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RNF220,HCA127和ASPP2在神经系统发育中的功能研究
孔清华
学位类型博士
导师毛炳宇
2009-06-30
学位授予单位中国科学院研究生院
学位授予地点北京
关键词非洲爪蟾 神经发育 泛素化 Rnf220 Hca127 Aspp2
摘要蛋白质降解对细胞生命活动的调节起着非常重要的作用,而泛素-蛋白酶体系统 (UPS)负责调控细胞内大部分蛋白的降解。UPS 与神经系统的功能有着密切的联系, 它可以控制神经的生长发育、调节突触的可塑性与功能,而其功能异常时会导致多种 神经退行性疾病。因此,在神经系统中研究泛素-蛋白酶体系统具有重要的生物学和 医学意义。 我们在非洲爪蟾大规模原位杂交中鉴定到一个在神经管腹侧特异表达的基因 xRNF220-like。同时我们也克隆了小鼠的mRNF220 和爪蟾的xRNF220 基因。它们都 在神经管的腹侧表达,且蛋白都具有E3 泛素连接酶的活性。为了寻找其降解的底物, 我们通过酵母双杂交技术进行筛选。我们发现mRNF220 可以与Sin3B 蛋白结合。进 一步实验发现Sin3B 受到UPS 的调控。而mRNF220 可以促进Sin3B 的泛素化修饰与 降解。在爪蟾胚胎中通过morpholino 阻断xRNF220-like 和Sin3B 的功能都可以导致 神经发育的异常。因此,在神经系统中mRNF220 可能通过控制Sin3B 的蛋白水平来 调节Sin3/HDAC 转录抑制复合体的活性,进而调节神经相关基因的表达。 我们还发现了一个可以与RNF220 及RNF220-like 结合的新基因HCA127。 HCA127 在进化上非常保守,在早期胚胎中特异地表达在神经管位置。在胚胎中阻断 其功能会影响神经的诱导与分化。生化实验表明HCA127 并不能被泛素化修饰。通 过对HCA127 进行酵母双杂交筛选我们发现它跟多个RING 泛素连接酶有相互作用, 还可以与一些泛素化通路中的调节蛋白结合,提示它可能是UPS 中的调节蛋白。将 其转染细胞发现可以降低细胞的泛素化水平。因此,HCA127 在神经系统中可能通过 调节蛋白的泛素化修饰来发挥功能。我们还对ASPP2 基因在早期发育中的功能进行了初步研究。ASPP2 可以促进p53 介导的细胞凋亡,并在体内抑制肿瘤的发生,但其在早期发育中的功能仍不清楚。我 们发现ASPP2 在神经系统有非常强的表达。阻断其功能可以导致眼睛、体轴和头部 的异常,并影响细胞增殖,还可以抑制神经分化标记基因N-tubulin 和神经嵴标记基 因Slug 的表达。我们初步揭示了ASPP2 在早期神经发育的功能。
其他摘要Protein degradation plays key roles in regulating cellular processes. The ubiquitin-proteasome system (UPS) is responsible for degrading most proteins. In the nervous system, the UPS has been implicated in the regulation of neuronal development, synaptic plasticity and function, and aberrations in the UPS lead to neurodegenerative diseases. Therefore, studying the roles of UPS in the nervous system is of great biological and medical significance. Through a large scale screening in Xenopus laevis by in situ hybridization, we identified the gene xRNF220-like that is expressed in ventral neural tube. The RNF220s of mouse and frog were also cloned. They were all expressed in the ventral neural tube and were proved to have E3 ubiquitin ligase activity. We performed a yeast two-hybrid screen to search for their substrates for degradation. Sin3B was identified to interact with mRNF220. Further experiments showed that Sin3B is regulated by UPS. mRNF220 could promote the ubiquitination and degradation of Sin3B. In Xenopus embryos inhibition of xRNF220-like and Sin3B by specific morpholinos led to defects in neural development. We proposed that RNF220 can regulate the activity of Sin3/HDAC suppressor complex by controlling Sin3B protein levels and thus regulate the expression of neural genes in the nervous system. We also identified a novel factor, HCA127, which interacts with RNF220 and RNF220-like. HCA127 is evolutionally conserved and expressed specifically in the nervous system. Inhibiting its function influenced neural induction and differentiation. HCA127 can’t be ubiquitinated. Through yeast two-hybrid screen we found that it interacts with a number of RING E3 ligases and several regulatory proteins of UPS, suggesting it may serves as an regulator of UPS. The ubiquitination level was reduced after transfection of HCA127 in cultured cells. We propose that HCA127 functions in the nervous system by regulating ubiquitination. The function of ASPP2 in embryonic development was studied. ASPP2 stimulates the apoptotic function of p53 and suppresses tumor formation in vivo. However its roles in early development are still obscure. We found that ASPP2 is strongly expressed in the nervous system. Loss of function led to abnormal axis, head and eye development. The mitosis was also inhibited. The expression of N-tubulin and Slug was suppressed in the morpholino injected sides. Our data suggest that ASPP2 functions in the regulation of both cell proliferation and cell differentiation in early neural development.
语种中文
文献类型学位论文
条目标识符http://ir.kiz.ac.cn/handle/152453/6359
专题科研部门_发育生物学(毛炳宇)
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孔清华. RNF220,HCA127和ASPP2在神经系统发育中的功能研究[D]. 北京. 中国科学院研究生院,2009.
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