摘要
肥胖的发生主要是因为体内脂肪组织中脂肪细胞数目的增多和脂肪细胞体积的增大。脂肪细胞分化过程中伴随许多调控因子表达量的动态变化,而这些调控因子表达量的变化则决定了脂肪细胞数目和功能的改变。目前,已经发现了大量调控脂肪细胞分化的调节因子,然而仍有许多新的调控因子可能参与调控脂肪细胞分化,其功能及调节机理还不清楚。
根据本实验室早期的基因芯片数据结合基因芯片数据库(GEO Datasets2659,2660和GEO Dataset2818),我们筛选出肪细胞分化过程中表达量变化较大的新基因Ogn和Prrx1,此前并无其在脂肪细胞中功能的相关报道。利用Real-time qPCR技术验证了Ogn和Prrx1mRNA在脂肪细胞分化过程中的表达规律,然后分别构建shRNA干扰载体,利用逆病毒感染体外培养的3T3-L1前体脂肪细胞,经过筛选、鉴定之后研究其在脂肪细胞分化中的功能,利用油红O染色和BODIPY染色技术显示候选基因对细胞内脂质积累的影响。随后,利用Real-time qPCR技术和Western blot技术研究在此过程中脂肪细胞分化标志基因以及相关信号通路基因和蛋白表达量的变化；并研究了所筛选因子在3T3-L1脂肪细胞葡萄糖吸收和甘油三脂分解中的功能。获得结果如下：
1Ogn通过Wnt/p-catenin信号通路调控3T3-L1脂肪细胞分化
1)根据基因芯片和数据库信息筛选发现Ogn在脂肪细胞分化过程中表达量逐渐降低,利用此外培养的3T3-L1前体脂肪细胞模型,Real-time qPCR技术结果显示,Ogn在脂肪细胞分化过程中的表达与基因芯片数据一致；此外,Ogn在小鼠白色脂肪组织基质细胞中的表达量高于在脂肪细胞中的表达,也与基因芯片数据一致。
2)利用逆病毒介导的Ogn shRNA干扰载体转染3T3-L1前体脂肪细胞,结果显示,抑制Ogn的表达促进了脂肪细胞分化,脂质积累增加,脂肪细胞分化标志因子PPARy和FABP4mRNA表达量及蛋白表达都显著增加。在3T3-L1前体脂肪细胞中干扰Ogn的表达抑制了基础状态下葡萄糖的吸收,而对脂质分解则没有影响。
3)3T3-L1前体脂肪细胞中过表达Ogn对脂肪细胞分化无明显影响,过表达Ogn的脂肪细胞中PPARy mRNA表达量有降低趋势,然而差异不显著。
4) Wnt3a能抑制脂肪细胞的分化,在3T3-L1前体脂肪细胞中干扰Ogn的表达时,Wnt3a对脂肪细胞分化的抑制作用明显减弱。进一步研究发现,Wnt3a能以时间依赖性方式促进β-catenin向细胞核转位,而抑制Ogn的表达则明显抑制此过程。可以推测,Ogn通过作用于β-catenin的转核过程从而参与脂肪细胞分化的调控。
总之,Ogn在脂肪细胞分化过程中表达下调,抑制Ogn的表达能促进脂肪细胞分化,上调脂肪细胞分化关键因子mRNA和蛋白的表达：Ogn能影响Wnt3a对β-catenin转核的调控作用,说明Ogn可能通过作用于Wnt/β-catenin信号通路来抑制脂肪细胞分化。
2Prrx1通过激活TGFβ通路抑制脂肪细胞分化
1)与基因芯片数据一致,Real-time qPCR结果显示,Prrxla, Prrx1b和Prrx2mRNA的表达在脂肪形成过程逐渐降低,Prrx1a和Prrx1b在WAT脂肪细胞中的表达要比基质细胞中低约80%,Prrx2在脂肪细胞中的表达几乎检测不到。
2)T扰Prrx1的表达能促进3T3-L1脂肪细胞中脂质积累：Real-time qPCR结果显示,抑制内源性Prrxl的表达显著上调了脂肪细胞中PPARγ、C/EBPa和FABP4mRNA的表达以及蛋白的表达。
3)过表达Prrx1a, Prrx1b和Prrx2对脂肪细胞分化没有影响：然而,Prrx1a, Prrx1b过表达载体瞬时转染HEK293T细胞能抑制PPARy启动子的活性。
4)利用Real-time qPCR技术研究发现,在3T3-L1前体脂肪细胞中干扰Prrx1的表达,能抑制Tgfb2和Tgf3的表达,提示Prrx1能在脂肪细胞中调控TGFβ言号通路；利用TGFβ通路特殊抑制剂SB431542处理脂肪细胞则能重现Prrx1对脂肪细胞分化的调控作用,表明Prrx1可能通过TGFβ信号通路调控脂肪细胞分化。
5) Wnt3a和TNFa均能以剂量依赖性方式抑制脂肪细胞分化,干扰3T3-L1脂肪细胞中Prrx1的表达对Wnt3a和TNFa通路调控脂肪细胞分化的作用没有影响。
6)给3周龄BL6和C3H小鼠饲喂高脂日粮构建了肥胖小鼠模型,取其腹股沟周围脂肪组织,提取总RNA, Real-time qPCR技术研究显示Prrx1a和Prrxlb mRNA在肥胖小鼠脂肪组织中的表达与Tgfb3的表达显著相关,提示体内Prrx1也与TGFβ信号通路密切相关。
综上,Prrx1a和Prrx1b在脂肪细胞分化过程中表达逐渐降低,抑制内源性Prrx1的表达能促进脂质积累,上调脂肪细胞分化标志因子mRNA和蛋白的表达;过表达Prrx1对脂肪细胞分化无明显作用,但能抑制PPARy启动子活性；Prrx1能通过激活TGFβ通路抑制脂肪细胞分化,并在肥胖小鼠脂肪组织中与Tgfb3的表达显著相关,提示Prrx1可能在体内体外均能作用于TGFβ通路来调控脂肪细胞分化。
Obesity is resulted from increasing of number and size of adipocytes. Development of adipocyte tissue is regulated by a lot of transcription regulator. Many genes go up or down during adipocytes differentiation regulating adipocytes differentiation and function. Although a lots of regulator were discovered, there are still many novel genes'function in adipocytes need to be uncovered, Their expressions were dynamically changed during adipocytes differentiation, which implied they may involved in the regulation of adipocytes differentiation.
