponectin mRNA in 165 d and BMAL1 mRNA in 180 d for betaineⅢgroups were significantly higher than those for controls (P<0.05). The expression levels of other genes did not reach significance among groups in different test stages (P>0.05).
Bisulfite sequencing analysis results revealed that LPL promoter region was hypomethylated in subcutaneous adipose tissue of 66 d broiler; percentages of global DNA methylation in control and betaine groups were 4.55% and 2.77%, respectively. However, PPARγ5′upstream regulatory region was in hypermethylation status in subcutaneous adipose tissue of 56 d broiler. And the global DNA methylation levels of PPARγdisplayed 76.7% and 71.7% in control and betaine groups, respectively. The DNA methylation level of LPL and PPARγwere not affected by betaine supplement in broiler subcutaneous adipose tissue (P>0.05).
Similar observations were made for 180 d laying hen, in which the LPL promoters was hypomethylated. The global DNA methylation levels were 3.20%, 4.31%, 1.83% and 3.65% among control and betaineⅠ(0.04% betaine addition),ⅡandⅢgroups, respectively, and no significant differences were found among groups (P>0.05). And, PPARγ5′upstream regulatory region was also hypermethylated in subcutaneous adipose tissue of 180 d laying hen. The global DNA methylation status indicated 89.6%, 85.7%, 85.7% and 71.2%, respectively. And the difference of methylation levels was significant between control and betaineⅢgroup (P<0.05).
引文
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