泛素蛋白酶體系統(UPS)是廣泛地存在於高等植物中，其參與在植物生長發育及抵抗環境逆境過程，然而proteasome在甘藷葉片老化中所扮演的角色尚未被釐清。已知在先前的研究中證實ethephon (ethylene-releasing compound)及NaCl逆境下會皆能促進甘藷葉片的老化。在本研究結果中分別在ethephon誘導三天及NaCl誘導六、九天與對照組相比都可以顯著促進葉片黃化，包括葉綠素降解、光合作用效率降低、增加NO/H2O2/MDA的含量及老化相關基因的表現，而藉由蛋白酶體抑制劑 MG132前處理能抑制在ethephon及NaCl逆境下包括(1) ethephon及NaCl分別在4小時及於24小時所產生的H2O2和NO二次信息因子、(2)訊息傳遞基因leucine-rich repeat receptor kinase SPLRR/calmodulin SPCAM/ethylene response factor SPERF1 6小時內的表現及(3)ethephon及NaCl分別在1天及 3天處理下乙烯合成相關基因ACC oxidase SPACO表現，減緩葉片老化。本實驗室從甘藷葉片中選殖出deleted in split hand/split foot DSS1-like protein SPDSS1全長，根據先前阿拉伯芥的研究指出其可能為26S proteasome的subunit。透過ethephon及NaCl誘導下葉片在老化前期可發現SPDSS1基因就有顯著的表現。而在外加SPDSS1融合蛋白前處理下可以延緩ethephon和NaCl所誘導的甘藷葉片老化，如果融合蛋白經失活處理(95℃ 5分鐘)後則會反轉延緩葉片老化現象，根據上述結果結論，不管是蛋白酶體抑制劑 MG132或SPDSS1融合蛋白都能延緩ethephon和NaCl所誘導的葉片老化，也建議26S proteasome在ethephon及NaCl誘導甘藷葉片老化過程扮演重要的角色。

Ubiquitin-proteasome system widely exists in higher plants, and has been reported to be associated with plant development and stress responses. The role of proteasomes in association with sweet potato leaf senescence mostly remains unclear. Ethephon, an ethylene-releasing compound, and NaCl stress have been demonstrated to promote leaf senescence in sweet potato. Significant changes including morphological leaf yellowing reduction of chlorophyll/Fv/Fm contents, elevation of nitric oxide (NO)/H2O2/malodialdehyde (MDA) levels, and promotion of senescence-associated gene expression were observed in ethephon-treated leaves on day 3 and NaCl-treated leaves on days 6 and 9, respectively, as compared to the untreated control. Both ethephon and NaCl stress-mediated effects were all alleviated by proteasome inhibitor MG132. Components induced by ethephon and NaCl, including (1) generation of H2O2 and NO second signal within 6 h for ethephon and 24 h for NaCl, (2) induction of leucine-rich repeat receptor kinase SPLRR/calmodulin SPCAM/ethylene response factor SPERF1 of signal transduction pathway within 6 h, and (3) expression of ACC oxidase SPACO of ethylene biosynthesis pathway starting from day 1 for ethephon and day 3 for NaCl, were all repressed by MG132 pretreatment and also correlated with the attenuation of induced leaf senescence. In our laboratory, a full-length cDNA SPDSS1 encoding a putative deleted in the split hand/split foot (DSS1)-like protein has been cloned from sweet potato leaves, which is a candidate subunit of 26S proteasome according to the previous report in Arabidopsis. Both ethephon and NaCl enhanced SPDSS1 expression level earlier than the induced leaf senescence. Exogenous SPDSS1 fusion protein also delayed ethephon and NaCl-induced leaf senescence in sweet potato, which could be reversed by boil pretreatment (95℃ for 5 min) of SPDSS1 fusion protein. Based on these results we conclude for the first time that both proteasome inhibitor MG132 and SPDSS1 fusion protein mitigate ethephon and NaCl-induced leaf senescence, suggesting a novel role of the 26S proteasome in association with ethephon and NaCl-induced leaf senescence in sweet potato.

