臺灣博碩士論文加值系統

Adducin（ADD）為細胞骨架蛋白肌動蛋白絲的結合蛋白，位於 spectrin-actin 交界處。其家族成員由 α、β及γ 三個相似的基因轉錄而成。ADD 蛋白間具有非常相似的胺基酸序列，ADD 蛋白皆擁有NH2 端的頭部、頸部及 COOH 端對蛋白酶敏感的尾部，在尾部含有典型的核定位序列（nuclear localization signal, NLS）。到目前為止，只有 ADD1 被報導，當上皮細胞失去細胞間附著時會進入細胞核。在本篇研究中發現，在細胞中表現 ADD 蛋白，ADD1 會分布於細胞質與細胞核，但 ADD2 及 ADD3 的分布只局限於細胞質。當在細胞中加入 CRM1 的抑制劑 leptomycin B，發現 ADD2 及 ADD3 會進入細胞核，暗示著 ADD2 及 ADD3 具有功能性的 NLS序列，使其能夠在核質穿梭。進一步在 ADD 蛋白接上 Simian virus 40 large T antigen 的 NLS 序列，我們發現 ADD2 與 ADD3 有較強的出核能力來拮抗SV40 NLS 的功能。且所有ADD 蛋白都具有多重出核訊號（nuclear export signal, NES）散佈在不同區域。另外，我們也證明ADD1 在頸部區域只含有一段富含 leucine的序列（377FEALMRMLDWLGYRT391），其為具功能性的 NES 序列，但 ADD2 及 ADD3 相對序列卻不是具有功能性的 NES 序列。接著，為了確定 ADD1 在核內的功能，利用尋找其核內的結合蛋白。利用免疫共沉澱，發現 ADD1會與轉錄因子 ZNF331 及 RNA 聚合酶 II 結合。綜合以上結果，我們得知，ADD 蛋白停留在細胞核的時間不同，是由於在演化結果造成其分布及功能的差異。

Adducin （ADD） proteins are actin binding proteins mainly localized at actin-spectrin junctions. The ADD family comprises three closely related genes（α, β and γ）. All ADD proteins are similar in their amino acid sequences and domain structures which consist of an amino-terminal head domain, a neck domain, and a carboxyl-terminal protease-sensitive tail domain within which a highly conserved bipartite nuclear localization signal（NLS）is present. Interestingly, only ADD1 has been reported to localize in the nucleus upon loss of cell-cell junctions. In this study, we found that transiently expressed ADD2 and ADD3 distributed only in the cytoplasm, whereas ADD1 distributed both in the cytoplasm and nucleus. However, ADD2 and ADD3 were partially sequestered in the nucleus in the presence of leptomycin, a CRM1 inhibitor, indicating that ADD2 and ADD3 possess functional NLS and actually shuttle between the cytoplasm and nucleus. Our results also suggest that ADD2 and ADD3 may be quickly transported to the cytoplasm upon entering the nucleus. Indeed, we found that ADD2 and ADD3 possess much higher potential than ADD1 to counteract the activity of the NLS derived from Simian virus 40 large T antigen. All ADD proteins appear to contain multiple nuclear export signals（NES）. A leucine-rich motif （377FEALMRMLDWLGYRT391） present in the neck domain was verified to be a functional NES for ADD1, but not for ADD2 and ADD3. To determine the function of ADD in the nucleus, we identify nuclear ADD binding proteins. Coimmunoprecipitation experiments indicated that transcription factor ZNF331 and RNA polymerase II are associated with ADD1.Taken together, our results provide an example to show that different members in the ADD family could have different potential to retain in the nucleus and suggest that ADD proteins may diversify their localization and function during the evolution.

I 文獻探討 1
第一章 核質運輸機制 1
第一節 核孔 1
第二節 核質運輸因子 2
第三節 蛋白之進核 4
第四節 蛋白之出核 6
第五節RanGTP的循環 7
第二章 Adducin 9
第一節ADD的發現 9
第二節ADD的結構與調控 9
第三節ADD的細胞功能 11
第四節ADD的生理功能 13
第五節ADD1的進核機制 14
第三章 真核生物的基因轉錄 16
第一節 基因轉錄 16
第二節 肌動蛋白核內功能 16
第三節 肌動蛋白結合蛋白與轉錄 18
第四節 鋅指蛋白331 的調控 20
II 研究動機 21
III 材料與方法 22
實驗材料 22
一、抗體 22
二、藥品 23
三、細胞株 24
四、質體 25
五、儀器 25
實驗方法 26
一、細胞培養 26
二、建立穩定表現外源蛋白的細胞株 26
三、質體的建立 26
四、短暫表現外源蛋白 30
五、Leptomycin B (LMB) 抑制 CRM1 30
六、蛋白免疫螢光染色 31
七、細胞蛋白質之萃取 31
八、免疫沉澱法 32
九、西方墨點轉漬法 32
IV 研究結果 34
一、ADD1分布於細胞核及細胞質而ADD2及ADD3只分布於細胞質 34
二、ADD1、ADD2 與ADD3 都具有進核的能力 34
三、ADD2 與 ADD3 較 ADD1 有更強的出核能力 35
四、ADD1 在頸部區域具有一段功能性的 NES 序列 36
五、ADD2 與 ADD3 出核機制異於 ADD1 36
六、ADD 家族具有多重 NES 序列散佈在不同區域 37
七、ADD 蛋白能透過不同 exportin 帶出細胞核 38
八、ADD1 會與核內蛋白結合 38
V 討論 40
VI 參考文獻 45
VII 附圖 67
VIII 附錄 79
Tables 79
Maps 82
ADD 序列比對 97

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