臺灣博碩士論文加值系統

被子植物由營養生長到開花的轉換過程，受到環境訊息、發育、荷爾蒙等因素縝密地調控。然而，部分被子植物在身處不利於開花的條件時，會出現稱為「成花反轉」的現象，使得花分生組織逆轉為營養性或花序特性。在自然情形下，苦苣苔科的俄氏草（Titanotrichum oldhamii）在花季晚期會經歷成花反轉、產生營養性的珠芽繁殖體，行使無性生殖。過去的研究指出俄氏草的成花反轉與花分生組織特性基因FLORICAULA / LEAFY（FLO/LFY）表現量下降有關，本研究卻發現ToFLO除了主要在花分生組織表現，也在珠芽序生成位置表現，暗示ToFLO表現降低並非成花反轉的必要條件。另一方面，花序分生組織特性基因TERMINAL FLOWER 1 / CENTRORADIALIS（TFL1/CEN）在模式物種中曾被報導與成花反轉相關。藉由qPCR，我發現ToCEN在莖頂高量表現，與成花反轉及珠芽序形成有關；ToCEN的原位雜合訊號，則顯示ToCEN的表現位在花序分生組織的中心區域及珠芽序原基中。將ToCEN在阿拉伯芥中異位表現，不但延遲開花，還促進花序分支生長，花序結構與珠芽序的形態極為相似；轉殖株的花也發生成花反轉為葉化枝條、花萼包圍的次生花序、雌蕊著生花苞的退化花等。此外，在野生型阿拉伯芥嫁接至35S::ToCEN轉殖株的實驗中，能在接穗中偵測到ToCEN的RNA，顯示ToCEN可能透過訊息傳輸抑制花發育、促進成花反轉以影響花序結構。總結而言，本研究揭示ToCEN可能調控俄氏草的成花反轉及珠芽發育，且透過訊息傳遞促進花序延長、抑制花發育和延遲開花時間。此外，ToCEN在俄氏草中可能獲得了珠芽序發育的新功能，使俄氏草能夠以此行使獨特的無性繁殖、完成生活史。

In angiosperms, the switch from vegetative growth to flowering is orchestrated by environmental signals, development, hormones, etc. However, few angiosperms go through “floral reversion”, floral meristems revert to vegetative or inflorescence identity, under conditions unfavorable for flowering. Naturally occurred Titanotrichum oldhamii (Gesneriaceae) reverting flowers to vegetative bulbils (propagules) in late flowering season for asexual reproduction. Previous studies suggested that floral reversion in Titanotrichum is related to floral meristem identity genes, FLORICAULA/LEAFY (FLO/LFY). In this study, however, ToFLO was found mainly expressed in typical floral meristem and the down-regulation expression of ToFLO during the initiation of bulbiliferous shoots suggest it may not be essential for floral reversion. In contrast, inflorescence meristem identity genes TERMINAL FLOWER 1/ CENTRORADIALIS (TFL1/CEN) were reported related to floral reversion in model plants. qPCR indicates the expression level of ToCEN is high in shoot apices, and this correlated to stages of floral reversion and initiation of bulbiliferous shoots. In situ hybridization signals of ToCEN demonstrate it was localized in central region of inflorescence meristem and bulbiliferous shoot primordia. Ectopic expression of ToCEN in Arabidopsis showed enhanced inflorescence architecture with extensive outgrowth of lateral shoot (coflorescence) branching, a phenotype greatly resembling bulbiliferous shoots, and delayed floral transition. Conversion of flowers into leaf‐like shoots, secondary inflorescence subtended by sepals and rudimentary flowers with buds on carpel are often found. In addition, in grafting of wild type Arabidopsis onto 35S::ToCEN stocks, I found that ToCEN RNA can be detected in grafted scions. This suggests ToCEN acts as transmissible signals to inhibit flower development and promote floral reversion to alter inflorescence architecture systematically. Together, this study revealed that ToCEN may regulate floral reversion and bulbil development of Titanotrichum, and also systematically functions in enhancing inflorescence outgrowth, inhibition of flower formation and delaying flowering time. ToCEN thus perhaps adopted a new role for bulbiliferous shoots formation in Titanotrichum, allowing it to incorporate this unique life history trait for asexual propagation.

