牛SHH基因通过PPARg通路调控脂肪生成
摘要
本研究利用定量PCR、基因克隆、原核表达、蛋白纯化、细胞培养以及真核表达等实验技术，综合分析了牛SHH基因和PPARg通路在脂肪生成中的调控机制，研究内容包括牛不同部位脂肪之间候选基因的表达差异，牛SHH基因在脂肪生成中的作用，牛SHH基因与PPARg通路的关系，PPARg基因与肌内脂肪生成的关系，PPARg基因启动子的克隆与活性鉴定以及PPARg基因的遗传变异分析等，本研究主要取得了以下结果：
1、前脂肪细胞的形成与分化影响不同部位脂肪组织的沉积
利用定量PCR分别检测了候选基因在牛皮下、肌间、胸腔和腹腔脂肪中的表达模式，结果表明HSL、CIDEC、Perilipin、Leptin和Glut4等与甘油三酯代谢相关的基因在肌间脂肪和腹腔脂肪的表达水平要显著高于皮下脂肪和胸腔脂肪（p <0.05），FABP4、PPARg、DLK1等与脂肪细胞分化相关的基因在牛皮下脂肪中的表达水平要低于肌间脂肪、腹腔脂肪和胸腔脂肪（p <0.05），说明牛不同部位脂肪组织的代谢水平、分化状态和分化潜能都存在差异，同时说明前脂肪细胞的形成与分化对不同部位脂肪的沉积有着重要的影响。
2、重组牛SHH蛋白能够抑制前脂肪细胞的分化
利用Overlap PCR的方法成功克隆了秦川牛的SHH基因，利用原核表达获得了重组牛SHH蛋白，并使用Western blotting验证了蛋白的特异性。将SHH蛋白按照1μg/mL的浓度添加到3T3-L1细胞的培养基中，能够抑制其向成熟脂肪细胞的分化，说明所获得的重组牛SHH蛋白具有抑制脂肪细胞分化的活性。
3、牛SHH基因通过PPARg通路调控脂肪生成
克隆了牛PPARg基因并构建了真核表达载体，在前脂肪细胞中过表达PPARg能够诱导脂肪细胞的分化，而同时过表达SHH和PPARg能够挽救SHH对脂肪细胞分化的抑制作用，说明牛SHH能够在早期通过PPARg通路调控脂肪细胞的分化。使用定量PCR分析表明，过表达SHH能够诱导Nr2f2的表达，而过表达Nr2f2能够抑制脂肪细胞分化过程中PPARg的表达，在细胞中同时过表达Nr2f2和PPARg能够挽救Nr2f2对脂肪细胞分化的抑制作用。说明SHH能够通过上调Nr2f2的表达水平抑制PPARg的表达进而抑制机体脂肪的生成。
4、克隆了牛PPARg基因的启动子并鉴定了其活性
通过将DsRed基因与EGFP基因克隆进同一个载体中构建了一种新的可以实时检测目标启动子的活性的双色荧光报告系统。利用该报告系统鉴定了PPARg基因启动子，PPARg的启动子位于转录起始位点-446bp到-196bp这段区域中，其在前脂肪细胞中表现出微弱的活性，而在脂肪细胞分化时活性得到显著增强。
5、牛原代肌肉细胞中过表达PPARg有助于肌内脂肪的生成
成功包装了PPARg基因的腺病毒过表达载体，在体外培养的牛原代肌肉细胞中过表达PPARg后，脂肪生成的正向调控基因FABP4、Glut4和AdPLA的表达水平分别上调至1.9、1.5和1.6倍，而脂肪生成的负向调控基因GATA2、HSL和Nf2f2的表达水平分别下调至0.5、0.3和0.6倍，说明过表达PPARg有助于体外培养的原代肌肉细胞中脂肪的生成。
6、Asp7Gly突变影响了PPARg的诱导脂肪分化能力
利用PCR-SSCP、PCR-RFLP和测序等方法检测了PPARg基因mRNA区域在秦川牛、南阳牛、郏县红牛和中国荷斯坦牛上的遗传变异，并分析了变异对牛体尺性状的影响。结果表明，在秦川牛、南阳牛和郏县红牛上共发现了4个SNPs（-110G>C，-27C>T，+20A>G和+1344G>T），其中-110G>C，-27C>T，+20A>G处于连锁状态与牛的体重、胸围和体长成负关联状态。PPARg的+20A>G突变引起了第7位氨基酸由Asp突变为Gly，被称为PPARg Asp7Gly，在前脂肪细胞中过表达PPARg的两种单倍型表明，Asp7Gly降低了PPARg在细胞内的转录活性，从而影响了其诱导脂肪细胞分化的能力，提示PPARg Asp7Gly可以作为一个分子标记进一步应用于牛的育种实践中。
Quantitative PCR, gene clone, prokaryotic expression, protein purification, cell culture andeukaryotic expression were used to analyze the regulatory mechanism of SHH gene andPPARg pathway comprehensively. The research contents includes expression patterns ofcandidate genes in fat tissue from different parts of cattle, effecting of cattle SHH inadipogenesis, relationship between SHH gene and PPARg pathway, effecting of PPARg inadipogenesis of intramuscular fat, identification the promoter of PPARg gene and the geneticvariation analysis of PPARg gene in cattle herds. The main results including:
