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系統識別號 U0026-1410201123500500
論文名稱(中文) 研究干擾素r所調控之annexin A2表現及其生理角色
論文名稱(英文) Study on the regulation of annexin A2 expression by IFN-r and its physiological roles
校院名稱 成功大學
系所名稱(中) 基礎醫學研究所
系所名稱(英) Institute of Basic Medical Sciences
學年度 100
學期 1
出版年 100
研究生(中文) 方宜婷
研究生(英文) Yi-Ting Fang
學號 s5894155
學位類別 博士
語文別 英文
論文頁數 139頁
口試委員 指導教授-林以行
召集委員-湯銘哲
口試委員-黎煥耀
口試委員-陳舜華
口試委員-胡承波
口試委員-賴明宗
中文關鍵字 干擾素r  annexin A2  p11  外泌體  自噬體  吞噬作用  肺上皮細胞 
英文關鍵字 IFN-r  annexin A2  p11  exosome  autophagosome  phagocytosis  lung epithelial cell 
學科別分類
中文摘要 Annexin A2 (p36) 能夠與S100A10 (p11) 形成heterotetramer之結構,而稱為annexin A2 complex,藉由鈣離子協助而黏附於細胞膜表面。Annexin A2具有多樣的生理角色,其功能根據座落在細胞中不同位置而有所不同。Annexin A2被發現富含於內皮細胞、肺部與腸道上皮細胞、以及一些癌細胞中。已有研究指出,annexin A2的表現以及移動與發炎反應的調控相當有關連,然而,經由發炎性細胞激素調控annexin A2表現之詳細機制目前尚未清楚。在抗磷脂質抗體症候群以及肺癌病人中已鑑定annexin A2為自體抗原,本實驗室過去針對SARS病毒感染的病人血清,偵測到抗SARS冠狀病毒棘蛋白區位二(S2)的抗體亦可交叉反應於人類肺上皮細胞中。利用蛋白質體分析技術檢定annexin A2為候選自體抗原之一,我們利用SARS病人血清及抗棘蛋白(S2)抗體再確認annexin A2為其交叉反應的抗原蛋白。進一步觀察到發炎性細胞激素IL-6以及IFN-可增加annexin A2表現於細胞表面,並且促進annexin A2分佈至與脂筏相關的區域,並且加強了抗棘蛋白抗體交叉反應於肺上皮細胞之能力。然而細胞在IFN-刺激下造成annexin A2外送至肺上皮細胞表面的機制仍然不清楚。目前已觀察到IFN-刺激可促使annexin A2以及p11於細胞表面的表現,並且兩者蛋白質會於細胞表面座落在一起。而IFN-刺激可經由JAK2/STAT1的訊息傳遞路徑而促進p11表現量的增加,然而annexin A2的蛋白質表現量不受刺激影響。值得注意的是,IFN-誘導的p11表現量上升對於後續的annexin A2外送細胞表面是不可或缺的,並且,p11所參與調控的annexin A2外送可能和外泌體的運送路徑有關。進一步地觀察到IFN-刺激所引發的自噬體生成和IFN-刺激導致annexin A2外送機制有關。在探討細胞表面的annexin A2其生理功能的部分,實驗結果顯示IFN-的刺激可增強肺上皮細胞吞噬凋亡細胞的能力,同時IFN-所促使annexin A2外送至細胞表面,將調節此吞噬凋亡細胞的能力。總結而言,此研究提供更多瞭解在IFN-刺激之下annexin A2外送的機制,及說明了它在細胞外扮演吞噬的重要角色。此研究中發炎反應下對於annexin A2表現的作用及其影響,亦可涉及在SARS疾病中所引發的自體免疫反應。
英文摘要 Annexin A2 (p36) is a phospholipid-binding, calcium-regulated, peripheral membrane protein that forms a heterotetramer with S100A10 (p11), called annexin A2 complex. Annexin A2 proteins have multiple biological functions depending on its cellular localization. Annexin A2 is widely expressed in many cell types, such as vascular endothelial cells, lung and intestine epithelial cells, and some tumor cells. Inflammation closely related with annexin A2 expression and relocation has been reported. However, the detailed mechanism of regulation of annexin A2 by inflammatory cytokine is still unclear. Annexin A2 has been identified as an autoantigen in patients of anti-phospholipid syndrome and lung cancer. In our previous study of SARS-CoV infection, we have demonstrated that antibodies against SARS-CoV spike protein domain 2 (S2) in SARS patient sera cross-react with human lung epithelial cells. The candidate autoantigen annexin A2 was identified by proteomics studies. We confirmed that SARS patient sera and anti-S2 antibodies both cross-react with annexin A2. We further observe an increase in surface expression and raft-associated distribution of annexin A2 after A549 epithelial cells are stimulated with inflammatory cytokines, interleukin-6 and interferon (IFN)-. The cytokine stimulation also enhances the binding capability of anti-S2 antibodies to human lung epithelial cells. However, the detailed mechanism of surface translocation of annexin A2 on lung epithelial cells after cytokine stimulation remains unclear. We have observed that both annexin A2 and p11 are increased and colocalized with each other on the surface of lung epithelial cells after IFN- stimulation. While total annexin A2 protein remains unchanged after IFN- treatment, the expression of p11 is upregulated via the IFN--activated JAK2/STAT1 signal pathway. Notably, IFN--induced p11 upregulation is required for annexin A2 translocation to the cell surface. We observe that p11-dependent surface expression of annexin A2 is associated with the exosomal secretion pathway. Furthermore, IFN--induced autophagosome formation is associated with extracellular secretion and surface expression of annexin A2. We further clarify the physiological function of surface annexin A2 in IFN--treated A549 cells. We observe that IFN- stimulation can enhance the phagocytic ability of lung epithelial cells to clear apoptotic T cells. The IFN--induced surface expression of annexin A2 mediates phagocytosis of apoptotic cells by lung epithelial cells. Taken together, this study provides insights into the surface translocation mechanism of annexin A2 and illustrates a pivotal function of surface annexin A2 in the phagocytic response to IFN-. It also has an implication for IFN--regulated annexin A2 with inflammation-associated SARS immunopathogenesis.
