臺灣博碩士論文加值系統

已知水稻T-DNA插入突變株M52048之早開花性狀是由於OsMADS14基因被活化造成。水稻OsMADS14基因與阿拉伯芥中參與開花調節之AP1為同源基因。本實驗室過去以ubiquitin啟動子大量表現OsMADS14，確實造成轉殖株極早開花與矮化，初步研究顯示OsMADS14持續表現，會促進開花調節路徑上游之開花調控基因Hd3a及RFT1提早表現且表現量持續增加。本研究旨在進一步了解大量表現OsMADS14會促進水稻早開花之可能分子機制；除了確認Hd3a與RFT1基因受到促進之外，想了解是否還有其它可能受OsMADS14影響而導致早開花之調節基因。於是搜尋受到阿拉伯芥AP1影響而導致早開花之水稻同源基因，包括開花抑制基因：RCN1、OsTEM1-Like及OsTOE1-Like；Cytokinins (CK) 生合成基因：OsLOG、OsLOG Like-2、OsLOG Like -3、OsCKX4、OsCKX5；及GA生合成基因：OsEUI1、OsGA13ox1等，進行分析。結果顯示開花基因Hd3a和RFT1在Ubi:OsMADS14轉殖株中皆提早表現；調控CK生合成之OsLOG基因受到抑制，OsCKX5基因受到活化，可能因而降低葉腋分生組織活性，促進開花。然而，開花抑制基因RCN1不僅沒有被抑制，反而表現量增加，這其中的調控機制，值得進一步探討。另外，GA生合成相關之OsEUI1和OsGA13ox1基因表現量增加，推測可能為導致水稻植株矮化重要原因之一。
本研究同時探討水稻中可能為對應於AtTFL1、AtTOE1、及AtTEM1之開花抑制相關基因：LOC_Os11g05470 (RCN1)，LOC_Os05g03040 (OsTOE1-Like)，LOC_Os01g49830 (OsTEM1-Like) 等之功能。自台灣水稻T-DNA插入突變庫中，獲得可能活化RCN1之T-DNA插入突變株M78020及可能活化OsTEM1-Like之突變株M89461，進行T-DNA插入位置確認及基因型與外表型之相關性分析。結果顯示T-DNA插入位正確，突變株M78020之野生型回復株與同質基因型植株，外表型及開花期沒有明顯差異，然而M89461同質基因型植株比異質基因型植株，呈現晚開花情形。突變株之目標OsTEM1-Like基因之表現仍需進一步分析確認。
本研究亦嘗試利用酒精誘導系統表達外源性基因，透過AlcR調節因子，調節目標基因表現。本酒精誘導系統，包含一個由35S 啟動子轉錄、對酒精敏感的AlcR轉錄因子及一個含有能受AlcR轉錄因子調節的啟動子AlcA結合下游目標基因的轉錄單位。這系統以抗殺草劑基因Bar為篩選標誌，與本研究室過去常使用Hyg篩選標誌之轉殖系統不同，所以需要建立篩選平台。篩選過程顯示，雖然有獲得再生植株，然而分析結果顯示所有植株皆非轉殖株，未來可調整抗性基因為Hyg，再嘗試建立以酒精誘導系統，表達外源性基因之水稻轉殖平台。

