臺灣博碩士論文加值系統

為了解蘭花早期生長發育是由哪些基因所調控，透過蝴蝶蘭V3 Sogo Yukidian花朵NGS資料庫的分析，選殖出在花朵分生組織 (floral meristem，FM) 及花苞中具有高表現量之基因PaSTM和PaMIP1。親緣演化樹分析顯示PaSTM和PaMIP1分別為阿拉伯芥AtSTM (KNOX gene family) 和AT4G37080 (ternary complex factor MIP1 leucine-zipper protein)、AT5G4260之同源基因。PaSTM在花分生組織及各時期花朵的子房中具有高表現量；而PaMIP1表現量隨花苞發育成熟逐漸累積，至花開後急遽下降。為探討PaSTM和PaMIP1的基因功能，透過模式植物阿拉伯芥基因轉殖進行功能性分析。35S::PaSTM和35S::PaSTM+VP16轉殖株葉片呈現短小、捲曲且有皺褶之形態；35S ::PaSTM+SRDX轉殖株則呈現細長葉片。在PaSTM轉殖株中，花瓣長度都較野生型短。35S::PaSTM和35S::PaSTM+VP16轉殖株中的花苞總數大於35S::PaSTM + SRDX轉殖株。35S ::PaSTM+SRDX轉殖株果莢長度較野生型短，且35S::PaSTM和35S::PaSTM+VP16轉殖株中促進莖頂分生組織 (SAM) 細胞增生之IPT7、GA2ox與KNAT6基因表現量上升，推測PaSTM可能為轉錄活化子，透過活化IPT7與GA2ox基因調控葉和花器的形態發育與KNAT6基因協調SAM形成與維持。此外，利用蝴蝶蘭病毒誘導基因靜默 (VIGS) 技術抑制PaSTM基因表現，結果使花苞總數減少、果莢減短，也印證了PaSTM調控分生組織發育的可能性。另外有發現花梗紅化與葉片黃化，猜測PaSTM可能與調控老化有相關。35S::PaMIP1和35S::PaMIP1 + SRDX轉殖株葉片面積較野生型大，而35S :: PaMIP1 + VP16轉殖株葉片面積則較小，推測PaMIP1在阿拉伯芥中以轉錄抑制子的特性調控葉片發育。

Early growth and development of orchids are regulated by genes. PaSTM and PaMIP1 are orchid genes with high transcription levels in the floral meristem (FM) and buds of Phalaenopsis, identified by referring to the Phalaenopsis V3 Sogo Yukidian flower NGS database. According to the phylogenetic analysis, PaSTM and PaMIP1 were homologous to Arabidopsis AtSTM (KNOX gene family) and AT4G37080 (ternary complex factor MIP1 leucine-zipper protein) / AT5G4260, respectively. PaSTM mRNAs are detected at a high level in the FM and ovaries of various flower stages. PaMIP1 mRNAs accumulate increasingly as floral buds mature and significantly decrease in the mature flowers. To investigate the gene functions of PaSTM and PaMIP1, the transgenic Arabidopsis system was applied. Transgenic plants carrying 35S::PaSTM and 35S::PaSTM+VP16 produced short, curl and wrinkled leaves, while that carrying 35S::PaSTM+SRDX produced slender leaves. In all PaSTM transgenic plants, the petal size was reduced. 35S::PaSTM and 35S::PaSTM+VP16 plants produced more floral buds than 35S::PaSTM+SRDX plants. 35S::PaSTM+SRDX transgenic plant has shorter fruit pods than the wild type. Besides, transcripts of IPT7, GA2ox and KNAT6, the genes promoting cell proliferation in SAM, were up-regulated in 35S::PaSTM and 35S::PaSTM+VP16 plants, which indicated that PaSTM plays as an activator to regulate the leaf and flower morphology by activating IPT7 and GA2ox, and coordinate SAM formation and maintenance by KNAT6. Furthermore, suppression of PaSTM activity in Phalaenopsis by VIGS technology reduced the floral bud formation and shortened the fruit pod, which corroborate the possibility that PaSTM regulates the meristem development. In addition, PaSTM suppressed plants displayed red peduncle and yellowish leaf. It is speculated that PaSTM may be related to the regulation of aging. 35S::PaMIP1 and 35S::PaMIP1+SRDX increased the leaf size, whereas 35S::PaMIP1+VP16 reduced the leaf size in transgenic plants, which implied that PaMIP1 plays as a repressor to regulate the leaf development.

