臺灣博碩士論文加值系統

本篇研究主要探討FYF和EDFs基因之間在調控花器老化脫落上的關係。EDF1/2/3/4::GUS 轉殖株中成熟花的GUS訊號較強，年輕的花中較弱。此情況和FYF::GUS相反。35S::EDF1/2/3/4 及EDF1/2/3/4+SRDX轉殖株都會提早花器的老化及脫落，並且可促老化相關基因SAG12、AtMCs和VPEs的表現，結果指出EDF1/2/3/4可作為一轉錄抑制子促進花器的老化及脫落。35S::EDF1/2/3/4+SRDX可以挽救etr1-1、ein2-1和35S::FYF植株中延緩花器老化及脫落的現象，並且在EDF1/2/3/4::GUS轉殖株中大量表現FYF可降低GUS基因的表現，顯示EDF1/2/3/4在乙烯訊息傳遞路中位於FYF的下游。這些結果都指出，FYF可藉由抑制轉錄抑制子EDF1/2/3/4來延緩花器的老化及脫落。
為了研究蘭花中FYF及SVP的同源基因，分別從Oncidium Gower Ramsey和Cattleya Intermedia選殖出OnSVP1/2、CaFYF1/2及CaSVP。35::OnSVP1/2+SRDX轉基因阿拉伯芥透過抑制EDF1/2/3/4、BOP2、IDA、HAESA延緩花器的老化和脫落。即使經過乙烯的處理，35::OnSVP1/2+SRDX轉基因阿拉伯芥仍然可延緩花器的老化及掉落，顯示轉基因植物為乙烯不敏感型植株。35::CaSVP+SRDX、35::CaSVP、35::CaFYF1/2+SRDX 、35::CaFYF1/2轉基因阿拉伯芥也可藉由抑制乙烯及離層訊息傳徑，達到延緩花器的老化及脫落。我們的結果顯示，從蘭花中選殖出的FYF及SVP同源基因在轉基因阿拉伯芥中可作為一轉錄抑制子延緩花器的老化及脫落。

In this study, we analyzed the relationships between Forever young flower (FYF) and Ethylene response DNA binding factors (EDFs) in regulating flower senescence/abscission. Strong GUS staining was detected in the mature flower and staining decreased in young flowers of the EDF1/2/3/4::GUS transgenic Arabidopsis. This result was opposite to FYF::GUS plants. 35S:EDF1/2/3/4 and 35S::EDF1/2/3/4+SRDX caused similar promotion of flower senescence/abscission and the activation of the senescence-associated genes SAG12, AtMCs and VPEs, indicating that EDF1/2/3/4 function as repressors in promoting senescence/abscission. EDF1/2/3/4 function downstream of FYF in the ethylene signaling pathway, and this finding was further supported by the observations that 35S::EDF1/2/3/4+SRDX rescue the delayed senescence/abscission phenotypes in the etr1 and ein2 mutants and in 35S::FYF plants. The 35S::FYF significantly reduced GUS expression in EDF1/2/3/4::GUS plants. These results indicate that FYF delayed flower senescence/abscission by repressing transcriptional repressor EDF1/2/3/4. To analyze the FYF and SVP orthologs from orchid, Oncidium short vegetative phase1/2 (OnSVP1/2), Cattleya forever young flower1/2(CaFYF1/2) and Cattleya short vegetative phase (CaSVP) were identified from Oncidium Gower Ramsey and Cattleya intermedia. 35::OnSVP1/2+SRDX delayed flower senescence/abscission by down-regulating EDF1/2/3/4, BOP2, IDA, HAESA in transgenic Arabidopsis. Furthermore, 35::OnSVP1/2+SRDX transgenic Arabidopsis were insensitive to ethylene. 35::CaSVP+SRDX, 35::CaSVP, 35::CaFYF1/2+SRDX and 35::CaFYF1/2 also delayed flower senescence/abscission by down-regulating ethylene and abscission signaling in transgenic Arabidopsis. Our results revealed that FYF and SVP orthologs from orchids function as a repressor to delay flower senescence/abscission in transgenic Arabidopsis.