We found a list of novel genes that have not been reported yet in adipocytes from microarray and gene expression data base(GEO Datasets2659,2660and GEO Dataset2818). Ogn and Prrxl were selected and their expression profile during timecourse of3T3-L1adipocytes differentiation were analyzed by Real-time qPCR. Then pSuper retrovirus shRNA RNAi vector for Ogn and Prrx1were constructed and transfected to HEK293T cells. The virous were collected and filtered before infecting3T3-L1preadipocytes. After selection with antibiotic, their function in adipocytes differentiation and function were studied. Oil red O staining and BODIPY staining were used to show lipid accumulation in adipocytes after shRNA knockdown. Realtime PCR and Western blot were used to study change of gene expression or protein expression of makers for adipocytes differentiation. Furthermore, their functions in glucose uptake and lipids lipolysis were also studied. Results are shown as follows:
1Ogn regulate3T3-L1adipocytes differentiation through Wnt/β-catenin signaling pathway
1) According to microarray data and gene expression data base, Ogn mRNA expression is decreased during adipocytes differentiation. Our results showed similar results in3T3-L1adipocytes. Ogn mRNA expression was higher in SVF than adipocytes fraction in WAT of mice.
2) Retrovirus mediated shRNA infection of3T3-L1preadipocytes showed that knockdown Ogn could promote adipogenesis, increase lipids accumulation as well as PPARy, FABP4mRNA and protein expression.
3) Over-expression of Ogn in3T3-L1preadipocytes could not effect adipocytes differentiation. PPARy mRNA was decreased in Ogn-overexpression adipocytes but not significantly.
4) Knockdown of Ogn in3T3-L1preadipocytes could decrease basal level of glucose uptake but not insulin stimulates glucose uptake.
5) Wnt3a could inhibit adipogenesis by stabling β-catenin in cytoplasm and promoting nuclear translocation. Knockdown of Ogn in3T3-L1preadipocytes could partially block this inhibition by decreasing β-catenin nuclear translocation. Thus Ogn could regulate adipocyte differentiation through wnt/β-catenin signaling pathway.
2Prrxl Inhibits Adipogenesis by Activating TGFβ Signaling
1) Prrx1a, Prrx1b and Prrx2are down-regulated during adipogenesis in consistence with microarray data and gene expression data base. Prrx1a or Prrx1b mRNA expression is about80%lower in adipocytes fraction than that of in SVF part in WAT of mice. Prrx2is hardly detected in adipocytes in WAT of mice.
2) Knockdown of Prrxl in3T3-L1preadipocytes could promote lipids accumulation shown by Oil red O staning or BODIPY staning, stimulate PPARγ、C/EBPa and FABP4mRNA or protein expression in adipocytes.
3) Over-expression of Prrx1a or Prrx1b in3T3-L1preadipocytes can suppression Γ promoter activity, but could not effect lipids accumulation or gene expression in adipocytes.
4) Real-time qPCR results showed that Tgfb2and Tgfb3mRNA expression was decreased in Prrx1knockdown in3T3-L1preadipocytes, which suggests Prrx1may regulating TGFP signaling pathway. In deed, using SB431542, a specific inhibitor of TGFβ signaling pathway, treats3T3-L1preadipocytes could phenocopy the knockdown of Prrxl. These data suggest that Prrxl could inhibit adipogenesis by activating TGFβ signaling pathway.
5) Wnt3a and TNFα can inhibit adipogenesis dose dependently in adipocytes. However, Knockdown of Prrxl in3T3-L1preadipocytes barely blocks this effect. This indicates that Prrxl regulate dipogenesis did not affect Wnt/β-catenin and TNFa signaling pathway.
6) By Feeding BL6and C3H mice with high fat diet from3weeks to15weeks, we constructed obese mouse model. In this model, qPCR data showed that Prrxla and Prrxlb were corelated with Tgfb3mRNA expression in WAT of mice.
引文
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