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研究生(外文):Ting-Yu Kuo
論文名稱(外文):The functional study of Bcl11A in neuronal morphogenesis
指導教授(外文):Yi-Ping Hsueh
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在目前已知的研究中,鋅指轉錄因子Bcl11A (或稱CTIP1)於B細胞發育過程中扮演了一個相當重要的角色。然而,除了在B細胞中所提供的功能以外,Bcl11A於腦組織中也有大量的表現,雖然目前為止對於它在腦組織裡的功能並不是非常清楚。在本篇論文中,我首次檢視了Bcl11A於鼠腦中各部區域以及細胞層級上分佈的情形。Bcl11A這一個基因,可利用選擇性剪接訊息RNA的方式來達到同ㄧ基因多種蛋白質表現的效果,而Bcl11A-S和Bcl11A-L便是兩種在鼠腦中較為多量表現的蛋白質。利用RNA干擾子的技術可以很精確地抑制Bcl11A-L蛋白質的表現量,藉由此方式,我發現Bcl11A-L本身具有調節神經細胞軸突分枝、生長甚至於促成多軸突表現的能力。而在本篇中,Bcl11A-S被發現可以干擾Bcl11A-L的聚合,達到降低Bcl11A-L調節基因表現的能力。在Bcl11A-L轉錄基因的功能上,在本篇研究發現DCC和MAP1b是兩個屬於Bcl11A-L下游所調控的基因。上述兩個基因對於Bcl11A-L的功能傳遞上是重要的,當人為地大量表現DCC時,便可除去Bcl11A-L RNA干擾子的作用。另外,關於神經細胞活性如何去調控神經突的分枝、生長的部份,本篇論文也提供了部份可能的機制。透過細胞膜上神經傳導物質受器所傳遞而來的神經活性,可以去影響Bcl11A-L蛋白質的表現量,進一步也影響了下游DCC以及MAP1b的蛋白質表現量。經由這個方式,神經活性的增加便可以造成神經細胞軸突形態上的改變,如同再使用干擾子技術所達到的效果一樣。本篇研究提供了關於Bcl11A-L在神經發育上的一些新的功能,以及神經活性如何調控神經網路的可能機制。
Bcl11A/CTIP1, a kruppel-like zinc finger transcription factor, plays an important role in B cell development. In addition to B lymphocytes, Bcl11A is also highly expressed in brain, although the function in brain is unclear. In this thesis, I first examined the regional and subcellular distribution of Bcl11A in rat brain. Using RNAi knockdown approach, I found that Bcl11A-L, one of Bcl11A isoform generated by alternative splicing, can control the neurite branching, outgrowth, and even promote multiple axon formation in cultured hippocampal neurons. Moreover, Bcl11A-S, another isoform of Bcl11A, could function as dominate negative molecule to disrupt the Bcl11A-L oligomerization and function in neuronal morphogenesis. Candidate search was then performed to identify the downstream effectors of Bcl11A-L. Both protein and RNA levels of DCC and MAP1b were regulated by Bcl11A-L. Overexpression of DCC in neurons can neutralize the Bcl11A-L knockdown effects. These data support that Bcl11A-L regulates expression of DCC and MAP1b and thus control axon branching and neurite outgrowth. Finally, the data of this thesis further show that NMDAR activation can regulate the proteins levels of Bcl11A-L, DCC and MAP1b. By regulating these molecule expressions, it provides a new mechanism why neuronal activity can regulate the axon branching, projection or turning during early development. In conclusion, my study suggests a new function of Bcl11A in neuronal morphogenesis and an important role of Bcl11A in NMDAR regulated axonal projection.
1. The MAGUK protein CASK is important for synaptogenesis and brain development 3
2. The CaMK-like and L27 domain of CASK 5
3. The PDZ domain of CASK 6
4. The SH3 domain and protein 4.1 binding motif of CASK 8
5. The GK domain of CASK 10
6. The zinc finger transcription factor Bcl11A is essential for lymphoid development 12
7. The function of BcL6 in B-cell development 13
8. The Bcl11A function in human fetal hemoglobin switching 15
9. The functional domains and isoforms of Bcl11A 17
10. The COUP-TF orphan nuclear receptors in brain development 18
11. The guidance molecule for axonal arborization 21
12. The cytoskeleton rearrangement in axonal arborization 23
13. Neuronal activity regulates neurite outgrowth, turning and branching

1. Bacteria culture 36
2. Bacterial transformation 36
3. Preparation of DNA 36
4. RNA preparation and RT-PCR 37
5. PCR program 37
6. Electrophoresis 38
7. Western Blotting 38
8. Biochemical fractionation of rat brain 39
9. Immunoprecipitation 40
10. Immunocytochemistry 41
11. Immunohistochemistry 42
12. Induction and purification of recombinant proteins 42
13. Antibody purification 43
14. Immobilizaiton of protein to solid support 44
15. Primary neuronal culture 45
16. Lentivirus production 46
17. Time-lapse imaging 46
18. Analysis of Neuronal Morphology 47


1. Characterization of Bcl11A antibodies 48
2. Distribution of Bcl11A in brain 49
3. Confocal analysis of Bcl11A distribution in neurons 49
4. Subcellular distribution of Bcl11A identified by biochemical fractionation 50
5. Interaction between Bcl11A-S and Bcl11A-L 51
6. Knockdown of Bcl11A upregulates neurite outgrowth 52
7. Downregulation of Bcl11A-L increase the arbor complexity of axons and dendrites 53
8. Overexpression of Bcl11A-L reduces axon and dendrite outgrowth 54
9. Time-lapse study of the effect of Bcl11A-L on axon outgrowth 55
10. Overexpression of oligomerization domain of Bcl11A-L mimics the effect of Bcl11A knockdown
11. DCC and MAP1b are downregulated by knockdown of Bcl11A-L 57
12. Overexpression of DCC rescues the axon phenotype induced by Bcl11A-L knockdown
13. Neuronal activity through NMDAR upregulates axon branching & dendrite outgrowth
14. NMDA treatment down regulates the neuronal morphogenesis related genes 61
15. Bcl11A-L overexpression prevents multiple axon and axonal branching induced by NMDA treatment
16. DCC overexpression neutralizes the effect of glutamate on axon arborization 62

1. The Bcl11A splicing forms have different subcellular distribution in neuron 63
2. The Bcl11A-L functions in brain development 63
3. Transcriptional regulation by Bcl11A-L in neuron 64
4. The function of Bcl11A-S in regulation of Bcl11A-L transcriptional activity 65
5. The cross-talk between neuronal activity and guidance molecules 65
6. Bcl11A-L and axonal regeneration 66
7. The function of notch and Bcl11A-L in early neuronal development 67
8. Regulation of Bcl11A-L by phosphorylation and CASK interaction 68

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