品种和母猪日粮蛋白水平对仔猪肝脏胆固醇代谢的影响及其机制
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摘要
胆固醇稳态是维持正常生命活动的保障。肝脏是胆固醇合成、转运、分泌和转化的场所,在维持机体胆固醇稳态方面是不可替代的。肝脏胆固醇代谢存在品种差异,但形成这种差异的分子机制尚不清楚。营养因素影响肝脏胆固醇代谢。食物中脂类和甾醇类对肝脏胆固醇合成、转运和转化的影响研究较多,而日粮蛋白水平对肝脏胆固醇代谢的影响鲜有报道。最近在大鼠上的研究表明,母体日粮蛋白限饲通过改变胆固醇转化关键酶表遗传修饰,影响子代肝脏胆固醇的转运和转化,而母体日粮蛋白水平对肝脏胆固醇合成的影响未见报道。本实验分两个部分。第一部分以肉脂型的二花脸猪和瘦肉型的大白猪为模型,研究肝脏胆固醇代谢的品种差异及其机制。第二部分以梅山猪为模型,研究母猪妊娠期和哺乳期饲喂传统的低蛋白日粮和NRC标准蛋白日粮对子代肝脏胆固醇代谢的影响及其表遗传机制。
1仔猪肝脏胆固醇代谢的品种差异
选取0日龄和25日龄的大白猪和二花脸猪各6头,称重采样,检测血清和肝脏的胆固醇水平和血清生化指标；Real-time PCR检测肝脏胆固醇代谢相关因子的表达；Western blotting检测肝脏胆固醇代谢调控因子和酶的蛋白含量。结果显示,0日龄和25日龄二花脸猪的体重极显著低于大白猪(P<0.01),0日龄和25日龄二花脸猪的肝重极显著低于大白猪的肝重(P<0.01),但两个日龄的肝体比在品种间无显著差异。0日龄二花脸仔猪血清总胆固醇和高密度脂蛋白胆固醇浓度极显著高于大白仔猪(P<0.01),血清低密度脂蛋白胆固醇浓度二花脸猪极显著低于大白猪(P<0.0J1),血清甘油三酯浓度无显著差异。0日龄仔猪肝脏胆固醇含量二花脸猪显著高于大白猪(P<0.05),肝脏甘油三酯含量在品种间无显著差异；25日龄二花脸仔猪血清总胆固醇、高密度脂蛋白胆固醇和低密度脂蛋白胆固醇,以及甘油三酯浓度均显著高于大白猪(P<0.01)。25日龄仔猪肝脏胆固醇含量(P<0.05)和甘油三酯含量(P<0.01)二花脸猪均显著高于大白猪。0日龄仔猪肝脏胆固醇代谢相关基因的表达差异表现为肝脏胆固醇转化中性途径的关键酶胆固醇-7α-羟化酶二花脸猪显著低于大白猪(P<0.01)；25日龄仔猪肝脏胆固醇调节元件结合蛋白2(SREBP2) mRNA表达和蛋白含量二花脸猪均显著高于大白猪(P<0.05)。
以上结果表明,二花脸猪肝脏和血清胆固醇含量高于大白猪。0日龄时主要与胆固醇-7α-羟化酶表达较低有关,提示新生二花脸猪肝脏胆固醇向胆汁酸的转化水平较低；25日龄主要表现为较高的SREBP2表达,提示二花脸猪肝脏胆固醇的合成能力较强。尽管0日龄和25日龄二花脸猪肝脏胆固醇含量均高于大白猪,但其分子机制具有年龄依赖性。
2母猪日粮蛋白水平对断奶仔猪肝脏胆固醇含量及相关基因和蛋白表达的影响
选用14头初产的小梅山母猪,随机分为两组,即标准蛋白组(SP)和低蛋白组(LP)。标准蛋白组妊娠期和哺乳期饲料中粗蛋白含量分别为12%和14%,低蛋白组妊娠期和哺乳期饲料中粗蛋白含量分别为6%和7%,饲料粗蛋白含量不同但能量相等。仔猪35d断奶。测定仔猪体重、肝重,采集血清和肝脏样品,检测血清激素水平、生化指标、肝脏甘油三酯和胆固醇含量；Real-time PCR检测肝脏胆固醇代谢相关因子的表达；Western blotting检测肝脏胆固醇代谢调控因子；酶活分析试剂盒检测肝脏胆固醇合戍关键酶的活性。结果显示：LP组断奶仔猪的体重和肝重显著低于SP组(P<0.05),但肝体比两组间没有差异；母体低蛋白日粮对断奶仔猪血清T3、T4没有影响,血清胰岛素水平LP组显著低于SP组(P<0.05)；母体低蛋白日粮显著降低了断奶仔猪血清和肝脏的胆固醇含量(P<0.05),肝脏甘油三酯有下降的趋势(P=0.08)；血清低密度脂蛋白胆固醇和高密度脂蛋白胆固醇,低密度脂蛋白胆固醇/高密度脂蛋白胆固醇比值在两组间无显著差异；LP组仔猪肝脏胆固醇调节元件结合蛋白2(SREBP2)的mRNA表达和蛋白含量显著升高(P<0.05),胰岛素诱导基因(Insig)和胆固醇调节元件结合蛋白裂解激活蛋白mRNA表达显著增加(P<0.05),3-羟-3-甲基戊二酸单酰CoA还原酶(HMGCR) mRNA表达和酶活显著增加(P<0.05),肝脏胆固醇降解经典途径的关键酶-CYP7α1mRNA表达LP组显著高于SP组(P<0.05),而肝脏胆固醇降解替代途径的关键酶-CYP27α1mRNA表达两组间没有差异。
