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dc.description.abstractYWHAZ, also known as 14-3-3ζ, belongs to the 14-3-3 family of proteins which mediate signal transduction by binding to phosphoserine-containing proteins. It's believed that YWHAZ plays an important role in cell signaling, cell cycle, and apoptotic death. YWHAZ is one of the major transforming growth factor-β-induced proteins, which can promote epithelial–mesenchymal transition of epithelial cells in cancer cell transformation. In our previous studies, we found that YWHAZ associates with TCF-4 and β-catenin in the nucleus and further promote EMT, cell invasion and migration by activating the YWHAZ/β-catenin axis in lung cancer. In order to investigate downstream genes of YWHAZ in lung cancer, Microarray and ChIP-seq assay were used to select candidate genes which may be regulated by YWHAZ. First, the genes that were up- or down-regulated by YWHAZ more than 1.5-fold and simultaneously contain TCF-4 binding motif in their 5'-upstream regulatory region determined by ChIP-seq were identified. We obtained fourteen candidate genes that might be regulated by YWHAZ. Next, we found that mRNA expression of nine candidate genes were regulated by knockdown of YWHAZ with siRNA or short hairpin RNA (shRNA) and recovered by re-expression of YWHAZ in the stable clones that harbor shRNA in CL1-5 cell line. Moreover, chromatin immunoprecipitation and quantitative PCR (ChIP-qPCR) assays demonstrated a significant interaction between the YWHAZ-β-catenin-TCF-4 complex and four candidate genes: ATF3, DMTF1, DUSP1 and PAK2. Electrophoretic mobility shift assay (EMSA) was performed that protein recognized and bound the TCF-4 binding motif on DNA of the promoter of four candidate genes. Taken together, our data reveal that YWHAZ can regulate these candidate genes through forming the regulatory complex with TCF-4 and β-catenin.en_US
dc.description.abstractYWHAZ蛋白,又稱為14-3-3ζ,屬於14-3-3蛋白家族的一員。14-3-3蛋白家族會藉由與含有磷酸化serine的蛋白(phosphoserine-containig protein)結合,來調控細胞的訊號傳遞(signal transduction)與生理功能。YWHAZ蛋白又為轉型生長因子β誘導蛋白(transforming growth factor-β-induced protein)之一,可促進癌細胞上皮細胞轉型間葉細胞過程(epithelial-mesenchymal transition;EMT)的發生,促使癌轉移的產生。