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研究生:洪毓婷
研究生(外文):Yu-Ting Hong
論文名稱:豆類金黃嵌紋病毒屬雙生病毒 (begomoviruses) C4蛋白影響寄主植物葉片捲曲趨向性分子機制之研究
論文名稱(外文):Studies on the molecular mechanisms modulating directions of leaf curling by C4 proteins of begomoviruses
指導教授:胡仲祺
指導教授(外文):Chung-Chi Hu
學位類別:碩士
校院名稱:國立中興大學
系所名稱:生物科技學研究所
學門:生命科學學門
學類:生物科技學類
論文種類:學術論文
論文出版年:2010
畢業學年度:98
語文別:中文
論文頁數:96
中文關鍵詞:雙生病毒C4蛋白豆類金黃嵌紋病毒屬
外文關鍵詞:geminivirusC4Begomovirus
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  • 被引用被引用:5
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雙生病毒為具有環狀單股 DNA 基因體的植物病毒,外型是雙球型病毒顆粒,可感染多種雙子葉植物,並造成經濟作物重大損失。不同的雙生病毒感染相同寄主植物,會造成不同類型的葉片捲曲病徵。本實驗室先前研究發現感染霍香薊黃脈病毒 (Ageratum Yellow Vein Virus, AYVV) 時,會造成圓葉菸草 (Nicotiana benthamiana) 葉片上捲;而感染番茄捲葉病毒 (Tomato Leaf Curl Virus, TLCV) 或南瓜捲葉病毒 (Squash Leaf Curl Virus, SqLCV) 時,會造成葉片下捲。並進一步經由 AYVV、TLCV 與 SqLCV 病毒基因交換重組試驗,發現不同病毒的 C4 蛋白,可以調控菸草葉片的捲曲方向。因此,本實驗以 AYVV、TLCV 和 SqLCV 之 C4 蛋白作為實驗材料,探討與病徵表現及調控葉片捲曲方向的分子機制。首先將上述三種雙生病毒之 C4 蛋白基因選殖於 pET21d ,經大腸桿菌大量表現並純化後,已製作出三種雙生病毒 C4 蛋白專一性抗血清。並證實可在感病植物中偵測到雙生病毒 C4 蛋白之表現。之後運用蛋白質二維電泳法 (Two Dimensional- Polyacrylamide Gel Electrophoresis, 2D-PAGE) 、遠西方墨點法 (Far-western blot analysis) 分析並尋找 N. benthamiana 可與雙生病毒 C4 蛋白發生交互作用的寄主因子。經由質譜分析與生物資訊學工具探究後,目前已發現與光合作用相關的 chloroplast photosynthetic oxygen-evolving 蛋白、植物抗病毒相關的 24K germin 類似蛋白、糖解作用和糖質新生路徑相關的 plastidic aldolase 與光合作用相關的 photosystem I light-harvesting chlorophyll a/b-binding protein 。另一方面也利用免疫共沉澱法尋找在原始構形下會與 C4 蛋白產生交互作用的水溶性植物蛋白,目前已發現與細胞的分裂、維持細胞的極性 (cell polarity) 有關的 Formin-like protein 、與蛋白質正確摺疊有關的 Luminal-binding protein 、與葉綠體中電子傳遞鏈的進行有關的 NADH:ubiquinone oxidoreductase 、與能量調控有關的 26S protease regulatory subunit 6A homolog B 、調控基因表現有關的 MAR-binding filament-like protein 與抗病相關的 Putative disease resistance protein RPP8-like protein 。而為探討 C4 蛋白單獨表現時對於寄主植物葉片捲曲病徵的影響,本研究分別將 AYVV、TLCV 與 SqLCV 的 C4 轉殖在 pBin35S 載體、竹嵌紋病毒衛星核酸表現載體 pCass-satBaMV 與竹嵌紋病毒表現載體 pCass-BaMV ,發現在植物體內 C4 mRNA 可以正常表現,但目前尚未觀察到 C4 蛋白的表現,也無法看到 C4 蛋白對植物病徵表現的影響,間接證實 C4 基因對於植物葉片捲曲的影響,是發生在蛋白質層次而非 RNA 抑制現象。未來將利用轉基因植物,在植物體內持續表現 C4 蛋白,以觀察 C4 蛋白對植物病徵所造成的影響。經由本研究,已發現可能參與受不同雙生病毒 C4 蛋白影響而調控植物葉片生長發育的相關基因,除可開發作為研究植物細胞分裂分化的模式系統,並期望未來可以應用於治療已感染雙生病毒的作物上,減少雙生病毒的危害與損失。

