本論文為首篇利用抑制性扣減雜交法 (suppression subtractive hybridization, SSH) 鑑別大型海洋綠藻裂片石蓴 (Ulva fasciata Delile) 銅逆境下差異表現基因之研究，並對於大型藻類面對高濃度重金屬銅的反應與其分子層面上可能之耐受機制以及基因調控有所綜合性的了解。抑制性扣減雜交法分析裂片石蓴於銅 (50 μM CuSO4) 處理6-12小時基因表現，共發現69個差異表現基因：55個正向調控及14個負向調控，依Gene Ontology (GO) Classification database進行功能分類，共可分為十大類：1. 轉錄作用 (Transcription)；2. 轉譯作用以及核醣體結構與生合成 (Translation, ribosomal structure and biogenesis)；3. 轉譯後修飾作用、蛋白質更新、伴隨蛋白 (Posttranslational modification, protein turnover, chaperones)；4. 光合作用 (Photosynthesis)；5. 細胞氧化還原平衡 (Cell redox homeostasis)；6. 逆境 (Stress)；7. 新陳代謝 (Metabolism)；8. 能量生產與轉換 (Energy production and conversion)；9. 運輸 (Transport)；10. 未知功能 (Function unknown)。本試驗結果可以推論裂片石蓴因應過量銅所產生之影響，主要藉由能量生產與轉換透過增加體內能量的生產，供應各種路徑所需的能量，並藉由控制鐵離子的恆定以及還原狀態之氫硫基 (-SH)，維持體內氧化還原之平衡，且藉由調控光合作用相關基因的表現，以維持光合作用之效率。除此之外，透過逆境相關基因表現的產物，去除活性氧族 (reactive oxygen species, ROS)，並且藉由其它類別之基因表現，促進修補受氧化之蛋白質、染色體與脂質。另外，經由新陳代謝類別之基因的協助，維持體內胺基酸的恆定以及增加體內proline的含量，以增強對銅逆境之耐受性。另外，我們從裂片石蓴於不同硫酸銅濃度 (0、5、10、20與50 μM) 經過四天處理的試驗得知，裂片石蓴透過維持體內抗氧化物質之恆定以及經由基因表現來調控抗氧化酵素活性，來應對銅所誘導之氧化逆境，但是此防禦能力並不足以減輕高濃度銅所造成之氧化傷害。然而，透過分析氧化還原相關與抗氧化酵素相關基因於外加銅以及活性氧族下的表現，我們得知ROS牽涉到正向調控抗氧化防禦相關基因的表現，裂片石蓴對於過量銅長時間的適應，係與抗氧化酵素相關基因以及UfMsrA (methionine sulfoxide reductase A) 的表現有關，而其對於過量銅短時間的馴化，反而與氧化還原相關基因以及UfGr (glutathione reductase) 的正向調控有關。啟動子在基因調控上扮演關鍵性之角色，我們透過分析UfMsr啟動子上之cis-acting elements，得知銅影響裂片石蓴基因之表現，應透過與防禦訊息相關之植物荷爾蒙以及其它逆境相關之訊息傳遞路徑，並且鈣離子與calmodulin應參與其中。總而言之，裂片石蓴可以透過分子層級上之調控來抵抗過量銅所造成之氧化逆境。

This is the first research by using suppression subtractive hybridization (SSH) to analysis the gene expression in marine macroalga Ulva fasciata Delile against excess copper toxicity, and it gives us a comprehensive understanding of the tolerant mechanism while macroalgae face to the excess copper. Suppression subtractive hybridization was used to identify genes differentially expressed following exposure to 50 μM CuSO4 for 6- 12h in a marine macroalga Ulva fasciata Delile. In this work, 69 genes were identified, of which 55 were up-regulated and 14 were down-regulated. According to the database of Gene Ontology (GO), these genes were classified into 10 categories as follows: 1. Transcription; 2. Translation, ribosomal structure and biogenesis; 3. Posttranslational modification, protein turnover, chaperones; 4. Photosynthesis; 5. Cell redox homeostasis; 6. Stress; 7. Metabolism; 8. Energy production and conversion; 9. Transport; 10. Function unknown. According to the results, we suggest that the responses of U. fasciata against excess copper toxicity are mainly through increase of the energy production for providing sufficient energy to many metabolic pathways, and control of the Fe homeostasis and redox form of thiol groups for maintaining the cellular redox homeostasis, moreover, expression of photosynthetic genes for letting the photosynthesis work. In addition, to scavenge the ROS is by expression of stress-related genes, meanwhile, the proteins, DNA and lipids damaged by ROS (reactive oxygen species) and copper are repaired by expression of the other categorical genes. Over and above, the genes expressing in the metabolism category might maintain the amino acids homeostasis and increase the purine content, and subsequently increase the tolerant capacity of U. fasciata against excess copper toxicity. In addition, the concentrations of antioxidants and the activities and gene expression of antioxidant enzymes were determined in Ulva fasciata Delile by a 4-day exposure to 0, 5, 10, 20 and 50 μM CuSO4. These results demonstrate that the maintenance of antioxidant homeostasis and the induction of activities of antioxidant enzymes via enhanced gene expression are used by U. fasciata to cope with the Cu-induced oxidative stress, but the defense capacity cannot sufficiently alleviate oxidative damage occurring under the condition of higher Cu concentrations. Moreover, according to the results from the expression of genes involved in the control of redox homeostasis and antioxidant defense was studied in macroalga Ulva fasciata Delile in response to CuSO4 (5 and 50 μM) and ROS (H2O2 and O2˙-), we suggest that ROS involved in up-regulation of antioxidant defense-related genes and the expression of genes of antioxidant defense enzymes and UfMsrA (methionine sulfoxide reductase A) are associated with long-term adaptation of U. fasciata to Cu excess and transcription of redox- related genes and UfGr (glutathione reductase) is up-regulated for short-term acclimation. Promoters play a key role in regulating gene expression. Based on the analysis of cis-acting elements on UfMsr promoters, we suggested that the signal transduction pathway of copper stress in U. fasciata is related to that of other stresses and of defense-related plant hormones, however, Ca2+ and calmodulin might participate in it. To sum up, U. fasciata could resist oxidative damage caused by excessive copper through the regulation on the molecular level.