以上结果说明,梅山母猪妊娠期和哺乳期饲喂低蛋白日粮,显著降低断奶仔猪血清和肝脏的胆固醇含量,并伴随肝脏胆固醇合成、转化相关基因表达模式的变化,提示母猪日粮蛋白水平通过调控相关功能基因表达,影响子代肝脏胆固醇代谢。
3母猪日粮蛋白水平影响子代肝脏胆固醇合成的表遗传机制
鉴于羟甲基戊二酸单酰辅酶A还原酶(HMGCR)为肝脏胆固醇合成的限速酶,母体日粮蛋白水平显著影响子代肝脏HMGCR的表达和活性,并且HMGCR的启动子序列清楚,因此选择HMGCR为靶基因研究母猪日粮蛋白水平影响子代肝脏胆固醇合成的表遗传机制。用5-甲基胞嘧啶抗体进行DNA免疫沉淀检测HMGCR启动子区的DNA甲基化程度；用组蛋白H3乙酰化抗体、组蛋白H3K9单甲基化抗体、组蛋白H3K27三甲基化抗体和组蛋白H3K4三甲基化抗体进行染色质免疫沉淀分别检测HMGCR基因启动子区组蛋白H3的乙酰化和甲基化修饰状况。此外,采用Real-time检测了以HMGCR为靶的miRNA的表达。结果表明,LP组HMGCR基因启动子区甲基化和组蛋白H3K9me水平显著低于SP组(P<0.01),而组蛋白组H3乙酰化显著升高(P<0.01),同时组蛋白H3K27me3有下降的趋势(P=0.08)；但以HMGCR为靶的miRNA的表达两组间无显著差异(P>0.08)。
以上结果表明梅山母猪妊娠期和哺乳期饲喂低蛋白日粮通过降低HMGCR基因启动子的甲基化水平,提高启动子区组蛋白H3乙酰化程度,并下调H3K9me和H3K27me3表遗传标记,从而使HMGCR基因表达上调,影响肝脏胆固醇稳态。
Cholesterol homeostasis is necessary for the maintenance of normal life activities. Liver is for the place where cholesterol is synthesized, transported, secreted and transformated, thus liver takes an irreplaceable role in cholesterol homeostasis. Hepatic cholesterol metabolism differs between breeds, yet the molecular mechanism is unclear. Nutrients, such as lipids and sterols in diets, are reported to effect on hepatic cholesterol transportation and transformation, while the effect of dietary protein is rarely reported. Recent study on rats show that maternal dietary protein restriction could change cholesterol homeostasis by epigenetic modification of key enzymes for cholesterol transportation and transformation in offspring livers, while the epigenetic programming on cholesterol synthesis in the liver is not explored yet. Our study divided into two parts. The first part took fat Erhualian and lean Large White as models to study the differences in hepatic cholesterol metabolism between breeds and the mechanism involved. The second part was done on Meishan pigs to study the effects of maternal protein (traditional low-protein diet and standard protein diet of NRC) during pregnancy and lactation on hepatic cholesterol metabolism of offspring and the epigenetic mechanisms.
