Feng Weiyi, Rosca Mariana, Fan Yuxuan, Hu Yufen, Feng Pingfu, Lee Hyoung-Gon, Monnier Vincent M, Fan Xingjun
First Affiliated Hospital, School of Medicine, Xi'an Jiaotong University, Xi'an, Shaanxi, P.R. China.
College of Medicine, Central Michigan University, Mount Pleasant, MI 48859, USA.
Hum Mol Genet. 2017 Apr 1;26(7):1376-1390. doi: 10.1093/hmg/ddx040.
Gamma glutamyl cysteine ligase (GCL) is the rate-limiting enzyme for intracellular glutathione (GSH) synthesis. The GSH concentration and GCL activity are declining with age in the central nervous system (CNS), and is accompanied by elevated reactive oxygen species (ROS). To study the biological effects of low GSH levels, we disrupted its synthesis both at birth by breeding a Gclc loxP mouse with a thy1-cre mouse (NEGSKO mouse) and at a later age by breeding with a CaMKII-ERT2-Cre (FIGSKO mouse). NEGSKO mice with deficiency of the Gclc in their entire CNS neuronal cells develop at 4 weeks: progressive motor neuron loss, gait problems, muscle denervation and atrophy, paralysis, and have diminished life expectancy. The observed neurodegeneration in Gclc deficiency is of more chronic rather than acute nature as demonstrated by Gclc targeted single-neuron labeling from the inducible Cre-mediated knockout (SLICK) mice. FIGSKO mice with inducible Gclc deficiency in the forebrain at 23 weeks after tamoxifen induction demonstrate profound brain atrophy, elevated astrogliosis and neurodegeneration, particularly in the hippocampus region. FIGSKO mice also develop cognitive abnormalities, i.e. learning impairment and nesting behaviors based on passive avoidance, T-Maze, and nesting behavior tests. Mechanistic studies show that impaired mitochondrial glutathione homeostasis and subsequent mitochondrial dysfunction are responsible for neuronal cell loss. This was confirmed by mitochondrial electron transporter chain activity analysis and transmission electron microscopy that demonstrate remarkable impairment of state 3 respiratory activity, impaired complex IV function, and mitochondrial swollen morphology in the hippocampus and cerebral cortex. These mouse genetic tools of oxidative stress open new insights into potential pharmacological control of apoptotic signaling pathways triggered by mitochondrial dysfunction.
γ-谷氨酰半胱氨酸连接酶(GCL)是细胞内谷胱甘肽(GSH)合成的限速酶。在中枢神经系统(CNS)中,GSH浓度和GCL活性会随年龄增长而下降,并伴有活性氧(ROS)水平升高。为了研究低GSH水平的生物学效应,我们通过将Gclc loxP小鼠与thy1-cre小鼠杂交(NEGSKO小鼠)在出生时破坏其合成,并在稍后通过与CaMKII-ERT2-Cre杂交(FIGSKO小鼠)来破坏其合成。在整个CNS神经元细胞中缺乏Gclc的NEGSKO小鼠在4周时出现:进行性运动神经元丧失、步态问题、肌肉去神经支配和萎缩、瘫痪,并且预期寿命缩短。如通过诱导型Cre介导的基因敲除(SLICK)小鼠的Gclc靶向单神经元标记所证明的,Gclc缺乏中观察到的神经退行性变具有更慢性而非急性的性质。在他莫昔芬诱导后23周时,在前脑具有诱导型Gclc缺乏的FIGSKO小鼠表现出严重的脑萎缩、星形胶质细胞增生和神经退行性变增加,特别是在海马区。FIGSKO小鼠还出现认知异常,即基于被动回避、T迷宫和筑巢行为测试的学习障碍和筑巢行为。机制研究表明,线粒体谷胱甘肽稳态受损及随后的线粒体功能障碍是神经元细胞丢失的原因。线粒体电子传递链活性分析和透射电子显微镜证实了这一点,其显示海马和大脑皮质中状态3呼吸活性显著受损、复合体IV功能受损以及线粒体肿胀形态。这些氧化应激的小鼠遗传工具为线粒体功能障碍触发的凋亡信号通路的潜在药理学控制开辟了新的见解。