The early flowering phenotype of a rice T-DNA insertion mutant M52048 had been demonstrated due to the activation of OsMADS14, an Arabidopsis AP1 homologous gene involved in flowering regulation. Overexpressing OsMADS14 with ubiquitin promoter construct Ubi:OsMADS14 in TNG67 revealed early flowering as observed in M52048. In addition, the early expression of two floregin genes Hd3a and RFT1 were observed and enhanced in Ubi:OsMADS14 transgenic rice. However, how the activation and overexpression of OsMADS14 in rice could result in early flowering as AP1 did in the Arabidopsis have not been characterized. The present study aims to understand the molecular mechanisms of early flowering that regulated by over-expressing OsMADS14. The rice homologous to Arabidopsis AP1-regulated genes that lead to early flowering and dwarf, such as floral repressors, RCN 1 (homologous to TFL1), OsTEM1-like (TEM1), OsTOE1-like (TOE1); cytokinin biosynthesis genes, such as OsLOG, OsLOG1-like, OsLOG2-like, OsLOG3-like, OsCKX4, OsCKX5; and GA biosynthesis related genes, such as OsEUI1 and OsGA13ox1, were identified and investigated. Results showed Hd3a and RFT1 expressed much earlier in Ubi:OsMADS14 transgenic rice, and OsLOG and OsCKX5 response similarly to their homologous genes LOG1 and CKX3 in Arabidopsis, which may reduce axillary meristem growth and promote flowering. Surprisingly, the flowering suppressor gene RCN1 was enhanced, response differently from that in Arabidopsis. Therefore, the role of RCN1 in rice required further investigation. Regarding to the GA biosynthesis-related genes, the expression level of OsEUI1and OsGA13ox1 were enhaced that could possibly explain the dwarf phenotype of Ubi:OsMADS14 transgenic rice.
The function of rice genes LOC_Os11g05470 (RCN1); LOC_Os05g03040 (OsTOE1-Like); LOC_Os01g49830 (OsTEM1-Like) homologous to Arabidopsis flowering suppressor genes AtTFL1, AtTOE1 and AtTEM1 were investigated. Two T-DNA insertion mutants, M78020 that may activate RCN1 and M89461 that may activate OsTEM1-like gene, were obtained from TRIM database for characterization. The insertion events of these two mutants were confirmed, and the mutant M89461 presente a late flowering phenotype, however mutant M78020 display no phenotype and no different in heading date to that of WT control. The links between the target genes expression and heading date phenotypes are under investigation.
Establishing an alcohol inducible transgenic platform is another project of this study. This alcohol inducible system contains an alcohol-sensitive transcription factor AlcR driven by 35S promoter, and a second transcription unit contains AlcA promoter-driven target gene that can be regulated by alcohol-bound AlcR transcription factor. This alcohol inducible system using herbicide resistant bar gene as selection marker which is different from the hygromycin selection system we used routinely. Therefore, the optimization of selection condition was performed. Although several regenerated rice plant were obtained, none of them contain the transgene. In order to set up the alcohol inducible transgenic platform, replacing herbicide with hygromycin selection is suggested.

摘要 i
Abstract ii
目錄 iv
表目次 vi
圖目次 vi
附表 vii
附圖 vii
附錄 vii
縮寫字對照表 viii
前言 1
前人研究 2
一、水稻基因功能之研究 2
二、參與花器形成之MADS-box基因研究 2
三、OsMADS14基因功能分析 4
四、開花時間之調控 4
五、Ubi:OsMADS14導致矮株與植物賀爾蒙吉貝素 (Gibberellin, GAs) 之生合成及 代謝路徑 6
六、酒精誘導外源基因表現平台之背景及應用 7
七、利用酒精誘導表現系統調節開花 8
八、利用酒精誘導表現系統調節植株矮化 9
材料與方法 10
一、儀器及設備、藥品、試驗材料 10
二、水稻轉殖表現載體之構築 10
三、水稻之轉殖方法 12
四、水稻轉殖基因分析 13
結果 17
一、水稻OsMADS14基因功能之探討 17
二、水稻RCN1、TEM1及TOE1等抑制開花相關基因之功能探討 20
三、建立誘導性轉殖基因表現平台的探討 22
討論 26
一、大量表現OsMADS14提早水稻開花抽穗，對花器之發育影響不大 26
二、開花激素Hd3a 和RFT1的表現可能受到MADS-box蛋白複合物調控 26
三、大量表現OsMADS14對調節開花相關之Cytokinins生合成基因表現之影響 27
四、大量表現OsMADS14 對GA代謝基因的影響 27
五、大量表現OsMADS14對開花抑制相關基因的影響 28
六、酒精誘導表現轉殖載體之構築及轉殖再生 28
七、農桿菌轉殖共培養後續處理 29
結論 31
參考文獻 32
表 40
圖 47
附表 61
附圖 64
附錄 70

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