摘要………………………………………………………………………………i
ABSTRACT…………………………………………………………………………...ii
前言…………………………………………………………………….……………1-1
材料方法…………………………………………………………………………….1-4
結果………………………………………………………………………………...1-12
一、 EDF1/2/3/4表現部位分析、分子選殖及載體構築……………......….1-12
二、利用EDF1/2/3/4啟動子驅動GUS報導基因表現，GUS蛋白活性隨著花朵的成熟而上升……………………………………………………………...1-12
三、異位表現EDF1/2/3/4和EDF1/2/3/4+SRDX促進花朵老化及脫落….1-13
四、 EDF1/2/3/4和EDF1/2/3/4+SRDX促進程式性細胞死亡相關基因導致花器提早老化脫落……………………………………………………………...1-13
五、異位表現EDF2+SRDX在雄不稔突變株dad1中，仍會促進花器老化及脫落，顯示此促進老化脫落之情況並非無法受孕所造成…………………1-14
六、異位表現EDF1/2/3/4+SRDX可挽救乙烯訊息傳遞突變株etr1-1所造成的花器延緩老化脫落的情況……………………………………………….. 1-14
七、異位表現EDF1/2/3/4+SRDX可挽救乙烯訊息傳遞突變株ein2-1所造成的花器延緩老化脫落的情況………………………………………...………1-15
八、異位表現EDF1/2+SRDX在35S::FYF轉基因植物中，可挽救因異位表現FYF所造成的延緩花器老化脫落的情況…………………………..……1-15
九、異位表現FYF 於EDF1/2/3/4 啟動子驅動GUS 報導基因的轉基因植物中，報導基因GUS 訊號受到抑制…………………………..………………1-15
討論………………………………………………………………………………...1-17
參考文獻…………………………………………………………………………...1-20
圖表………………………………………………………………………………...1-24
表1-1.本篇文章所使用之PCR引子(primer)序列…………………….….....1-24
表1-2.異位表現EDF1/2/3/4花朵老化脫落統計……………………….......1-26
表1-3.異位表現EDF1/2/3/4+SRDX花朵老化脫落統計……………….......1-26
表1-4.etr1-1突變株異位表現EDF1/2/3/4+SRDX花朵老化脫落統計.........1-27
表1-5.ein2-1突變株異位表現EDF1/2/3/4+SRDX花朵老化脫落統計........1-27
表1-6.異位表現FYF之轉基因植物，額外異位表現EDF1/2+SRDX花朵老化脫落統計……………………………………………………………………...1-38
圖1-1.EDF1/2/3/4於野生型阿拉伯芥中部位表現情況…………………....1-39
圖1-2.EDF1/2/3/4啟動子驅動GUS報導基因之轉基因植物，GUS活性偵測……………………………………………………………...........................1-30
圖1-3.EDF1/2/3/4之表現隨著花朵成熟而上升…………............................1-31
圖1-4.35S::EDF1/2/3/4和35S::EDF1/2/3/4+SRDX轉基因植物，花器提早老化脫落，並且有無法受孕的情況………….....................................................1-32
圖1-5.異位表現EDF1/2/3/4造成老化指標基因SAG12上升......................1-33
圖1-6.異位表現EDF1/2/3/4+SRDX造成老化指標基因SAG12上升..........1-34
圖1-7.異位表現EDF1/2/3/4和EDF1/2/3/4+SRDX造成程式性細胞死亡相關基因表現量上升………...................................................................................1-35
圖1-8.異位表現EDF2+SRDX在雄不稔突變株dad1中，仍會促進花器老化及脫落………...................................................................................................1-36
圖1-9.異位表現EDF1/2/3/4+SRDX可挽救etr1-1突變株花器延緩老化脫落的情況……………...........................................................................................1-37
圖1-10.異位表現EDF1/2/3/4+SRDX可挽救ein2-1突變株花器延緩老化脫落的情況 .....1-48
圖1-11.異位表現EDF1/2+SRDX可促進35S::FYF轉基因植物花朵的老化脫落，以EDF1/2/3/4啟動子驅動GUS報導基因之轉基因植物，GUS活性可因額外異位表現FYF而降低 .....1-39
圖1-12.FYF可藉由促進FUF1基因，抑制EDF1/2/3/4延緩花器老化脫落.1-40
附圖1-1.目標基因載體構築示意圖................................................................1-41
