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|標題:||The effect of NbUbE3R1 on the infection cycle of Bamboo mosaic virus in Nicotiana benthamiana
菸草差異性表現基因 NbUbE3R1 對於竹嵌紋病毒生命週期的影響
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當病毒感染植物時，可能影響寄主基因的表現，了解寄主基因在病毒的生活史中所扮演的角色，可有助於抗病毒技術的發展，在前人的研究中利用了 cDNA amplified fragment polymorphism (AFLP)之技術篩選出一些在竹嵌紋病毒感染菸草(Nicotiana benthamiana)時，有差異性表現的基因片段。其中一個基因片段ACGT2-1 在感染病毒後的 5 至 7 天可見基因表現量的增加，而利用病毒引發基因靜默技術(virus-induced gene silencing，VIGS)使得 ACGT2-1 基因表現量下降後 可發現竹嵌紋病毒外鞘蛋白在葉片的累積量顯著增加至 135% 推測 ACGT2-1，基因可能與病毒的防禦機制有關。接著想進一步了解 ACGT2-1 基因是參與在病毒的複製還是移動的階段，因此將處理 VIGS 後的原生質體接種病毒，發現在竹嵌紋病毒外鞘蛋白的累積量增加，然而利用北方墨點法偵測發現病毒正股及負股RNA 均有顯著增加的情形，顯示 ACGT2-1 可能參與病毒早期複製之階段。另外利用 Rapid amplification of cDNA ends (RACE)技術獲得了 ACGT2-1 的全長基因 NbACGT2-1 經由其序列比對後 發現它具有 RING domain 以及 transmembrane，region，已有研究指出 RING domain 可作為 E2 轉移泛蛋白給受質的一個平台，於細胞調節 protein ubiquitination 扮演重要角色，我們將之命名為 NbUbE3R1,若將此基因於菸草植物過量表現後，再進行 BaMV 的感染，發現竹嵌紋病毒外鞘蛋白的累積量減少 以上實驗結果明確的顯示此基因扮演了抑制竹嵌紋病毒的功能，而當此蛋白去除transmembrane region 後，可發現竹嵌紋病毒外鞘蛋白的累積量增加，失去限制病毒之功能。而 NbUbE3R1 如何與病毒蛋白結合使得具有防禦的功能，以及其進行防禦的機制，則有待進一步的研究了解。
The expression profile of host genes could be altered by viral infection. To understand how these differentially expressed host genes during virus infection involved in viral infection cycle could help the development of antiviral strategies. In a previous study, our lab has identified 90 differentially expressed genes in Bamboo
mosaic virus (BaMV)-inoculated plants by cDNA-AFLP screening method. One of the upregulated genes ACGT2-1 was further characterized. Tobacco rattle virus (TRV)-based virus induced gene silencing (VIGS) was used to knock down ACGT2-1 expression in Nicotiana benthamiana plant. The results showed that coat protein accumulation of BaMV was enhanced to 135%. This result suggests that ACGT2-1 might play a defense role against BaMV infection. Furthermore, to determine if this gene displayed interference in BaMV replication or cell-to-cell movement, BaMV RNA was inoculated into protoplasts derived from the knockdown and control plants.
In the plasmodesmata-removal system the coat protein accumulation was enhanced compared to that of the control protoplasts. Northern blot analysis indicated that the
accumulation of BaMV plus- and minus- strand RNAs was also enhanced. The results demonstrated that ACGT2-1 might be involved in the replication step of BaMV
infection cycle. In addition, rapid amplification of cDNA ends (RACE) technique was used to obtain the full-length of ACGT2-1. Furthermore, the accumulation of BaMV was reduced when ACGT2-1 was transiently expressed in plants. This result clearly demonstrates that ACGT2-1 plays a role in inhibiting the accumulation of BaMV. BLAST analysis of the ACGT2-1 showed that the ORF contained a RING domain and transmembrane region. The RING domain-containing proteins were demonstrated to act as scaffolds and promote the direct transfer of ubiquitin from the E2 conjugating enzyme to the substrate, and also play an important role in cell modulating protein ubiquitination. ACGT2-1 could be a putative C3HC4-type zinc finger ubiquitin E3 ligase; we then designated this protein as NbUbE3R1. Further studies will focus on whether NbUbE3R1 can interact with the viral protein to inhibit its function and the possible mechanism for the defense.
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