第一章 蝴蝶蘭中PaSTM基因之功能性分析 Functional analysis of PaSTM gene in Phalaenopsis 1
前言 2
材料和方法 8
一、研究材料 8
二、阿拉伯芥種植 8
三、阿拉伯芥總體核糖核酸 (RNA) 萃取 8
四、蝴蝶蘭V3總體核糖核酸 (RNA) 萃取 9
五、阿拉伯芥反轉錄增幅反應 (Reverse transcription PCR) 9
六、蘭花反轉錄聚合酶酵素連鎖反應 9
七、即時定量聚合酶連鎖反應 (Real-time quantitative PCR) 9
八、聚合酶連鎖反應 (Polymerase Chain Reaction，PCR) 10
九、瓊脂凝膠膠片 (Agarose gel) 之配製 10
十、膠體電泳 (Electrophoresis) 10
十一、DNA片段回收與純化 11
十二、接合反應 (Ligation) 11
十三、大腸桿菌勝任細胞製備 (Competent cell preparation) 11
十四、大腸桿菌轉型作用 (Transformation into E.coli) 12
十五、轉形細菌之菌落聚合酶連鎖反應篩選 (Colony PCR) 12
十六、抽取高純度小量質體DNA (Preparation of plasmid DNA) 12
十七、限制酶酵素裁切 (Digestion of restriction enzyme) 13
十八、DNA定序與序列比對 13
十九、農桿菌 (Agrobacterium tumefaciens) 勝任細胞備製 13
二十、農桿菌轉型作用 13
二十一、阿拉伯芥之轉殖 13
二十二、轉基因阿拉伯芥之篩選 14
二十三、Gateway 基因重組選殖系統 (Gateway clone) 14
二十四、蝴蝶蘭病毒誘導基因靜默 (Virus induced gene silencing，VIGS) 14
二十五、阿拉伯芥植株葉片長寬統計與相對生長素率計算 15
二十六、甲苯胺藍 (Toluidine Blue Solution) 染色 15
二十七、科學製圖方法 15
結果 16
一、蝴蝶蘭V3基因PaSTM的基因選殖與序列分析 16
二、蝴蝶蘭V3中PaSTM的表現量分析 16
三、PaSTM基因大量表現、C端接上抑制子區域SRDX及C端接上活化子區
域VP16與病毒誘導基因靜默 (Virus induced gene silencing，VIGS) 技術之重組載體構築 17
四、大量異位表現PaSTM、PaSTM+SRDX及PaSTM+VP16於阿拉伯芥中影
響葉片型態、花器異常、果莢型態及種子數量 18
五、大量異位表現PaSTM、PaSTM+SRDX及PaSTM+VP16於阿拉伯芥轉殖
株中之外源性基因PaSTM及下游基因IPT7、GA2ox及KNAT6的表現量 19
六、利用病毒誘導基因靜默 (Virus induced gene silencing，VIGS) 技術抑制蝴蝶蘭V3 內生性基因STM影響花器之型態及花梗紅化 20
七、利用病毒誘導基因靜默 (Virus induced gene silencing，VIGS) 技術抑制蝴蝶蘭V3 內生性基因STM抑制花苞產量、開花速度與果莢長度 21
八、蝴蝶蘭葉片內生性基因STM基因的表現量分析 21
九、利用病毒誘導基因靜默 (Virus induced gene silencing，VIGS) 技術抑制蝴蝶蘭葉片內生性基因STM基因導致葉片黃化死亡 22
討論 23
參考文獻 27
圖表 33
表1-1、本實驗蝴蝶蘭基因PaSTM所使用之引子 (primer) 33
圖1-1、PaSTM 之編碼序列與胺基酸序列 34
圖1-2、PaSTM 與各物種KNOX之演化樹分析 35
圖1-3、PaSTM與阿拉伯芥中STM之胺基酸序列比對分析 36
圖1-4、蝴蝶蘭中PaSTM各時期各部位之表現量分析 37
圖1-5、PaSTM之選殖與大量表現PaSTM、PaSTM+SRDX及PaSTM+VP16構
築 38
圖1-6、大量異位表現PaSTM、PaSTM+SRDX及PaSTM+VP16於阿拉伯芥
中影響葉片型態之分析 39
圖1-7大量異位表現PaSTM、PaSTM+SRDX及PaSTM+VP16於阿拉伯芥中
影響花瓣面積大小和花苞數量之性狀分析 40
圖1-8大量異位表現PaSTM+SRDX於阿拉伯芥中影響果莢長度及種子數量
減少 41
圖1-9、大量異位表現PaSTM、PaSTM+SRDX及PaSTM+VP16於阿拉伯芥轉
殖株中之外源性基因PaSTM及下游基因IPT7、GA2ox與KNAT6的表現量42
圖1-10、蝴蝶蘭中PaSTM利用VIGS影響花器的結構及表現量分析 43
圖1-11、蝴蝶蘭中PaSTM利用VIGS使花瓣產生裂痕 44
圖1-12、蝴蝶蘭中PaSTM利用VIGS使花梗紅化 45
圖1-13、蝴蝶蘭中PaSTM利用VIGS使花苞數量減少及早期凋落 46
