臺灣博碩士論文加值系統

近年來由於極端氣候影響，造成耕地土壤乾旱鹽化問題，嚴重衝擊農業生產及糧食供需平衡。因此，改進作物逆境耐受性特別在乾旱逆境方面是重要的世界性議題。Ubiquitin-Proteasome System (UPS)廣泛存在於不同植物，並且在甘藷非生物性逆境例如高鹽逆境耐受性扮演重要角色，然而在乾旱逆境耐受性方面其生理功能則尚未清楚。乾旱處理12及15天相較於未處理的對照組顯著促進甘藷葉片老化及褐化死亡，而且在形態、生化、及代謝上顯著增加老化相關指標變化包括形態黃化/褐化死亡、葉綠素及Fv/Fm含量減少、抗氧化酵素含量及活性下降、NO/H2O2/ MDA含量上升、mitogen-activated protein kinase蛋白質含量及磷酸化程度增加、及cysteine protease SPCP1 表現量增加。這些乾旱造成的影響皆受蛋白酶體抑制劑MG132減緩，顯示蛋白酶體活性與甘藷乾旱逆境耐受性有關。本實驗室先前從甘藷葉片選殖的一個全長cDNA SPDSS1，其基因解碼產生一個deleted in split hand/split foot 1 (DSS1)-like protein，為酸性蛋白質具有75個胺基酸(228核苷酸)及三個保守的aspartic/glutamic acid-rich domains，根據先前阿拉伯芥報導為19S 調節性蛋白酶體(19S regulatory proteasome)的次單元體。乾旱會誘導SPDSS1表現且其時間早於葉片老化，外加SPDSS1融合蛋白質會加速乾旱誘導的葉片老化/褐化死亡及增加老化相關指標變化提早至處理後第9及12天。這些外加SPDSS1融合蛋白質加速乾旱的影響亦受蛋白酶體抑制劑MG132減緩。依據上述結果結論蛋白酶體抑制劑MG132及19S 調節性蛋白酶體次單元體deleted in split hand/split foot 1 (DSS1)-like protein SPDSS1 參與調控甘藷乾旱逆境耐受性，也建議Ubiquitin-Proteasome system與乾旱逆境耐受性的生理角色有關

In recent year drought and soil salinization of cultivated lands under extreme climate severely result in the unbalance between agricultural crop production and demand-provision systems. Therefore, improvement of crop stress tolerance especially for drought is an important world-wide issue. Ubiquitin-Proteasome System (UPS) widely exists in plants and plays a role in association with abiotic salt stress tolerance in sweet potato. However, its physiological function under drought stress is unclear. In sweet potato, drought promoted senescence and necrosis in leaves. Significant morphological, biochemical and metabolic changes of senescence-associated markers, including morphological yellowing/necrosis, reduction of chlorophyll and Fv/Fm contents, decrease of antioxidant enzyme/activity levels, elevation of NO/H2O2/MDA amounts, increase of mitogen-activated protein kinase/phosphorylation levels, and enhancement of cysteine protease SPCP1 expression were observed in treated leaves on days 12 and 15 as compared to the untreated control. Drought-mediated effects were all alleviated by the proteasome inhibitor MG132, suggesting an association of proteasome activities in drought stress tolerance. In our laboratory, a cloned full-length cDNA SPDSS1 from sweet potato leaves encodes a putative deleted in split hand/split foot 1 (DSS1)-like protein, which is an acid protein with 75 amino acids (228 nucleotides) and three conserved aspartic/glutamic acid-rich domains in its open reading frame and is a candidate subunit of the 19S regulatory proteasome according to the previous report in Arabidopsis. Drought enhanced SPDSS1 expression level earlier than the induced leaf senescence. Exogenous SPDSS1 fusion protein accelerated drought-induced senescence/necrosis and enhanced changes of senescence-associated markers on day 9 and 12 in treated leaves, which could be inhibit and reversed by the proteasome inhibitor MG132. Based on these results we conclude that proteasome inhibitor MG132 and the 19S regulatory proteasome subunit deleted in split hand/split foot 1 (DSS1)-like protein SPDSS1 participate in the modulation of drought stress tolerance in sweet potato leaves. A novel physiological role of the Ubiquitin-Proteasome system in association with drought stress tolerance is also suggested.

