Pontarin Giovanna, Ferraro Paola, Valentino Maria L, Hirano Michio, Reichard Peter, Bianchi Vera
Department of Biology, University of Padova, I-35131 Padova, Italy.
J Biol Chem. 2006 Aug 11;281(32):22720-8. doi: 10.1074/jbc.M604498200. Epub 2006 Jun 13.
Mitochondrial (mt) neurogastrointestinal encephalomyopathy (MNGIE) is an autosomal recessive disease associated with depletion, deletions, and point mutations of mtDNA. Patients lack a functional thymidine phosphorylase and their plasma contains high concentrations of thymidine and deoxyuridine; elevation of the corresponding triphosphates probably impairs normal mtDNA replication and repair. To study metabolic events leading to MNGIE we used as model systems skin and lung fibroblasts cultured in the presence of thymidine and/or deoxyuridine at concentrations close to those in the plasma of the patients, a more than 100-fold excess relative to controls. The two deoxynucleosides increased the mt and cytosolic dTTP pools of skin fibroblasts almost 2-fold in cycling cells and 8-fold in quiescent cells. During up to a two-month incubation of quiescent fibroblasts with thymidine (but not with deoxyuridine), mtDNA decreased to approximately 50% without showing deletions or point mutations. When we removed thymidine, but maintained the quiescent state, mtDNA recovered rapidly. With thymidine in the medium, the dTTP pool of quiescent cells turned over rapidly at a rate depending on the concentration of thymidine, due to increased degradation and resynthesis of dTMP in a substrate (=futile) cycle between thymidine kinase and 5'-deoxyribonucleotidase. The cycle limited the expansion of the dTTP pool at the expense of ATP hydrolysis. We propose that the substrate cycle represents a regulatory mechanism to protect cells from harmful increases of dTTP. Thus MNGIE patients may increase their consumption of ATP to counteract an unlimited expansion of the dTTP pool caused by circulating thymidine.
线粒体神经胃肠性脑肌病(MNGIE)是一种常染色体隐性疾病,与线粒体DNA(mtDNA)的缺失、耗竭及点突变相关。患者缺乏功能性胸苷磷酸化酶,其血浆中胸苷和脱氧尿苷浓度很高;相应三磷酸酯水平的升高可能会损害正常的mtDNA复制和修复。为了研究导致MNGIE的代谢事件,我们使用在接近患者血浆浓度(比对照组高100多倍)的胸苷和/或脱氧尿苷存在下培养的皮肤和肺成纤维细胞作为模型系统。这两种脱氧核苷使皮肤成纤维细胞的线粒体和胞质dTTP库在增殖细胞中增加近2倍,在静止细胞中增加8倍。在静止成纤维细胞与胸苷(而非脱氧尿苷)长达两个月的孵育过程中,mtDNA降至约50%,且未出现缺失或点突变。当我们去除胸苷但维持静止状态时,mtDNA迅速恢复。培养基中存在胸苷时,静止细胞的dTTP库以取决于胸苷浓度的速率快速周转,这是由于胸苷激酶和5'-脱氧核糖核苷酸酶之间底物(即无效)循环中dTMP降解和再合成增加所致。该循环以ATP水解为代价限制了dTTP库的扩张。我们提出底物循环是一种保护细胞免受dTTP有害增加影响的调节机制。因此,MNGIE患者可能会增加ATP消耗,以抵消循环胸苷导致的dTTP库无限制扩张。