Curr Opin Neurobiol:REST/NRSF在神经退行性疾病中的调节作用

2018-07-10 Ella 大医编

研究者已发现NRSF/REST含有9个锌指结构域,在其氨基端及羧基端均可以募集相关蛋白形成复合体从而引起靶基因转录的表观修饰沉默。

NRSF即是神经元限制性沉默因子(neuron-restrictive silence factor),又称为REST(抑制元件1-沉默转录因子,repressor element-1-silencing transcription factor),可结合到DNA的神经元限制性沉默元件(neuron-restrictive silence element, NRSE)上,抑制含有NRSE元件的神经元特异性基因在非神经元中的转录。研究者已发现NRSF/REST含有9个锌指结构域,在其氨基端及羧基端均可以募集相关蛋白形成复合体从而引起靶基因转录的表观修饰沉默。

REST多在非神经细胞、胚胎干细胞和神经干细胞中高表达,以维持神经元特异性基因的转录抑制状态及干细胞的“干性”;在神经组细胞、神经元前体细胞、神经元中,其含量逐渐降低,相应的靶基因转录水平则在整体上表现为逐渐升高的趋势。NRSF的下调是神经元及胶质细胞分化形成的基础,受到NRSF/REST转录调控的基因包括编码众多功能蛋白(如离子通道、神经递质及合成酶、突触囊泡蛋白、细胞黏附份子、激素、神经营养因子、细胞骨架蛋白、细胞外基质蛋白,等等)的mRNA,也包括shRNAs和长链ncRNAs。

越来越多的研究证实,NRSF/REST是维持正常脑功能所必需的因子,其异常表达和分布不仅参与了缺血性中风导致的神经细胞死亡进程,也与神经退行性病、癫痫等的病理机制有密切关联。研究发现,由缺血导致的神经元REST的升高抑制了包括GluR2、mu阿片受体、NCX1以及miR-132等的基因表达,而下调REST的水平则可挽救受损神经元免于死亡。在阿尔茨海默患者及转基因小鼠模型中,研究人员均发现有脑内NRSF/REST的含量升高,进一步的研究表明,AD的早期阶段出现核内NRSF/REST的下降,而在亨廷顿病中,神经元细胞核内则有NRSF/REST的聚集。在帕金森病中,MPP+可引起SH-SY5Y细胞中核REST的表达及转位,而在条件性敲除NRSF/REST的小鼠中,MPTP导致的黑质TH神经元丢失明显加剧。在癫痫的发生发展中,也发现脑内NRSF/REST水平的升高以及一系列靶基因转录的下调。因此,NRSF/REST的变化都与脑功能的改变及神经元的存亡息息相关。

因此,小编将向大家介绍一下这个神奇的转录因子在神经退行性病变中的作用。

RET与神经退行性疾病

REST调节异常与神经系统疾病有关。在胚胎发生期间REST表达的改变引起细胞凋亡,分化异常。在分化的神经元中,REST是静止的,但是可被缺血和癫痫等神经元损伤激活,在这些疾病中,REST和REST依赖的表观遗传重塑的增加与神经元死亡有因果关系。神经元损伤的引起的REST表达增加不仅会抑制突触后受体和转运蛋白,还会抑制突触前囊泡蛋白如突触蛋白I,突触素,突触结合蛋白II,IV,VI和VII,突触小泡蛋白 II和SNARE蛋白 SNAP25。因此,不仅成体神经元中的神经传递依赖于极低的REST表达水平,而且PC12细胞和胰腺β细胞中的刺激诱导的胞吐作用同样也依赖于极低的REST表达水平。相反,在老化的神经元,REST的低水平和REST的损失与相关的老年痴呆症发病有密切关联。同时,REST和其靶基因的失调还参与的癫痫,亨廷顿氏病,帕金森氏病和SMCX,X连锁的形式智力低下等发病过程。

REST和缺血

最近的研究结果表明REST表达在成熟的分化神经元中被激活以响应神经元损伤,并且REST依赖的靶基因沉默对于神经元死亡是必需的。在大鼠全脑缺血模型中,观察到有CK1和β- TRCP的减少和REST蛋白质丰度的增加,在CA1 锥体细胞中层,REST被募集至靶基因的一个子集,将其与近端启动子的辅阻遏物复合物结合,编排重塑。这些发现均提示神经变性疾病中的表观遗传失调。而HDAC抑制剂 TSA对缺血诱导的神经元死亡有实质性的保护作用,表明表观遗传重塑和缺血诱导的神经元死亡之间的因果关系。在活体动物的大脑中表达的针对REST或显性阴性REST的shRNA的表达在临床相关的全局缺血模型中提供了对CA1神经元的强有力保护。这些发现提供了强有力的证据,即海马神经元兴奋性神经元中REST的激活与神经元死亡偶然相关。REST在缺血性神经元死亡中的类似作用已在短暂性大脑中动脉闭塞(tMCAO)中得到证实。

CHIP-CHIP分析和生物信息学分析表明,在缺血后神经元中,REST协调了“转录反应”靶基因集合的沉默,包括编码AMPA受体亚基GluA2,NMDAR亚基NR1,神经元乙酰胆碱受体亚基的那些基因。 β2,毒蕈碱乙酰胆碱受体M4,NF-κB亚基2, TRPV1和SNARE蛋白突触结合蛋白6,其中编码GluA2的基因被高度排名。这一点很重要,因为GluA2亚基是AMPARs的“离子门控器”。而缺乏GluA2的AMPAR可渗透Ca2+和Zn 2+并且表现出明显的内向整流,GluA2在异聚AMPA受体中的存在使得通道对Ca 2+和Zn 2+不可渗透并且是线性的。GluA2的存在还影响通道动力学,电导和靶向突触位点的靶向。这一点同样很重要,因为Ca 2+可渗透的AMPA受体与缺血诱导的神经元死亡有随机相关性。

