臺灣博碩士論文加值系統

篩選自Taiwan Rice Insertional Mutant library (TRIM library)中的T-DNA突變株M52048具有矮株、早開花、節彎曲和穗無法完全抽出之特殊性狀，根據順向遺傳學證明含有加強子的T-DNA插在水稻第三號染色體上，活化T-DNA插入位兩側的OsMADS14、OsMADS34和一putative cysteine protease，後稱為OsCP7。OsMADS14和OsMADS34皆屬於MADS-box基因家族，為調控水稻開花機制及花器生長發育的轉錄因子。OsMADS14屬於MADS-box家族的A群基因，作用在光週期調控下游，可促使水稻開花，走向生殖生長期，其調控機制已被研究得相當透徹；OsMADS34則屬於MADS-box家族的E群基因，可結合其他MADS-box基因以調控穗及花器的發育。OsCP7經序列比對為一cysteine protease，並經活性測試確定具有protease功能，但其在水稻中所扮演的生理功能仍未明朗。
目前實驗室已有前人構築分別大量表現上述三個基因的轉殖株，分別為Ubi:OsMADS14、Ubi:OsMADS34和OsCP7:OsCP7。Ubi:OsMADS14造成轉殖株矮株和極早開花，此性狀與過去對其功能研究的結果相符合。而在Ubi:OsMADS34轉殖株則出現有早開花、穗無法完全抽出的性狀，穗無法正常抽出主要是由於第一節間縮短所造成，過去有研究指出乾旱環境造成ABA的累積可導致穗無法完全抽出之性狀，且RNA microarray顯示Ubi:OsMADS34中與乾旱逆境有關之基因，如DREB1A，DREB1E及EATB，表現量有所提升，因此進一步對轉殖株施予外加ABA inhibitor 與GA的處理，發現並無法有效延長其第一節間長度而恢復正常抽穗，顯示ABA累積並非造成Ubi:OsMADS34第一節間無法抽長之主因。透過對於節間GA生合成及代謝有關基因與細胞壁生合成相關基因的分析發現，第一節間中GA13ox1 (將GA生合成路徑導向GA1，相對減少GA4，導致植株半矮)表現量上升，同時第一節中EATB (可抑制GA生合成基因，OsCPS2的表現)和EUI1 (可將活性GA去活化)表現量提升，表示節間延長受阻可能和這些調控節間延長相關基因有關，但OsMADS34大量表現導致節間縮短的機制仍然需要更進一步研究。而Ubi:OsMADS34也造成雄蕊抽長較緩慢，顯示OsMADS34亦可能在轉殖株中發揮其class E MADS-box基因之功能，和其他MADS-box基因共同調控花器生長，未來也能往此方向著手，藉由分析OsMADS34交互作用的蛋白質來推測受其調控的基因為何。
擁有典型cysteine protease結構的OsCP7經前人以Ubiquitin及CaMV 35S啟動子構築大量表現OsCP7但無法得到穩定存活之轉殖株後，改採OsCP7上游1.6kb序列作為啟動子構築得到OsCP7:OsCP7轉殖株，其性狀有略晚抽穗和種子黑斑且稔實率低下，之後透過找出T-DNA插入位證實上述性狀與基因型為正相關，驗證了OsCP7:OsCP7-5的性狀與基因型具有相關性，造成這些性狀之機制須更進一步研究。

