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完整後設資料紀錄
DC 欄位 | 值 | 語言 |
---|---|---|
dc.contributor.advisor | 王俊能(Chun-Neng Wang) | |
dc.contributor.author | JenYu Chang | en |
dc.contributor.author | 張仁育 | zh_TW |
dc.date.accessioned | 2021-06-15T04:15:49Z | - |
dc.date.available | 2020-01-01 | |
dc.date.copyright | 2010-01-21 | |
dc.date.issued | 2009 | |
dc.date.submitted | 2010-01-09 | |
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dc.identifier.uri | http://tdr.lib.ntu.edu.tw/jspui/handle/123456789/45355 | - |
dc.description.abstract | 部分被子植物已分化的花芽或花序分生組織,會逆轉回營養芽或營養性莖頂,這種現象前人統稱為「成花反轉」。一般推測,成花反轉現象能保障該類植物具有多年性生活史、或具有假性胎生(如產生珠芽等無性繁殖體)繁殖策略。臺灣原生之苦苣苔科植物俄氏草(Titanotrichum oldhamii)於其一年為週期之生活史中具獨特的成花反轉(花序轉換)現象:成花花序於花期後轉變為珠芽枝,即由重複產生側生的珠芽(枝)分生組織取代花芽分生組織以進行假性胎生。為檢視俄氏草花序轉換的可能機制,本研究中釣取與花序及成花相關的四個候選調節基因:協調花芽形成的花芽分生組織特性基因ToFLO、概與其拮抗之花序分生組織特性基因ToCEN、莖頂分生組織幹細胞特性基因ToROA,及花芽分化末期抑制幹細胞活性的花器特性基因ToPLE,並相互比較其在不同發育時期基因表現與形態上的關聯。 各候選基因中唯獨花序分生組織ToCEN的表現趨勢吻合發育上的轉換:原先未在分化花芽中表現之ToCEN卻於將成為珠芽枝的分生組織中持續表現,此即暗示俄氏草花序反轉與分生組織額外獲賦予營養型生長之花序特性有關;另外花芽分生組織特性基因ToFLO仍可表現在側生之珠芽(枝)分生組織中,推測其表現下降時非花序轉換發生時之必要條件,而ToFLO表現亦非步向花芽分化之充分條件,故尚無證據支持俄氏草花序轉換係與花芽分生組織失去成花特性有關。本研究初步顯示俄氏草花序轉換概與花序分生組織特性基因ToCEN表現相關,預計可再進行功能性分析以釐清ToCEN所扮演的角色,及明瞭俄氏草如何藉此機制調節花序轉換。 | zh_TW |
dc.description.abstract | In certain angiosperms, vegetative growth, such as leaves and asexual propagules formation on inflorescence, may reappear after floral commitment. This phenomenon is named “floral reversion”. Floral reversion is probably essential for pseudovivpary plants’ survival for its life history, i.e. sexual reproduction can be replaced by asexual propagation, such as bulbils. Titanotrichum oldhamii (Gesneriaceae), one of pseudoviviparous plant, is native in Taiwan. Its flowering inflorescence can reverse to the bulbiliferous shoot after flowering. What molecular mechanism controls the floral reversion in Titanotrichum is of particular interest. Four candidate genes including floral meristem identity gene ToFLO, inflorescence meristem identity gene ToCEN, floral organ identity gene ToPLE and shoot apical meristem stem cell identity gene ToROA were isolated in this study. Expression patterns of these genes were compared among developmental stages: before and during the inflorescence transition. Putative inflorescence meristem identity gene ToCEN is the only candidate gene whose expression was alternated when inflorescence transition occurred. During the floral differentiation, ToCEN is not expressed in the floral meristems but in meristems that are potentially transferred to the bulbiliferous shoot. It is known that overexpression of TFL1/CEN homolog in Arabidopsis and rice promotes vegetative growth and branching of inflorescence. Thus, inflorescence transition in Titanotrichum may be associated with ToCEN in obtaining vegetative meristem identity. In contrast to ToCEN, the floral meristem identity gene ToFLO could somehow be expressed in bulbiliferous meristem. This result might imply that down-regulation of ToFLO is not necessary for inflorescence transition, and ToFLO is not sufficient to provide floral differentiation as well. Thus, there is not enough evidence to support that inflorescence transition is due to loss of floral meristem identity. In conclusion, I propose the inflorescence meristem identity gene ToCEN is a good candidate for studying the inflorescence transition in Titanotricum. Further functional study would be helpful to ensure the role of ToCEN in inflorescence transition. | en |
