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研究生:郭慕涵
研究生(外文):Mu-Han Kuo
論文名稱:雞傳染性鼻炎菌RTX毒素疫苗之研發
論文名稱(外文):Development of the RTX-toxin as the vaccine against Avibacterium paragallinarum
指導教授:張伯俊
口試委員:陳秋麟謝快樂
口試日期:2016-05-27
學位類別:碩士
校院名稱:國立中興大學
系所名稱:微生物暨公共衛生學研究所
學門:獸醫學門
學類:獸醫學類
論文種類:學術論文
論文出版年:2016
畢業學年度:104
語文別:中文
論文頁數:77
中文關鍵詞:雞傳染性鼻炎菌RTX毒素重組次單位疫苗
外文關鍵詞:Avibacterium paragallinarumRTX toxinRecombinant subunit vaccine
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雞傳染性鼻炎菌 (Avibacterium paragallinarum)為格蘭氏陰性菌,會引起雞傳染性可利查 (Infectious coryza, IC),其為一種雞隻急性上呼吸道疾病,此疾病會造成產蛋率下降與生長遲緩。雞傳染性鼻炎菌可分為A、B、C三種血清型,目前市售的不活化疫苗僅對同血清型提供保護力,但血清型之間則沒有交叉保護力。儘管目前有全菌不活化疫苗,但此疾病仍造成經濟上重大損失。RTX毒素是許多陰性菌所產生的外毒素,是一個能在膜上形成孔洞的蛋白,會使得鈣離子通透性增加,最後造成細胞破裂而死亡。雞傳染性鼻炎菌的RTX毒素由avxCABD所組成,AvxA為毒素蛋白的主要結構,AvxC是作為脂肪酸修飾,AvxB、AvxD則是將毒素基因表現後運送至菌體外。在本篇研究中,利用Targetron gene knockout system構築AvxA突變株,並比較野生株與突變株之差異,以動物攻毒試驗評估是否降低細菌的致病能力,藉此探討其毒力因子是否在致病中扮演重要的角色。另外,亦構築avxCA互補株,並觀察是否回復原本性狀產生蛋白。藉由銀染和西方墨點法分析A. paragallinarum野生株、突變株與互補株表達AvxA蛋白的情形,結果顯示野生株與互補株皆有全長250 kDa蛋白的表現,突變株則沒有。此外,利用Escherichia coli表現AvxA毒素的C端重組蛋白對雞隻進行免疫,再以A. paragallinarum進行眼窩下竇攻毒試驗,其結果顯示此重組蛋白能提供保護力,具有作為次單位疫苗的潛力。

Avibacterium paragallinarum is a Gram-negative bacterium that causes infectioun coryza (IC), an important acute respiratory disease of chickens. Although the whole-cell inactivated vaccine is available, the disease still causes significant economic loss worldwide. RTX (Repeat in Toxin) is a pore-forming exotoxin produced by many Gram-negative bacteria. It may induce calcium flux in host cells and lead to cell lysis. The rtx gene from A. paragallinarum is composed of CABD genes. AvxA is the main structural protein toxin, which is activated by AvxC. AvxB and AvxD are type I secretion proteins that secrete AvxA into extracellular medium. Many studies showed that RTX toxin is an important virulence factor of pathogenic bacteria. The purpose of this study is to develope a vaccine based on AvxA. In this study, AvxA mutant is constructed by using Targetron gene knockout system to determine the role of RTX toxin in the pathogenesis of A. paragallinarum. Furthermore, AvxCA complement was constructed in A. paragallinarum to restore the function of the AvxA. Moreover, a recombinant protein containing the C-terminal half of AvxA was produced in Escherichia coli. Immunization of chickens with this recombinant protein confers chickens protection against challenge infection with A. paragallinarum. This is the first report showing that the recombinant RTX toxin could be a vaccine antigen against A. paragallinarum.

