探討第一型小窩蛋白在細菌與脂多醣及半乳糖氨所引發之炎症反應中所扮演的角色與分子機制

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研究生：	蔡宗晃 Tsai, Tsung-Huang
論文名稱：	探討第一型小窩蛋白在細菌與脂多醣及半乳糖氨所引發之炎症反應中所扮演的角色與分子機制 The molecular mechanism of caveolin -1 in bacteria- and lipopolysaccharide/D-galactosamine-induced inflammatory response
指導教授：	江安世 Chiang, Ann-Shyn 徐松錕 Shyue, Song-Kun
口試委員:
學位類別：	博士 Doctor
系所名稱：	生命科學暨醫學院 - 生物科技研究所 Biotechnology
論文出版年：	2010
畢業學年度：	98
語文別：	英文
論文頁數：	90
中文關鍵詞：	第一型小窩蛋白、巨噬細胞、吞噬能力、細菌清除力、發炎反應、急性肝衰竭
外文關鍵詞：	Caveolin-1, Macrophage, phagocytosis, bacterial killing, immune response, acute liver injury
相關次數：	點閱：2 下載：0
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Part I

An overwhelming immune response, particularly from macrophages, with Gram-negative bacteria-induced sepsis plays a critical role in survival of and organ damage in infected patients. Caveolin-1 (Cav-1) is essential and the major protein of caveolae structure, which is responsible for proteins trafficking and regulating of enzymatic activities of many interacting proteins. Cav-1 level is induced during the differentiation of monocyte to macrophage. However, how Cav-1 regulates macrophage activity remains unclear. In this study, we examined the vital role of Cav-1 in the response of macrophage and mice to bacteria and LPS exposure. We found that deletion of Cav-1 decreased the expression of CD14 and CD36 during macrophage differentiation and suppressed their phagocytotic ability. Administration of CD36 and CD14 antiserums synergistically inhibited bacterial engulfment by macrophages, suggesting that the suppressed phagocytosis in Cav-1-/- macrophages is due to the downregulation of CD36 and CD14. As well, the ability to kill bacteria was inhibited in Cav-1-/- macrophages and mice peritoneal cavity, tissue and plasma, which was partly attributed to hindered expression of inducible nitric oxide synthase (iNOS) induced by bacteria or LPS. Furthermore, deletion of Cav-1 attenuated the expression of Toll-like receptor 4 (TLR4) and myeloid differentiation factor 88 (MyD88) and the activation of nuclear factor-□B (NF-kB), all of which impeded the production of inflammatory cytokines in response to bacterial exposure in Cav-1-/- macrophages and mice. In addition, disruption of the lipid raft by cholesterol depletion reduced bacterial phagocytosis and clearance, and iNOS expression. Thus, Cav-1 participates in the regulation of CD14, CD36, TLR4 and MyD88 protein expression and is crucial for the immune response of macrophages to bacterial infection. Cav-1 may be a therapeutic target in the treatment of sepsis.

Part II

Acute liver injury, a major contributor to disease-related mortality, often occurs as a consequence of inflammatory disorders such as nonalcoholic and alcoholic hepatitis, sepsis, and drug-induced liver injury. Bacterial lipopolysaccharide (LPS) is implicated in the pathogenesis of acute liver injury and several chronic inflammatory liver diseases. To investigate the role of Cav-1 in LPS-induced acute liver injury, we test a standard experimental method of LPS/GaLN-induced acute liver injury by injection of low dose of LPS in combination with D-galactosamine (GalN). LPS/GalN treatment increased hepatic Cav-1 mRNA accumulation at 1.5 to 4 h in WT mice. Deletion of Cav-1 markedly reduced mortality, serum aspartate aminotransferase and alanine aminotransferase levels, and less liver tissue injury including parenchymal hemorrhage, neutrophilic infiltration, and hepatic apoptosis and necrosis as compared with WT mice. Western blot and IHC staining of the liver tissues showed that LPS/GalN treatment for 5 h induced higher cleavaged caspase-3, caspase-8 in WT mice than in Cav-1-/- mice. Increased hepatic expression of cytokine, chemokine, and iNOS in WT mice suggest that up-regulation of cytokine, chemokine, and iNOS may represent a potential mechanism for Cav-1-mediated exacerbation of liver injury. Deletion of Cav-1 attenuated LPS/GalN-induced acute liver injury and hepatocyte apoptosis with prolonged survival as compared with WT mice. In conclusion, our results suggest that Cav-1 contributes to the LPS/GalN-induced hepatic inflammation, apoptosis, and subsequent mortality. Therefore, suppression or disruption of Cav-1 may represent a potential approach for acute liver injury.

