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研究生:劉命如
研究生(外文):Minq-Ru Liou
論文名稱:紅龍果嵌紋病毒生物特性及全長度核酸序列之研究
論文名稱(外文):Characterization of Pitaya Mosaic Virus ans Analysis of Its Full-length Sequence
指導教授:劉瑞芬劉瑞芬引用關係
指導教授(外文):Ruey-Fen Liou
學位類別:碩士
校院名稱:國立臺灣大學
系所名稱:植物病理學研究所
學門:農業科學學門
學類:植物保護學類
論文種類:學術論文
論文出版年:2000
畢業學年度:88
語文別:中文
論文頁數:89
中文關鍵詞:紅龍果嵌紋病毒生物特性全長度核酸序列紅龍果
外文關鍵詞:Pitaya Mosaic VirusBiologyFull-length SequenceHylocereus undatus
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紅龍果(Hylocereus undatus Britt. & Rose)屬於仙人掌科(Cactaceae),三角柱屬(Hylocereus)之多年生攀緣性肉質植物,為極具經濟價值之新興熱帶栽培果樹,也常做為庭園美化、綠籬及砧木使用。國內外並無相關報告提到紅龍果病毒病害之發生,本實驗於1999年,在新竹縣關西鎮亞森觀光農場,發現紅龍果之三角莖上有淡綠色嵌紋病徵,初步由電子顯微鏡觀察,發現有絲狀病毒粒子存在,長度約為483~517 nm。病原病毒可經機械及扦插繁殖方式傳播,以汁液接種法接種至紅藜(Chenopodium amaranticolor)並經三次單斑分離後,再接種至七科二十一種指示植物上,發現會在雞冠花(Celosia cristata)、千日紅(Gomphrena globosa)及奎藜(Chenopodium quinoa)之葉片產生局部性病斑,回接至紅龍果也產生先前所觀察到之嵌紋病徵。自奎藜病葉純化之病毒粒子,其A260/A280之比值為1.20。以純化病毒為免疫抗原,施打於紐西蘭大白兔製備多元抗體,所得之抗體除了用於建立酵素聯結免疫吸附反應(enzyme-linked immunosorbent assay, ELISA)外,還用以對Bamboo mosaic virus (BaMV)、Cactus virus X (CVX)及Papaya mosaic virus (PapMV)等potexviruses進行免疫擴散反應,結果顯示此病毒會與CVX抗血清及紅龍果嵌紋病毒抗血清產生白色的沉澱帶。純化病毒經12 % SDS-聚丙醯胺膠體電泳分析,鞘蛋白分子量約為26 kDa。由純化病毒萃取viral RNA,經1.2 %變性瓊脂膠體電泳分析,可觀察到一條RNA亮帶,估算其基因組大小約為6-7 kb。以上各項實驗結果顯示,自紅龍果三角莖上所分離到的病毒病原,極可能屬於Potexvirus,並將其暫時命名為紅龍果嵌紋病毒(Pitaya mosaic virus, PtMV)。為了進一步探討病毒的分子特性,並嘗試構築病毒的infectious clone,乃利用rapid amplification of cDNA ends (RACE)進行病毒全長度核酸序列選殖與分析。目前已經確定之PtMV病毒核酸序列共包含6605個核酸(不包含poly (A) tail),5’端非轉譯區(untranslated regions, UTR)有65個核酸,尚缺在5’端一小段序列,3’-UTR有101個核酸。PtMV之核酸序列主要含有5個開放解讀框(open reading frame, ORF),分別是ORF1 (175 kDa)、ORF2 (25 kDa)、ORF3 (12 kDa)、ORF4 (7 kDa)、ORF5 (24 kDa),其中ORF1含有病毒複製所需之replicase之功能基,ORF2、ORF3及ORF4組成triple gene block,與病毒移行所需的相關基因類似,ORF5則為鞘蛋白基因。在ORF1中還發現ORF6 (15 kDa)及ORF7 (8 kDa)兩個開放解讀框。各個ORF與BaMV O strain、Cymbidium mosaic virus (CymMV)、Foxtail mosaic virus (FoMV)、PapMV、Plantago asiatica mosaic virus (PlAMV)及PVX HB strain等Potexvirus屬病毒之基酸序列都有相當之保守性。以PtMV之ORF1及ORF5之基酸序列,分別與上述6種Potexvirus屬病毒做類緣關係分析,結果均顯示與PapMV病毒最接近。
Pitaya (Hylocereus undatus Britt. & Rose), which belongs to Cactaceae, has become an important tropic fruit in Taiwan in recent years. Pitaya showing mosaic symptoms on the pads was collected from fields in the Kawnshi area in 1999. Indicator plant tests indicated that the causal agent of the mosaic symptoms was mechanically transmissible, and local lesions developed on the leaves of Chenopodium amaranticolor 5 days after inoculation. After three successive single lesion transfers on the same plants, a virus isolate was obtained. In addition to C. amaranticolor, this virus is capable of infecting C. quinoa, Celosia argentea, and Gomphrena globosa. A back inoculation assay was performed to ensure that the virus, named Pitaya mosaic virus (PtMV) was indeed the causal agent of the mosaic symptom observed on pitaya. Electron microscopic examination of negatively stained virus particles from pad extracts of the diseased plants revealed that the virus has a flexuous rod-shaped morphology with a length of 483 to 517 nm. Purified viruses had an A260/A280 ratio of 1.20. Antiserum raised against PtMV reacted positively with infected plants and was used for detection of PtMV by ELISA. The molecular mass of the coat protein (CP) was estimated to be 26 kDa by SDS-polyacylamide gel electrophoresis. Analysis of viral RNA by formaldehyde agarose gel electrophoresis revealed the presence of a single species of RNA approximately 6-7 kb in length. Molecular cloning of the viral genome was subsequently carried out by rapid amplification of cDNA ends (RACE). Sequences of PtMV analyzed so far contained 6605 nucleotides (excluding 3’ poly(A) tail), which included a total of 7 putative open reading frames (ORF). Of the 7 ORFs being identified, ORF1 encodes RNA dependent RNA polymerase (RdRp), ORF2, ORF3, and ORF4 encode proteins which are involved in virus movement, ORF5 encodes coat protein. Each of these proteins displays significant homology to the corresponding ORFs of other potexviruses, indicating that genome organization of PtMV is basically similar as those of other potexviruses. Phylogenetic analysis based on multiple sequence alignments of RdRp and CP reveals a closest relationship of PtMV to Papaya mosaic virus.
