臺灣博碩士論文加值系統

端粒是真核染色體末端的特殊結構，協助DNA的完整複製、保護染色體不被nuclease分解，以及不被細胞的DNA確認點系統視為DNA損害。酵母菌溫度敏感變異株cdc13-1在30度C時，具有不正常的單股端粒累積，且細胞週期會被停止在G2/M期，而Gbp2，一個在in vitro狀態下可與單股端粒結合的蛋白，在過量表現時可以彌補cdc13-1在30度C時的生長缺陷，並恢復cdc13-1細胞的端粒至正常株相仿的長度。為了尋找Gbp2影響端粒的機制，我們利用酵母菌雙雜交系統篩選到三個能與Gbp2作用的蛋白，分別是Hrp1、Nfi1和Rlr1，而Hrp1與Rlr1在in vitro狀態下可與Gbp2直接作用。分析這些蛋白對端粒機制的相關表型影響，發現Gbp2與Hrp1的過量表現可以影響端粒位置效應，且Gbp2的單股端粒結合能力對於端粒位置效應的影響是需要的；而RLR1刪除株則呈現端粒變長的情形。進ㄧ步分析GBP2與HRP1之間的相關性，發現gbp2刪除株與hrp1-8、hrp1-7在25度C時具有近合成致死的關係，而這樣的生長缺陷並非來自於RNA的合成以及輸出上的缺失。分析gbp2刪除株與hrp1的單一變異株以及雙重變異株後，發現gbp2刪除株結合了hrp1的變異後，對於端粒、HM loci以及rDNA三處的緘默效應的影響力不同。而從已知的Sir2、Sir3以及Sir4在染色體上的分布及對緘默效應的調控，顯示細胞同時具有gbp2刪除株與hrp1的變異時，可能會影響Sir蛋白的分布，而改變了緘默效應之間的調節。而由Rap1與Sir蛋白群間的相關性、GBP2刪除株會改變Rap1蛋白在細胞核內的分布情形，以及Gbp2和Hrp1過量表現會專一性影響端粒位置效應，表示Gbp2和Hrp1可能是透過Rap1與Sir蛋白群而影響端粒機制。

The telomeres of Saccharomyces cerevisiae are composed of ~250-300 bp double-stranded (TG1-3)n/(C1-3A)n and a single-stranded TG1-3 tail. They maintain chromosome integrity and stability. Gbp2p is identified as a single-strand TG1-3 DNA binding protein in vitro. It affects telomere localization since the localization of Rap1p in nuclei was altered in cells deleted of GBP2. Gbp2p overexpression restores the growth arrest and telomere length phenotype in cdc13-1, a temperature-sensitive mutant of an essential telomere-binding protein, Cdc13p. In gbp2-deleted cells, however, the telomere length and telomere position effect are not affected. To elucidate the role of Gbp2p in telomere function, a yeast two-hybrid system was used to identify its interacting proteins. Hrp1p was identified that interacted with Gbp2p and bound directly in vitro. Hrp1p is a nuclear polyadenylated RNA-binding protein involving in the RNA processing. Mutations on both GBP2 and HRP1 caused growth defect, further supports the interactions between these two proteins. However, mutations on either GBP2 and/or HRP1 did not affect telomere length, suggesting that these two genes are not involved in telomere length regulation. The silencing loci of yeast, including telomere silencing locus, were affected differently in gbp2 hrp1 mutants. These effects were not caused by RNA processing since the localization of mRNA or the level of several mRNAs were not affected in gbp2 and/or hrp1 mutants. Together, our results define a novel role of Gbp2 and Hrp1p in modulating silencing loci of yeast cells.

目錄 頁次
目錄……………………………………………………………I
圖次目錄………………………………………………………II
附表目次………………………………………………………IV
縮寫表…………………………………………………………V
中文摘要………………………………………………………1
英文摘要………………………………………………………2
緒論……………………………………………………………3
材料方法………………………………………………………13
實驗結果………………………………………………………35
討論……………………………………………………………56
參考文獻………………………………………………………64
附圖……………………………………………………………73
附表……………………………………………………………105
附錄……………………………………………………………108

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