口試委員會審定書……………………………………I
誌謝……………………………………………………II
中文摘要………………………………………………………VI
英文摘要………………………………………………VIII
圖表目錄………………………………………………X
縮寫字對照………………………………………………………XVIII
第一章 總論………………………………………1
壹、 前言………………………………………2
貳、 研究背景…………………………………4
一、 海洋大型綠藻 (裂片石蓴)………………4
二、 重金屬與植物之關係……………………5
三、 銅逆境對生物之影響……………………7
1. 過量銅逆境對動物之影響………………7
2. 過量銅逆境對植物之影響………………8
3. 植物隔離游離態銅之防禦………………11
四、 植物之氧化逆境…………………………13
1. 活性氧族之來源與角色…………………13
2. 植物之抗氧化機制………………………17
3. 銅與氧化逆境之關係……………………20
五、 抑制性扣減雜交…………………………22
參、 本論文研究方向與實驗架構……………25
肆、 參考文獻…………………………………26
第二章 利用抑制性扣減雜交法鑑定銅害下裂片石蓴
差異表現之基因…………………………61
壹、 中文摘要…………………………………62
貳、 英文摘要…………………………………64
參、 文獻回顧…………………………………66
肆、 材料與方法………………………………69
伍、 結果………………………………………84
一、 抑制性扣減雜交與基因庫之建立………84
二、 利用點雜合反應鑑別之差異表現基因…84
三、 DNA序列與生物資訊資料庫基因比對之結
果…………………………………………85
四、 利用北方墨點法確認差異表現之基因…86
陸、 討論………………………………………88
柒、 參考文獻…………………………………97
第三章 過量銅逆境下裂片石蓴抗氧化酵素活性與基
因表現之關係……………………………126
壹、 中文摘要…………………………………127
貳、 英文摘要…………………………………128
參、 文獻回顧…………………………………129
肆、 材料與方法………………………………131
伍、 結果………………………………………148
一、 過氧化氫 (H2O2)、TBARS、ascorbate
(ASC) 與 glutathione (GSH) 含量……148
二、 MnSOD、FeSOD、APX、GR以及CAT基因
全長選殖…………………………………148
三、 MnSOD、FeSOD、APX、GR與CAT基因表
現量與酵素活性之關係…………………149
陸、 討論………………………………………150
柒、 參考文獻…………………………………153
第四章 過量銅逆境下裂片石蓴氧化還原平衡與抗氧
化酵素相關基因之表現…………………187
壹、 中文摘要…………………………………188
貳、 英文摘要…………………………………189
參、 文獻回顧…………………………………190
肆、 材料與方法………………………………192
伍、 結果………………………………………198
一、 UfMsrA、UfTrx、UfCyp與UfFer cDNAs之表
徵…………………………………………198
二、 銅處理短時間與長時間下對於基因表現之影
響…………………………………………198
三、 銅處理短時間下H2O2之含量以及外加H2O2
與O2˙- (menadione) 對於基因表現之影響…
……………………………………………199
陸、 討論………………………………………201
柒、 參考文獻…………………………………206
第五章 裂片石蓴氧化調節相關基因UfMsrA與
UfMsrB啟動子之研究…………………233
壹、 中文摘要…………………………………234
貳、 英文摘要…………………………………235
參、 文獻回顧…………………………………236
肆、 材料與方法………………………………239
伍、 結果………………………………………244
一、 UfMsrA與UfMsrB之啟動子DNA序列…244
二、 生物資訊資料庫比對搜尋cis-acting
elements之結果…………………………244
陸、 討論………………………………………247
柒、 參考文獻…………………………………254
第六章 綜合討論與未來研究方向………………289
壹、 綜合討論與未來研究方向………………290
一、 銅害下裂片石蓴分子層面之耐受機制…290
二、 裂片石蓴基因表現之調控………………292
貳、 參考文獻…………………………………295
附錄……………………………………………………304

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Chapter 2
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宋明軒。(2006) 裂片石蓴protease 基因表現與高鹽誘導氧化逆境及蛋白質氧化之關係。國立中山大學海洋生物研究所碩士論文。台灣，中華民國。
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