Roos Wynand, Baumgartner Manuela, Kaina Bernd
Division of Applied Toxicology, Institute of Toxicology, Obere Zahlbacher Str. 67, D-55131 Mainz, Germany.
Oncogene. 2004 Jan 15;23(2):359-67. doi: 10.1038/sj.onc.1207080.
Various tumor-therapeutic drugs and environmental carcinogens alkylate DNA inducing O(6)-methylguanine (O(6)MeG) that provokes cell death by apoptosis. In rodent fibroblasts, apoptosis triggered by O(6)MeG is executed via the mitochondrial damage pathway. Conversion of O(6)MeG into critical downstream lesions requires mismatch repair (MMR). This is thought to signal apoptosis upon binding to O(6)MeG lesions mispaired with thymine. Alternatively, O(6)MeG lesions might be processed by MMR giving rise to DNA double-strand breaks (DSBs) during replication that finally provoke apoptosis. To test this, we examined apoptosis triggered by O(6)MeG in human peripheral lymphocytes in which O(6)-methylguanine-DNA methyltransferase (MGMT) had been inactivated by O(6)-benzylguanine (O(6)BG) and which were not proliferating or proliferating upon CD3/CD28 stimulation. Treatment with N-methyl-N'-nitro-N-nitrosoguanidine (MNNG) or the anticancer drug temozolomide induced apoptosis only in proliferating, but not resting cells. With exceptional high alkylation doses (>/=15 microM of MNNG), apoptosis was also observed in resting lymphocytes, albeit at a lower level than in proliferating cells. This response was not affected by O(6)BG, suggesting that replication-independent apoptosis at high dose levels is caused by lesions other than O(6)MeG. O(6)MeG-triggered apoptosis in proliferating lymphocytes was preceded by a wave of DSBs, which coincided with p53 and Fas receptor upregulation, while Fas ligand, Bax and Bcl-2 expression was not altered. Treatment with anti-Fas neutralizing antibody attenuated MNNG-induced apoptosis in MGMT-depleted proliferating lymphocytes. The data suggest that O(6)MeG is converted by MMR and DNA replication into DSBs that trigger apoptosis by p53 stabilization and Fas/CD95/Apo-1 upregulation. This is supported by the finding that ionizing radiation, inducing DSBs on its own, provokes apoptosis in lymphocytes in a replication-independent way. The strict proliferation dependence of apoptosis triggered by O(6)MeG may explain the specific killing response of MGMT-deficient proliferating cells, including tumors, to O(6)MeG generating anticancer drugs and suggests that tumor proliferation rate, Fas responsiveness, MGMT and MMR status are important prognosis parameters.
多种肿瘤治疗药物和环境致癌物会使DNA发生烷基化,诱导产生O(6)-甲基鸟嘌呤(O(6)MeG),进而通过凋亡引发细胞死亡。在啮齿动物成纤维细胞中,由O(6)MeG触发的凋亡是通过线粒体损伤途径执行的。将O(6)MeG转化为关键的下游损伤需要错配修复(MMR)。这被认为在与胸腺嘧啶错配的O(6)MeG损伤结合时发出凋亡信号。或者,O(6)MeG损伤可能由MMR处理,在复制过程中产生DNA双链断裂(DSB),最终引发凋亡。为了验证这一点,我们检测了在人外周淋巴细胞中由O(6)MeG触发的凋亡,其中O(6)-甲基鸟嘌呤-DNA甲基转移酶(MGMT)已被O(6)-苄基鸟嘌呤(O(6)BG)灭活,这些细胞在CD3/CD28刺激下不增殖或增殖。用N-甲基-N'-硝基-N-亚硝基胍(MNNG)或抗癌药物替莫唑胺处理仅在增殖细胞中诱导凋亡,而在静止细胞中不诱导。在异常高的烷基化剂量(≥15μM的MNNG)下,在静止淋巴细胞中也观察到凋亡,尽管水平低于增殖细胞。这种反应不受O(6)BG的影响,表明高剂量水平下与复制无关的凋亡是由O(6)MeG以外的损伤引起的。在增殖淋巴细胞中,O(6)MeG触发的凋亡之前有一波DSB,这与p53和Fas受体上调同时发生,而Fas配体、Bax和Bcl-2表达未改变。用抗Fas中和抗体处理可减弱MNNG诱导的MGMT缺失的增殖淋巴细胞中的凋亡。数据表明,O(6)MeG通过MMR和DNA复制转化为DSB,通过p53稳定和Fas/CD95/Apo-1上调触发凋亡。这得到了以下发现的支持:电离辐射自身诱导DSB,以与复制无关的方式在淋巴细胞中引发凋亡。O(6)MeG触发的凋亡对增殖的严格依赖性可能解释了MGMT缺陷的增殖细胞(包括肿瘤细胞)对产生O(6)MeG的抗癌药物的特异性杀伤反应,并表明肿瘤增殖率、Fas反应性、MGMT和MMR状态是重要的预后参数。