1. The differences on hepatic cholesterol metabolism between Erhualian and Large White piglets
Large White and Erhualian piglets (at birth and25-day-old) were used in the study,6each time point per breed. Animals were weighted, liver and serum were collected for the analysis. Cholesterol concentration and other lipid parameters were measured by commercial kits, hepatic cholesterol metabolism-related gene expressions were tested by Real-time PCR. The protein of the key regulatory factors and enzyme were investigated by Western blotting. These results suggest that Erhualian piglets had significantly lower body and liver weight compared with Large White (P<0.01), but the liver/body weight ratio showed no difference between the two breeds, no matter which time point tested. On the birth piglets, Erhualian had significantly higher serum total cholesterol (Tch) as well as higher density lipoprotein cholesterol (HDL-C) and liver cholesterol content (P<0.01), but significantly lower serum low density lipoprotein cholesterol (LDL-C) concentration compared with Large White; serum and liver triglyceride (TG) concentrations were similar between breeds (P>0.05) together with significantly lower CYP7A1mRNA in the Erhualian piglets. On the25-day-old piglets, cholesterol and triglyceride concentrations in serum (P<0.01) and liver (P<0.05) were significantly higher in Erhualian than those in Large White, with significantly higher SREBP2mRNA and protein levels in Erhualian.
In conclusion, Erhualian had significantly higher liver and serum cholesterol than Large White. The lower CYP7A1expression in birth Erhualian indicated a lower conversion rate from cholesterol to bile acids in Erhualian birth, while the higher content of SREBP2in liver result in a stronger cholesterol synthesis in Erhualian of25-day-old, indicating that the age-depended of the mechanism effect on cholesterol concentration in breed-specific in this study.
2. Effects of maternal protein on liver cholesterol, cholesterol metabolism related gene and protein expression in g Little Meishan weanin piglets
14primiparous Little Meishan sow were randomly divided into two groups, treated with standard protein (SP) and low-protein (LP) during pregnancy and lactation respectively. Crude protein concentration in SP, according to NTC, was12%during pregnancy and14%during lactation and those of LP were6%and7%, respectively; the energy content was balanced to the same. The piglets were under35-day weaning program, slaughtered at day35. Serum and liver were collected for measurement of hormones, serum lipid parameters and liver triglyceride and cholesterol analysis. Hepatic cholesterol metabolism-related gene expressions were tested by Real-time PCR. The protein of the key regulatory factors and enzyme were investigated by Western blotting. The activities of key enzymes in liver cholesterol synthesis were evaluated by commercial kits. The results indicated that body weight and liver weight of weaning piglets in LP group were significantly lower than that of SP group (P<0.05), while the liver/body weight ratio were comparable. Seum T3and T4were not changed by maternal protein, but circulating insulin, cholesterol and liver cholesterol were significantly lower in piglets from LP fed sows(P<0.05), with a tendency to down of liver TG(P=0.08). The serum HDL, LDL and HDL/LDL ratio were not changed. Piglets from LP fed sows had significant higher mRNA and protein of SREBP2, higher transcription of Insig and HMGCR and lower levels of CYP7al transcript(P<0.05). These results indicated that by feeding Little Meishan sows with low-protein diet during pregnancy and lactation, had lower serum and liver cholesterol level, accompanied with change on cholesterol synthesis-and transformation-related genes and/or proteins, suggesting that the liver cholesterol metabolism in the offspring was programmed by maternal diet protein through changing functional gene and/or protein expression.
3. Maternal dietary protein level affects offspring hepatic cholesterol synthesis through epigenetic regulation of gene expression
HMGCR is the rate-limiting enzyme in hepatic cholesterol synthesis and its promoter region is well displaied, so it is an ideal target to explore the epigenetic modifications involved in the programming mechanism on the offsprings.
