Leaner Virna D, Donninger Howard, Ellis Chad A, Clark Geoffrey J, Birrer Michael J
National Cancer Institute, 9610 Medical Center Dr., Room 300, Rockville, MD 20850-3300, USA.
Mol Cell Biol. 2005 Apr;25(8):3324-37. doi: 10.1128/MCB.25.8.3324-3337.2005.
The c-Jun/AP-1 transcription complex is associated with diverse cellular processes such as differentiation, proliferation, transformation, and apoptosis. These different biological endpoints are likely achieved by the regulation of specific target gene expression. We describe the identification of Ras guanine nucleotide exchange factor 1, Ras-GRF1, by microarray analysis as a c-Jun/AP-1 regulated gene essential for anchorage-independent growth of immortalized rat fibroblasts. Increased Ras-GRF1 expression, in response to inducible c-Jun expression in Rat1a fibroblasts, was confirmed by both real-time PCR and Northern blot analysis. We show that c-Jun/AP-1 can bind and activate the Ras-GRF1 promoter in vivo. A 75-kDa c-Jun/AP-1-inducible protein, p75-Ras-GRF1, was detected, and the inhibition of its expression with antisense oligomers significantly blocked c-Jun-regulated anchorage-independent cell growth. p75-Ras-GRF1 expression occurred with a concomitant increase in activated Ras (GTP bound), and the activation of Ras was significantly inhibited by antisense Ras-GRF1 oligomers. Moreover, p75-Ras-GRF1 could be coprecipitated with a Ras dominant-negative glutathione S-transferase (GST) construct, GST-Ras15A, demonstrating an interaction between p75-Ras-GRF1 and Ras. A downstream target of Ras activation, Elk-1, had increased transcriptional activity in c-Jun-expressing cells, and this activation was inhibited by dominant-negative Ras. In addition, c-Jun overexpression resulted in an increase in phospho-AKT while phosphorylation of ERK1/2 remained largely unaffected. The inhibition of phosphatidylinositol 3-kinase (PI3K)-AKT signal transduction by Ly294002 and wortmannin significantly blocked c-Jun-regulated morphological transformation, while inhibition of basal MEK-ERK activity with PD98059 and U0126 had little effect. We conclude that c-Jun/AP-1 regulates endogenous p75-Ras-GRF1 expression and that c-Jun/AP-1-regulated anchorage-independent cell growth requires activation of Ras-PI3K-AKT signal transduction.
c-Jun/AP-1转录复合体与多种细胞过程相关,如分化、增殖、转化和凋亡。这些不同的生物学终点可能是通过调控特定靶基因的表达来实现的。我们通过微阵列分析鉴定出Ras鸟嘌呤核苷酸交换因子1(Ras-GRF1)是c-Jun/AP-1调控的基因,对永生化大鼠成纤维细胞的锚定非依赖性生长至关重要。通过实时PCR和Northern印迹分析证实,在Rat1a成纤维细胞中,响应于可诱导的c-Jun表达,Ras-GRF1表达增加。我们表明,c-Jun/AP-1在体内可结合并激活Ras-GRF1启动子。检测到一种75 kDa的c-Jun/AP-1诱导蛋白p75-Ras-GRF1,用反义寡聚体抑制其表达可显著阻断c-Jun调控的锚定非依赖性细胞生长。p75-Ras-GRF1表达伴随着活化Ras(结合GTP)的增加,反义Ras-GRF1寡聚体可显著抑制Ras的活化。此外,p75-Ras-GRF1可与Ras显性负性谷胱甘肽S-转移酶(GST)构建体GST-Ras15A共沉淀,表明p75-Ras-GRF1与Ras之间存在相互作用。Ras激活的下游靶点Elk-1在表达c-Jun的细胞中转录活性增加,这种激活被显性负性Ras抑制。此外,c-Jun过表达导致磷酸化AKT增加,而ERK1/2的磷酸化基本不受影响。Ly294002和渥曼青霉素对磷脂酰肌醇3-激酶(PI3K)-AKT信号转导的抑制显著阻断了c-Jun调控的形态转化,而用PD98059和U0126抑制基础MEK-ERK活性的作用不大。我们得出结论,c-Jun/AP-1调节内源性p75-Ras-GRF1表达,且c-Jun/AP-1调控的锚定非依赖性细胞生长需要Ras-PI3K-AKT信号转导的激活。