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研究生:李惟宇
研究生(外文):Wei-Yu Lee
論文名稱:類趨化激素CXCL17之基因調控機制與受體下游訊息傳遞後之功能探討
論文名稱(外文):Characterization of the gene regulatory profile and receptor signaling of an orphan chemokine-like protein CXCL17
指導教授:羅清維
指導教授(外文):Ching-Wei Luo
學位類別:碩士
校院名稱:國立陽明大學
系所名稱:生命科學暨基因體科學研究所
學門:生命科學學門
學類:生物訊息學類
論文種類:學術論文
論文出版年:2010
畢業學年度:98
語文別:英文
論文頁數:81
中文關鍵詞:趨化激素類趨化激素CXCL17免疫單核球巨嗜細胞
外文關鍵詞:ChemokineCXCL17immunitystomachmonocytemacrophage
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趨化激素17 (CXCL17)目前已知的功能有: 吸引免疫細胞的移動、幫助癌細胞的生長與促進血管的新生。但是對於趨化激素17蛋白質的體內型態、傳遞訊息的受體或是其在免疫上所扮演的角色等所知卻有限。大部分CXC 趨化激素家族的成員,其蛋白質的序列上具有四個半胱氨酸 (cysteine),但是,趨化激素 17又再多出了兩個高度保留的半胱氨酸,全長總共含有六個半胱氨酸。另一方面,藉由序列比對的方式,我們發現在不同物種中其第二個與第三個半胱氨酸間皆含有兩個高度保留的轉化酶 (convertase) 切位,說明了趨化激素17在蛋白成熟的過程當中,有可能會受到轉化酶的剪切,使得成熟型的趨化激素17最後只保留四個半胱氨酸在其蛋白質序列中。在我們本次的實驗中證明,無論是全長的趨化激素17或是成熟型的趨化激素 17,皆具有活化早期反應基因(early response gene) c-fos 表現的能力;更進一步的,我們也證明這些活化的訊息傳遞路徑包括了 ERK1/2 和p38 MAPK的磷酸化反應途徑。在免疫的功能上,我們首先觀察到的是趨化激素17具有吸引單核球細胞與巨噬細胞移動的能力,我們進一步的觀察到這些受到趨化激素17刺激的單核球細胞,在IL-8與VEGF的基因表現上有增加的趨勢,這兩個細胞激素目前已知的功能包含了免疫細胞的吸引、組織的修復與血管的新生等。更進一步的,我們觀察到受到趨化激素 17初級刺激的巨噬細胞,在發炎反應的活化上會受到部分的抑制,因此可以避免過度的發炎反應對組織可能產生的傷害。在表現的位置上,先前我們的實驗室已經證明趨化激素 17主要是表現在胃部;而在本論文中我們更進一步的證明,當胃部細胞受到酒精或細菌脂多醣 (LPS) 等不同的外來物質刺激時,趨化激素17的基因表現會有增加的情況。綜合本研究的結果,我們推論:當胃部細胞受到外來物質刺激時,趨化激素17的表現量會增加,進而吸引並刺激免疫細胞產生特定的反應,諸如藉由抑制過度的發炎反應與增加組織修復相關的細胞激素釋放等方式,來展現趨化激素17在胃部所扮演的組織保護角色。
Chemokine (C-X-C motif) ligand 17 (CXCL17) has the potentialities to attract leukocytes, promote tumor growth and induce angiogenesis. However, its protein nature, receptor identity and detailed immune action remain poorly defined. Unlike regular CXC chemokines with four cysteines in sequence, CXCL17 contains six highly conserved cysteines. Of interest, in comparison with CXCL17 in different species, two conserved convertase cutting sites were found between cysteine 2 and 3, which potentially lead to the formation of processed CXCL17 with four-cysteine nature. Here we demonstrated that 4-cys and 6-cys forms of CXCL17 have the same effects to elevate the c-fos expression as well as the ERK and p38 phosphorylation in CXCL17-responding immune cells. We also demonstrated that CXCL17 can stimulate and attract THP-1 monocytes and J774 macrophage cells. CXCL17 up-regulated the mRNA expression of IL-8 and VEGF in treated THP-1. Such factors have been well known to involve in tissue repair and angiogenesis. Furthermore, CXCL17-primed J774 cells showed hyporesponse to inflammation, which is important for tissue protection. During tissue screening, CXCL17 showed the highest level in the stomach, where its expression is further up-regulated under the contact with stimuli such as ethanol or LPS. Therefore, taking together with its expression pattern and immune functions, we conclude CXCL17 likely acts as a protective chemokine in the stomach by suppressing macrophage-driven inflammation and subsequently promoting the release of tissue repair factors.