Whitefly-transmitted geminiviruses, members of the genus Begomovirus, can infect a variety of dicotyledonous plants and inflict different types of symptoms, causing severe economical losses. Previous studies in our laboratory revealed that Nicotiana benthamiana plants infected by Ageratum Yellow Vein Virus (AYVV) display severe upward leaf curling symptoms. In contrast, the leaves of those infected by Tomato Leaf Curl Virus (TLCV) or Squash Leaf Curl Virus (SqLCV) showed distinct downward curling. Further studies using recombinant viruses have identified C4 protein coding region as the possible determinant of the directions of leaf curling. However, the host factors involved in leaf curling symptoms remain elusive. Therefore, the objectives of this study are to identify the host factors and underlying mechanisms controlling the directions of leaf curling, using the C4 proteins of AYVV, TLCV and SqLCV as materials. The C4 gene of AYVV, TLCV, and SqLCV were amplified and cloned in a pET21d vector. The respective C4 proteins were over-expressed in Escherichia coli, purified, and used as the antigens to raise specific sera against C4 proteins. The bacterially expressed C4 proteins were used as baits to identify the interacting host proteins of N. benthamiana using two dimensional- polyacrylamide gel electrophoresis (2D-PAGE), far-western blot analyses, and co-immunopricipitation assays. At least four host factors interacting with various C4 proteins were identified as membrane-bound proteins: chloroplast photosynthetic oxygen-evolving protein, 24K germin-like protein, plastidic aldolase, and photosystem I light-harvesting chlorophyll a/b-binding protein. Six additional host factors were identified as soluble proteins: Formin-like protein 19, Luminal-binding protein, NADH-ubiquinone oxidoreductase, 26S protease regulatory subunit 6A homolog B, MAR-binding filament-like protein 1-1 and Putative disease resistance protein RPP8-like protein. The biological functions of individual C4 proteins were further assayed in N. benthamiana by cloning the C4 genes of different begomoviruses in the transient expression vector, pBin35S, Bamboo mosaic virus satellite RNA expression vector, pCass-satBaMV, and the Bamboo mosaic virus expression vector, pCass-BaMV, followed by subsequent inoculations. Although C4 mRNA could be clearly detected, neither C4 proteins nor abnormal symptoms were observed in the inoculated plants, confirming that C4 gene functions at the protein level instead of the RNA level. Taken together, this study has identified several host factors involved in energy metabolism or cell proliferation which can interact with C4 proteins of different geminiviruses. It is expected that through the understanding of the underlying mechanisms, more insights could be provided into the development and differentiation of plant cells, and effective measures may be developed for the “curing” of diseased caused by geminiviruses.

前言 ………………………………………………………………………..…..……..1
前人研究………………………………………………………………………........…3
一、 雙生病毒簡介…………………………………………………………………3
(一) 經濟重要性…………………………………………...…………………3
(二) 分類地位………………………………………………...………………3
(三) 基因體構造………………………………………...……………………4
二、 影響葉片發育的植物因子……………………………………………………5
(一) 基因異常…………………………………………………..………….…5
(二) 小分子RNA異常……………………………………………..…………6
(三) 荷爾蒙異常……………………………………………….……………..6
三、 雙生病毒影響植物病徵表現的因子…………………………………………7
(一) 病毒基因體…………………………………………...…………………7
(二) 病毒組織向性…………………………………..………………...……10
四、 雙生病毒C4蛋白與宿主分子間的交互作用………………………..…..…11
五、 實驗室前人研究………………………………………………………..……12
六、 本研究目的……………………………………………………………..……12
材料與方法…………………………………………………………..………………14
一、 試驗材料……………………………………………………..…………..…..14
二、 在E. coli表現系統中誘導表現並純化C4蛋白……………………….……14
三、 利用遠西方墨點法,尋找與C4蛋白交互作用的宿主因子……………..…17
四、 利用免疫共沉澱,尋找與C4蛋白交互作用的宿主因子………………..…20
五、 利用不同的表現載體,在植物中大量表現C4蛋白,探討C4蛋白獨立存在的生物功能…………………………………………...……………..…….21
(一) 利用pBin19-35S載體表現C4蛋白………………………...……….21
(二) 利用pCass-satBaMV載體表現C4蛋白…………………….....……23
(三) 利用pCass-BaMV載體表現C4蛋白………………………...…..…25
結果……………………………………………………………………………..……27
一、 C4蛋白之表現與純化,並製作專一性抗血清………………………...……27
二、 尋找與C4蛋白交互作用的宿主因子…………………………………...….28
三、 利用不同的表現載體,在植物中大量表現C4蛋白,探討C4蛋白獨立存在的生物功能………………………………………………………….....….34
(一) 利用pBin19-35S載體在植物中表現C4蛋白……………….….…..34
(二) 利用pCass-satBaMV載體在植物中表現C4蛋白………….…....…36
(三) 利用pCass-BaMV載體在植物中表現C4蛋白………………….…38
討論…………………………………………………………………………......……41
一、 C4蛋白之表現與純化,並製作專一性抗血清……………………….....…..41
二、 與C4蛋白交互作用的宿主因子…………………………………………....42
三、 C4蛋白獨立存在的生物功能…………………………………………...…..49
四、 未來研究方向與應用…………………………………………………..……51
五、 結論…………………………………………………………………..………52
參考文獻…………………………………………………………………………..…53
圖表、附錄……………………………………………………………..…………..…63


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