圖1-14、蝴蝶蘭中PaSTM利用VIGS抑制花苞產量及開花速度的統計 47
圖1-15、蝴蝶蘭中PaSTM利用VIGS使果莢長度減少及統計 48
圖1-16、蝴蝶蘭中PaSTM 葉子各時期之表現量分析 49
圖1-17、蝴蝶蘭中PaSTM 利用VIGS使葉子凋亡及表現量分析 50
圖1-18、PaSTM基因功能性的假想模型圖 51
附錄 52
附圖1-1、莖頂分生組織的結構和組織示意圖 52
附圖1-2、pGEM®-T Easy vector之圖譜 (3015 bp) 53
附圖1-3、pEpyon-22K之圖譜 (10293 bp) 54
附圖1-4、pEpyon-2aK之圖譜 (9801 bp) 55
附圖1-5、pEpyon-2bK之圖譜 (9801 bp) 56
第二章 蝴蝶蘭中PaMIP1基因之功能性分析 Functional analysis of PaMIP1 gene in Phalaenopsis 57
前言 58
材料和方法 61
結果 62
一、蝴蝶蘭V3基因PaMIP1的基因選殖與序列分析 62
二、蝴蝶蘭V3中PaMIP1的表現量分析 63
三、PaMIP1基因大量表現、C端接上抑制子區域SRDX及C端接上活化子區域VP16之重組載體構築 63
四、大量異位表現PaMIP1、PaMIP1+SRDX及PaMIP1+VP16於阿拉伯芥中
影響葉片 64
五、大量異位表現PaMIP1、PaMIP1+SRDX及PaMIP1+VP16於阿拉伯芥中
影響葉片型態 64
六、PaMIP1的兩個相似阿拉伯芥基因AT4G37080及AT5G42690於阿拉伯芥中影響葉片型態 65
七、利用病毒誘導基因靜默 (Virus induced gene silencing，VIGS) 技術抑制蝴蝶蘭V3內生性基因MIP1的表現量 65
討論 66
參考文獻 69
圖表 73
表2-1、本實驗蝴蝶蘭基因PaMIP1所使用之引子 (primer) 73
圖2-1、PaMIP1之編碼序列與胺基酸序列 74
圖2-2、PaMIP1 與各物種之演化樹分析 75
圖2-3、PaMIP1保守序列Lzipper-MIP1及DUF547與阿拉伯芥基因之演化樹分析 76
圖2-4、AT4G37080之編碼序列與胺基酸序列 77
圖2-5、AT5G42690之編碼序列與胺基酸序列 78
圖2-6、PaMIP1與阿拉伯芥基因AT4G37080及AT5G42690之胺基酸序列比對分析 79
圖2-7、蝴蝶蘭中PaMIP1各時期各部位之表現量分析 80
圖2-8、PaMIP1之選殖與大量表現PaMIP1、PaMIP1+SRDX及PaMIP1+VP16之構築 81
圖2-9、AT4G37080之選殖與大量表現AT4G37080、AT4G37080+SRDX及
AT4G37080+VP16之構築 82
圖2-10、AT5G42690之選殖與大量表現AT5G42690之構築 83
圖2-11、大量異位表現PaMIP1、PaMIP1+SRDX及PaMIP1+VP16於阿拉伯芥中植株的相對生長速率比較 84
圖2-12、大量異位表現PaMIP1、PaMIP1+SRDX及PaMIP1+VP16於阿拉伯芥中影響葉片型態之分析 85
圖2-13、大量異位表現AT4G37080、AT4G37080+SRDX及AT4G37080+VP16於阿拉伯芥中影響葉片型態之分析 86
圖2-14、大量異位表現AT5G42690於阿拉伯芥中影響葉片型態之分析 87
圖2-15、蝴蝶蘭中PaMIP1利用VIGS之表現量分析 88
圖2-16、PaMIP1基因功能性假想模型圖 89
附錄 90
附圖2-1、pGEM®-T Easy vector之圖譜 (3015 bp) 90
附圖2-2、pEpyon-22K之圖譜 (10293 bp) 91
附圖2-3、pEpyon-2aK之圖譜 (9801 bp) 92
附圖2-4、pEpyon-2bK之圖譜 (9801 bp) 93
附圖2-5、葉片橢圓形面積公式 94
附圖2-6、eFP Browser 中阿拉伯芥AT4G37080之表現趨勢 95
附圖2-7、eFP Browser 中阿拉伯芥AT5G42690之表現趨勢 96

第一章
許巍瀚，阿拉伯芥中調控細胞分裂與配子體發育相關基因之功能性分析，博士論文，國立中興大學生物科技學研究所，臺中 (2012)。
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第二章
許巍瀚，阿拉伯芥中調控細胞分裂與配子體發育相關基因之功能性分析，博士論文，國立中興大學生物科技學研究所，臺中 (2012)。
趙子翔，蝴蝶蘭中抑制生長速度之yippee like genes PaYIP B-1 及 PaYIPB-2之功能性分析，碩士論文，國立中興大學生物科技學研究所，臺中 (2019)。
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