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研究生：	李書宏 Lee, Shu-Hong
論文名稱：	甲硫胺酸硫氧化物還原酶B7與B8對氧化逆境反應之研究 Functional study of cytosolic methionine sulfoxide reductases B7 (MSRB7) and MSRB8 in response to oxidative stress
指導教授：	詹明才 Chan, Ming-Tsair
學位類別：	博士 Doctor
系所名稱：	生物科學與科技學院 - 生物科技研究所 Institute of Biotechnology
論文出版年：	2014
畢業學年度：	103
語文別：	英文
論文頁數：	145
中文關鍵詞：	阿拉伯芥、甲硫胺酸硫氧化物還原酶、氧化逆境、谷胱甘肽s-轉移酶、巴拉刈、質譜儀、非抗生素篩選系統
外文關鍵詞：	A. thaliana, methionine sulfoxide reductase B, oxidative stress, glutathione transferase, methyl viologen, LC-MS/MS, non-antibiotic selection
相關次數：	點閱：119 下載：1
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植物在生物性逆境與非生物性逆境下會產生過氧化物 (reactive oxygen species, ROS)，而過量的ROS會氧化甲硫胺酸 (methionine, Met) 形成甲硫胺酸硫氧化物 (methionine sulfoxide, MetO)，造成蛋白質結構改變並影響其活性。而甲硫胺酸硫氧化物還原酶 (MetO reductases, MSR) 可將MetO還原為Met，因此MSR在氧化逆境下能保護蛋白質免於氧化。本研究發現受到氧化逆境誘導表現且表現於細胞質的MSRB7與MSRB8，均具有還原R-form MetO的酵素活性，並利用TRXH3 (Thioredoxins) 與TRXH5形成氧化還原循環 (redoxin cycle)。為了研究MSRB7與MSRB8的生化特性與生理功能，我們轉殖降低表現 (B7i與B8i) 與持續表現 (B7ox與B8ox) 的阿拉伯芥。降低表現MSRB7/8的阿拉伯芥對於殺草劑巴拉刈 (methyl viologen, MV) 較野生型與B7/8ox敏感，反之，B7/8ox能提高轉殖植物對氧化逆境的耐受度，且B7/8ox植株中抗氧化酵素GST (glutathione-S-transferases) 活性較高。為了鑑定MSRB的受質，我們以MSRB7作為模式蛋白質，利用溴化氰水解比較蛋白質體 (Comparative proteomic analysis using cyanogen bromide digestion) 分析，發現兩個受MV誘導表現的GSTF2與GSTF3在B7ox植株中的含量較高。進一步研究發現MSRB7分別與GSTF2與GSTF3結合，並還原GSTF2/3重要的氨基酸Met。MSRB7除了維持GSTF2/3的酵素活性同時也增加蛋白質穩定性。因此，植物利用MSRB7與GST的交互作用維持GSTF2與GSTF3的酵素活性與蛋白質穩定性，提高對氧化逆境的耐受度。
植物基因轉殖技術對於基礎研究與農業應用科技十分重要。為了快速篩選擬轉植植株，大多利用轉入耐受抗生素或殺草劑基因，並利用抗生素或殺草劑作為篩選劑進行篩選。已知持續表現阿拉伯芥MSRB7、MSRB8與MSRB9可提高植物對MV的抗性，因此其具有成為非抗生素篩選基因的潛力。本研究成功在阿拉伯芥與番茄建立MV-MSRB篩選系統，MSRBs不但可以做為非抗生素篩選系統，篩選得到的植株亦可提高對氧化逆境的抗性。

Methionine sulfoxide reductases (MSRs) catalyse the reduction of oxidized methionine residues, thereby protecting proteins against oxidative stress. Accordingly, MSRs have been associated with stress responses, disease and senescence in a taxonomically diverse array of organisms. However, the cytosolic substrates of MSRs in plants remain largely unknown. Both of MSRB7 and MSRB8 were methyl viologen (MV)-inducible and could reduce R-form MetO to Met. Both of them are reduced by TRXH3 (Thioredoxins) and TRXH5 through redoxin cycle. Here, we used a proteomic analysis strategy to identify MSRB7 substrates. We showed that two glutathione transferases (GSTs), GSTF2 and GSTF3, had fewer oxidized methionine (MetO) residues in MSRB7-overexpressing A. thaliana plants than in wild-type plants. Conversely, GSTF2 and GSTF3 were highly oxidized and unstable in MSRB7-knockdown plants. MSRB7 was able to restore the MetO-GSTF2M100/104 and MetO-GSTF3M100 residues produced during oxidative stress. Furthermore, both GSTs were specifically induced by the oxidative stress inducer, methyl viologen. Our results indicate that specific GSTs are substrates of MSRs, which together provide a major line of defense against oxidative stress in A. thaliana.
Plant transformation is an important tool for basic research and agricultural biotechnology. MSRB7 transgenic Arabidopsis and tomato plants were successfully obtained by Agrobacterium-mediated transformation and selection on medium supplemented with MV. We suggest that specific MSRB genes that are overexpressed in transgenic plants may constitute a new non-antibiotic selection system with increased tolerance to oxidative stress concomitant with MV treatment.