REST和阿尔茨海默病

Yankner及其同事发表的一篇具有里程碑意义的论文报道,REST在衰老大脑中具有神经保护作用。作者表明,由Wnt信号通路激活的REST表达是正常衰老的一个突出特征,REST的丢失与轻度或严重的认知障碍和AD有关。对来自正常老化人类受试者的尸检组织的检查显示 ,在具有轻度或严重认知障碍AD的受试者中,前额皮质和海马 CA1和CA3 的神经元的细胞核中的REST显着增加。SH-SY5Y细胞中的ChIP-seq实验显示REST的占用主要是参与神经元死亡的靶点(p38 MAPK,FAS,FADD ,TRADD ,BAX,BID,BBC3(也称为PUMA),线粒体通透性转换孔蛋白和细胞色素c)和AD病理学(γ-分泌酶) ,早老素 2,早老素增强子-2和CDK5R1)。因此,具有减少的REST的衰老小鼠表现出增强的对氧化应激的易感性。与此相一致的是,提高REST表达的REST的错义变体可预防与阿尔茨海默病和神经变性相关的海马萎缩。患有轻度认知障碍的患者。REST在老化的大脑中提供的神经保护超越AD,因为患有额颞痴呆和路易体痴呆的受试者的组织中的REST也被耗尽。此外,在黑质神经元中升高的REST表达在帕金森病的动物模型中是保护性的。这些研究结果表明老年神经元中的REST表达赋予神经保护作用,并将REST作为轻度和重度认知障碍AD的潜在治疗靶点。

总之,过去十年见证了新发现,这些发现共同指出了REST在神经退行性疾病中的调节作用。虽然REST与缺血后成熟神经元的神经元死亡有因果关系,但在衰老的神经元中,它可提供神经保护作用。

值得进一步探索的新兴概念是由REST调节的靶基因集合以细胞类型依赖性和上下依赖性方式变化。这些研究结果将REST作为正常和病理条件下神经元基因的多方面调节器。但是,虽然目前的研究在理解REST在生理和病理条件下如何在神经元中发挥作用方面取得了重大进展,但许多问题仍未得到解决。例如,装配TET3和NSD3是REST激活基因转录的唯一机制,还是其他蛋白质与REST组装以促进基因表达?是NMDA受体和氯离子转运的唯一目标基因通过REST依赖调节后生在产后发育的关键时期进行重塑,还是还有其他与突触可塑性相关的基因,这些基因在发育的这个时期由REST调节?还有尚未发现的小分子抑制剂或REST激活剂可能为开发新的治疗策略打开大门,以改善与神经退行性疾病和疾病相关的神经元死亡和受损认知吗?在老化神经元,剩下的由参与抑制基因,得到的神经保护的细胞凋亡和氧化应激包括许多Bcl2家族成员。还有其他任何环境可以让REST抑制这个基因集合吗?

总之,许多未得到解答的问题表明,多年来关于REST和神经元基因表观遗传重塑的富有成效的研究正在进行中。

原始出处:Hwang J, Zukin RS. Rest, a master transcriptional regulator in neurodegenerative disease. Curr Opin Neurobiol. 2018;48:193-200

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level=null, likeNumber=71, replyNumber=0, topicName=null, topicId=null, topicList=[], attachment=null, authenticateStatus=null, createdAvatar=https://cdnapi.center.medsci.cn/medsci/head/2018/04/12/cd30eefc465f70dc4dad89bd57479781.jpg, createdBy=02be2318568, createdName=一天没事干, createdTime=Tue Jul 10 06:38:47 CST 2018, time=2018-07-10, status=1, ipAttribution=)]
    2019-06-04 sunylz
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  4. 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  5. 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level=null, likeNumber=71, replyNumber=0, topicName=null, topicId=null, topicList=[], attachment=null, authenticateStatus=null, createdAvatar=https://cdnapi.center.medsci.cn/medsci/head/2018/04/12/cd30eefc465f70dc4dad89bd57479781.jpg, createdBy=02be2318568, createdName=一天没事干, createdTime=Tue Jul 10 06:38:47 CST 2018, time=2018-07-10, status=1, ipAttribution=)]
    2018-07-12 jihuaijun1112

    学习学习学习

    0

  6. 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  7. 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    2018-07-10 1ddf0692m34(暂无匿称)

    学习了.长知识

    0

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level=null, likeNumber=71, replyNumber=0, topicName=null, topicId=null, topicList=[], attachment=null, authenticateStatus=null, createdAvatar=https://cdnapi.center.medsci.cn/medsci/head/2018/04/12/cd30eefc465f70dc4dad89bd57479781.jpg, createdBy=02be2318568, createdName=一天没事干, createdTime=Tue Jul 10 06:38:47 CST 2018, time=2018-07-10, status=1, ipAttribution=)]
    2018-07-10 龙胆草

    学习谢谢分享

    0

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    2018-07-10 zb1235672

    学习了!!!!!

    0

  10. 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    2018-07-10 一天没事干

    很好的学习机会

    0

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