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研究生:詹月瑄
研究生(外文):Yueh-Hsuan Chan
論文名稱:轉錄抑制因子B細胞誘發成熟因子1於樹突細胞發育及功能之調控
論文名稱(外文):A Role for the Transcriptional Repressor Blimp-1 in Dendritic Cell Homeostatic Development and Function
指導教授:林國儀
指導教授(外文):Kuo-I Lin
學位類別:博士
校院名稱:國防醫學院
系所名稱:生命科學研究所
學門:生命科學學門
學類:生物學類
論文種類:學術論文
論文出版年:2009
畢業學年度:98
語文別:英文
論文頁數:115
中文關鍵詞:樹突細胞轉錄抑制因子B 細胞誘發成熟因子 1
外文關鍵詞:dendritic celltranscriptional repressorBlimp-1
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樹突細胞 (Dendritic cell) 對於免疫反應的誘發及調控是相當重要的。但目前對於調控樹突細胞發育的分子機制卻不甚清楚。在這篇論文裡,我們證明一個對於漿細胞分化和 T 淋巴球的恆定及功能相當重要的轉錄抑制因子,B 細胞誘發成熟分子 1 (Blimp-1),會抑制樹突細胞發育的平衡。在小鼠的造血系統中剔除 B 細胞誘發成熟因子 1 會導致平衡狀態下的傳統樹突細胞 (conventional DC) 及漿細胞樣樹突細胞 (plasmacytoid DC) 增加。剔除 B 細胞誘發成熟因子 1會專一地導致 CD8 陰性傳統樹突細胞的立即前軀細胞增加,進而造成 CD8 陰性傳統樹突細胞的增加。骨髓樹突細胞在接受刺激成熟後會透過核因子 B (NF-B) 和 p38 裂質素活化蛋白激酶 (p38 MAPK) 的路徑使 B 細胞誘發成熟分子 1 的表現量增加,此外,樹突細胞在體內和體外的成熟都是需要 B 細胞誘發成熟分子 1。B 細胞誘發成熟分子 1 會直接抑制促發炎細胞激素/ 趨化激素基因,例如細胞白介素第六因子 (Il-6) 和單核細胞趨化蛋白 1 (Ccl2)。在樹突細胞株中大量表現 B 細胞誘發成熟分子 1,並利用基因晶片的分析方法發現 165 個基因會被抑制、而 74 個基因會被誘發。此外,B 細胞誘發成熟分子 1 在漿細胞樣樹突細胞對於雙股核糖核酸和非甲基化去氧核糖核酸所產生的干擾素 (interferon) 是不可或缺的。總而言之,我們發現 B 細胞誘發成熟分子 1 對於樹突細胞發育的平衡扮演負向的調控,但對樹突細胞的成熟卻是正向的調控因子。
Dendritic cells (DCs) are important for the initiation and regulation of immune responses. However, the molecular mechanisms that govern DC development are largely unclear. In this thesis, we show that the transcriptional repressor, B lymphocyte-induced maturation protein 1 (Blimp-1), which is critical for regulation of plasma cell differentiation and T cell homeostasis and function, negatively regulates DC homeostatic development. Deletion of Prdm1, the gene encoding Blimp-1, in mouse hematopoietic lineages resulted in an increase in the steady-state number of conventional DCs (cDCs) and plasmacytoid DCs (pDCs). Specifically, Prdm1 deletion increased immediate CD8– cDC precursors in peripheral lymphoid organs, causing selective expansion of the CD8– cDC population. Blimp-1 is up-regulated in bone marrow-derived DCs via the p38 MAPK and NF-B pathways upon stimulus-induced maturation, and is required for DC maturation in vitro and in vivo. Blimp-1 directly suppresses the proinflammatory cytokine/chemokine genes, Il-6 and Ccl2. Using Affymetrix microarray analysis, we identified 165 genes are down-regulated, whereas 74 genes are up-regulated by enforced Blimp-1 expression in a murine DC line. Moreover, Blimp-1 is indispensable for type I interferon production in response to double-strand RNA and unmethylated DNA in pDCs. Taken together, our findings reveal that Blimp-1 acts as a negative regulator for select DC subsets homeostasis and a positive regulator during DC maturation.