Chinese abstract (中文摘要)	 I
English abstract	 II
Acknowledgements	 III
Contents	 IV
List of tables	 VIII
List of figures	 IX
Abbreviation	 XII
Chapter 1. Oxidative stress and methionine sulfoxide reductase	 1
1 Introduction	 1
1.1  Oxidative stress	 1
1.2  Methionine sulfoxide reductase	 1
Chapter 2. Functional analysis of MSRB7 and MSRB8	 4
1 Introduction	 4
2 Materials and methods	 5
2.1  Plant materials and growth conditions	 5
2.2  RNA isolation and gene expression analysis	 5
2.3  Plasmid constructions	 5
2.4  GUS histochemical staining and activity	 6
2.5  Immunoblot analysis	 6
2.6  Production of recombinant proteins	 6
2.7  MSRB enzyme activity	 7
2.8  Immunoprecipitation	 7
2.9  Yeast two-hybrid assay	 7
2.10 Bimolecular fluorescence complementation (BiFC) and protoplast transient assay	 8
2.11 Determination of chlorophyll content	 8
2.12 Ion leakage rate	 8
2.13 Catalase activity	 8
2.14 Peroxidase activity	 9
2.15 In vivo GST activity	 9
2.16 Hydrogen Peroxide Staining	 9
2.17 Statistical analyses	 9
3 Results	 10
3.1  Cytosolic MSRB7 and MSRB8 are relatively abundant expression in the root and can be induced by oxidative stress	 10
3.2  Activity of MSRB7 and MSRB8	 10
3.3  Redox-dependent interacted between MSRB7/8 and TRXH3/5	 11
3.4  Transgenic Arabidopsis plants overexpressing MSRB7/B8 have shorter roots	 12
3.5  Cytosolic MSRB7 and MSRB8 knockdown lines display sensitivity to oxidative stress, whereas overexpression lines exhibit tolerance	 12
3.6  MSRB7 and MSRB8 transgenic plants cultivated in soil exhibited resistance to MV spraying	 13
3.7  MSRBOx plants accumulate less H2O2 and exhibit higher glutathione S-transferase activity under MV-induced oxidative stress	 13
3.8  MSRB7 is induced by methyl viologen in aerial and root part	 13
4 Summary	 14
Chapter 3. Identification and characterization the substrates of MSRB7	 15
1 Introduction	 15
2 Materials and methods	 16
2.1  Comparative proteomic analysis using cyanogen bromide digestion	 16
2.2  Methionine residues in GSTs and their differential oxidation as analysed by mass spectrometry	 17
2.3  BiFC and protoplast transient assay	 17
2.4  H2O2 content	 17
2.5  In vitro GST activity	 18
3 Results	 18
3.1  Identification of putative MSRB7 substrates by comparative proteomic analysis using cyanogen bromide digestion	 18
3.2  GSTF2, and GSTF3 are induced by methyl viologen	 19
3.3  Methionine residues in GSTF2 and GSTF3 are repaired by MSRB7 in vivo	 19
3.4  GSTF2, GSTF3 and GSTF8 interact with MSRB7	 20
3.5  MSRB7 can restore the activities of oxidized GSTF2 and GSTF3 in vitro	 20
3.6  MSRB7 maintains the stability of GSTF2 and GSTF3 in vivo	 21
3.7  The Met residues of GSTF2 and GSTF3 are important for maintaining GST activity, and their oxidized states are reduced by MSRB7	 21
3.8  Reverse genetic evidence verifies the preservation of activity and stability of GSTF2 and GSTF3 by MSRB7 under oxidative stress	 22
4 Summary	 23
Chapter 4. Establishment of non-antibiotic selection system	 24
1 Introduction	 24
2 Materials and methods	 24
2.1  Arabidopsis selection	 24
2.2  Transformation and regeneration of tomato	 25
2.3  Southern blot analysis	 25
2.4  Reverse transcription-polymerase chain reaction (RT-PCR)	 26
3 Results	 26
3.1  Comparison of selection efficiencies of MV and hygromycin in Arabidopsis	 26
3.2  Transgenic Arabidopsis selected by MV expressed high levels of MSRB mRNA	 27
3.3  Transgenic tomato plants overexpressing MSRB7 confers tolerance to oxidative stress	 27
3.4  MSRB7 overexpressing tomato plants show normal phenotype	 28
3.5  MV concentration test on tomato cotyledons	 28
3.6  Evaluation of MV-MSRB selection system for tomato transformation	 28
5 Summary	 29
Chapter 5. Discussion	 30
1   Overexpression of cytosolic MSRB genes confers tolerance to oxidative stress	 30
2   MSRBs play important roles in redox homeostasis	 30
3   The activities of MSRB7 and MSRB8	 31
4   Regeneration of MSRB7/8 activity by the TRXH3/5	 31
5   ROS accumulation is reduced in transgenic MSRB plants	 32
6   Comparative proteomic analysis using cyanogen bromide digestion as an efficient strategy for the identification of MSRB7 substrates	 33
7   MSRB7 maintains the activity and stability of the substrates, GSTF2 and GSTF3	 34
8   Establishment of MV-MSRB selection system	 36
9   Conclusions	 37
References	 38
Appendix formulations	 106
                                    

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校外：2016-11-28公開

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