臺灣博碩士論文加值系統

由Taiwan Rice Insertional Mutant (TRIM) library 中得到T-DNA插入突變株M52048，具有矮株、提早開花、節彎曲以及穗抽出不完全等多種外表型。T-DNA以順向的方式插入水稻第三號染色體中，同時也藉由T-DNA所攜帶的35S 增強子活化附近的三個基因OsMADS34、OsMADS14與OsCP7 (putative cysteine protease)。OsMADS34與OsMADS14屬於MADS-box基因家族，在水稻中主要參與花器形成與開花時間的調控，OsCP7屬於C1A cysteine protease家族的一員。本研究將探討OsMADS14與OsCP7在水稻中所具有的功能。
過去研究指出大量表現OsMADS14會造成水稻早開花，因此為了解開花調節基因的表現是否受OsMADS14影響，於是分析OsGI、OsMADS50、Ehd2、Hd1、Ehd1、Hd3a、RFT1、OsMADS14、OsMADS18等開花調節基因在突變株M52048與Ubi:OsMADS14、Ubi:OsMADS34、OsCP7:OsCP7轉殖株中的表現情形。實驗結果顯示OsMADS14上游的兩個開花激素基因Hd3a與RFT1在M52048與Ubi:OsMADS14中皆提早表現，而在異位表現OsMADS34與OsCP7之轉殖株中則否。證實Hd3a與RFT1亦受到OsMADS14的表現而活化，然而其調控機制仍需要進一步的研究。
以玉米ubiquitin或花椰菜嵌紋病毒CaMV 35S啟動子大量表現OsCP7，皆無法得到穩定的轉殖株；而以OsCP7上游1.6 kb的原生性啟動子，成功得到OsCP7:OsCP7轉殖株，其性狀為略矮、略晚抽穗、穎花黑斑、稔實率低等。相對於在TNG67的水稻穗中無法偵測到OsCP7表現，OsCP7:OsCP7的葉片與水稻穗在RNA以及蛋白質層次皆可發現OsCP7被大量表現，顯示這些外表性狀確實與OsCP7有關聯。經由質體救援得知OsCP7:OsCP7其一株系的T-DNA插入位，因此得以藉由基因型分析區別轉殖株為同質或異質品系。雖然同質品系的外表性狀較為嚴重，但其RNA與蛋白表現量卻無法與異質品系辨別出差異。而異位表現之OsCP7蛋白主要表現在水稻穗中，因此為了探討大量表現OsCP7基因所造成的穎花黑斑與低稔實率，於是觀察OsCP7:OsCP7的花器組織結構，與TNG67相比並沒有明顯的差異，而經由花粉活性分析證實了異位性表現OsCP7影響花粉的發育。偵測OsCP7於水稻穗的表現情形，發現穎花黑斑與OsCP7的表現量呈現正相關，而且黑斑的產生可能是緣於mature OsCP7蛋白提升促使水稻進行計畫性細胞凋亡所導致的結果。