目 錄

論文審定書.................................................................................................i
誌謝..............................................................................................................ii
中文摘要………………………...………………...……….…...…….…iii
英文摘要……………………......…………..……………......…..………iv
目錄…...…………………………………………………….…..…….…...v
圖次……………………………..………….....…….……....………......ix
縮寫表......................................................................................................xiii
壹、 緒論………………………..……………………….………………1
甘藷.............…......………………………………………………1
葉片老化……………….....………………………………………1
乙烯訊息傳導途徑和葉片老化………...........…………………..2
鹽分逆境誘導的葉片老化…………........………………………4
Ubiquitin-proteasome system…………………...........…………...6
DSS1 (deleted in split hand/split foot)…………………………..9
甘藷葉片老化研究…………………......……………………….10
研究目的 (Specific aims)…..………………………………….11
貳、 研究材料和方法…………......………………………...………...12
(一) 實驗材料……………..…………………………..……………..12
甘藷………………………………………..………...12
基因………………………………………………..…………...12
實驗藥品……………………………..…………….……………13
(二) 實驗方法…………………………..………………..……………14
實驗流程圖..........................………..……………..…………….14
Ethephon處理………………………..……………….................15
NaCl處理...................................………..……………………….15
Effector 或 inhibitor前處理……….....…..…………………….16
葉片外觀型態...............................................................................19
葉綠素含量測定……………………………..…….....................19
Fv/Fm 測量…………………….…………………...………......20
甘藷SPDSS1表現載體構築、融合蛋白表現及純化………..…20
重組SPDSS1表現載體構築…………………...........……….....20
SPDSS1融合蛋白表現.................................................................23
蛋白質濃度測定............…….......................................................23
SDS-PAGE……………………………….......………………….24
SPDSS1融合蛋白純化………………..………………………...25
DAB染色…...........……………………..……………………….25
H2O2含量測定………….……………..………………………...26
NO含量測定………………………..…………………………...27
MDA含量測定.............................................................................28
甘藷相關基因表現分析...............................................................29
RNA萃取......................................................................................29
RT-PCR反應................................................................................30
PCR...............................................................................................31
甘藷相關基因的primers..............................................................32
參、 結果...................................................................................................35
Part A. 蛋白酶體抑制劑MG132延緩ethephon誘導之葉片老
化...................................................................................................35
Ethephon促進甘藷葉片老化.......................................................35
蛋白酶體抑制劑MG132前處理延緩ethephon誘導之
葉片老化及H2O2/MDA/NO含量的累積....................................35
蛋白酶體抑制劑MG132前處理延緩ethephon誘導之H2O2和
NO二次信息因子產生及葉片老化.............................................37
蛋白酶體抑制劑MG132前處理抑制ethephon可誘導之信息傳
遞因子、乙烯生合成及葉片老化相關基因的表
現...................................................................................................39
Part B. 蛋白酶體抑制劑MG132延緩NaCl誘導之葉片老
化...................................................................................................40
NaCl逆境促進甘藷葉片老化......................................................40
蛋白酶體抑制劑MG132前處理延緩NaCl誘導之葉片老化及
H2O2/MDA/No含量的累積..........................................................41
蛋白酶體抑制劑MG132前處理延緩NaCl誘導之H2O2和NO
二次信息因子產生及葉片老化...................................................42
蛋白酶體抑制劑MG132前處理抑制NaCl可誘導之信息傳遞因
子、乙烯生合成及葉片老化相關基因的表現.............................44
Part C. 甘藷SPDSS1延緩ethephon及NaCl誘導的葉片老化...............45
甘藷SPDSS1基因表現受ethephon及NaCl所誘導...................45
外加SPDSS1融合蛋白延緩ethephon誘導之葉片老化及
H2O2/MDA/NO含量累積.............................................................46
外加SPDSS1融合蛋白延緩NaCl誘導之葉片老化及
H2O2/MDA/NO含量累積.............................................................48
單獨外加SPDSS1融合蛋白並無顯著影響黑暗下誘導之葉片老
化...................................................................................................49
肆、 討論...................................................................................................50
參考文獻...................................................................................................56

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