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研究生:何政勳
研究生(外文):Cheng-Hsun Ho
論文名稱:探討EB病毒對第三號誘餌受體調控機制及其對鼻咽癌轉移之影響
論文名稱(外文):Identification of Epstein-Barr Virus-mediated Decoy Receptor 3 upregulation and its effect on nasopharyngeal carcinoma metastasis
指導教授:陳紀如
指導教授(外文):Chi-Ju Chen
學位類別:博士
校院名稱:國立陽明大學
系所名稱:微生物及免疫學研究所
學門:生命科學學門
學類:微生物學類
論文種類:學術論文
畢業學年度:97
語文別:中文
論文頁數:137
中文關鍵詞:第三號誘導受體鼻咽癌轉移
外文關鍵詞:Decoy receptor 3DcR3Epstein-Barr virusEBVnasopharyngeal carcinomaNPCmetastasis
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EB病毒屬於人類疱疹病毒,其感染已發現和許多惡性腫瘤的生成有相關性,然而EB病毒如何逃避人類免疫系統的監控及促使腫瘤生成仍不清楚。第三號誘餌受體 (DcR3) 是一種分泌型的腫瘤壞死因子受體,可在一些惡性腫瘤的組織中發現到其有相當高的表現量。第三號誘餌受體已知道可藉著抑制細胞凋亡及擾亂宿主免疫系統來幫助腫瘤細胞的生存。先前的報導發現到第三號誘餌受體在EB病毒相關的淋巴瘤細胞中有大量表現,但在非EB病毒相關的淋巴瘤細胞表現量甚少,暗示著EB病毒調控第三號誘餌受體的可能性。
測試不同的EB病毒基因後,我們發現到EB病毒潛伏期穿膜蛋白一號 (LMP1)及溶裂期轉錄活化子R (Rta) 分別在EB病毒潛伏期及溶裂期扮演著正調控第三號誘餌受體表現的角色。在沒有EB病毒的細胞中,大量表現潛伏期穿膜蛋白一號或溶裂期轉錄活化子R可以刺激第三號誘餌受體表現。而第三號誘餌受體的表現量在有EB病毒感染的細胞中也隨著潛伏期穿膜蛋白一號或溶裂期轉錄活化子R的表現量多寡而有正相關性。進一步地分析發現到在潛伏期穿膜蛋白一號或溶裂期轉錄活化子R的蛋白質C端活化區域對於第三號誘餌受體的調控是相當重要的。調控機制上,溶裂期轉錄活化子R主要是藉由結合到第三號誘餌受體的啟動子上並增強其活性,而潛伏期穿膜蛋白一號則是透過傳遞多種訊息路徑來調控第三號誘餌受體的表現。在功能探討方面,我們發現第三號誘餌受體可以增進鼻咽癌細胞移動及侵入能力,暗示著第三號誘餌受體對鼻咽癌轉移的重要性。最後,在鼻咽癌檢體中我們偵測到潛伏期穿膜蛋白一號及第三號誘餌受體的共同表現。
綜合上述結果,我們推測EB病毒可藉著潛伏期穿膜蛋白一號或溶裂期轉錄活化子R來活化第三號誘餌受體表現,進而促進鼻咽癌細胞的轉移。
Epstein-Barr virus (EBV) belongs to human γ-herpesvirus and its infection is known to be associated with various malignancies. However, how EBV escapes host immune surveillance and promotes tumorigenesis are not well elucidated. Decoy receptor 3 (DcR3) is a soluble tumor necrosis factor receptor found to be overexpressed in several tumor types and its overexpression has been implicated in tumor cells surviving through inhibiting apoptosis and interfering immunity. A previous study reported that amplification of DcR3 is detected in EBV-associated lymphomas but rare in non-EBV associated lymphomas, suggesting the role of EBV in DcR3 upregulation. After screening serial EBV genes, we found EBV latent membrane protein 1 (LMP1) and lytic transactivator Rta are responsible for upregulating DcR3 in EBV latent and lytic stages, respectively. Overexpression of Rta or LMP1 into EBV-negative cell lines enhanced DcR3 expression in a dose-dependent manner. Additionally, the amounts of Rta or LMP1 was correlated with DcR3 expression levels in EBV-infected cell lines. Further dissection revealed that C-terminal activation domain of Rta or LMP1 was essential for DcR3 stimulation. Rta induced DcR3 expression mainly through binding to Rta-responsive element (RRE) on DcR3 promoter, while LMP1-mediated DcR3 elicitation was via triggering diverse signaling pathways. Functionally, DcR3 was shown to enhance cell motility and invasiveness in nasopharyngeal carcinoma (NPC) cells, implicating the importance of DcR3 on promoting tumor metastasis. Finally, we showed the co-expression of DcR3 and LMP1 in NPC biopsies. Taken together, EBV may enhance DcR3 expression by LMP1 or Rta, which contributes to NPC metastasis.