附圖1-2.EDF1/2/3/4 cDNA分子選質與構築之示意膠圖.............................1-42
附圖1-3.EDF1/2/3/4啟動子選質與構築之示意膠圖....................................1-43
附圖1-4.EDF1/2/3/4於eFP browser中各部位表現情況...............................1-44
附圖1-5.EDF1/2/3/4 cDNA序列比對結果....................................................1-45
附圖1-6.pGEM®-T Easy vector之圖譜............................................................1-46
附圖1-7.pEpyon-01K之圖譜...........................................................................1-47
附圖1-8.pEpyon-22K之圖譜..........................................................................1-58
附圖1-9.pEpyon-2aK之圖譜...........................................................................1-59
附圖1-10.pEpyon-2bK之圖譜........................................................................1-50
附圖 1-11.Gen-KB DNA Ladder.....................................................................1-51
前言…………………………………………………………………….……………2-1
材料方法…………………………………………….................................................2-4
結果……………………………………………………………………………….....2-7
一、從嘉德麗雅蘭中選殖出FYF同源基因………………...…………….…..2-7
二、CaFYF1/2於嘉德麗雅蘭中各部位分析………………………………….2-7
三、異位表現CaFYF1/2、CaFYF1/2+SRDX和CaFYF1/2+VP16促進轉基因植物提早開花…. ……………………………………………………..…….....2-7
四、異位表現CaFYF1/2、CaFYF1/2+SRDX延緩轉基因植物花朵老化脫落，異位表現CaFYF1/2+VP16促進轉基因植物花朵老化脫落………………...2-8
五、異位表現CaFYF1/2、CaFYF1/2+SRDX轉基因植物，為乙烯不敏感型植株………………………………………………….………………..…...….......2-8
六、異位表現CaFYF1/2、CaFYF1/2+SRDX轉基因植物，抑制乙烯訊息傳遞路徑及花器凋落訊息傳遞路徑之基因……………………………..…..…… 2-8
七、從嘉德麗雅蘭及文心蘭中選殖出SVP同源基因………………….....…..2-8八、利用即時定量PCR偵測CaSVP基因於嘉德麗雅蘭各部位表現情況….2-9九、利用即時定量PCR偵測OnSVP1/2基因於文心蘭各部位表現情況…....2-9
十、異位表現AtSVP、OnSVP1/2+SRDX、CaSVP和CaSVP+SRDX抑制花器老化脫落，異位表現OnSVP1 /2∆II+VP16和CaSVP+VP16促進花器老化脫落…………………………………………………………..………..…….........2-9
十一、異位表現AtSVP、OnSVP1/2+SRDX、CaSVP和CaSVP+SRDX，為乙烯不敏感型植株………………………………………………………..….....2-10
十二、異位表現AtSVP抑制乙烯訊息傳遞路徑及離層分化起始傳遞路徑之基因………………………………………………………………………….. 2-10
十三、異位表現OnSVP1/2+SRDX抑制乙烯訊息傳遞路徑及離層分化起始傳遞路徑之基因………………………………………………………………...2-10
十四、異位表現CaSVP和CaSVP+SRDX抑制乙烯訊息傳遞路徑及離層分化起始傳遞路徑之基因………………………………………………………...2-10
討論…………………...…………………………………………………………... 2-11
參考文獻…………………………………………………………………………...2-15
圖表………………………………………………………………………………...2-17
表2-1.本篇文章所使用之PCR引子(primer)序列…………………….….....2-17
表2-2.異位表現CaFYF1、CaFYF1+SRDX和CaFYF1+VP16轉基因植物開花時間及葉片數統計…………………………………………………….......2-19
表2-3.異位表現CaFYF2、CaFYF2+SRDX和CaFYF2+VP16轉基因植物開花時間及葉片數統計…………………………………………………….......2-19
表2-4.異位表現CaFYF1、CaFYF1+SRDX和CaFYF1+VP16花朵老化脫落朵數統計..........………………………………………..………………….......2-20
