臺灣博碩士論文加值系統

硝酸鹽是植物體生長不可或缺重要營養源之一，同時亦可作為訊息分子調控植物的側根發育、開花時間及胺基酸之生合成。因此，植物如何調控基因表現來適應環境硝酸鹽濃度的波動便是一個重要的問題，然而對於硝酸鹽訊息路徑的研究不多，僅有參與調控訊息路徑的轉錄因子ANR1、SPL9、LBD37/38/39，及NLP7 (NIN-like protein 7)等蛋白分子被報導。AtNLP7屬於NIN-like protein (NLPs) 家族，其家族共9個成員，NLPs具有兩個重要的功能區塊，其一為RWP-RK domain，功能為DNA結合功能區塊，目前已知AtNLPs 家族利用RWP-RK domain結合在Nitrate-response cis-element並活化下游基因表現，藉此調控硝酸鹽訊息相關基因表現；其二，PB1 (Phox and Bem 1) domain，功能為蛋白質與蛋白質間交互作用，此功能區塊常見於真核細胞內參與訊息傳遞的蛋白質中。
本研究主要是以生物物理分析研究NLP7的PB1 domain之特性。在蛋白質純化過程中發現，此蛋白質會出現聚集現象(aggregation)形成多聚體(oligomer)。在經過數種緩衝液的改善下，成功讓NLP7 PB1蛋白質在溶液條件20 mM Tris-HCl pH 8.8，0.5 M NaCl，10 % glycerol，5 mM GSH，1 mM TCEP下，以二聚體(dimer)形式穩定存在。利用小角度X光散射(Small-angle X-ray scattering)分析，得到NLP7 PB1蛋白質二聚體之Rg值 (25.46A)、Dmax值 (71.14A)，再將重建蛋白質結構外殼與同源性結構模型進行疊合。進一步發現NLP7 PB1蛋白質在最適溶液中，呈現不同的聚合形式，如蛋白質濃度在1 mg/ml時為二聚體、2 mg/ml時為四聚體、高於2 mg/ml時皆為多聚體。經分析其模擬結構及序列比對，NLP7 PB1同時具有Type I和Type II的特徵，在以定點突變，分別對Type I (D909/D911) 以及 Type II (K867)特徵胺基酸進行突變後，其所得純化之目標蛋白質皆呈現單體形式，證明NLP7 PB1屬於Type I/II PB1 domain (第三型)，NLP7 PB1藉由自身K867殘基和OPCA motif (D909, D911, E913, D922) 之間交互作用形成同多聚體(homo-oligomer)。本研究表明NLP7可能藉由多聚體與單體之轉換來參與硝酸鹽訊息傳遞路徑或以PB1區塊與其他調控路徑中的蛋白交互作用，達成相關基因表現的調控。

中文摘要 i
Abstract iii
目錄 iv
圖目錄 vii
表目錄 ix
附錄 x
縮寫對照表 xi
Chapter 1 前言 1
1.1 硝酸鹽為植物必需之營養源並可作為訊息分子 1
1.2 初級硝酸鹽反應 (Primary Nitrate Response, PNR) 2
1.3 RWP-RK 轉錄因子家族 (RWP-RK transcription factor family) 2
1.4 NIN-like proteins (NLPs) family 4
1.5 NIN-like protein 7為硝酸鹽訊息路徑之主要調控者 5
1.6 NLP7 之 PB1 (Phox and Bem1p) domain 功能區塊 6
1.7 研究目標 7
Chapter 2 材料與方法 9
2.1 NLP7 PB1基因 9
2.2 NLP7 PB1 domain 表現載體之構築 9
2.3 勝任細胞(Competent cell)之製備 10
2.4 轉形作用 10
2.5 重組蛋白之大量表現 10
2.6 破菌 11
2.7 重組蛋白質之純化 11
2.8 蛋白質濃縮與定量 11
2.9 蛋白質保存緩衝液置換 12
2.10 SDS-聚丙烯醯胺膠體電泳(SDS-PAGE) 12
2.11 西方墨點法(Western blotting) 12
2.12 粒徑排阻層析分析(Size-exclusion chromatography, SEC) 13
2.13 動態光散射分析(Dynamic light scattering, DLS) 13
2.14 小角度散射實驗(Small-angle X-ray scattering) 13
2.15 同源性結構模型建立與結構比對分析 14
2.16 蛋白質結晶實驗 14
2.16.1 Pre-crystallization test (PCT測試) 14
2.16.2 蛋白質結晶條件篩選 15
2.16.3 結晶條件之微調 15
Chapter 3 實驗結果 16
3.1 NLP7 PB1蛋白質表現與純化 16
3.2 NLP7 PB1蛋白質粒徑排阻層析分析(SEC) 16
3.3 NLP7 PB1聚合型態分析 17
3.4 NLP7 PB1突變型之純化結果與聚合型態分析 17
3.5 NLP7 PB1重組蛋白質小角度X光散射(SAXS)分析 18
3.6 蛋白質養晶實驗 19
Chapter 4 討論 21
4.1 NLP7 PB1蛋白質純化策略 21
4.2 NLP7 PB1蛋白質於不同緩衝溶液多聚體型態比較 21
4.3 NLP7 PB1蛋白質多聚體型態與樣品濃度關係比較 22
4.4 NLP7 PB1蛋白質藉由K867與OPCA motif的交互作用形成多聚體 22
4.5 NLP7可能以多聚體形式參與硝酸鹽訊息路徑調控 23
Chapter 5 結論 25
圖表 26
參考文獻 56
附錄 66

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