Thesis Validation Letter i
Acknowledgment ii
Chinese Abstract iii
English Abstract iv
Table of Content v
Table of Figures viii
List of Abbreviations x
Chapter 1. Introduction 1
1.1 Drought Stress and Leaf Senescence 1
1.2 Leaf Senescence 2
1.3 MAPK Cascade 3
1.4 Antioxidant-related Enzymes 4
1.5 Senescence-associated Cysteine Proteases 4
1.6 Proteasome 5
1.7 Deleted in Split Hand/Split Foot 1 Protein (DSS1) 7
1.8 Sweet potato 8
1.9 Related Research of Leaf Senescence in Sweet Potato 9
1.10 Specific Aims 10
Chapter 2. Materials and Methods 11
2.1 Experimental Flow Chart 11
2.2 Materials 12
2.2.1 Sweet Potato Leaves 12
2.2.2 Related Genes and Antibodies 12
2.2.3 Chemicals 13
2.3 Methods 14
2.3.1 Drought Treatment 14
2.3.2 Effector and Inhibitor Treatment 14
2.3.3 Induction and Purification of SPDSS1 Fusion Protein 15
2.3.4 Leaf Morphology 16
2.3.5 Chlorophyll Content Quantification 16
2.3.6 Fv/Fm Determination 17
2.3.7 DAB Staining 17
2.3.8 H2O2 Measurement 17
2.3.9 Nitric Oxide Detection 17
2.3.10 MDA Content Assessment 18
2.3.11 Catalase Activity Assay 19
2.3.12 Ascorbate Peroxidase Activity Assay 19
2.3.13 SDS PAGE 20
2.3.14 Western Blot Hybridization 21
2.3.15 Statistical Analysis 22
Chapter 3. Result 23
3.1 Drought promote leaf senescence 23
3.1.1 Drought promotes changes of senescence-associated markers and cell necrosis in leaves 23
3.1.2 Expression and protein phosphorylation of mitogen-activated protein kinase SPMAPK are significantly enhanced by drought 23
3.1.3 Antioxidant protein and enzymatic activity levels are drastically reduced in drought-induced necrotic leaves 24
3.1.4 Expression of senescence-associated cysteine protease SPCP1 is remarkably elevated in drought-induced necrotic leaves 25
3.2 Deleted-in-split hand/split foot 1 protein SPDSS1 accelerate drought-mediated leaf senescence 25
3.2.1 Expression of Deleted-in-Split Hand/Split Foot 1 (DSS1)-like protein SPDSS1 increases at the early stage of drought treatment 25
3.2.2 Exogenous SPDSS1 fusion protein enhances drought-mediated changes of senescence-associated markers and cell necrosis in leaves 26
3.2.3 Exogenous SPDSS1 fusion protein accelerates drought-mediated induction and protein phosphorylation of mitogen-activated protein kinase SPMAPK 27
3.2.4 Exogenous SPDSS1 fusion protein enhances drought-mediated decrease of antioxidant protein and enzymatic activity levels in leaves 28
3.2.5 Exogenous SPDSS1 fusion protein promotes drought-mediated induction of senescence-associated cysteine protease SPCP1 in leaves 29
3.3 Proteasome inhibitor MG132 alleviates drought-mediated leaf senescence 29
3.3.1 Proteasome inhibitor MG132 alleviates drought-mediated changes of senescence-associated markers and cell necrosis in leaves 29
3.3.2 Proteasome inhibitor MG132 mitigates drought-mediated induction and protein phosphorylation of mitogen-activated protein kinase SPMAPK 31
3.3.3 Proteasome inhibitor MG132 delays drought-mediated decrease of antioxidant protein and enzymatic activity levels in leaves 31
3.3.4 Proteasome inhibitor MG132 attenuates drought-mediated induction of senescence-associated cysteine protease SPCP1 in leaves 32
3.4 Effects of SPDSS1 on drought-mediated leaf senescence can be attenuated by proteasome inhibitor MG132 33
3.4.1 Proteasome inhibitor MG132 alleviates SPDSS1 fusion protein effects on drought-mediated changes of senescence-associated markers and cell necrosis in leaves 33
3.4.2 Proteasome inhibitor MG132 delays SPDSS1 fusion protein effects on drought-mediated decrease of antioxidant protein and enzymatic activity levels in leaves 34
3.4.3 Proteasome inhibitor MG132 attenuates SPDSS1 fusion protein effects on drought-mediated induction of senescence-associated cysteine protease SPCP1 in leaves 35
Chapter 4. Discussion 36
Chapter 5. Conclusion 40
Chapter 6. References 41

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