The T-DNA mutant M52048 identified from Taiwan Rice Insertional Mutant (TRIM) library showed dwarf, early flowering, node bending and impaired in panicle exertion. Three flanking genes, OsMADS34, OsMADS14 and OsCP7 (putative cysteine protease 7) were activated in this mutant. Both OsMADS34 and OsMADS14 belong to MADS-box gene family that may participate in regulation of flowering time and the identity of floral organ. OsCP7 is a member of C1A cysteine proteases. In this study, the function of OsMADS14 and OsCP7 were further investigated.
Previous study has demonstrated that over-expression of OsMADS14 could cause early flowering. To understand whether or not any other flowering regulatory genes were affected by the expression of OsMADS14. The flowering regulatory genes including OsGI, OsMADS50, Ehd2, Hd1, Ehd1, Hd3a, RFT1, OsMADS14 and OsMADS18 were investigated in mutant M52048 and Ubi:OsMADS14, Ubi:OsMADS34 and OsCP7:OsCP7 transgenic rice respectively. Results showed two florigen genes, Hd3a and RFT1, expressed much earlier in M52048 and Ubi:OsMADS14 but not in Ubi:OsMADS34 and OsCP7:OsCP7, suggesting that Hd3a and RFT1 were regulated by the expression of OsMADS14. The mechanism how the expression of OsMADS14 could regulate florigen genes requires further investigation.
Expression of OsCP7 driven by the maize ubiquitin promoter or the CaMV 35S promoter in transgenic rice cannot be obtained successfully. However transgenic rice, OsCP7:OsCP7 using 1.6 kb of OsCP7 promoter could be easily obtained and OsCP7:OsCP7 transgenic rice plants revealed slightly shorter in plant height, delayed flowering, lower fertility and lesion-like spots on spikelet. In contrast to the wild-type where no OsCP7 was detected in panicles, the RNA and protein expressions of OsCP7 in OsCP7:OsCP7 transgenic rice were detected in leaves at all development stages and panicles, and their expressions in transgenic rice correlated to the observed phenotypes. In addition, the phenotypes of segregated homozygous plants showed more significant than those of heterozygous plants within the same transgenic line, suggesting the dosage effect of transgene. However the expression levels of RNA and protein cannot be differentiated between homo- and hetero-zygous lines. To unravel the causes that lead to lower fertility of OsCP7:OsCP7, the floral organ and pollen viability were investigated. The floral organ showed no obvious differences between wild-type and OsCP7:OsCP7, but the pollen viability of OsCP7:OsCP7 was lower than that of wild-type, indicating that continuing expression of OsCP7 influence pollen development. Further investigation also indicated that the lesion-like spots on spikelet was correlated with the expression levels of OsCP7 and the lesions could possibly due to the programmed cell death caused by the activity of increased mature OsCP7 present in spikelet.

中文摘要 i
英文摘要 ii
目錄 iv
圖表目次 vi
縮寫對照表 vii
前言 1
前人研究 2
一、水稻基因功能之探討 2
二、突變株M52048 2
三、MADS-box基因家族 3
四、花器形成之ABCDE模式與A群基因的功能 3
五、開花時間調控機制 4
六、OsMADS14對於開花時間之影響 6
七、Cysteine protease 6
八、Papain-like cysteine proteases 6
九、PLCP參與計畫性細胞凋亡 8
十、OsCP7可能參與計畫性細胞凋亡 9
材料與方法 11
一、儀器與設備 11
二、實驗藥品 11
三、研究材料 11
四、水稻轉殖基因分析 11
五、T-DNA插入點之生物資訊分析 14
六、水稻花粉活性分析 14
結果 15
壹、OsMADS14促進早開花可能的分子機制探討 15
一、開花調控基因於Ubi:OsMADS14轉殖株於各時期表現分析 15
二、開花調控基因於突變株M52048於各時期表現分析 16
三、開花調控基因於Ubi:OsMADS34與OsCP7:OsCP7轉殖株於各時期表現分析 16
貳、OsCP7基因研究 16
一、OsCP7轉殖株外表性狀分析 16
二、OsCP7:OsCP7-5轉殖株基因型分析 17
三、OsCP7基因於不同轉殖株之表現量分析 17
四、OsCP7基因於OsCP7:OsCP7-5轉殖株不同基因型之表現量分析 17
五、OsCP7基因於TNG67與OsCP7:OsCP7-5同質品系轉殖株在不同時期與不同部位之表現量分析 18
六、OsCP7基因於OsCP7:OsCP7-5同質品系穎花中之表現量分析 18
七、穎花黑斑嚴重程度不同之OsCP7:OsCP7-5同質品系表現量分析 18
八、水稻花器觀察與花粉活性分析 19
九、35S:OsCP7-6x His tag mutant之表現量分析與農藝性狀調查 19
十、35S:OsCP7 RNAi之農藝性狀調查 20
討論 21
一、OsMADS14為促進水稻開花的正向調控子 21
二、開花激素RFT1與Hd3a的表現可能受MADS-box蛋白複合物調控 21
三、OsMADS34可能也參與開花時間調控機制 22
四、OsCP7可能參與計畫性細胞凋亡，促使水稻產生黑斑以及低稔實率，甚至導致死亡 22
五、OsCP7可能參與老化機制 23
結論 25
參考文獻 26
表 35
圖 38
附表 51
附圖 53

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