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研究生：	毛姝涵 Mao, Su-Han
論文名稱：	小分子核糖核酸196a在細胞模式以及動物模式對神經型態的影響 Effects of MiR-196a on Neuronal Morphology in vitro and in vivo
指導教授：	楊尚訓 Yang, Shang-Hsun
學位類別：	碩士 Master
系所名稱：	醫學院 - 生理學研究所 Department of Physiology
論文出版年：	2016
畢業學年度：	104
語文別：	英文
論文頁數：	84
中文關鍵詞：	小分子核醣核酸196a 、神經細胞發育、神經型態、Ranbp10
外文關鍵詞：	miR-196a, neuronal development, neuronal morphology, Ranbp10
相關次數：	點閱：86 下載：2
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小分子核醣核酸由小片段且不會轉譯成蛋白質的核醣核酸所組成，其對基因的調控在神經細胞發育扮演很重要的腳色，神經的發育階段包含了神經軸突前端生長錐的建構、神經樹突上神經突的形成、以及神經軸突的生長。在我們實驗室先前的研究指出其中一種小分子核醣核酸｢小分子核醣核酸196a｣在細胞、基因轉殖小鼠以及從患者身上取得的誘導性多功能幹細胞等亨丁頓舞蹈症模型中，皆發現此疾病有被改善的情形，然而目前對於小分子核醣核酸196a在亨丁頓舞蹈症的神經保護機制仍是不清楚的，而在近年的研究指出維持好的神經型態是神經保護的機制之一。我們推測小分子核醣核酸196a可能透過提升神經的型態來改善亨丁頓舞蹈症因神經退化產生的症狀，在本篇研究中，我們發現小分子核醣核酸196a在細胞模式中能夠加強神經軸突的生長以及促進細胞內的順行運輸，此外，在小分子核醣核酸196a基因轉殖小鼠模式中，基因轉殖小鼠有較好的神經軸突的生長以及在海馬迴有較強的長期增益效應現象，並且有較優秀的學習記憶行為。為了更深入探討小分子核醣核酸196a的作用機制，我們利用生物資訊學找到了其中一個在血小板中與微管建構平衡有關的蛋白質Ran-binding protein 10 (Ranbp10)，我們發現小分子核醣核酸196a能夠藉由抑制Ranbp10的表現來促進beta-微管蛋白的表現及建構。相對的，大量表現Ranbp10會阻斷beta-微管蛋白的建構，並且抑制神經軸突的生長以及細胞內逆行運輸。為了能夠進一步在動物模式證實Ranbp10對神經型態的影響，我們產出高度表現Ranbp10的基因轉殖小鼠，並發現此基因小鼠有較差的神經軸突的生長趨勢。綜合我們的研究結果，我們發現小分子核醣核酸196a能夠藉由抑制Ranbp10表現來提升神經軸突的生長，而使得神經有較好的型態，也因此我們期許在未來這個發現能夠提供亨丁頓舞蹈症的治療策略方向。

MicroRNAs (miRNAs), non-protein-coding RNA molecules, play an important role at different stages of neuronal development, including growth cone organization, spine formation, and neurite outgrowth. Our previous data indicates that a specific miRNA, miR-196a, ameliorates phenotypes of Huntington’s disease (HD). However, the protective mechanism of miR-196a in HD is still unknown. Maintaining the neuronal morphology is one of the potential neuroprotective mechanisms. We hypothesize miR-196a might promote neuronal morphology and regulate resulting phenotypes. In this study, we discovered miR-196a enhanced neurite outgrowth in both neuroblastoma cell and transgenic mouse models. In addition, miR-196a promoted anterograde transport. Furthermore, miR-196a transgenic mice not only expressed stronger long-term potentiation in hippocampal CA1 but also appeared more advantageous functions of learning and memory than the non-transgenic mice. To understand the mechanisms, we used bioinformatics and identified a target gene, Ran-binding protein 10 (Ranbp10), which is related to microtubule equilibrium in platelet. We discovered that miR-196a suppressed the expression of Ranbp10 and promoted beta-tubulin organization. On the contrary, overexpression of Ranbp10 impaired the expression profiling of beta-tubulin and decreased neurite outgrowth and retrograde transport. Furthermore, we generated Ranbp10 transgenic mice and discovered the suppression of neurite outgrowth in the mice. In summary, our results suggest miR-196a improves neurite outgrowth through alteration of Ranbp10. Therefore, we anticipate this study will provide a potential therapeutic direction for HD.

