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系統識別號 U0026-2411201411281900
論文名稱(中文) 探討fur在幽門桿菌游動能力之角色
論文名稱(英文) Roles of fur in Helicobacter pylori motility
校院名稱 成功大學
系所名稱(中) 分子醫學研究所
系所名稱(英) Institute of Molecular Medicine
學年度 103
學期 1
出版年 104
研究生(中文) 李艾芸
研究生(英文) Ai-yun Lee
電子信箱 T16014072@mail.ncku.edu.tw
學號 T16014072
學位類別 碩士
語文別 中文
論文頁數 111頁
口試委員 指導教授-吳俊忠
口試委員-何漣漪
口試委員-鄧景浩
口試委員-賴信志
中文關鍵字 幽門桿菌  游動性  Fur  鞭毛結構  趨化反應  群體感應  AI-2 
英文關鍵字 Helicobacter pylori  motility  Fur  flagellar structure  chemotaxis  quorum sensing  AI-2 
學科別分類
中文摘要 幽門桿菌 (Helicobacter pylori) 是常見的人類致病菌,全球人口中約有50%受之感染,而感染後會增加罹患胃部疾病甚至是胃癌的風險。游動能力為幽門桿菌成功拓殖於人體胃黏膜的重要因素。然而幽門桿菌之所以能在胃部如此惡劣的環境中生存,乃因具有良好的適應能力,鐵攝取調控因子 (ferric uptke regulator, Fur) 即為其中重要的調控因子之一。過去在轉錄基因體學及蛋白質體學分析結果中皆發現Fur會影響部分鞭毛基因之表現,然而其中的詳細機制尚未明瞭,因此我擬定探討Fur在幽門桿菌的游動能力中所扮演的角色,並試圖了解其中的調控機制。首先,利用半固態培養基確認fur突變株之游動性,並於結果中發現fur突變後確實會影響幽門桿菌之游動能力,使之游動性降低。針對此問題我分成三個方向進行探討,其中包括鞭毛的生合成、趨化反應以及群體感應。於鞭毛的生合成的部分,雖然fur突變後其鞭毛素蛋白FlaA及FlaB的表現量皆增加,但其鞭毛結構與野生株相比並沒有明顯差異,故排除可能為fur突變後影響鞭毛生合成而導致其游動力降低的可能性。在趨化系統方面,我發現趨化系統相關基因之表現量於fur突變後也並無顯著差異。於缺鐵環境的刺激下會使得cheY失去大量表現的能力,但在半固態培養基的趨化實驗發現,經趨化物刺激後fur突變株之游動圈仍較小,且沒有產生變化,因此推測趨化系統亦非影響fur突變株游動力下降的原因。而群體感應的結果發現,fur突變後其培養液中AI-2的含量與野生株相比,於對數期早期較低,且此差異會隨細菌的生長而減少、消失;然而其luxS的表現量與野生株相比,於對數期晚期 (18 hr) 及穩定期 (24 hr) 皆無明顯差異。總結以上,推測Fur可能會藉由LuxS-independent的方式影響AI-2的產量,或是直接影響AI-2分泌的能力,來調控幽門桿菌J99於對數期早期的游動能力。
英文摘要 Helicobacter pylori infects 50% of the population worldwide, it can increase the risk for developing gastroduodenal diseases even the gastric cancer. Motility is essential for H. pylori to successfully colonize in human gastric mucosa persistently. One of the proteins that contribute to the remarkable adaptability is the ferric uptake regulator (Fur) which functions as a global regulator of gene expression. Transcriptional profile and proteomic analysis showed that Fur regulates the expression of some flagella-related genes in H. pylori 26695 strain. However, the detail mechanisms are still unclear. In this study, we aimed to investigate the roles of Fur in H. pylori motility, and try to clarify its regulatory mechanism. First, the motility of H. pylori J99 fur mutant was examined by the soft-agar motility assay. Our data showed mutant of fur decreased the motility of H. pylori. Thus we further divided into three sections including flagella formation, chemotaxis and quorum-sensing to study the effects of decreasing motility. Although increased expression in flagellins FlaA and FlaB, fur mutant J99 had a normal flagella structures compared with wild-type by using transmission electron microscopy. Further, we checked the expression of chemotaxis-related genes. The results showed that there was no significant difference on the chemotaxis-related genes between fur mutant J99 and wild-type under normal cultural condition. Although the expression of cheY in fur mutant significantly lose the ability to be increased under iron depletion treatment, the motility zone of fur mutant J99 was still smaller than wild-type and it seemed not affected in chemotaxis soft agar test. Furthermore, we considered whether quorum-sensing was involved. With AI-2 bioassay, the results showed that the AI-2 of fur mutant was dramatically reduced compared with wild-type J99 in the early-log phase. However, the RNA level of LuxS, served as AI-2 synthase, had no significant difference between fur mutant and wild-type J99 on both late-log phase (18 hr) and stationary phase (24 hr). In conclusion, these results suggested that Fur might regulate H. pylori J99 motility in log phase through LuxS-independent modulating AI-2 production or affecting the amount of AI-2 secretion.