Contents
中文摘要 part I………………………………………………………1
中文摘要 part II…………………………… ………………………2
Abstract part I………………………………………………………3
Abstract part II……………………………… ……………………4

I.    Introduction
Part I
1.1.1  Macrophages regulate innate immune response
       against microbial infection…………………………… 5
1.1.2  Macrophages recognize microbes with several
       membrane receptor systems……………………………… 5
1.1.3  Toll-like receptor 4 and Myeloid differentiation
       protein 88……………………………………………………6
1.1.4  Nuclear factor-kB (NF-kB)…………………… …………7
1.1.5  Caveolin-1……………………………………………………8
1.1.6  Formation and function of Caveolae……………………9
1.1.7  Caveolae are associated with the entry of various
       pathogens……………………………………………………10
1.1.8  The role of Caveolin-1 in immunity………………… 10
1.1.9  Study aim ………………………………………………… 11
Part II
1.2.1  LPS/GalN model for acute liver injury………………12
1.2.2  Cytokine and chemokine regulate liver injury…… 13
1.2.3  Neutrophil infiltration is a key for liver injury15
1.2.4  Apoptotic mechanism …………………………………… 16
1.2.5  iNOS regulates the hepatotoxic effect………………16
1.2.6  Study aim ………………………………………………… 17

II.    MATERIALS AND METHODS
2.1  Reagents……………………………………………………… 18
2.2  Cell culture………………………………………………… 18
2.3  Mice and bone marrow-derived macrophages (BMDMs)… 19
2.4  Preparation of peritoneal macrophages…………………19
2.5  Preparation of green fluorescence protein (GFP)-
     expressing bacteria…………………………………………20
2.6  Flow cytometry……………………………………………… 20
2.7  Recombinant adenovirus construction……………………20
2.8  RT-PCR and Real-time PCR………………………………… 21
2.9  Preparation of lipid rafts……………………………… 22
2.10 Immunoblotting……………………………………………… 22
2.11 Phagocytosis, bacterial killing and colony-forming-
     unit (CFU) assay…………………………………………… 23
2.12 Phagocytosis and clearance of E. coli within the
     peritoneal cavity……………………………………………24
2.13 Determination of NF-kB activity…………………………24
2.14 Quantification of cytokines………………………………24
2.15 Mice hepatic damage and Mortality study………………24
2.16 Histopathology……………………………………………… 25
2.17 Immunohistochemistry……………………………………… 25
2.18 Blood chemistry………………………………………………25
2.19 MPO activity………………………………………………… 26
2.20 TUNEL assay……………………………………………………26
2.21 Statistical Analysis……………………………………… 26


III.    Results
PartI
3.1.1  Suppressed phagocytosis in Cav-1-/- macrophages
       in vitro and in……………………………………………28
3.1.2  Suppressed CD36 and CD14 expression in Cav-1-/-
       macrophages…………………………………………………29
3.1.3  Cav-1 deletion attenuates bacterial killing in
       vitro and in ………………………………………………30
3.1.4  Impaired bacterial killing of Cav-1-/-
       macrophages is implicated in part in impeded iNOS
       expression………………………………………………… 31
3.1.5  Cav-1-/- macrophages attenuated other protein
       beside iNOS for bacterial killing……………………31
3.1.6  Suppressed TLR4 and MyD88 expression and
       signaling in Cav-1-/- macrophages ………………… 32
3.1.7  Cav-1-/- macrophages attenuated NF-κB activity
       though TLR4-MyD88 signaling……………………………32
3.1.8  Cav-1 deletion inhibits inflammatory cytokine
       production………………………………………………… 33
3.1.9  Conclusion………………………………………………… 33
Part II
3.2.1  LPS/GalN induces Cav-1 expression……………………34
3.2.2  Cav-1-/- mice attenuates LPS/GalN-induced lethality
       rate and liver injury ………………………………… 34
3.2.3  Cav-1-/- mice reduces LPS/GalN-induced apoptosis of
       hepatocyte cells………………………………………… 35
3.2.4  Cav-1-/- mice reduces LPS/GalN-induced Cytokines and
       Chemokines expression……………………………………36
3.2.5  Cav-1-/- mice reduces LPS/GalN-induced hepatic iNOS
       and NO production…………………………………………36
3.2.6  Conclusion………………………………………………… 37
IV.    Discussion
Part I
4.1.1  Impaired phagocytotic ability of Cav-1-/-
       macrophages…………………………………………………38
4.1.2  Macrophage recognized pathogen and trigger immune
       response with membrane-associated proteins……… 38
4.1.3  Deletion of Cav-1 attenuates bacterial clearance
       in vitro and in vivo…………………………………… 39
4.1.4  Deletion of Cav-1 or disruption of caveolae
       suppresses iNOS expression and NO production…… 40
4.1.5  Bacterial killing is associated with TLR4-MyD88-
       NF-kB signaling in macrophages……………………… 41
4.1.6  Attenuated NF-kB activation due to impaired TLR4-
       MyD88 signaling in Cav-1-/- macrophages……………41
4.1.7  Cav-1 deletion attenuates bacteria-induced
       inflammatory response……………………………………42
4.1.8  Future direction………………………………………… 43
Part II
4.2.1  Cav-1 contributes to LPS/GalN-induced mortality and
       acute liver injury……………………………………… 44
4.2.2  Cav-1 deletion markedly attenuates GalN/LPS-induced
       mortality and serum AST/ALT upregulation………… 45
4.2.3  Deletion of Cav-1 attenuates neutrophil
       infiltration, apoptosis is associated with reduced
       cytokine and chemokine expression in response to
       LPS/GalN challenge……………………………………… 45
4.2.4  Deletion of Cav-1 suppresses iNOS expression through
       impairing TNF-a expression…………………………… 47
4.2.5  Future direction………………………………………… 48

