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研究生:曹譯友
研究生(外文):I-Yu Tsao
論文名稱:白腹叢蚊原酚氧化酵素基因分子選殖,特性與功能分析
論文名稱(外文):Molecular cloning, characterization and functional analyses of prophenoloxidase genes from mosquito Armigeres subalbatus
指導教授:陳正成陳正成引用關係
指導教授(外文):Cheng-Chen Chen
學位類別:博士
校院名稱:國立陽明大學
系所名稱:臨床醫學研究所
學門:醫藥衛生學門
學類:醫學學類
論文種類:學術論文
論文出版年:2009
畢業學年度:97
語文別:中文
論文頁數:167
中文關鍵詞:蚊子原酚氧化酵素雙股核醣核酸干擾調控子分析形態發育超微結構
外文關鍵詞:mosquitoprophenoloxidasedouble-stranded RNA interferencepromoter analysismorphogenesisultrastructure
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酚氧化酵素(phenoloxidase, PO)早已被認為參與昆蟲許多重要生理功能,例如表皮骨骼化(cuticular sclerotization)、傷口癒合(wound healing)、卵殼硬化(tanning of egg chorion)與黑化包囊反應(melanization)。本研究室先前自白腹叢蚊(Armigeres subalbatus, Ar. subalbatus)選殖出兩個白腹叢蚊原酚氧化酵素,並分別命名為白腹叢蚊原酚氧化酵素I (Ar. subalbatus prophenoloxidase I, As-pro-POI) 與白腹叢蚊原酚氧化酵素II (Ar. subalbatus prophenoloxidase II, As-pro-POII)。本研究則更進一步選殖出六個新的原酚氧化酵素,分別命名為白腹叢蚊原酚氧化酵素III-VIII (Ar. subalbatus prophenoloxidase III-VIII, As-pro-POIII-VIII)。As-pro-POIII、IV與VI mRNA表現在成蚊皆呈持續表現且不受吸血、微絲蟲感染及細菌感染等刺激影響,惟這些酚氧化酵素在幼蟲及蛹期表現有變動,而As-pro-POV則在吸血後才表現。本研究進一步探討As-pro-PO III功能。結果發現當蚊蛹As-pro-PO III表現被雙股核醣核酸(double-stranded RNA,dsRNA)擊低(knockdown)後,大部分蚊蛹會死亡,僅少量能羽化為畸形成蟲並死亡。而超微結構研究結果發現,注射雙股核醣核酸As-pro-PO III導致蚊蛹新形成內表皮(endocuticle)與預成蟲表皮(pharate adult cuticle)結構不完整,如薄層(lamellae)層數明顯變少、幾丁質微絲所架構的半螺旋結構(helicoidal structure)消失,且在表皮蛋白基質出現許多電子疏鬆(electron-lucent)空洞結構。而蚊蛹在注射雙股核醣核酸As-pro-PO III後,所羽化的成蟲亦觀察到相同影響。基因調控活性試驗結果證實As-pro-PO III基因由生長發育相關調控子Zeste進行調控。前人研究已證實酚氧化酵素作用產生的類醌(quinone)為表皮蛋白交互作用重要物質,因此我們認為當As-pro-PO III表現被擊低後,導致鍵結表皮結構的類醌不足,進而使表皮結構不正常與基質流失,因而影響新表皮形成。我們的研究證實As-pro-PO III在蚊子表皮形成扮演重要角色,此為酚氧化酵素的新功能。
It has long been suggested that phenoloxidases (POs) play key roles in various physiological functions in insects, e.g., cuticular sclerotization, wound healing, egg tanning and melanizatic encapsulaction of pathogens. Two POs, namely Armigeres subalbatus prophenoloxidase I (As-pro-PO I) and Ar. subalbatus prophenoloxidase II (As-pro-PO II), were previously identified from mosquito Ar. subalbatus. In this study, we further isolated six pro-POs, designated As-pro-PO III, IV, V, VI,VII and VIII, from Ar. subalbatus. Expression profiles analysis revealed that As-pro-PO III, IV and VI mRNA were persistently expressed in adult mosquitoes and were not significantly affected by blood feeding, microfilaria inoculation, or Escherichia coli inoculation, but expression levels of these pro-POs fluctuated in larval and pupal stages. On the contrary, As-pro-POV transcript level was significantly increased after blood feeding. The function of As-pro-POIII was furthered illustrated in this study. Knockdown of As-pro-PO III expression using double-stranded RNA (dsRNA) resulted in high pupal mortality and deformed adults that subsequently died following emergence. Further ultrastuctural study revealed that inoculation of As-pro-POIII dsRNA resulted in the incomplete formation of nascent pupal endocuticle and pharate adult cuticle, i.e. a significantly fewer cuticular lamellae were deposited, the helicoidal patterns of chitin microfibrils were disorganized, and numerous electron-lucent spaces were formed in cuticular protein matrix. Similar disruptions were also observed in cuticle of adult derived from As-pro-POIII dsRNA-inoculated pupae. Promoter activity analysis suggested that the As-pro-POIII was positively regulated by a putative Zeste motif, a developmental regulatory element. It has been long suggested that the quinone, generated by PO-catalyzed oxidation reactions, function as cross-linking agents; therefore, it seems reasonable to suggest that the loss of As-pro-POIII-mediated protein-protein linkages causes morphological abnormalities in protein matrix. Our findings suggested that As-pro-POIII plays a role in cuticle formation in mosquitoes, a novel function for phenol oxidizing enzymes.
中文摘要------------------------------------------------------------ 4
英文摘要------------------------------------------------------------ 5
致謝---------------------------------------------------------------- 7
緒論---------------------------------------------------------------- 8
壹、 昆蟲免疫反應----------------------------------------------- 9
貳、 黑色素生成路徑與生理功能----------------------------------- 14
參、 昆蟲的體壁與結構------------------------------------------- 16
肆、 原酚氧化酵素----------------------------------------------- 19
伍、 負向控制基因表達------------------------------------------- 21
研究動機----------------------------------------------------------- 23
研究流程----------------------------------------------------------- 24
材料與方法--------------------------------------------------------- 25
壹、 生物材料--------------------------------------------------- 25
貳、 白腹叢蚊原酚氧化酵素互補去氧核醣核酸選殖------------------- 26
參、 白腹叢蚊原酚氧化酵素基因篩選與調控分析--------------------- 29
肆、 白腹叢蚊原酚氧化酵素轉錄作用分析--------------------------- 32
伍、 白腹叢蚊原酚氧化酵素功能分析------------------------------- 33
陸、 實驗數據統計與分析----------------------------------------- 35
結果--------------------------------------------------------------- 36
壹、 白腹叢蚊原酚氧化酵素cDNA選殖------------------------------ 36
貳、 白腹叢蚊原酚氧化酵素分子mRNA表現分析---------------------- 39
參、 白腹叢蚊原酚氧化酵素基因序列結構及上游序列調控分析--------- 40
肆、 白腹叢蚊原酚氧化酵素III功能分析--------------------------- 41
討論--------------------------------------------------------------- 46
參考文獻----------------------------------------------------------- 54
圖、表及附錄說明--------------------------------------------------- 75
表----------------------------------------------------------------- 76
圖----------------------------------------------------------------- 80
附錄--------------------------------------------------------------- 106
論文常用中英對照及縮寫一覽表--------------------------------------- 120
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