Please use this identifier to cite or link to this item: http://hdl.handle.net/11455/90083
標題: The study on the viral RNA intracellular trafficking and the involvement of autophagy pathway in the infection of Bamboo mosaic virus in Nicotiana benthamiana
追蹤竹嵌紋病毒在細胞內的移動以及自噬作用對竹嵌紋病毒的影響之探討
作者: 黃盈屏
Ying-Ping Huang
關鍵字: 竹嵌紋病毒;細胞內移動;自噬作用;Bamboo mosaic virus;Intracellular trafficking;Autophagy
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摘要: 
在研究植物病毒時,我們首先關切的幾項議題不外乎是病毒的感染、複製、移動以及病毒和宿主之間的相互作用。因為竹嵌紋病毒的感染會造成國內竹筍產量削減,使得農產經濟降低,藉由研究竹嵌紋病毒與宿主之間的相關性,幫助我們更加了解竹嵌紋病毒感染的機制。目前已知竹嵌紋病毒是屬於Flexivividae科Potexvirus屬,為單一正股的RNA 病毒,除了基因體核酸之外,我們還可以偵測到兩個主要的次基因體核酸,並且可以偵測到衛星核酸。本篇論文主要針對病毒在細胞中的移動受到宿主蛋白的調控並且追蹤病毒在細胞中的所在位置,讓我們了解病毒複製時的位置。另外,病毒感染時是否刺激宿主產生防禦機制來抑制病毒複製與移動也是本篇論文想要探討的議題。
竹嵌紋病毒RNA於細胞內的位置,已經利用免疫金標定核酸的定位法發現病毒會座落在細胞核、粒腺體、葉綠體中,但還是無法確認病毒核酸是座落在細胞的何處進行複製。由於之前發現竹嵌紋病毒的3端會與葉綠體磷酸甘油酸激酶(phosphoglycerate kinase; PGK)結合,這更增加了竹嵌紋病毒座落在葉綠體的可能性。因此在這個研究中,我們利用噬菌體MS2品系所產生的MS2鞘蛋白會專一性與MS2核酸序列結合的特性,將病毒核酸重組上MS2序列,然後在細胞內共同表現病毒核酸以及MS2鞘蛋白-核蛋白-螢光蛋白複合體,利用共軛焦顯微鏡觀察病毒的位置,若是MS2鞘蛋白沒有和MS2核酸結合並且移動到病毒所在的位置,那麼螢光蛋白則會座落於細胞核中。實驗中,我們構築了MS2序列插入BaMV於複製酵素與移動蛋白基因之間的突變病毒,另外也重新構築含進核訊息-MS2鞘蛋白-螢光蛋白的融合蛋白。實驗結果發現,我們確實可以在葉綠體中偵測到螢光的訊號,另外由抽取病毒感染葉子的葉綠體,我們也可以偵測到病毒的正負股核酸基因體,這些結果更有利於證實竹嵌紋病毒核酸基因體進入葉綠體中複製的可能性。
當植物遭受非生物性或是生物性的逆境時,自噬作用扮演了將細胞中養分重新分解再利用的重要功能或提供適當的防禦機制。然而有些病毒卻可以逃脫自噬作用,甚至是利用自噬作用來幫助它們在細胞中複製。這裡我們想探討竹嵌紋病毒受到自噬作用的影響。當竹嵌紋病毒感染植物時,我們發現植物自噬作用相關基因的表現量都增加了,顯示病毒感染時有可能會誘導自噬作用。當我們使用抑制劑(wortmannin和3-MA)抑制自噬作用時,竹嵌紋病毒的鞘蛋白及病毒核酸累積量都下降了。然而,處理3-MA時,馬鈴薯病毒X胡瓜嵌紋病毒及菸草嵌紋病毒的鞘蛋白表現量均不受影響,這顯示3-MA藉由抑制第三型PI3Ks而影響自噬體的後期成形會影響竹嵌紋病毒在細胞中的累積。接著,我們發現在自噬作用相關基因NbATG8靜默的植物中,與控制組相較下,竹嵌紋病毒的鞘蛋白及病毒核酸在1357天的表現量降低了,而在NbATG8靜默的原生質體中,竹嵌紋病毒的鞘蛋白及病毒核酸在24小時的累積量也降低了。另外,在NbATG8大量表現的原生質體中,竹嵌紋病毒的鞘蛋白會增加,與靜默的實驗結果相符。這些結果顯示,自噬體對於竹嵌紋病毒的複製具正向調控。然而其詳細的機制仍有待釐清。

The cDNA-amplified fragment length polymorphism technique was applied to isolate the differentially expressed genes during Bamboo mosaic virus (BaMV), a single-stranded positive sense RNA virus, infection on Nicotiana benthamiana plants. One of the upregulated genes was cloned and predicted to contain a TBC domain designated as NbRabGAP1 (Rab GTPase activation protein 1). No significant difference was observed in BaMV accumulation in the NbRabGAP1-knockdown and the control protoplasts. However, BaMV accumulation was 50% and 2% in the inoculated and systemic leaves, respectively, of the knockdown plants compared to those of the control plants. By measuring the spreading area of BaMV infection foci in the inoculated leaves, we found that BaMV moved less efficiently in the NbRabGAP1-knockdown plants than in the control plants. Transient Over-expression of the wild type NbRabGAP1, but not the disable NbRabGAP1 activity mutant, significantly increaseincreases BaMV accumulation in N. benthamiana. These results suggest that NbRabGAP1 with a functional Rab-GAP activity is involved in virus movement.
