Pikkarainen Sampsa, Kennedy Robert A, Marshall Andrew K, Tham El Li, Lay Kenneth, Kriz Thomas A, Handa Balvinder S, Clerk Angela, Sugden Peter H
National Heart and Lung Institute Division, Faculty of Medicine, Imperial College London, London SW7 2AZ, United Kingdom.
J Biol Chem. 2009 Oct 2;284(40):27195-210. doi: 10.1074/jbc.M109.037887. Epub 2009 Jul 28.
The Mdm2 ubiquitin ligase is an important regulator of p53 abundance and p53-dependent apoptosis. Mdm2 expression is frequently regulated by a p53 Mdm2 autoregulatory loop whereby p53 stimulates Mdm2 expression and hence its own degradation. Although extensively studied in cell lines, relatively little is known about Mdm2 expression in heart where oxidative stress (exacerbated during ischemia-reperfusion) is an important pro-apoptotic stimulus. We demonstrate that Mdm2 transcript and protein expression are induced by oxidative stress (0.2 mm H(2)O(2)) in neonatal rat cardiac myocytes. In other cells, constitutive Mdm2 expression is regulated by the P1 promoter (5' to exon 1), with inducible expression regulated by the P2 promoter (in intron 1). In myocytes, H(2)O(2) increased Mdm2 expression from the P2 promoter, which contains two p53-response elements (REs), one AP-1 RE, and two Ets REs. H(2)O(2) did not detectably increase expression of p53 mRNA or protein but did increase expression of several AP-1 transcription factors. H(2)O(2) increased binding of AP-1 proteins (c-Jun, JunB, JunD, c-Fos, FosB, and Fra-1) to an Mdm2 AP-1 oligodeoxynucleotide probe, and chromatin immunoprecipitation assays showed it increased binding of c-Jun or JunB to the P2 AP-1 RE. Finally, antisense oligonucleotide-mediated reduction of H(2)O(2)-induced Mdm2 expression increased caspase 3 activation. Thus, increased Mdm2 expression is associated with transactivation at the P2 AP-1 RE (rather than the p53 or Ets REs), and Mdm2 induction potentially represents a cardioprotective response to oxidative stress.
Mdm2泛素连接酶是p53丰度和p53依赖性凋亡的重要调节因子。Mdm2表达常常受p53-Mdm2自调节环调控,即p53刺激Mdm2表达,进而导致其自身降解。尽管在细胞系中已得到广泛研究,但对于心脏中Mdm2的表达了解相对较少,而氧化应激(在缺血再灌注期间加剧)是心脏中一种重要的促凋亡刺激因素。我们证明,氧化应激(0.2 mM H₂O₂)可诱导新生大鼠心肌细胞中Mdm2转录本和蛋白表达。在其他细胞中,组成型Mdm2表达受P1启动子(外显子1上游5'端)调控,诱导型表达受P2启动子(内含子1中)调控。在心肌细胞中,H₂O₂增加了来自P2启动子的Mdm2表达,该启动子包含两个p53反应元件(REs)、一个AP-1 RE和两个Ets REs。H₂O₂未显著增加p53 mRNA或蛋白的表达,但确实增加了几种AP-1转录因子的表达。H₂O₂增加了AP-1蛋白(c-Jun、JunB、JunD、c-Fos、FosB和Fra-1)与Mdm2 AP-1寡脱氧核苷酸探针的结合,染色质免疫沉淀分析表明,它增加了c-Jun或JunB与P2 AP-1 RE的结合。最后,反义寡核苷酸介导的H₂O₂诱导的Mdm2表达降低增加了半胱天冬酶3的激活。因此,Mdm2表达增加与P2 AP-1 RE(而非p53或Ets REs)的反式激活相关,Mdm2的诱导可能代表了对氧化应激的一种心脏保护反应。