中文摘要 …………………………………………… 
英文摘要 …………………………………………… 
目次 ………………………………………………… 
表次 ………………………………………………… 
圖次 ………………………………………………… 
第一章 緒論 ……………………………………… 1
第二章 文獻探討 ………………………………… 2
第一節 Avibacterium paragallinarum之歷史背景 ……… 2
第二節 A. paragallinarum之生物學基本特性 …………… 2
第三節 A. paragallinarum之生化特性 …………………… 3
第四節 A. paragallinarum之血清型分型 ………………… 4
第五節 A. paragallinarum之流行病學 …………………… 5
第六節 A. paragallinarum之致病性 ……………………… 5
6-1 感染與傳播途徑 …………………………………… 5
6-2 臨床症狀與組織病理變化 ………………………… 6
6-3 診斷與鑑定方法 …………………………………… 6
6-4 管理與防治 ………………………………………… 8
第七節 A. paragallinarum之毒力因子 ……………… 8
7-1 血球凝集蛋白 (Hemagglutinin, HA) …………… 9
7-2 莢膜 (Capsule) …………………………………… 9
7-3 脂多醣 (Lipopolysaccharide, LPS) …………… 10
7-4 RTX (Repeat in toxin)毒素蛋白………………… 11
7-5 其他毒力因子 ……………………………………… 12
第八節 A. paragallinarum之疫苗研發 ……………… 13
第九節 實驗架構 ………………………………………… 14
第三章 材料與方法 ……………………………………… 15
第一節 A. paragallinarum菌株之分離 ……………… 15
第二節 A. paragallinarum菌株之鑑定 ……………… 15
2-1 A. paragallinarum菌株增殖與型態鑑定 ……… 15
2-2 PCR鑑定 …………………………………………… 15
第三節 菌株保存 ……………………………………… 16
3-1 A. paragallinarum菌株保存 …………………… 16
3-2 操作用A. paragallinarum菌株 ………………… 16
3-3 Escherichia coli菌株保存 ……………………… 17
第四節 DNA核酸萃取 …………………………………… 17
4-1 A. paragallinarum菌株之genomic DNA萃取 ………… 17
4-2 A. paragallinarum菌株之plasmid DNA萃取 ………… 18
4-3 E. coli菌株之plasmid DNA萃取 ……………………… 18
第五節 A. paragallinarum之RTX基因缺損株之構築 ……… 19
5-1 Targetron基因突變系統 …………………………… 19
5-2 Target site selection …………………………… 19
5-3 PCR增幅intron re-targeting 片段 ……………… 19
5-4 Re-targeting PCR產物純化與回收 (Clean-up)……… 20
5-5 Targetron載體構築之限制酶切割 (Enzyme digestion) … 20
5-6 膠體萃取 (Gel extraction) ……………………………… 21
5-7 Targetron載體構築之接合反應 (Ligation) ……………… 21
5-8 E. coli勝任細胞製備與熱休克轉型作用 (Transformation)…22
5-9 嵌入複製起始點與啟動子 …………………………………… 22
5-10 A. paragallinarum勝任細胞製備與電穿孔轉型作用
(Electroporation) ……………………………………… 23
5-11 突變株之挑選 (colony PCR) ………………………… 24
第六節 A. paragallinarum之avxCA基因互補株之構築 ……… 25
6-1 PCR增幅avxCA基因 ……………………………………… 25
6-2 限制酶切割 (Enzyme digestion)、接合反應 (Ligation)與轉型作用(Transformation) ……………………………………… 26
6-3 質體快篩選殖與限制酵素切割之分析 ………………… 27
6-4 質體甲基化作用(Methylation)與DNA萃取(DNA extraction)…27
6-5 A. paragallinarum勝任細胞製備與電穿孔轉型作用
(Electroporation) ………………………………………… 28
第七節 A. paragallinarum野生株、突變株、互補株之蛋白分析與毒性分析………………………………………………………… 29
7-1 TCA蛋白沉澱 (Trichloroacetic acid protein precipitation)與銀染分析 (Silver staining) …………………………… 29
7-2 西方墨點法分析 (Western blot) …………………… 30
7-3 A. paragallinarum野生株與RTX突變株之攻毒試驗 …… 31
第八節 AvxA-C-ter次單位疫苗之重組蛋白製備 ……………… 32
8-1 PCR增幅avxA基因之C端片段 ……………………………… 32
8-2 限制酶切割 (Enzyme digestion)、接合反應 (Ligation) 與轉型作用 (Transformation) …………………………………… 32
8-3 質體之快篩選殖 ………………………………………… 33
8-4 以原核系統表現重組蛋白 (Protein expression) …… 33
8-5 重組蛋白之純化 (Purification) ……………………… 34
8-6 重組蛋白之電泳分析 (SDS-PAGE analysis) ………… 35
8-7 重組蛋白之西方墨點法 (Western blot) …………… 35
8-8 重組蛋白之濃縮與置換 ………………………………… 36
8-9 重組蛋白濃度測定 …………………………… 36
第九節 RTX重組次單位疫苗與A. paragallinarum AvxCA互補株不活化疫苗之保護效力評估 …………………………… 36
9-1 實驗動物 ……………………………………… 36
9-2 免疫策略與攻毒計畫流程 …………………… 37
9-3 檢測免疫後雞隻血清對抗原之抗體反應 ………………… 37
9-4 重組次單位疫苗之製備 …………………………………… 38
9-5 A. paragallinarum AvxCA互補株不活化疫苗之製備 …… 38
9-6 攻毒病原菌之製備 ………………………………………… 39
9-7 實驗動物免疫攻毒後之釣菌與鑑定 ……………………… 39
第四章 結果 ………………………………………………… 40
第一節 A. paragallinarum之鑑定 ……………………… 40
第二節 A. paragallinarum avxA基因突變株之構築 ……… 40
2-1 增幅intron re-targeting片段與電泳產物確認 ……… 40
2-2 嵌入A. paragallinarum複製起始點與啟動子之確認 … 40
2-3 A. paragallinarum avxA基因突變株之挑選 ………… 41
第三節 野外株與突變株之攻毒試驗 ……………………… 41
第四節 A. paragallinarum互補株之構築 ……………… 42
4-1 增幅全長avxCA基因與電泳產物確認 ……………… 42
4-2 互補株之質體確認 …………………………… 42
第五節 A. paragallinarum AvxA蛋白表現 ……………… 42
第六節 AvxA-C-ter重組蛋白之製備 ……………………… 43
6-1 增幅avxA之C端基因與電泳產物確認 ………………… 43
6-2 E. coli轉型株獲得重組載體之確認 ………………… 43
6-3 以原核系統表現重組蛋白與確認 …………………… 44
6-4 重組蛋白之純化 ……………………………………… 44
第七節 重組次單位疫苗與A. paragallinarum AvxCA互補株不活化疫苗保護效力之評估 ………………………………………… 44
7-1 檢測免疫後雞隻血清對抗原之抗體反應 …………… 44
7-2 動物免疫攻毒試驗 …………………………………… 45
7-3 實驗動物免疫攻毒試驗後之釣菌與鑑定 …………… 47
第五章 討論 …………………………… 65
第六章 參考文獻 ……………………… 69


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