臺灣博碩士論文加值系統

竹嵌紋病毒 ( Bamboo mosaic virus, BaMV ) 為單股正極的RNA病毒，其會使竹葉產生嵌紋狀病徵。有的BaMV分離株還包含衛星核酸 (satellite RNA)，簡稱 satBaMV。SatBaMV為次病毒分子，必需依賴BaMV的幫助才能複製、移動及包被。由親源關係的分析發現，自然界的satBaMV分離株可概分為兩群，且在其5’端非轉譯區 (untranslated region, UTR)含有一高度變異區 (hypervariable region, HV region)。雖然此區核酸序列的歧異度高達20%，但大多能摺疊成一保守的二級結構，稱之為apical hairpin stem loop (AHSL)。有的satBaMV分離株會干擾BaMV的複製並減輕其在宿主植物所引起之病徵，例如BSL6；有的則影響不大，例如BSF4。而決定此生物特性之因子位於satBaMV 5’ UTR。在本研究中，我們進一步確認了BSL6干擾BaMV複製的主要決定因子位於其HV region，且與AHSL結構的維持有關。根據生物特性的分析結果，亦歸納出能夠干擾BaMV複製之satBaMV分離株在其AHSL上具有相同結構與序列的中環 (internal loop)。除了AHSL的結構外，中環上特定之核苷酸 (nucleotides) 在satBaMV干擾BaMV複製過程中扮演著重要的角色。分析satBaMV之間以及satBaMV與BaMV的競爭關係得知，具干擾能力之satBaMV有較大的競爭優勢。有趣的是，在BaMV 5’端亦發現與satBaMV類似AHSL之結構 (AHSL-like structure)。因此，我們推測AHSLs結構可能與BaMV及satBaMV的複製有關， BSL6則是藉由其AHSL結構對複製所需之因子的高競爭力，進而影響BaMV的複製。另外，以satBaMV為載體表現基因靜默抑制蛋白 (gene silencing suppressor) 並不影響其干擾BaMV複製的能力。

Satellite RNAs associated with Bamboo mosaic virus (satBaMV) are subviral agents depending on BaMV for replication, encapsidation and systemic movement. Natural satBaMV isolates can be categorized into 2 major phylogenetic groups. A hypervariable (HV) region with divergence of up to 20% in the 5’ untranslated region (UTR) of satBaMV isolates, folds into a conserved apical hairpin stem loop (AHSL) structure containing an apical loop and 2 internal loops. Among the satBaMV isolates, BSL6 reduces the accumulation of BaMV RNAs and attenuates the BaMV-induced symptoms in co-inoculated plants, whereas BSF4 does not. The determinants of the downregulation of BaMV replication were mapped to the 5’ UTR of BSL6. In this study, we confirmed the predicted secondary structure of BSL6 5’ UTR by enzymatic probing and demonstrated that the major determinants of satellite RNA-mediated interference were in the HV region. The integrity of the AHSL structure of interfering satBaMV was essential for the interference of BaMV accumulation. Concurrent analyses of natural satBaMV isolates revealed that all of the interfering isolates contained the same structures and sequences in the internal loops. Refined analyses indicated that, besides the AHSL structure, C60 and 81UGC83 in the internal loops play a crucial role in the downregulation. Given that interfering satBaMV is more competent than noninterfering satBaMV and BaMV during replication, and the 5’ end of BaMVs harboring satBaMV co-evolved the AHSL-like structure with satBaMV, we propose that the 5’ AHSL of BSL6 may be a potent determinant for recruiting the limited quantities of cellular and/or viral factors required for the replication of BaMV and satBaMV. Moreover, the downregulation event by satBaMV was not affected by a viral gene-silencing suppressor inserted into BaMV or satBaMV RNA.

正文目錄 (Contents)
中文摘要 ………………………………………………………………………… 1
英文摘要 (Abstract) …………………………………………………………… 2
緒言 (Introduction) …………………………………………………………… 3
材料與方法 (Materials and Methods) …………………………………………… 10
結果 (Results)
Section I. Downregulation of BaMV requires the 5’ AHSL structure and sequence of satBaMV. …………………………………………………………………… 21
Section II. Replication competence of BaMV and satBaMV variants requires the conserved AHSL structure in their 5’ UTR. …………………………………… 36
Section III. SatBaMV-mediated interference with BaMV replication in the presence of viral suppressor …………………………………………………………… 43
討論 (Discussion) ……………………………………………………………… 45
參考文獻 (References) …………………………………………………………… 52

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