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研究生:戴國偉
研究生(外文):kao-Wei Dai
論文名稱:雙生病毒中病徵決定因子之研究分析
論文名稱(外文):Studies on the symptom determinants of geminivirus
指導教授:胡仲祺
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
畢業學年度:96
語文別:中文
論文頁數:91
中文關鍵詞:雙生病毒嵌合病毒
外文關鍵詞:geminiviruschimeric virus
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豆類金黃嵌紋病毒屬之病毒為感染雙子葉植物的雙生病毒,是熱帶及亞熱帶地區主要的植物病毒,其病毒顆粒為雙球型,內含一條或兩條環狀單股DNA。當雙生病毒在感染植物後會使得已分化的植物細胞再次進入S期細胞循環,以利病毒本身的DNA進行複製,並因不同的病毒與寄主植物而產生不同類型的病徵。然而會造成植物發育異常,除了植物本身的因素以外,病毒也會造成植物的生長發育的異常,但是其病徵是由病毒的哪個蛋白所引起仍然眾說紛紜。先前本研究室發現菸草(Nicotiana benthamiana)在感染霍香薊黃脈病毒南投株(Ageratum Yellow Vein Virus Nan-Tou Strain, AYVV-NT)時會造成葉片上捲,而在感染番茄捲葉病毒(Tomato Leaf Curl Virus Tai- Nan strain, TLCV-TN)或南瓜捲葉病毒(Squash Leaf Curl Virus Yi-Lan strain, SqLCV-YL)時會造成葉片下捲,所以本實驗以AYVV和TLCV作為實驗材料,探討其基因體中是否含有影響病徵的蛋白或是序列。首先利用重組的方式,置換兩種病毒的基因體片段,再利用滾環複製放大反應(rolling circle amplification, RCA),來構築具有感染力的infectious clones,再利用已構築的infectious clones接種N. benthamiana,結果發現當置換了nt 1968~138這段區域後,病徵會隨著這個片段的改變而不同,因此可以知道這個區域是會影響病徵的位置,然而這個區域包含了五分之三的Rep蛋白、AC4、IR region,其中發現AC4與前人研究一致,是一個會影響病徵的因素。為進一步的研究AC4蛋白功能,本實驗利用序列比對分析發現AYVV-NT與TLCV-TN的AC4蛋白,在胺基酸序列上只有45 %的相似度,因此利用基因剔除的方式把AC4蛋白移除,來觀察病徵捲曲的方向,發現不論哪ㄧ個AC4蛋白被移除的嵌合病毒,都無法有捲曲病徵的表現,因此AC4蛋白極可能是影響捲曲方向的因子之ㄧ;然而從目前的結果顯示出nt 1968~138這段區域對於病徵有決定性的影響。另外本研究亦採用DNA shuffling的方式進行體外逢機重組,與混合接種的方式進行體內同源重組,以進一步探討雙生病毒造成病徵的機制,從混合接種的試驗發現病徵的捲曲方向,是由該種病毒的數量多寡所決定,與古典遺傳學相似的數量遺傳,優勢族群決定植物病徵的表現型;而在體外重組的部份仍需找尋最佳化條件,以利後續研究。因此未來將繼續以人工突變方式進行精確定位(fine mapping),以明瞭雙生病毒影響植物細胞分裂並造成捲葉病徵之分子機制。
Geminiviruses are characterized by circular single-stranded DNA genomes with one or two components which are encapsidated in geminate semi-icosahedral particles and replicate via a double-stranded DNA intermediate. Previous studies have revealed the ability of geminiviruses to alter the cell-cycle processes and inflict various type of symptoms on different host plants, indicating that geminiviruses possess the potential to interact with host factor and regulate the symptom expression in a species-specific manner. However, the responsive viral protein and the regulatory mechanism remained to be elucidated. It has been shown by our group that Nicotiana benthamiana plants infected with Ageratum yellow vein virus Nan-Tou Strain (AYVV-NT) display severe upward leaf curling symptoms. In contrast, the leaves of those infected by Tomato leaf curl virus Tai-Nan strain (TLCV-TN) or Squash leaf curl virus Yun-Lin strain (SqLCV-YL) curl downward. Therefore, the aim of this study is to identify and characterize sequences or genes in geminivirus genome invplved in symptom determination. Using the infectious clone of AYVV-NT and TLCV-TN as the experiment materials, recombinant viruses were constructed to locate the viral sequence or genes that influence the disease phenotype. In the first approach, chimeric viruses were constructed by exchanging fragments of the viral genomes generated by restriction enzymes. The resulted chimeric monomers were used as the templates to construct infectious clones using the simplified method developed previously in our laboratory. Results of the inoculation assays on N. benthamiana revealed that symptoms were determined by viral genome spanning nt 1968 ~ 138, which include the intergenic region (IR) 、AC4 gene and 5’-portion of Rep gene. By loss-of-function mutations, it was found that AC4 is likely involved in the leaf curl symptoms, although further fine mapping is required to identify the exact symptom determinant. The second approach is the co-inoculation of AYVV-NT and TLCV-TN in N. benthamiana plants. No recombinant virus was isolated from the co-inoculated plants. The result revealed that the direction of leaf curling was controlled by the difference in population sizes of the two viruses, similar to the control of phenotypes in classic quantitative genetics. The third approach is to generate random recombinants by DNA shuffling, by which no infectious recombinant virus was obtained to date. Collectively, the results revealed the primary viral determinants of the direction of leaf curling symptom. It is expected that through the use of the infectious recombinant geminiviruses, the underlying mechanism for the control symptoms will be elucidated.
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) 賀爾蒙異常 6
(三) 小分子RNA異常 7
三 、 雙生病毒影響病徵表現的因子 7
(一) 病毒基因體 7
(二) 病毒組織向性 12
四、 實驗室前人研究 13
五 、 本研究目的 13
材料與方法 14
一 、 試驗材料 14
二 、 試驗方法 14
(一) 基因體置換之試驗 14
(二) 構築具有感染力之質體 17
(三) 接種菸草( Nicotiana benthamiana )試驗 19
(四) 接種病毒的植物分析與檢測 20
(五) 南方墨點法分析 (Southern blot assay) 22
(六) 病毒外鞘蛋白表現之偵測 24
(七) 剔除(knockout)AC4基因試驗 25
(八) 體內重組試驗 27
(九) 體外重組試驗(DNA shuffling) 28
結果 30
一 、 AYVV與TLCV基因體片段交換 30
二 、構築具有感染力之嵌合病毒 31
三 、病徵研究與分析 31
四 、AC4基因剔除之分析 33
五 、天然重組研究與分析 34
六 、人為重組研究與分析 35
討論 37
一、病毒基因體置換不影響病毒複製 37
二、小分子RNA的影響 38
三、組織向性的影響 39
四、嵌合病毒的探討 40
五、體內重組之分析 41
六 、 結論 42
參考文獻 43
圖表與附錄 67
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