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 encode a putative cysteine protease, belongs to C1A cysteine protease family, its function remains unknown. In this study the function of OsMADS34 and OsCP7 were further investigated.
Previous study showed that over-expression of OsMADS34, Ubi:OsMADS34 transgenic rice, could cause slightly early flowering and impaired in panicle exertion. Morphological dissection indicated that the impaired in panicle exertion was mainly caused by shortening the first internode (peduncle). Inhibition of the peduncle elongation caused by drought stress and ABA accumulation has been reported. In the present study, some drought-related genes, such as DREB1A, DREB1E and EATB, were regulated in Ubi:OsMADS34 transgenic rice and therefore hypothesized that the phenotype of transgenic rice may regulated by drought stress or plant hormones. However, treated transgenic rice with ABA inhibitor and/or GA could not improve the peduncle elongation and panicle exertion, suggested that the shortened peduncle and impaired in panicle exertion in Ubi:OsMADS34 transgenic rice might not cause by ABA accumulation. Analysis of GA biosynthesis related genes and the cell elongation promoting genes, at the internodes, revealed high expression levels of EATB, EUI1, GA13ox1, GA20ox2, OsPK1 and lower expression of XTH28, suggested that the shortened internode might due to the imbalance expression of these genes. However the mechanism how these genes involved in internode elongation remain to be elucidated. In addition to the shortened peduncle, the anther development was also affected in Ubi:OsMADS34, suggesting that OsMADS34 function as an E class MADS-box gene may interact with other MADS-box genes to regulate the floral organ development. Further study by searching OsMADS34 interaction proteins will help us to unravel the possible function of OsMADS34.
For the study of OsCP7 gene, we were unable to obtained stable transgenic rice lines with constitutive promoter constructs, suggesting that constitutively ectopic expression of OsCP7 might cause lethal. Instead, transgenic lines with a 1.6 kb of OsCP7 promoter construct, OsCP7:OsCP7, were successfully obtained. OsCP7:OsCP7 revealed slightly dwarf, delayed flowering, lesion-like spots on panicles and lower fertility, and these phenotypes are correlated to the expression of OsCP7 gene. The possible mechanisms that cause these aberrant panicle developments were under investigated.

中文摘要 i
英文摘要 ii
目錄 iii
表目次 vi
圖目次 vi
附表 vii
附圖 vii
縮寫字對照表 viii

前言 1
前人研究 2
一、水稻基因功能探討 2
二、MADS-box基因 2
三、ABCDE類型之MADS-box基因 3
四、OsMADS34相關研究 4
五、MADS-box基因與植物賀爾蒙之關係 4
六、水稻節間延長 5
七、Cysteine protese之分類 6
八、Papain-like cysteine protease (PLCP) 6
九、Cysteine protease與植物的計畫性細胞凋亡 (Programmed cell death, PCD) 7
十、OsCP7相關研究 7
材料與方法 9
一、實驗藥品 9
二、儀器與設備 9
三、T-DNA插入水稻突變株之栽種 9
四、水稻轉殖株之構築 9
五、水稻轉殖基因分析 11
六、花粉活性測試 14
七、節間切片觀察 14
結果 15
壹、水稻OsMADS34之基因研究 15
一、 Ubi:OsMADS34之外表性狀觀察 15
二、 TNG67與Ubi:OsMADS34之稻穗結構比較 15
三、 TNG67與Ubi:OsMADS34之節間延長分析 15
四、TNG67與Ubi:OsMADS34 之稔實率與花粉活性分析 16
五、TNG67與Ubi:OsMADS34 之花器形態觀察 16
六、TNG67與Ubi:OsMADS34之節間切片觀察 17
七、Ubi:OsMADS34之節間延長相關基因表現分析 17
八、以生物資訊方式分析可供OsMADS34結合之cis-element 18
九、Ubi:HA-OsMADS34 構築及分析 18
貳、水稻OsCP7之基因研究 19
一、35S:OsCP7 之分析 20
二、35S:OsCP7-6xHis tag之分析 20
三、OsCP7:OsCP7 轉殖株之分析 21
討論 23
一、大量表現OsMADS34提早水稻抽穗 23
二、大量表現OsMADS34對稻穗結構無明顯影響 23
三、大量表現OsMADS34造成節間生長異常 23
四、大量表現OsMADS34影響稔實、花粉活性及穎花生長 24
五、大量表現OsMADS34影響節間結構 26
六、大量表現OsMADS34影響細胞延長的相關酵素表現 26
七、大量表現OsMADS34影響GA代謝基因的調控 27
八、可能受OsMADS34蛋白結合之cis-element分析 28
九、Ubi:HA- OsMADS34轉殖株確認大量表現HA- OsMADS34 28
十、過度表現OsCP7造成水稻死亡 29
十一、異位表現OsCP7造成水稻穎花黑斑及稔實率低下 30
十二、OsCP7:OsCP7-5之外表型與基因型之間具有正相關性 30
結論 32
參考文獻 33

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