論文目次 中文摘要 i
Extended abstract ii
致謝 vi
目錄 viii
表目錄 xii
圖目錄 xiii
符號及縮寫 xiv
緒論 1
一、幽門桿菌 1
1. 發現的歷史 1
2. 分類與特性 1
3. 傳染與流行病學調查 2
二、 幽門桿菌之毒力因子 3
1. 尿素酶 (Urease) 4
2. 鞭毛素蛋白 (Flagellin) 5
3. 黏附因子 (Adhesin) 5
a. BabA (blood-antigen binding protein A) 6
b. SabA (sialic acid-binding adhesin) 7
4. 空泡毒素 (Vacuolating cytotoxin A, VacA) 7
5. 細胞毒素攜帶抗原A (Cytotoxin associated gene A, CagA) 9
三、幽門桿菌感染及致病的進程 10
四、影響細菌游動能力之因素 11
1. 鞭毛構造的形成 (Flagella formation) 11
2. 鞭毛馬達的活性 (Motor activity) 12
3. 趨化系統(Chemotaxis system) 13
4. 群聚感應(Quorum sensing) 14
a. 同種細菌間的對話 (intraspecies communication) 14
b. 不同種細菌間的對話 (inter-species specific communication) 15
五、鐵攝取調控因子(ferric uptke regulator, Fur) 16
1. Fur superfamily的特徵 16
2. Fur為global regulator 17
3. Fur對幽門桿菌致病力的影響 18
六、研究方向 20
材料與方法 21
一、實驗藥品及溶液配方 21
二、菌種及其培養條件與保存 21
三、細菌DNA之萃取 22
1. 大腸桿菌質體 (Plasmid) 22
2. 幽門桿菌染色體DNA (Genomic DNA) 22
四、聚合酶連鎖反應 (Polymerase chain reaction, PCR) 23
五、核酸凝膠電泳 (Agarose gel electrophoresis) 23
六、限制酶酵素切割及DNA接合反應 23
七、大腸桿菌勝任細胞 (Competent cells) 製備 24
八、大腸桿菌細胞轉型作用 (Transformation) 24
九、幽門桿菌自然轉型作用 (Natural transformation) 25
十、 南方墨點法 (Southern blotting) 25
1. 探針 (Probe) 之製備 25
2. 染色體DNA之限制酶切割反應 26
3. DNA轉漬 (Transfer) 26
4. 雜交反應 (Hybridization) 27
5. 清洗 (Wash) 與呈色作用 (Detection) 27
6. 探針剝離反應 (Stripping) 27
十一、幽門桿菌於培養液中的生長曲線分析 28
十二、鐵離子比色法 (Colorimetric Fe analysis) 28
十三、膠體游動能力分析 (Soft agar motility assay) 28
十四、穿透式電子顯微影像 (Transmission electron microscopy, TEM) 29
1. 準備電顯片 (鍍碳銅網) 29
2. 細菌樣品製備以及負染色作用 29
十五、AI-2 bioassay 30
十六、RNA實驗之操作 30
1. 幽門桿菌RNA之萃取 30
2. 反轉錄-聚合酶連鎖反應 (Reverse transcription PCR, RT-PCR) 31
3. 定量即時聚合酶鏈鎖反應 (Real-time quantitative RT-PCR, RT-qPCR) 32
十七、蛋白質實驗之操作 33
1. 幽門桿菌的蛋白質製備 33
a. 總體蛋白 (Total lysate) 33
b. 層析法 (Fractionation) 33
2. 蛋白質濃度量 34
3. 蛋白質膠體電泳 (SDS-PAGE) 34
4. 西方點墨法 (Western blotting) 35
5. 重組蛋白質之大量誘導表現 36
6. 蛋白質純化 36
a. 超音波破菌 (Sonication) 36
b. 鎳離子親合管柱 (Nickel column) 37
c. 蛋白質濃縮 38
d. 膠體過濾層析管柱 (Gel-filtration column) 38
十八、Anti-Fur老鼠多株抗體製備 38
1. 抗體製備 38
2. 抗體效價分析 39
十九、生物資訊分析工具 40
1. 全基因體調控子分析工具 - Vitrual Footprint version 3.0 40
2. 比較核苷酸與胺基酸序列之工具 40
3. 統計學分析 40
結果 41
一、構築J99 fur突變株 (SW842) 及復原株 (SW843) 41
二、J99野生株與fur突變株於幽門桿菌游動能力之差異 42
三、fur突變株對鞭毛蛋白表現量之影響 43
四、以穿透式電子顯微鏡 (TEM) 觀察鞭毛結構 45
五、Fur對趨化系統造成之影響 46
六、Fur可能參與調控cheY之表現 46
七、Fur影響幽門桿菌對數期 (log phase) 之AI-2分泌量 48
八、Fur可能並非藉由調控luxS之表現量來影響幽門桿菌CFS (cell free supernatant) 中AI-2的含量 49
討論 50
一、fur突變後改變典型鞭毛轉錄階層的調控? 50
二、fur復原株於鞭毛相關基因表現量的影響 51
三、於半固態培養基上進行趨化能力測試實驗的可信度 52
1. 鐵離子螯合劑DPP的添加是否會影響細菌的生長? 52
2. fur突變株其游動圈之表現型態不受DPP影響? 53
四、Fur可能藉由非LuxS路徑來影響幽門桿菌CFS (cell free supernatant) 中AI-2的含量 53
五、半固態培養基上呈現的細菌游動圈之表現型態 54
1. 游動圈內外圈缺乏明確的定義 54
2. 以AI-2的含量來解釋fur突變株游動圈表現形態的時態性變化 55
六、Fur會干擾正常鞭毛基因表現的時效性? 與群體感應有關? 55
七、總結 57
參考文獻 59
圖表 80
附錄 101
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