Abstract in English……………...…………………………………………………….1
Abstract in Chinese……………………………………………………………...........2
Introduction………………………………………………………………..………….3
1. Chemokines and their subfamilies………………………………………………….3
2. C-X-C ligand 17 (CXCL17) chemokine…………………………………………....4
3. Protein post-translation by pro-protein convertases (PCs)………………………….6
4. Chemokines and G protein-coupled receptors (GPCRs)……………………………7
5. GPCRs and their downstream signaling pathways…………………………………7
6. Chemokines and inflammation……………………………………………………..8
7. Chemokines in mucosal inflammation……………………………………………..9
8. Chemokines and tumors…………………………………………………………...10
9. The role of monocytes in innate immune responses………………………………11
10. The role of monocytes in tumor development……………………………………12
11. The role of macrophages in immune response…………………………………...12
12. The many faces of macrophage activation……………………………………….13
12.1. Priming……………………………………………………………………13
12.2. Classically activated macrophages (M1 macrophages)…………………...14
12.3. Alternatively activated macrophages (M2a macrophages)……………….14
12.4. Type II activated macrophages (M2b macrophages)……………………..15
12.5. Deactivated macrophages (M2c macrophages)…………………………...15
12.6. Tumor-associated macrophages (TAMs)…………………………………15
13. Chemokines and macrophage activation…………………………………………16
14. Specific aims……………………………………………………………………..17
Materials and methods……………………………………………………………...18
A) Materials
1. Cell lines……………………………………………………………………….18
2. Luria-Bertani (LB) medium…………………………………………………...18
3. Cell culture media……………………………………………………………..18
4. Plasmids……………………………………………………………………….19
5. Antibodies……………………………………………………………………..20
B) Methods
1. Protein induction and expression…………………………………………….21
2. Extraction of crude recombinant protein……………………………………..21
3. Protein purification…………………………………………………………...21
4. Protein quantification………………………………………………………...22
5. Protein refolding……………………………………………………………...22
6. Sodium dodecyl sulfate polyacrylamide gel electrophoresis (SDS-PAGE)…23
7. Protein transfer……………………………………………………………….24
8. Western blotting……………………………………………………………...24
9. Cell transfection……………………………………………………………...25
10. Luciferase and β-galactosidase reporter assay……………………………...25
11. Cell cultures…………………………………………………………………26
12. Primary cell cultures………………………………………………………...26
13. Cell treatments………………………………………………………………27
14. RNA extraction …………………………………………………………….29
15. cDNA synthesis……………………………………………………………..29
16. Quantitative real-time polymerase chain reaction…………………………..30
17. Transwell invasion assay……………………………………………………31
18. Nitric oxide (NO) assay……………………………………………………..31
Results…………………………………………………………………………...…..33
1. Production of the recombinant human CXCL17 proteins…………………………33
2. Bioactivity comparison between 4-cys and 6-cys CXCL17 by c-fos induction…..33
3. The receptor for CXCL17 is still unknown………………………………………..34
4. Treatment with CXCL17 activate the phosphorylation of ERK1/2 and p38 but not JNK in J774 macrophages………………………………………………………… 35
5. CXCL17 is a chemoattractant for monocyte and macrophage cell lines………….35
6. Ethanol can regulate CXCL17 mRNA expression in rat stomach…………………36
7. LPS induces CXCL17 gene expression in rat glandular stomach primary cells…..37
8. CXCL17 induces IL-8 and VEGF gene expression in THP-1 monocytes………...38
9. CXCL17-activated THP-1 monocyes cannot further differentiate into macrophages
......…………………………………………………………………………………38
10. CXCL17 cannot induce IL-6, TNF-α and iNOS mRNA expression in J774 cells
……………………………………………………………………………………..39
11. CXCL17 acts as an anti-inflammatory factor to suppress proinflammatory gene expression in LPS-stimulated J774 cells…………………………………………40
12. CXCL17 pretreatment suppresses NO (nitrite oxide) production in LPS stimulated-J774 cells……………………………………………………………..40
Discussion……………………………………………………………………………42
1. Bioactivity comparison between 4-cys and 6-cys CXCL17………………………42
2. CXCL17, receptors and signal transduction………………………………………43
3. The biological function of CXCL17 in the stomach………………………………44
4. The biological function of CXCL17 in monocytes (THP-1 cells)………………...46
5. The biological function of CXCL17 in macrophages (J774 cells)………………...47
References…………………………………………………………………………50
Table 1…………………………………………………………………………58
Figures……………………………………………………………………………..59

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