在實驗室先前的研究發現,YWHAZ蛋白會與β-catenin蛋白和轉錄因子T-cell factor 4 (TCF4)在核中結合,形成一複合體(complex),並促進EMT、癌細胞侵襲(invasion)和移動(migration)的發生。為了進一步研究在肺癌細胞中是否有其他基因受YWHAZ所調控,進而促進肺癌進程(lung cancer progression)的發生,我們利用微陣列分析(microarray)和染色質免疫沉澱配合高通量定序(Chromatin immunoprecipitation sequencing;ChIP-seq))去尋找可能受調控的基因。首先,我們挑選出14個有潛力的基因。接著,我們挑選能穩定靜默YWHAZ蛋白表現之細胞株,並進行細胞移動(migration)與侵襲(invasion)能力測試,發現YWHAZ的抑制會降低肺癌細胞移動與侵襲能力。再利用此細胞株進行即時定量聚合酶連鎖反應(QPCR)並與微陣列分析(microarray)結果比對後,確認有9個可能受YWHAZ調控之基因。而且在靜默YWHAZ表現之細胞株中再次表現YWHAZ後,可能受調控之基因表現有回復(recover)之情形。此外,進行染色質免疫沉澱配合即時定量聚合酶連鎖反應(ChIP-QPCR)實驗發現,ATF3、DMTF1、DUSP1和PAK2確實與YWHAZ-β-catenin-TCF-4蛋白複合體有交互作用;電泳移動分析結果顯示,ATF3、DMTF1、DUSP1和PAK2基因的啟動子上確實有TCF-4的結合序列。綜合以上結果,YWHAZ蛋白會經由與β-catenin和TCF-4蛋白形成的複合體來調控下游基因之表現。zh_TW
dc.description.tableofcontents目錄 中文摘要 i 英文摘要 ii 目錄 iii 實驗結果表圖目錄 vi 第一章 序論 1 第一節 前人研究 1 一、肺癌 1 二、癌轉移 3 三、YWHAZ蛋白 4 四、YWHAZ-β-catenin-TCF-4複合體調控之下游目標基因篩選 9 第二節 研究目的及架構 12 第二章 實驗材料與方法 14 第一節 材料 14 第二節 細胞株及細胞培養 14 第三節 細胞冷凍保存及解凍 15 第四節 細胞蛋白質之萃取與定量 15 第五節 免疫沉澱 16 第六節 蛋白質凝膠電泳分析 17 第七節 西方墨點法 18 第八節 細胞RNA萃取 19 第九節 反轉錄聚合酶連鎖反應 20 第十節 聚合酶連鎖反應 20 第十一節 質體DNA萃取 21 第十二節 DNA片段回收之純化與黏接反應 21 第十三節 勝任細胞製備與轉形作用 22 第十四節 菌種甘油保存 23 第十五節 菌落聚合酶連鎖反應 23 第十六節 大量質體萃取 23 第十七節 轉染作用 24 第十八節 慢病毒屬感染 25 第十九節 加藥篩選建立穩定表現細胞株 26 第二十節 基因靜默 26 第二十一節 利用生物資訊方法篩選出潛在的YWHAZ-TCF4蛋白 複合體調控目標基因 27 第二十二節 染色質免疫沉澱分析 35 第二十三節 細胞侵襲能力分析 36 第二十四節 細胞遷移能力分析 37 第二十五節 分離萃取核質蛋白 37 第二十六節 電泳移動分析 38 第二十七節 統計分析結果 40 第三章 實驗結果 41 第一節 於CL1-5中靜默YWHAZ表現並建立穩定靜默細胞株 41 第二節 觀察靜默YWHAZ表現對肺癌細胞型態的影響 41 第三節 靜默YWHAZ表現能降低肺癌細胞的移動能力 41 第四節 靜默YWHAZ表現能降低肺癌細胞的侵襲能力 42 第五節 篩選受YWHAZ-TCF-4蛋白複合體調控之下游目標基 因 42 第六節 確認下游目標基因在YWHAZ shRNA作用的細胞株內 的訊息核醣核酸 43 第七節 確認下游目標基因在YWHAZ siRNA作用的細胞株內 的訊息核醣核酸 44 第八節 重新表現YWHAZ基因可回復受YWHAZ靜默影響之 下游基因 44 第九節 重新表現YWHAZ能再次促進肺癌細胞移動的能力 45 第十節 重新表現YWHAZ能再次促進肺癌細胞侵襲的能力 46 第十一節 上調控基因個別與YWHAZ、β-catenin和TCF-4蛋白有 交互作用 46 第十二節 下調控基因個別與YWHAZ、β-catenin和TCF-4蛋白有 交互作用 47 第十三節 上調控基因與YWHAZ-β-catenin-TCF-4蛋白複合體有 交互作用 47 第十四節 下調控基因與YWHAZ-β-catenin-TCF-4蛋白複合體有 交互作用 48 第十五節 ATF3、DMTF1、DUSP1和PAK2基因的啟動子上確實 有TCF-4的結合序列 48 第四章 討論 50 第五章 結論 56 實驗結果表圖 57 附表 79 附圖 82 附錄 84 參考文獻 88zh_TW
dc.subjectLUNG CANCERen_US
dc.titleStudy of the role of YWHAZ in lung cancer and identification of its downstream genesen_US
dc.typeThesis and Dissertationen_US
item.fulltextwith fulltext-
item.openairetypeThesis and Dissertation-
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