致謝 i
中文摘要 ii
Abstract in English iii
Table of Contents v
List of Figures ix
List of Tables xi
Abbreviations xii
1. Introduction 1
1.1. Floral reversion 1
1.2. Pseudovivipary and bulbils: an alternative reproductive strategy 3
1.3. Titanotrichum oldhamii, a bulbiliferous plant with floral reversion 4
1.4. Regulation of floral transition, meristem identity and maintenance 7
1.5. Inflorescence meristem identity genes: TFL1/CEN 10
1.6. Aim of this study 14
2. Materials and Methods 16
2.1. Plant materials and growth condition 16
2.2. Molecular study 16
2.2.1. Genomic DNA extraction 16
2.2.2. Total RNA extraction 18
2.2.3. DNase treatment and advanced purification of total RNA 19
2.2.4. First-strand cDNA synthesis 20
2.2.5. Standard polymerase chain reaction (PCR) 21
2.2.6. Agarose gel electrophoresis 22
2.2.7. Quantitative real-time PCR (qPCR) 23
2.3. RNA in situ hybridization (ISH) 26
2.3.1. Fixation, dehydration and embedding 26
2.3.2. Microtomy 27
2.3.3. Production of large amount of plasmid for RNA probes 28
2.3.4. DIG-labeling in vitro transcription 29
2.3.5. Dot-blotting for quantification of RNA probes 31
2.3.6. In situ hybridization 32
2.3.7. Reaction termination and Imaging 35
2.4. Functional study of ToCEN using transgenic Arabidopsis 36
2.4.1. Selection of transformed seeds 36
2.4.2. Aseptic sowing and further cultivation on soil 36
2.4.3. Scanning electron microscope (SEM) 37
2.4.4. Statistical analysis 37
2.4.5. Grafting 38
3. Results 39
3.1. Field observation 39
3.2. RT-PCR for ToCEN, ToTFL and ToFLO in Titanotrichum 42
3.2.1. Design of RT-PCR 42
3.2.2. RT-PCR results of ToCEN, ToTFL and ToFLO 42
3.3. qPCR for ToCEN and ToTFL in Titanotrichum 45
3.3.1. Design of qPCR 45
3.3.2. qPCR results for ToCEN and ToTFL 45
3.4. RNA in situ hybridization results of ToCEN in Titanotrichum 48
3.5. Functional study of ToCEN overexpression in Arabidopsis 52
3.5.1. Selection of 35S::ToCEN transgenic plants via GFP and PCR 52
3.5.2. Weak and strong phenotype of 35S::ToCEN T2 plants 54
3.5.3. Effects on morphology in 35S::ToCEN T2 plants 58
3.5.4. Effects on floral transition in 35S::ToCEN T2 plants 61
3.6. Grafting of wild-type Arabidopsis on 35S::ToCEN T2 plants 66
4. Discussion 70
4.1. ToCEN expression was linked to floral reversion, while ToTFL was perhaps related to later stages of bulbil development 70
4.2. Spatial pattern of ToCEN expression is similar to other TFL1/CEN-like genes, but additionally associated with bulbil development 72
4.3. ToFLO expression may also acquire new functions in inflorescence maintenance and bulbiliferous shoot formation 73
4.4. ToCEN functions in controlling inflorescence architecture, inhibition of flower formation, and delaying floral transition 75
4.4.1. ToCEN regulated inflorescence architecture by affecting the length of main axis and lateral branching 75
4.4.2. Flower formation was also inhibited by ToCEN, leading to flower and inflorescence reversion 77
4.4.3. Ectopic expression of ToCEN in A. thaliana delayed floral transition 78
4.4.4. ToCEN may regulate floral reversion by graft-transmissible signals 79
Future aspects 82
References 84
Appendices 92
Appendix 1. Gene sequence used in this study 92
Appx. 1.1. Sequence of ToCEN (CDS and 3’-UTR) 92
Appx. 1.2. Sequence of ToTFL (CDS and 3’-UTR) 93
Appx. 1.3. Sequence of ToFLO (CDS and 3’-UTR) 94
Appendix 2. Phylogenetic reconstruction of TFL1/CEN-like genes 96
Appx. 2.1. Phylogenetic reconstruction of TFL1/CEN-like genes 96
Appx. 2.2. List of TFL1/CEN-like genes used in phylogeny 97
Appendix 3. Primer used in this study 99
Appx. 3.1. Primer used in RT-PCR 99
Appx. 3.2. Primer used in qPCR 100
Appx. 3.3. Primer used for in situ hybridization probe synthesis 101
Appx. 3.4. Primer used for identifying transgenic Arabidopsis 101
Appendix 4. qPCR primer efficiency test 102
Appx. 4.1. ToTubA1 102
Appx. 4.2. ToCEN 103
Appx. 4.3. ToTFL 104
Appx. 4.4. AtTubB 105
Appendix 5. In situ hybridization results for ToFLO in Titanotrichum 106
Appendix 6. Design of pFAST-G02 fusion with 35S::ToCEN 108
Appendix 7. Phenotypes of 35S::ToCEN T1 Arabidopsis 109
Appendix 8. Solution recipes for molecular study 110
Appx. 8.1. Experiments except for ISH 110
Appx. 8.2. Solution recipes for ISH 111
Appendix 9. qPCR of ToCEN expression in 35S::ToCEN T2 seedlings 112

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