Methylation level of HMGCR promoter region was measured by DNA immunoprecipitation by using5-methylcytosine antibody. Anti-histone H3acetylation, histone H3K9monomethyl, histone H3K27trimethyl and histone H3K4trimethyl antibodies were used to measure histone H3acetylation and methylation in HMGCR promoter region by chromatin immunoprecipitation. In addition, expression of miRNAs targeting HMGCR was measured by Real-time PCR. Results indicated methylation in HMGCR promoter region and histone H3K9me enrichment in the LP group was significantly lower than that in SP group (P<0.01), together with significantly higher histone H3acetylation (P<0.01) but lower histone H3K27me3(P=0.08) enrichment on HMGCR promoter region. The expression of liver miRNAs targeting HMGCR showed no significant difference between LP and SP (P>0.05). In summary, the low-protein diet during pregnancy and lactation period of Little Meishan sows could enhance HMGCR expression in the offspring livers through reducing DNA methylation of HMGCR promoter region, increasing acetylation of histone H3, and down regulation epigenetic modifications, such as H3K9me and H3K27me3.
1仔猪肝脏胆固醇代谢的品种差异
选取0日龄和25日龄的大白猪和二花脸猪各6头,称重采样,检测血清和肝脏的胆固醇水平和血清生化指标；Real-time PCR检测肝脏胆固醇代谢相关因子的表达；Western blotting检测肝脏胆固醇代谢调控因子和酶的蛋白含量。结果显示,0日龄和25日龄二花脸猪的体重极显著低于大白猪(P<0.01),0日龄和25日龄二花脸猪的肝重极显著低于大白猪的肝重(P<0.01),但两个日龄的肝体比在品种间无显著差异。0日龄二花脸仔猪血清总胆固醇和高密度脂蛋白胆固醇浓度极显著高于大白仔猪(P<0.01),血清低密度脂蛋白胆固醇浓度二花脸猪极显著低于大白猪(P<0.0J1),血清甘油三酯浓度无显著差异。0日龄仔猪肝脏胆固醇含量二花脸猪显著高于大白猪(P<0.05),肝脏甘油三酯含量在品种间无显著差异；25日龄二花脸仔猪血清总胆固醇、高密度脂蛋白胆固醇和低密度脂蛋白胆固醇,以及甘油三酯浓度均显著高于大白猪(P<0.01)。25日龄仔猪肝脏胆固醇含量(P<0.05)和甘油三酯含量(P<0.01)二花脸猪均显著高于大白猪。0日龄仔猪肝脏胆固醇代谢相关基因的表达差异表现为肝脏胆固醇转化中性途径的关键酶胆固醇-7α-羟化酶二花脸猪显著低于大白猪(P<0.01)；25日龄仔猪肝脏胆固醇调节元件结合蛋白2(SREBP2) mRNA表达和蛋白含量二花脸猪均显著高于大白猪(P<0.05)。
以上结果表明,二花脸猪肝脏和血清胆固醇含量高于大白猪。0日龄时主要与胆固醇-7α-羟化酶表达较低有关,提示新生二花脸猪肝脏胆固醇向胆汁酸的转化水平较低；25日龄主要表现为较高的SREBP2表达,提示二花脸猪肝脏胆固醇的合成能力较强。尽管0日龄和25日龄二花脸猪肝脏胆固醇含量均高于大白猪,但其分子机制具有年龄依赖性。
2母猪日粮蛋白水平对断奶仔猪肝脏胆固醇含量及相关基因和蛋白表达的影响
选用14头初产的小梅山母猪,随机分为两组,即标准蛋白组(SP)和低蛋白组(LP)。标准蛋白组妊娠期和哺乳期饲料中粗蛋白含量分别为12%和14%,低蛋白组妊娠期和哺乳期饲料中粗蛋白含量分别为6%和7%,饲料粗蛋白含量不同但能量相等。仔猪35d断奶。测定仔猪体重、肝重,采集血清和肝脏样品,检测血清激素水平、生化指标、肝脏甘油三酯和胆固醇含量；Real-time PCR检测肝脏胆固醇代谢相关因子的表达；Western blotting检测肝脏胆固醇代谢调控因子；酶活分析试剂盒检测肝脏胆固醇合戍关键酶的活性。结果显示：LP组断奶仔猪的体重和肝重显著低于SP组(P<0.05),但肝体比两组间没有差异；母体低蛋白日粮对断奶仔猪血清T3、T4没有影响,血清胰岛素水平LP组显著低于SP组(P<0.05)；母体低蛋白日粮显著降低了断奶仔猪血清和肝脏的胆固醇含量(P<0.05),肝脏甘油三酯有下降的趋势(P=0.08)；血清低密度脂蛋白胆固醇和高密度脂蛋白胆固醇,低密度脂蛋白胆固醇/高密度脂蛋白胆固醇比值在两组间无显著差异；LP组仔猪肝脏胆固醇调节元件结合蛋白2(SREBP2)的mRNA表达和蛋白含量显著升高(P<0.05),胰岛素诱导基因(Insig)和胆固醇调节元件结合蛋白裂解激活蛋白mRNA表达显著增加(P<0.05),3-羟-3-甲基戊二酸单酰CoA还原酶(HMGCR) mRNA表达和酶活显著增加(P<0.05),肝脏胆固醇降解经典途径的关键酶-CYP7α1mRNA表达LP组显著高于SP组(P<0.05),而肝脏胆固醇降解替代途径的关键酶-CYP27α1mRNA表达两组间没有差异。