V.    References……………………………………………… 49
VI.    Figures
Fig.1. Deletion of Cav-1 suppresses macrophage
       phagocytosis……………………………………………… 63
Fig.2. Over-expression of Cav-1 increases phagocytotic
       ability for macrophage………………………………… 64
Fig.3. Deletion of Cav-1 impairs Mac3 and CD206
       expression………………………………………………… 65
Fig.4. Disruption of the lipid raft hinders phagocytosis in
       macrophages…………………………………………………66
Fig.5. Decreased expression of CD14 and CD36 in Cav-1-/-
       macrophages is associated with attenuated
       phagocytosis……………………………………………… 67
Fig.6. Deletion of Cav-1-/- suppresses CD14 and CD36
       protein levels in peritoneal macrophages………… 68
Fig.7. Expression of CD14 and CD36 was examined in
       peritoneal macrophages treated with MβCD………… 69
Fig.8. CD14 and CD36 antibodies inhibit macrophage
       phagocytosis……………………………………………… 70
Fig.9. Deletion of Cav-1 suppresses bacterial killing in
       vitro and in vivo…………………………………………71
Fig.10.Deletion of Cav-1 suppresses bacterial killing in
       vivo………………………………………………………… 72
Fig.11.Deletion of Cav-1 impedes LPS- or bacteria-
      induced iNOS expression and NO production………… 73
Fig.12.Disruption of the lipid rafts hinders LPS-induced
       iNOS expression and NO production ………………… 74
Fig.13.Effects of Cav-1 and iNOS deletion in bacterial
       killing in peritoneal macrophages……………………75
Fig.14.Attenuated TLR4 and MyD88 expression in Cav-1-/-
       macrophages…………………………………………………76
Fig.15.Deletion of Cav-1 impairs NF-kB activation……… 77
Fig.16 Decreased production of IL-1b, IL-6 and TNF-a in
       Cav-1-/- macrophages and mice…………………………78
Fig.17.Model proposing how Cav-1 regulates protein
       expression related to bacteria- and LPS-induced
       signaling……………………………………………………79
Fig.18.Cav-1 mRNA accumulation in response to LPS/GalN
       administration in WT mice………………………………80
Fig.19.Attenuated mortality and liver injury in Cav-1-/-
       mice treated with LPS/GalN…………………………… 81
Fig.20.Treatment with LPS/GalN results in a higher liver
       injury and Neutrophils infiltration in WT than in
       Cav-1-/- mice………………………………………………82
Fig.21.Treatment with LPS/GalN results in a higher
       frequency of Liver Neutrophils infiltration in WT
       than in Cav-1-/- mice……………………………………83
Fig.22.In situ TUNEL stain of tissue section of liver from
       Cav-1-/- and WT mice after LPS/GalN challenge……84
Fig.23.Section of liver stained for active caspase-3 and
       caspase-8 in WT and Cav-1-/- mice challenged with
       LPS/GalN…………………………………………………… 85
Fig.24.Cav-1-/- mice attenuated caspase-3 and caspase-8
       activation in liver 6 h after LPS/GalN treatment 87
Fig.25.Inflammatory cytokine and chemokine are attenuated
       in Cav-1-/- mice challenged with LPS/GalN…………88
Fig.26.Attenuated cytokine and chemokine mRNA levels
       after LPS/GalN treatment in Cav-1-/-……………… 89
Fig.27.Cav-1-/- mice attenuate iNOS expression and NO
       production in liver tissue at 5 h after LPS/GalN
       administration…………………………………………… 90

                                

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