中文摘要…………………………………………………………………1
英文摘要…………………………………………………………………3
壹、前言…………………………………………………………………5
貳、前人研究……………………………………………………………7
一、仙人掌科植物相關之病毒病害……………………………………7
(一)、Cactus virus 2 (CV-2)……………………………………7
(二)、Cactus virus X (CVX) ……………………………………7
(三)、Saguaro cactus virus (SgCV)……………………………8
(四)、Zygocactus symptomless virus (ZSLV)…………………9
二、 Potexvirus屬病毒之特性 ………………………………………10
(一)、5’端非轉譯區………………………………………………11
(二)、3’端非轉譯區………………………………………………11
(三)、開放解讀框1…………………………………………………12
(四)、開放解讀框2、3及4…………………………………………13
(五)、開放解讀框5…………………………………………………13
(六)、開放解讀框6…………………………………………………14
參、材料與方法…………………………………………………………15
一、病毒來源………………………………………………………15
二、病毒之分離……………………………………………………15
三、指示植物接種試驗……………………………………………15
四、電子顯微鏡觀察………………………………………………16
五、病毒純化………………………………………………………16
六、血清學試驗……………………………………………………17
(一)、抗血清的製備………………………………………………17
(二)、病毒免疫球蛋白(Immunoglobulin G, IgG)之純化 ……17
(三)、一次抗體結合體(IgG conjugate)之製備 ………………18
(四)、免疫擴散反應(Immunodiffusion) ………………………18
(五)、免疫電顯(Immuno-electron microscopy) ……………18
(六)、雙三明治抗體酵素聯結免疫反應(DAS-ELISA) …………19
七、西方轉漬法分析鞘蛋白大小…………………………………20
(一)、SDS-聚丙醯膠體電泳分析(Sodium dodecyl sulfate polyacrylamide gel electrophoresis, SDS-PAGE)………………20
(二)、Coomassie blue染色分析…………………………………20
(三)、西方轉漬分析法(Western blotting analysis) ………21
八、病毒全核醣核酸之抽取………………………………………21
九、北方雜合分析(Northern hybridization analysis) ……22
(一)、RNA電泳分析 ………………………………………………22
(二)、RNA轉漬 ……………………………………………………23
(三)、核酸探針製備………………………………………………23
(四)、前置雜合反應(Prehybridization)………………………23
(五)、雜合反應(Hybridization) ………………………………24
(六)、以酵素連結免疫反應與鹼性去磷酸呈色反應偵測雜合訊息 ……………………………………………………………24
十、基因全序列的選殖………………………………………………25
(一)、第一股合成…………………………………………………25
(二)、第二股合成…………………………………………………25
(三)、Adaptor ligation…………………………………………26
(四)、5’RACE及3’RACE…………………………………………26
(五)、回收PCR擴增結果之DNA …………………………………27
(六)、將PCR擴增產物選殖至T-vector …………………………27
(七)、質體小量製備………………………………………………28
(八)、核酸定序及序列分析………………………………………29
肆、結果…………………………………………………………………30
一、紅龍果被感染病毒之病徵 ……………………………………30
二、指示植物接種試驗 ……………………………………………30
三、紅龍果嵌紋病毒之純化 ………………………………………31
四、血清學試驗 ……………………………………………………31
(一)、免疫擴散反應………………………………………………31
(二)、免疫電顯……………………………………………………31
(三)、雙三明治抗體酵素聯結免疫反應(DAS-ELISA) …………32
五、以西方轉漬法分析病毒鞘蛋白…………………………………32
六、以甲醛變性瓊脂膠體電泳分析病毒核酸………………………32
七、病毒全長度核酸序列選殖與定序………………………………33
(一)、全長度核酸序列選殖結果…………………………………33
(二)、全長度核酸序列定序結果…………………………………34
(三)、以北方雜合法分析病毒核酸………………………………36
伍、討論…………………………………………………………………37
陸、參考文獻……………………………………………………………41
柒、圖……………………………………………………………………54
捌、表……………………………………………………………………119
附錄一、試劑配方………………………………………………………123
附錄二、引子序列………………………………………………………129
附錄三、病毒之命名原則………………………………………………130
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