以上结果说明,梅山母猪妊娠期和哺乳期饲喂低蛋白日粮,显著降低断奶仔猪血清和肝脏的胆固醇含量,并伴随肝脏胆固醇合成、转化相关基因表达模式的变化,提示母猪日粮蛋白水平通过调控相关功能基因表达,影响子代肝脏胆固醇代谢。
3母猪日粮蛋白水平影响子代肝脏胆固醇合成的表遗传机制
鉴于羟甲基戊二酸单酰辅酶A还原酶(HMGCR)为肝脏胆固醇合成的限速酶,母体日粮蛋白水平显著影响子代肝脏HMGCR的表达和活性,并且HMGCR的启动子序列清楚,因此选择HMGCR为靶基因研究母猪日粮蛋白水平影响子代肝脏胆固醇合成的表遗传机制。用5-甲基胞嘧啶抗体进行DNA免疫沉淀检测HMGCR启动子区的DNA甲基化程度；用组蛋白H3乙酰化抗体、组蛋白H3K9单甲基化抗体、组蛋白H3K27三甲基化抗体和组蛋白H3K4三甲基化抗体进行染色质免疫沉淀分别检测HMGCR基因启动子区组蛋白H3的乙酰化和甲基化修饰状况。此外,采用Real-time检测了以HMGCR为靶的miRNA的表达。结果表明,LP组HMGCR基因启动子区甲基化和组蛋白H3K9me水平显著低于SP组(P<0.01),而组蛋白组H3乙酰化显著升高(P<0.01),同时组蛋白H3K27me3有下降的趋势(P=0.08)；但以HMGCR为靶的miRNA的表达两组间无显著差异(P>0.08)。
以上结果表明梅山母猪妊娠期和哺乳期饲喂低蛋白日粮通过降低HMGCR基因启动子的甲基化水平,提高启动子区组蛋白H3乙酰化程度,并下调H3K9me和H3K27me3表遗传标记,从而使HMGCR基因表达上调,影响肝脏胆固醇稳态。
Cholesterol homeostasis is necessary for the maintenance of normal life activities. Liver is for the place where cholesterol is synthesized, transported, secreted and transformated, thus liver takes an irreplaceable role in cholesterol homeostasis. Hepatic cholesterol metabolism differs between breeds, yet the molecular mechanism is unclear. Nutrients, such as lipids and sterols in diets, are reported to effect on hepatic cholesterol transportation and transformation, while the effect of dietary protein is rarely reported. Recent study on rats show that maternal dietary protein restriction could change cholesterol homeostasis by epigenetic modification of key enzymes for cholesterol transportation and transformation in offspring livers, while the epigenetic programming on cholesterol synthesis in the liver is not explored yet. Our study divided into two parts. The first part took fat Erhualian and lean Large White as models to study the differences in hepatic cholesterol metabolism between breeds and the mechanism involved. The second part was done on Meishan pigs to study the effects of maternal protein (traditional low-protein diet and standard protein diet of NRC) during pregnancy and lactation on hepatic cholesterol metabolism of offspring and the epigenetic mechanisms.