Abstract.................................I
中文摘要.................................II
Acknowledgments..................................III
Contents.......................................IV
Figure Contents...................................VII
Chapter 1.   Introduction..............................1
1.1    Neuronal morphology and cytoskeleton................1
1.1.1    Neuronal polarization......................2
1.1.2    Neurite outgrowth.........................4
1.1.3    Spine formation.........................5
1.1.4    Synaptic plasticity.........................6
1.1.5    Transport............................7
1.2    The regulatory genes of neuronal morphology.............9
1.2.1    Neuronal disease-related genes.................9
1.2.2    Cytoskeleton-related gene....................10
1.3 The regulatory microRNAs of neuronal morphology.........11
1.3.1    Neurite and spine structure-related miRNAs..........12
1.3.2    Plasticity-related miRNAs...................12
1.3.3    MiR-196a and neuronal morphology..............13
Chapter 2.   Objective and specific aims......................15
2.1     Research rationales and hypothesis................15
2.2     Specific aims.............................16
Chapter 3.   Materials and methods......................17
3.1     DNA construction..........................17
3.2     DNA transformation..........................17
3.3     Plasmid DNA extraction .......................18
3.4     N2a Cell culture............................19
3.5     Primary cortical neuron culture...................19
3.6     Transfection..............................20
3.7     Microtubule polymerization.....................21
3.8     Western blotting............................22
3.9     Immunofluorescence staining.....................24
3.10    Transgenic mice...........................24
3.11    Genotyping for transgenic mice....................25
3.12    Golgi stain...............................26
3.13    Animal behavior test.........................27
3.14    Electrophysiology...........................28
3.15    Live cell imaging...........................29
3.16    Statistical analysis..........................30
Chapter 4.   Results...............................31
4.1     The effects of miR-196a on neurite outgrowth in vitro and in vivo..31
4.2     The effects of miR-196a on microtubule polymerization in vitro...32
4.3     The effects of miR-196a on intracellular transport in vitro......34
4.4     The effects of miR-196a on synaptic plasticity...........34
4.5     The effect of miR-196a on learning and memory functions......35
4.6     The suppression of Ranbp10 by miR-196a..............36
4.7     The suppression of neurite outgrowth by overexpression of Ranbp10 in vitro.................................37
4.8     The effects of Ranbp10 on microtubule polymerization........38
4.9     The suppression of intracellular transport by overexpression of Ranbp10...............................39
4.10    Production of Ranbp10 transgenic mice...............40
4.11    The suppression of neurite outgrowth by Ranbp10 in vivo.....41
Chapter 5.   Discussion..............................42
5.1     New findings from this thesis.....................42
5.2  The effects of miR-196a and Ranbp10 on microtubule polymerization ............................42
5.3     The effects of miR-196a and Ranbp10 on intracellular transport...44
5.4     The effects of miR-196a on synaptic plasticity and behavior function...............................45
5.5     The future work...........................46
Chapter 6.   Conclusion.............................48
Chapter 7.   Reference..............................49
Chapter 8.   Figures..............................61
Figure 1.  The effects of miR-196a on neurite outgrowth in primary cortical neurons...............................61
Figure 2.  The effects of miR-196a on neurite outgrowth in miR-196a 
transgenic mice...........................63
Figure 3.  The effects of miR-196a on microtubule polymerization in N2a cells
....................................65
Figure 4.  The effects of miR-196a on intracellular transport in N2a cells
....................................67
Figure 5.  The effects of miR-196a on hippocampus CA-1 LTP
....................................69
Figure 6.  The effects of miR-196a on memory function through the tests of 
T-maze........ ........................70
Figure 7.  The effects of miR-196a on learning memory function through the 
tests of novel objective recognition.................71
Figure 8.  The suppression of Ranbp10 and the promotion of -tubulin by 
miR-196a in N2a cells.........................72
Figure 9.  The suppressing effect of Ranbp10 on neurite outgrowth in N2a cells.................................74
Figure 10. The suppressing effect of Ranbp10 on neurite outgrowth in primary cortical neurons...........................76
Figure 11. The effects of Ranbp10 on microtubule polymerization in N2a cells
....................................77

Figure 12. The suppressing effect of Ranbp10 on transport in N2a cells
....................................79
Figure 13. Expression of Ranbp10 in Ranbp10 transgenic mice
....................................81
Figure 14. The effects of Ranbp10 on neurite outgrowth in Ranbp10 transgenic mice..................................82
                                    

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