Koyama Tatsuya, Nakaoka Yoshikazu, Fujio Yasushi, Hirota Hisao, Nishida Keigo, Sugiyama Shoko, Okamoto Kitaro, Yamauchi-Takihara Keiko, Yoshimura Michihiro, Mochizuki Seibu, Hori Masatsugu, Hirano Toshio, Mochizuki Naoki
Department of Structural Analysis, National Cardiovascular Center Research Institute, 5-7-1 Fujishirodai, Suita, Osaka, Japan.
J Biol Chem. 2008 Aug 29;283(35):24234-44. doi: 10.1074/jbc.M803907200. Epub 2008 Jun 23.
Grb2-associated binder 1 (Gab1) coordinates various receptor tyrosine kinase signaling pathways. Although skeletal muscle differentiation is regulated by some growth factors, it remains elusive whether Gab1 coordinates myogenic signals. Here, we examined the molecular mechanism of insulin-like growth factor-I (IGF-I)-mediated myogenic differentiation, focusing on Gab1 and its downstream signaling. Gab1 underwent tyrosine phosphorylation and subsequent complex formation with protein-tyrosine phosphatase SHP2 upon IGF-I stimulation in C2C12 myoblasts. On the other hand, Gab1 constitutively associated with phosphatidylinositol 3-kinase regulatory subunit p85. To delineate the role of Gab1 in IGF-I-dependent signaling, we examined the effect of adenovirus-mediated forced expression of wild-type Gab1 (Gab1(WT)), mutated Gab1 that is unable to bind SHP2 (Gab1(DeltaSHP2)), or mutated Gab1 that is unable to bind p85 (Gab1(Deltap85)), on the differentiation of C2C12 myoblasts. IGF-I-induced myogenic differentiation was enhanced in myoblasts overexpressing Gab1(DeltaSHP2), but inhibited in those overexpressing either Gab1(WT) or Gab1(Deltap85). Conversely, IGF-I-induced extracellular signal-regulated kinase 1/2 (ERK1/2) activation was significantly repressed in myoblasts overexpressing Gab1(DeltaSHP2) but enhanced in those overexpressing either Gab1(WT) or Gab1(Deltap85). Furthermore, small interference RNA-mediated Gab1 knockdown enhanced myogenic differentiation. Overexpression of catalytic-inactive SHP2 modulated IGF-I-induced myogenic differentiation and ERK1/2 activation similarly to that of Gab1(DeltaSHP2), suggesting that Gab1-SHP2 complex inhibits IGF-I-dependent myogenesis through ERK1/2. Consistently, the blockade of ERK1/2 pathway reversed the inhibitory effect of Gab1(WT) overexpression on myogenic differentiation, and constitutive activation of the ERK1/2 pathway suppressed the enhanced myogenic differentiation by overexpression of Gab1(DeltaSHP2). Collectively, these data suggest that the Gab1-SHP2-ERK1/2 signaling pathway comprises an inhibitory axis for IGF-I-dependent myogenic differentiation.
Grb2相关结合蛋白1(Gab1)协调多种受体酪氨酸激酶信号通路。尽管骨骼肌分化受某些生长因子调控,但Gab1是否协调肌源性信号仍不清楚。在此,我们研究了胰岛素样生长因子-I(IGF-I)介导的肌源性分化的分子机制,重点关注Gab1及其下游信号传导。在C2C12成肌细胞中,IGF-I刺激后,Gab1发生酪氨酸磷酸化,并随后与蛋白酪氨酸磷酸酶SHP2形成复合物。另一方面,Gab1与磷脂酰肌醇3激酶调节亚基p85组成性结合。为了阐明Gab1在IGF-I依赖性信号传导中的作用,我们研究了腺病毒介导的野生型Gab1(Gab1(WT))、无法结合SHP2的突变型Gab1(Gab1(DeltaSHP2))或无法结合p85的突变型Gab1(Gab1(Deltap85))的强制表达对C2C12成肌细胞分化的影响。在过表达Gab1(DeltaSHP2)的成肌细胞中,IGF-I诱导的肌源性分化增强,但在过表达Gab1(WT)或Gab1(Deltap85)的成肌细胞中受到抑制。相反,在过表达Gab1(DeltaSHP2)的成肌细胞中,IGF-I诱导的细胞外信号调节激酶1/2(ERK1/2)激活被显著抑制,但在过表达Gab1(WT)或Gab1(Deltap85)的成肌细胞中增强。此外,小干扰RNA介导的Gab1敲低增强了肌源性分化。催化失活的SHP2的过表达对IGF-I诱导的肌源性分化和ERK1/2激活的调节与Gab1(DeltaSHP2)相似,表明Gab1-SHP2复合物通过ERK1/2抑制IGF-I依赖性肌生成。一致地,ERK1/2通路的阻断逆转了Gab1(WT)过表达对肌源性分化的抑制作用,并通过Gab1(DeltaSHP2)的过表达抑制了ERK1/2通路的组成性激活对增强的肌源性分化的抑制。总体而言,这些数据表明Gab1-SHP2-ERK1/2信号通路构成了IGF-I依赖性肌源性分化的抑制轴。