論文目次 English Abstract I
Chinese Abstract III
誌謝 IV
Contents VI
Table List X
Figure List XI
Abbreviations XIV
Chapter 1-Introduction 1
I. The annexins superfamily 2
II. Annexin A2 3
II.1. Ca2+-dependent and -independent phospholipid binding 4
II.2. p11 (annexin A2 natural ligand) 5
II.3. Multiple functions of annexin A2 6
II.3.1. Intracellular annexin A2 7
II.3.2. Extracellualr annexin A2 9
II.4. Regulation of annexin A2 expression 11
II.5. Surface annexin A2 and human disease 12
III. Severe acute respiratory syndrome (SARS) 14
III.1. SARS immunophathogenesis 14
III.2. Interferon (IFN) and SARS 17
IV. Inflammation 18
IV.1. Inflammatory cytokine and autoimmunity 18
IV.2. Inflammatory cytokine IFN-18
V. Unconventional secretion pathway 20
V.1. Unconventional secretion pathway 21
V.2. Biogenesis of exosome 21
V.3. Autophagy 22
VI. Phagocytosis of apoptotic cells 23
VI.1. Phagocytosis of apoptotic cells (Efferocytosis) 23
VI.2. Annexin and phagocytosis 24
Chapter 2-Specific aims 26
Chapter 3-Materials and methods 31
I. Materials 32
I.1. Cell lines 32
I.2. Patient sera 32
I.3. Drugs 32
I.4. Recombinant SARS S2 protein and anti-S2 preparation 34
I.5. Kits 36
I.6. Antibodies 36
I.7. Vectors 37
I.8. Consumables 37
I.9. Instruments 38
II. Methods 39
II.1. Cell cultures 39
II.2. Cell treatment 39
II.3. Cell cytosol and membrane preparation 39
II.4. 2-DGE and LC-MS/MS analysis 39
II.5. Sequence database search 41
II.6. Protein binding assay 41
II.7. Cell surface elution 42
II.8. LDH assay 42
II.9. Cell biotinylation 43
II.10. Exosome isolation 43
II.11. Plasmid transfection 43
II.12. Short interfering RNA (siRNA) treatment 44
II.13. Generation of lentivirus shRNA and infection of A549 cells 44
II.14. Western blotting 45
II.15. Flow cytometry analysis 46
II.16. Immunofluorescence 46
II.17. Phagocytosis assay 47
II.18. Statistics 47
Chapter 4-Results 48
I. To investigate whether IFN-induces surface expression of autoantigen
annexin A2 on human lung epithelial cells 49
II. To clarify the mechanism of surface translocation of annexin A2 by IFN-
regulation 52
III. To analyze the effect of autophagy on the surface translocation of annexin
A2 via exosome secretion pathway by IFN- 57
IV. To study the physiological effect of annexin A2 surface expression by
IFN-stimulation 58
Chapter 5-Discussion 61
Part I 62
I.1. Autoimmune response in SARS-CoV infection 62
I.2. Annexin A2 and autoimmunity 63
Part II 64
II.1. Regulation of protein expression of p11 and annexin A2 64
II.2. Exosome secretion pathway and annexin A2 65
II.3. The hypothetic mechanism of annexin A2 escaping from exosome to
plasma membrane 66
II.4. Post-translational modification and surface translocation of annexin A2 67
II.5. Other regulated pathway for IFN--induced surface translocation of
annexin A2 67
Part III 68
III.1. Autophagy and unconventional exocytosis of annexin A2 68
III.2. Ubiquitination of annexin A2 and selective autophagy 69
Part IV 69
IV.1. Role of lung epithelial cells in clearance of apoptotic cells 69
IV.2. IFN--enhanced phagocytosis of apoptotic cells 70
IV.3. SARS and phagocytosis of apoptotic cells 70
IV.4. Role of annexin A2 in phagocytosis of apoptotic cells 71
References 72
Tables 94
Figures and Figure legends 96
Appendix 133
Publications 136
Curriculum Vitae 137
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