CONTENT
Chapter 1. Introduction 1
1.1. Dendritic cells (DCs) 1
1.2. Conventional DCs 1
1.3. Plasmacytoid DCs 3
1.4. The Origin of steady-state DCs 4
1.5. Differentiation of DCs from hematopoietic progenitors 5
1.6. Transcriptional regulation of DC development 7
1.7. B lymphocyte-induced maturation protein 1 (Blimp-1) 9
1.8. Blimp-1 in B cell terminal differentiation 10
1.9. Blimp-1 in T cell biology 11
1.10. Blimp-1 in myeloid lineage differentiation 12
1.11. Aims of this thesis 13
Chapter 2. Materials and Methods 15
2.1. Mice 15
2.2. Cell lines 15
2.3. Preparation of cells from lymphoid organs 16
2.3.1. Splenic B cells cultures 16
2.3.2. Splenic DC cultures 16
2.3.3. Bone marrow-derived DC (BMDC) cultures 17
2.4. Antibodies and Flow cytometry 18
2.5. Splenic immediate DC precursor analysis 19
2.6. Mixed lymphocyte reaction (MLR) 19
2.7. In vivo DC maturation 20
2.8. In vivo BrdU labeling 20
2.9. Lentivirus preparation and transduction 20
2.10. Cytometric bead array (CBA) and ELISA 21
2.11. RNA isolation and Real time PCR 21
2.12. Nuclear and cytosolic extract preparation 23
2.13. Western blotting 24
2.14. Immunofluorescence staining 25
2.15. DNA pull-down assay 25
2.16. Electrophoretic Mobility Shift Assay (EMSA) 26
2.17. Chromatin immunoprecipitation (ChIP) assay 27
2.18. Luciferase Assay 28
2.19. Microarray analysis 29
2.20. Statistics 30
Chapter 3. Results 31
3.1. Deletion of Prdm1, gene encoding Blimp-1, in hematopoietic lineage 31
3.2. Blimp-1 negatively regulates CD8– cDC development in spleen 32
3.3. Deficiency of Blimp-1 leads to increased CD8– cDC immediate precursors 33
3.4. Stimulus-induced Blimp-1 expression during DC maturation is mediated by NF-kB and p38 MAPK pathways 35
3.5. Blimp-1 is required for DC maturation 36
3.6. Il-6 and Ccl2 are suppressed directly by Blimp-1 38
3.7. Microarray analysis of Blimp-1 targets in a DC line 41
3.8. Blimp-1 negatively regulates pDC homeostatic development 45
3.9. Blimp-1 regulates gene expression profile of pDCs 46
3.10. Blimp-1 is indispensable for type I IFN production 47
Chapter 4. Discussion 48
Chapter 5. Conclusion 56
Chpater 6. References 58
TABLE CONTENT
Table 1. List of immune response-associated genes significantly altered in JAWS II cells after Blimp-1 transduction. 81
Table. 2 List of genes involved in DC migration that were significantly changed in JAWS II cells after Blimp-1 transduction. 82
Table 3. List of cell death, cell proliferation, and cell stress-associated genes significantly altered in JAWS II cells after Blimp-1 transduction. 83
Table 4. List of transcription factors significantly altered in JAWS II cells after Blimp-1 transduction. 84
Table 5. List of signaling molecules significantly changed in JAWS II cells after Blimp-1 transduction. 85
Table 6. List of metabolic genes significantly changed in JAWS II cells after Blimp-1 transduction. 86
FIGURE CONTENT
Figure 1. The generation of CKO mice. 87
Figure 2. Phenotype of Ctrl and CKO B cells. 88
Figure 3. Phenotype of Ctrl and CKO T cells. 89
Figure 4. Blimp-1 negatively regulates homeostasis of CD11chi cDCs. 90
Figure 5. Blimp-1 negatively regulates CD8– cDC homeostasis. 91
Figure 6. Bone marrow DC progenitors in Ctrl and CKO mice. 92
Figure 7. Increased percentage of CD8– cDC immediate precursors in CKO spleens. 93
Figure 8. The expression of Blimp-1 mRNA in various splenic cDC subsets. 94
Figure 9. Transcripts of IRF family transcription factors in Ctrl and CKO splenic cDC subsets. 95
Figure 10. Ctrl and CKO splenic cDC subsets have similar proliferation and survival rates. 96
Figure 11. Blimp-1 expression is increased upon maturation. 97
Figure 12. Blimp-1 induction is mediated by NF-kB and p38 MAPK pathways. 98
Figure 13. Impaired DC maturation in CKO mice. 99
Figure 14. Defects in CKO DC maturation are not due to reduced expression of TLRs or impaired TLR-mediated signaling. 100
Figure 15. Impaired T cell proliferation elicited by CKO BMDCs. 101
Figure 16. CKO cDCs are refractory to maturation in vivo. 102
Figure 17. Increased IL-6 and MCP-1 production in LPS-stimulated CKO BMDC culture. 103
Figure 18. IL-6 and MCP-1 transcripts are increased in CKO DC cultures. 104
Figure 19. Blimp-1 binds to Il-6 and Ccl2 sites in vivo. 105
Figure 20. Blimp-1 binds to Il-6 and Ccl2 sites in vitro. 106
Figure 21. Endogenous Blimp-1 binds to Il-6 and Ccl2 sites in vitro. 107
Figure 22. Ectopic Blimp-1 suppresses the endogenous expression and secretion of IL-6 and MCP-1 in JAWS II cells. 108
Figure 23. Blimp-1 suppresses promoter activity of Il-6 and Ccl2. 109
Figure 24. Hierarchical clustering of genes subjected to changes by Blimp-1 expression in JAWS II cells. 110
Figure 25. Confirmation of selected microarray results by RT-QPCR. 111
Figure 26. Blimp-1 expresses in pDCs and is required for maintaining pDC homeostasis. 112
Figure 27. In vitro pDC development of Ctrl and CKO bone marrow progenitors. 113
Figure 28. Blimp-1 regulates genes involved in B cell and pDC development or function. 114
Figure 29. IFNa and IFNb production by TLR ligands-stimulated Ctrl or CKO splenic pDCs. 115

APPENDIX CONTENT
Appendix 1. Qualification for Ph. D. Candidate.
Appendix 2. Progress report.
Appendix 3. Published paper.
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