1. The formation and differentiation of preadipocytes affect the fat deposition from differentparts of body.
Quantitative PCR were used to test the expression patterns of candidate genes in thesubcutaneous fat, abdominal fat, pleural fat and intermuscular fat. The results showed that theexpressions of triglyceride metabolism-related genes including HSL, CIDEC, Perilipin, Leptinand Glut4were higher in the abdominal fat and intermuscular fat than the pleural fat andsubcutaneous fat, and the expressions of adipose differentiation-related genes includingFABP4, PPARg and DLK1are lower in the subcutaneous than the intermuscular fat,abdominal fat and pleural fat，suggesting that the adipose metabolism level, differentiationstatus and differentiation potential are different from different parts of cattle, and alsosuggenting that the formation and differentiation of preadipocytes affect the fat depositionfrom different parts of body.
2. Recombinant bovine SHH protein can inhibit the differentiation of preadipocytes
Overlap PCR was used to clone the SHH gene of Qinchuan cattle, and the specificity ofrecombinant SHH protein purified from prokaryotic expression was tested by western blotting.Recombinant SHH protein could inhibit the differentiation of3T3-L1cells at theconcentration of1μg/mL, indicating that the recombinanat bovine SHH protein processes theinhibiting activity on adipocyte differentiation.
3. Cattle SHH inhibits adipocyte differentiation through PPARg pathway
Cattle PPARg gene was cloned. Overexpression of PPARg could induce the adipocytedifferentiation, and co-overexpression of SHH and PPARg could rescue the inhibition of SHHon adipocyte differentiation, suggesting that SHH inhibits adipocyte differentiation throughPPARg pathway. Quantitative PCR analysis showed that overexpression of SHH could induce the expression of Nr2f2, while overexpression of Nr2f2could inhibit the expression of PPARgduring adipocyte differentiation. Co-overexpression of Nr2f2and PPARg could rescue theinhibition of Nr2f2on the adipocyte differentiation, suggesting that SHH is able to inhibit theexpression of PPARg by increasing the expression of Nr2f2, thereby inhibiting adipocytedifferentiation.
4. Cloning and activity identification of PPARg promoter
A novel dual color fluorescent reporter system was established by cloning a Discosoma redfluorescent protein gene and a green fluorescent protein gene into a single vector. This dualcolor fluorescent reporter system allows real time quantitative monitoring of promoter activity.Transcriptional activity of PPARg promoter was identified, the results showed that the PPARgpromoter located at the-446bp to-196bp from the transcriptional start site. This promoterdrives weak transcriptional activity in preadipocyte, and its transcriptional activity increasessignificantly during the adipocyte differentiation.
5. Overexpression of PPARg in the bovine primary muscle cells contributes to the formationof intramuscular fat
Adenovirus vector of PPARg (Ad-PPARg) was packed successfully, and infection with theAd-PPARg in the cattle primary muscle cells showed that, the expressions of adipogenesisup-regulating genes including FABP4, Glut4and AdPLA increased1.9,1.5and1.6times,respectively, while the expression of adipogenesis down-regulating genes including GATA2,HSL and Nr2f2decreased0.5,0.3and0.6times, suggesting that overexpression of PPARg inthe bovine primary muscle cells contributes to the formation of intramuscular fat.
6. Asp7Gly mutation damages the inducing adipogenesis of PPARg
PCR-SSCP, PCR-RFLP and sequencing were used to detect the genetic variations of PPARggene in Qinchuan cattle, Nanyang cattle, Jiaxian Red cattle and China Holstein cattle, and theassociations between the PPARg variations and cattle body measurements were analyzed. Theresults showed that four SNPs (-110G>C,-27C>T,+20A>G and+1344G>T) were detected inthe PPARg gene in Qinchuan cattle, Nanyang cattle and Jiaxian Red cattle rather than ChinaHolstein cattle. The mutations-110G>C,-27C>T and+20A>G under linkage disequilibriumwere associated with smaller body weight, heart girth and body length of cattle. The mutation+20A>G changed the7thamino acid of PPARg from Asp to Gly, so called PPARg Asp7GLy.Overexpression of the two haplotypes of PPARg in preadipocytes showed that, Asp7Glydamages the transcriptional activity of PPARg, thereby decreases its inducing adipogenesis.These findings may explains the associations results of the mutations. PPARg Asp7Gly can beused as a molecular marker in cattle breeding practices.