中文摘要……………………………………………………………………...……...1
Abstract………………………………………………………………………..……..2
Introduction……………………………………………………………………….....3
1. Epstein-Barr virus (EBV)……………………………………………………….3
1.1 Characteristics of EBV………………………………………………….......3
1.2 Prospected route of EBV infection………………………………………….3
1.3 Life cycle of EBV…………………………………………………………...4
1.4 Latent membrane protein 1 (LMP1)………………………………………...7
1.5 Transactivator R (Rta)…………………………………………………….....9
1.6 EBV-associated diseases…………………………………………………...11
2. Decoy Receptor 3 (DcR3)…………………………………………………...…12
2.1 Discovery and characteristics of DcR3…………………………………….12
2.2 Expression profiles of DcR3……………………………………………….13
2.3 Functions of DcR3…………………………………………………………14
2.3.1 Decoy receptor………………………………………………………...14
2.3.2 Immune modulator…………………………………………………….16
2.4 Endogenous and exogenous regulation of DcR3…………………………..17
2.5. Correlation between EBV infection and DcR3 overexpression…………..18
3. Epithelial-mesenchymal transition (EMT)…………………………………….19
3.1 Discovery and characteristics of EMT……………………………….…….19
3.2 Regulation of EMT………………………………………………………...19
3.3 EMT and tumor metastasis………………………………………………...21
3.4 EMT and NPC metastasis…………………………………………….……22
Objectives…………………………………………………………………………...24
Material & Methods…………………………………………………….…………26
1. Bacteria culture………………...........................................................................26
2. Bacteria transformation………………………………………………………...26
3. Cell lines and culture…………………………………………………………..26
4. Plasmids and siRNA duplex…………………………………………………...27
5. Antibodies and reagents………………………………………………………..28
6. Purification of plasmid DNA…………………………………………………..29
7. Transient transfection…………………………………………………………..30
8. Purification of human PBMCs and primary B cells…………...........................31
9. Purification of EBV virion……………………………………………………..32
10. Western blot……..……………………………………………………………33
11. Enzyme-linked immunosorbent assay (ELISA)……………………………...34
12. Total RNA purification………………………………………………………..34
13. Reverse transcriptional polymerase chain reaction (RT-PCR)……………….35
14. Reporter assay………………………………………………………………...36
15. Isolation of nuclei…………………………………………………………….36
16. Electrophoretic mobility shift assay (EMSA)………………………………...36
17. Chromatin immunoprecipitation (ChIP)……………………………………...37
18. Immunohistochemistry (IHC) staining……………………………………….39
19. Generation of EBV-infected cell lines………………………………………..39
20. Generation of HONE-1-DcR3 stable lines…………………………………...40
21. Transwell migration assay and Matrigel-invasion assay……………………..40
22. Statistic analysis………………………………………………………………41
Results……………………………………………………………………………....42
1. EBV infection increases DcR3 expression…………………………………….42
2. DcR3 overexpression is detected in EBV type III latency…………………….42
3. LMP1 mainly regulates DcR3 expression during EBV latency……………….43
4. LMP1 alone induces DcR3 expression………………………………………...44
5. Both CTAR1 and CTAR2 of LMP1 are responsible for DcR3 stimulation……44
6. LMP1 enhances DcR3 expression via NF�羠 and PI3-K signaling pathways…45
7. Co-expression of LMP1 and DcR3 in NPC biopsies…………………………..47
8. Pro-inflammatory cytokines also induces DcR3 expression…………………..48
9. DcR3 expression is increased upon EBV reactivation……...…………………49
10. Rta, but not Zta, is responsible for DcR3 upregulation……...……………….50
11. Rta induces DcR3 expression in EBV-latent LCLs…………………………..50
12. C-terminal region of Rta is important for DcR3 activation………..................51
13. Rta binds RRE on putative DcR3 promoter………………………………......52
14. Rta induces DcR3 expression mainly through binding pDcR3-RRE……...…53
15. PI3-K signaling plays a minor role in Rta-mediated DcR3 upregulation….....54
16. CBP co-operates with Rta to induce DcR3 expression….....…………………56
17. Higher DcR3 expression in metastatic NPC than in primary NPC…………..57
18. DcR3 enhances LMP1-mediated cellular migration and invasion…………...57
19. DcR3-Fc increases NPC cells migration and invasion……………………….58
20. Significantly enhancement of cellular metastatic potential in DcR3-
stably expressed NPC cells…………………………………………………...59
21. DcR3 induced EMT of NPC cells is occurred under serum-deprivation……..60
Discussion…………………………………………………………………………..62
Reference…………………………………………………………………...………71
Figures…………………………………………………………………………..…..90
Tables………………………………………………………………………………129
Appendix…………………………………………………………………………..131
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