Since chloroplast pPhosphoglycerate kKinase (PGK) was reported to interact with the very 3' end of BaMV RNA, it displayed a high possibility of viral RNA localized at chloroplasts. To observe the localization of viral RNA in infected cells, we employed the interaction of NLS-GFP-MS2 (phage MS2 coat protein) with the modified BaMV RNA containing the MS2 coat protein binding sequence in live plant cells. In the experiment, we have constructed the eight repeats of MS2 binding sequence at the intergenic region of between BaMV replication gene and the movement protein gene. Furthermore, we have reconstructed the fluorescence fusion protein to display a stronger nucleus location signaling (MS2-NLS-GFP was changed to NLS-MS2-OFP) in cells. Using confocal microscopy, we observed that the genomic RNA of BaMV was localized at the chloroplasts of the infected cells. Furthermore, we can also detect the minus-strand RNA in the isolated chloroplasts from BaMV infected leaves. Taken together, these results suggested that BaMV genomic RNA targeting to the chloroplasts as a replicate site is the most recommended.
Autophagy plays an important role in nutrient recycling for plant homeostasis while plants suffer the abiotic or biotic stresses including the response to the pathogen infection. In this study, we would like to investigate the effect of autophagy on the infection cycle of BaMV. Initially, we examined the relative gene expression of some autophagy-associated genes (ATGs) and found out that these genes were upregulated post BaMV infection. The coat protein and viral RNA accumulation levels of BaMV were reduced when the cells were treated with the autophagy inhibitors, wortmannin and 3-MAMethyladenine. Nevertheless, the coat protein accumulation of PVXPotato virus X, CMVucumber mosaic virus and TMVobacco mosaic virus was also decreased when treated with wortmannin but not with 3-MA, an inhibitor through blocking the classIII PI3Ks which was critical during the late stage of vesicle expansion. Furthermore, we also found that the coat protein and viral RNA accumulation levels of BaMV were reduced in the NbATG8-silenced protoplasts and plants. As well, the accumulation of BaMV coat protein was increased in NbATG8-transiently expressed protoplasts. Altogether, these results suggest that BaMV utilizes the autophagosome for replication in some way albeit the overall mechanism of the interaction between autophagosome and virus was still unclear.
URI: http://hdl.handle.net/11455/90083
Rights: 同意授權瀏覽/列印電子全文服務,2018-07-29起公開。
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