1、前脂肪细胞的形成与分化影响不同部位脂肪组织的沉积
利用定量PCR分别检测了候选基因在牛皮下、肌间、胸腔和腹腔脂肪中的表达模式，结果表明HSL、CIDEC、Perilipin、Leptin和Glut4等与甘油三酯代谢相关的基因在肌间脂肪和腹腔脂肪的表达水平要显著高于皮下脂肪和胸腔脂肪（p <0.05），FABP4、PPARg、DLK1等与脂肪细胞分化相关的基因在牛皮下脂肪中的表达水平要低于肌间脂肪、腹腔脂肪和胸腔脂肪（p <0.05），说明牛不同部位脂肪组织的代谢水平、分化状态和分化潜能都存在差异，同时说明前脂肪细胞的形成与分化对不同部位脂肪的沉积有着重要的影响。
2、重组牛SHH蛋白能够抑制前脂肪细胞的分化
利用Overlap PCR的方法成功克隆了秦川牛的SHH基因，利用原核表达获得了重组牛SHH蛋白，并使用Western blotting验证了蛋白的特异性。将SHH蛋白按照1μg/mL的浓度添加到3T3-L1细胞的培养基中，能够抑制其向成熟脂肪细胞的分化，说明所获得的重组牛SHH蛋白具有抑制脂肪细胞分化的活性。
3、牛SHH基因通过PPARg通路调控脂肪生成
克隆了牛PPARg基因并构建了真核表达载体，在前脂肪细胞中过表达PPARg能够诱导脂肪细胞的分化，而同时过表达SHH和PPARg能够挽救SHH对脂肪细胞分化的抑制作用，说明牛SHH能够在早期通过PPARg通路调控脂肪细胞的分化。使用定量PCR分析表明，过表达SHH能够诱导Nr2f2的表达，而过表达Nr2f2能够抑制脂肪细胞分化过程中PPARg的表达，在细胞中同时过表达Nr2f2和PPARg能够挽救Nr2f2对脂肪细胞分化的抑制作用。说明SHH能够通过上调Nr2f2的表达水平抑制PPARg的表达进而抑制机体脂肪的生成。
4、克隆了牛PPARg基因的启动子并鉴定了其活性
通过将DsRed基因与EGFP基因克隆进同一个载体中构建了一种新的可以实时检测目标启动子的活性的双色荧光报告系统。利用该报告系统鉴定了PPARg基因启动子，PPARg的启动子位于转录起始位点-446bp到-196bp这段区域中，其在前脂肪细胞中表现出微弱的活性，而在脂肪细胞分化时活性得到显著增强。
5、牛原代肌肉细胞中过表达PPARg有助于肌内脂肪的生成
成功包装了PPARg基因的腺病毒过表达载体，在体外培养的牛原代肌肉细胞中过表达PPARg后，脂肪生成的正向调控基因FABP4、Glut4和AdPLA的表达水平分别上调至1.9、1.5和1.6倍，而脂肪生成的负向调控基因GATA2、HSL和Nf2f2的表达水平分别下调至0.5、0.3和0.6倍，说明过表达PPARg有助于体外培养的原代肌肉细胞中脂肪的生成。
6、Asp7Gly突变影响了PPARg的诱导脂肪分化能力
利用PCR-SSCP、PCR-RFLP和测序等方法检测了PPARg基因mRNA区域在秦川牛、南阳牛、郏县红牛和中国荷斯坦牛上的遗传变异，并分析了变异对牛体尺性状的影响。结果表明，在秦川牛、南阳牛和郏县红牛上共发现了4个SNPs（-110G>C，-27C>T，+20A>G和+1344G>T），其中-110G>C，-27C>T，+20A>G处于连锁状态与牛的体重、胸围和体长成负关联状态。PPARg的+20A>G突变引起了第7位氨基酸由Asp突变为Gly，被称为PPARg Asp7Gly，在前脂肪细胞中过表达PPARg的两种单倍型表明，Asp7Gly降低了PPARg在细胞内的转录活性，从而影响了其诱导脂肪细胞分化的能力，提示PPARg Asp7Gly可以作为一个分子标记进一步应用于牛的育种实践中。
Quantitative PCR, gene clone, prokaryotic expression, protein purification, cell culture andeukaryotic expression were used to analyze the regulatory mechanism of SHH gene andPPARg pathway comprehensively. The research contents includes expression patterns ofcandidate genes in fat tissue from different parts of cattle, effecting of cattle SHH inadipogenesis, relationship between SHH gene and PPARg pathway, effecting of PPARg inadipogenesis of intramuscular fat, identification the promoter of PPARg gene and the geneticvariation analysis of PPARg gene in cattle herds. The main results including:
1. The formation and differentiation of preadipocytes affect the fat deposition from differentparts of body.
Quantitative PCR were used to test the expression patterns of candidate genes in thesubcutaneous fat, abdominal fat, pleural fat and intermuscular fat. The results showed that theexpressions of triglyceride metabolism-related genes including HSL, CIDEC, Perilipin, Leptinand Glut4were higher in the abdominal fat and intermuscular fat than the pleural fat andsubcutaneous fat, and the expressions of adipose differentiation-related genes includingFABP4, PPARg and DLK1are lower in the subcutaneous than the intermuscular fat,abdominal fat and pleural fat，suggesting that the adipose metabolism level, differentiationstatus and differentiation potential are different from different parts of cattle, and alsosuggenting that the formation and differentiation of preadipocytes affect the fat depositionfrom different parts of body.
2. Recombinant bovine SHH protein can inhibit the differentiation of preadipocytes
Overlap PCR was used to clone the SHH gene of Qinchuan cattle, and the specificity ofrecombinant SHH protein purified from prokaryotic expression was tested by western blotting.Recombinant SHH protein could inhibit the differentiation of3T3-L1cells at theconcentration of1μg/mL, indicating that the recombinanat bovine SHH protein processes theinhibiting activity on adipocyte differentiation.
3. Cattle SHH inhibits adipocyte differentiation through PPARg pathway
Cattle PPARg gene was cloned. Overexpression of PPARg could induce the adipocytedifferentiation, and co-overexpression of SHH and PPARg could rescue the inhibition of SHHon adipocyte differentiation, suggesting that SHH inhibits adipocyte differentiation throughPPARg pathway. Quantitative PCR analysis showed that overexpression of SHH could induce the expression of Nr2f2, while overexpression of Nr2f2could inhibit the expression of PPARgduring adipocyte differentiation. Co-overexpression of Nr2f2and PPARg could rescue theinhibition of Nr2f2on the adipocyte differentiation, suggesting that SHH is able to inhibit theexpression of PPARg by increasing the expression of Nr2f2, thereby inhibiting adipocytedifferentiation.
4. Cloning and activity identification of PPARg promoter
A novel dual color fluorescent reporter system was established by cloning a Discosoma redfluorescent protein gene and a green fluorescent protein gene into a single vector. This dualcolor fluorescent reporter system allows real time quantitative monitoring of promoter activity.Transcriptional activity of PPARg promoter was identified, the results showed that the PPARgpromoter located at the-446bp to-196bp from the transcriptional start site. This promoterdrives weak transcriptional activity in preadipocyte, and its transcriptional activity increasessignificantly during the adipocyte differentiation.
5. Overexpression of PPARg in the bovine primary muscle cells contributes to the formationof intramuscular fat
Adenovirus vector of PPARg (Ad-PPARg) was packed successfully, and infection with theAd-PPARg in the cattle primary muscle cells showed that, the expressions of adipogenesisup-regulating genes including FABP4, Glut4and AdPLA increased1.9,1.5and1.6times,respectively, while the expression of adipogenesis down-regulating genes including GATA2,HSL and Nr2f2decreased0.5,0.3and0.6times, suggesting that overexpression of PPARg inthe bovine primary muscle cells contributes to the formation of intramuscular fat.
6. Asp7Gly mutation damages the inducing adipogenesis of PPARg
PCR-SSCP, PCR-RFLP and sequencing were used to detect the genetic variations of PPARggene in Qinchuan cattle, Nanyang cattle, Jiaxian Red cattle and China Holstein cattle, and theassociations between the PPARg variations and cattle body measurements were analyzed. Theresults showed that four SNPs (-110G>C,-27C>T,+20A>G and+1344G>T) were detected inthe PPARg gene in Qinchuan cattle, Nanyang cattle and Jiaxian Red cattle rather than ChinaHolstein cattle. The mutations-110G>C,-27C>T and+20A>G under linkage disequilibriumwere associated with smaller body weight, heart girth and body length of cattle. The mutation+20A>G changed the7thamino acid of PPARg from Asp to Gly, so called PPARg Asp7GLy.Overexpression of the two haplotypes of PPARg in preadipocytes showed that, Asp7Glydamages the transcriptional activity of PPARg, thereby decreases its inducing adipogenesis.These findings may explains the associations results of the mutations. PPARg Asp7Gly can beused as a molecular marker in cattle breeding practices.
引文
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