1. The differences on hepatic cholesterol metabolism between Erhualian and Large White piglets
Large White and Erhualian piglets (at birth and25-day-old) were used in the study,6each time point per breed. Animals were weighted, liver and serum were collected for the analysis. Cholesterol concentration and other lipid parameters were measured by commercial kits, hepatic cholesterol metabolism-related gene expressions were tested by Real-time PCR. The protein of the key regulatory factors and enzyme were investigated by Western blotting. These results suggest that Erhualian piglets had significantly lower body and liver weight compared with Large White (P<0.01), but the liver/body weight ratio showed no difference between the two breeds, no matter which time point tested. On the birth piglets, Erhualian had significantly higher serum total cholesterol (Tch) as well as higher density lipoprotein cholesterol (HDL-C) and liver cholesterol content (P<0.01), but significantly lower serum low density lipoprotein cholesterol (LDL-C) concentration compared with Large White; serum and liver triglyceride (TG) concentrations were similar between breeds (P>0.05) together with significantly lower CYP7A1mRNA in the Erhualian piglets. On the25-day-old piglets, cholesterol and triglyceride concentrations in serum (P<0.01) and liver (P<0.05) were significantly higher in Erhualian than those in Large White, with significantly higher SREBP2mRNA and protein levels in Erhualian.
In conclusion, Erhualian had significantly higher liver and serum cholesterol than Large White. The lower CYP7A1expression in birth Erhualian indicated a lower conversion rate from cholesterol to bile acids in Erhualian birth, while the higher content of SREBP2in liver result in a stronger cholesterol synthesis in Erhualian of25-day-old, indicating that the age-depended of the mechanism effect on cholesterol concentration in breed-specific in this study.
2. Effects of maternal protein on liver cholesterol, cholesterol metabolism related gene and protein expression in g Little Meishan weanin piglets
14primiparous Little Meishan sow were randomly divided into two groups, treated with standard protein (SP) and low-protein (LP) during pregnancy and lactation respectively. Crude protein concentration in SP, according to NTC, was12%during pregnancy and14%during lactation and those of LP were6%and7%, respectively; the energy content was balanced to the same. The piglets were under35-day weaning program, slaughtered at day35. Serum and liver were collected for measurement of hormones, serum lipid parameters and liver triglyceride and cholesterol analysis. Hepatic cholesterol metabolism-related gene expressions were tested by Real-time PCR. The protein of the key regulatory factors and enzyme were investigated by Western blotting. The activities of key enzymes in liver cholesterol synthesis were evaluated by commercial kits. The results indicated that body weight and liver weight of weaning piglets in LP group were significantly lower than that of SP group (P<0.05), while the liver/body weight ratio were comparable. Seum T3and T4were not changed by maternal protein, but circulating insulin, cholesterol and liver cholesterol were significantly lower in piglets from LP fed sows(P<0.05), with a tendency to down of liver TG(P=0.08). The serum HDL, LDL and HDL/LDL ratio were not changed. Piglets from LP fed sows had significant higher mRNA and protein of SREBP2, higher transcription of Insig and HMGCR and lower levels of CYP7al transcript(P<0.05). These results indicated that by feeding Little Meishan sows with low-protein diet during pregnancy and lactation, had lower serum and liver cholesterol level, accompanied with change on cholesterol synthesis-and transformation-related genes and/or proteins, suggesting that the liver cholesterol metabolism in the offspring was programmed by maternal diet protein through changing functional gene and/or protein expression.
3. Maternal dietary protein level affects offspring hepatic cholesterol synthesis through epigenetic regulation of gene expression
HMGCR is the rate-limiting enzyme in hepatic cholesterol synthesis and its promoter region is well displaied, so it is an ideal target to explore the epigenetic modifications involved in the programming mechanism on the offsprings.
Methylation level of HMGCR promoter region was measured by DNA immunoprecipitation by using5-methylcytosine antibody. Anti-histone H3acetylation, histone H3K9monomethyl, histone H3K27trimethyl and histone H3K4trimethyl antibodies were used to measure histone H3acetylation and methylation in HMGCR promoter region by chromatin immunoprecipitation. In addition, expression of miRNAs targeting HMGCR was measured by Real-time PCR. Results indicated methylation in HMGCR promoter region and histone H3K9me enrichment in the LP group was significantly lower than that in SP group (P<0.01), together with significantly higher histone H3acetylation (P<0.01) but lower histone H3K27me3(P=0.08) enrichment on HMGCR promoter region. The expression of liver miRNAs targeting HMGCR showed no significant difference between LP and SP (P>0.05). In summary, the low-protein diet during pregnancy and lactation period of Little Meishan sows could enhance HMGCR expression in the offspring livers through reducing DNA methylation of HMGCR promoter region, increasing acetylation of histone H3, and down regulation epigenetic modifications, such as H3K9me and H3K27me3.
引文
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