表2-5.異位表現CaFYF2、CaFYF2+SRDX和CaFYF2+VP16花朵老化脫落朵數統計.........………………………………………..…………………........2-20
表2-6.異位表現AtSVP、 AtSVP ∆II、 OnSVP1/2+SRDX、OnSVP1/2 ∆II +VP16、CaSVP、CaSVP+SRDX和CaSVP+VP16花朵老化脫落朵數統計………....2-21
圖2-1.嘉德麗雅蘭(Cattleya Intermedia)、文心蘭(Oncidium Gower Ramsey)及阿拉伯芥(Arabidopsis thaliana)中FYF基因的親緣演化關係，序列相似度比對……………………………………………………………...........................2-22
圖2-2.CaFYF1/2於嘉德麗雅蘭各部位表現情況…………..........................2-23
圖2-3.異位表現CaFYF1/2、CaFYF2/1+SRDX和CaFYF1/2+VP16造成植株早開花…………......……………………………………….............................2-24
圖2-4.異位表現CaFYF1/2、CaFYF2/1+SRDX之轉基因植物延遲花器老化脫落，為乙烯不敏感型植株，異位表現CaFYF2/1+VP16促進花器老化脫落………….......................................................................................................2-25
圖2-5.異位表現CaFYF1/2、CaFYF1/2+SRDX之轉基因植物，抑制乙烯訊息傳遞路徑下游基因EDF1/2/3/4和脫落訊息傳遞路徑中BOP1/2、IDA基因.......................................................................................................................2-26
圖2-6.嘉德麗雅蘭、文心蘭及阿拉伯芥中SVP基因的親緣演化關係，序列相似度比對………...........................................................................................2-27
圖2-7.CaSVP於嘉德麗雅蘭各部位表現情況………....................................2-38
圖2-8.OnSVP1/2於文心蘭南茜各部位表現情況……………......................2-29
圖2-9.異位表現AtSVP、OnSVP1/2+SRDX、CaSVP和CaSVP+SRDX延遲花器老化脫落，異位表現OnSVP1/2 ∆II+VP16和CaSVP+SRDX促進花器老化脫落……………...............................................................................................2-30
圖2-10.異位表現AtSVP、OnSVP1/2+SRDX、CaSVP及CaSVP+SRDX之轉基因植物，為乙烯不敏感型植株.....................................................................2-31
圖2-11.異位表現AtSVP之轉基因植物，抑制乙烯訊息傳遞路徑下游基因ERF1、EDF1/3和脫落訊息傳遞路徑中IDA、HAESA和HSL2基因.......2-32
圖2-12.異位表現OnSVP1/2+SRDX之轉基因植物，抑制乙烯訊息傳遞路徑下游基因ERF1、EDF1/2/3/4和脫落訊息傳遞路徑中BOP2、IDA和HAESA基因...................................................................................................................2-33
圖2-13.異位表現CaSVP+SRDX之轉基因植物，抑制乙烯訊息傳遞路徑下游基因EDF1/3和脫落訊息傳遞路徑中IDA基因.............................................2-34
圖2-14.假說圖示..............................................................................................2-35
附圖2-1.目標基因構築流程電泳圖................................................................2-36
附圖2-2.目標基因構築流程電泳圖................................................................2-37
附圖2-3.CaFYF1 核甘酸序列及胺基酸序列................................................2-48
附圖2-3.CaFYF2 核甘酸序列及胺基酸序列................................................2-39
附圖2-3.CaSVP 核甘酸序列及胺基酸序列..................................................2-40

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