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dc.description.tableofcontents | 謝辭……………...………………………………………………………….i
Abstract in Chinese ……………...………………………………………iii Abstract in English ………………………..……………………………..iv Table of Contents ………………………………………………………...vi List of Figures ………………………………………………………………………….x List of Tables ………………………………………………………………………….xii Abbreviations …………………………………………………………..xiii 1. Introduction ...…………………………………………………………1 1.1. General background of floral reversion and pseudovivipary in Titanotrichum oldhamii……...………………………………………………….1 1.1.1. Floral reversion: Vegetative growth after a period of flowering process ….………………..1 1.1.2. Floral reversion may serve the function of perenniality and pseudovivipary in life history of plants …………………………………………………………………………………………1 1.1.3. Pseudovivipary, an asexual reproductive strategy in plants growing in patch and limited habitat or extreme climates ……………………………………………………………………..2 1.1.4. Titanotrichum oldhamii, the native pseudoviviparous species in Taiwan…………………….3 1.1.5. Hypothesis of floral reversion mechanism: loss of floral maintenance and / or gain of inflorescence / vegetative meristem identity……………….…………………………………...4 1.2. Choices of candidate genes …………………………………………………….6 1.2.1. Signaling pathways of flowering ……………………………………………………………….6 1.2.2. Flowering integrators …………………………………………………………………………...6 1.2.2.1. FLOWERING LOCUS C (FLC) ……………………………………………………………6 1.2.2.2. TERMINAL FLOWER2 (TFL2)/ LIKE HETEROCHROMATIN PROTEIN 1 (LHP1) ...7 1.2.2.3. FLOWERING LOCUS T (FT) …………………………………………………………...8 1.2.2.4. SUPPRESSOR OF OVEREXPRESSION OF CONSTANS 1 (SOC1) ………………….9 1.2.3. Meristem identity genes ………………………………………………………………………...9 1.2.3.1. The inflorescence meristem identity gene: TERMINAL FLOWER1 (TFL1) in Arabidopsis and CENTRORADIALIS (CEN) in Antirrhinum ……………………………9 1.2.3.2. The floral meristem identity gene: LEAFY (LFY) in Arabidopsis and FLORICAULA (FLO) in Antirrhinum …………………………………………………………..…………11 1.2.4. Floral meristem determinacy genes …………………………………………………………..12 1.2.5. Meristem identity and determinacy genes serve as good candidates in preliminary study of floral reversion …………………………………………………………………………………13 1.2.6. Finding out the robust candidate that correlated to inflorescence transition in Titanotrichum …………………………………………………………………………………..14 2. Materials and Methods ……………………………………………...16 2.1. Plant materials ………………………………………………………………..16 2.2. Histological study ……………………………………………………………..16 2.2.1. Paraffin section ………………………………………………………………………………..16 2.2.1.1. Samples fixation ………………………………………………………………………….16 2.2.1.2. Paraffin embedding ………………………………………………………………………17 2.2.1.3. Section ……………………………………………………………………………………..17 2.2.1.4. Staining of the samples on slides …………………………………………………………18 2.2.1.5. Mounting and taking the photos …………………………………………………………19 2.3. Molecular study ………………………………………………………………….19 2.3.1. Preparation of basic DNA / cDNA resource ……………………………………………….....19 2.3.1.1. Genomic DNA extraction ……………………………………………………………………...19 2.3.1.2. RNA extraction ………………………………………………………………………………...20 2.3.1.3. First-strand cDNA synthesis …………………………………………………………………..21 2.3.2. Gene cloning-related methods ………………………………………………………………...22 2.3.2.1. Standard polymerase chain reaction (PCR) ………………………………………………..22 2.3.2.2. Thermal asymmetric interlaced PCR (TAIL-PCR) …………………………………………23 2.3.2.3. Rapid amplification of 3' cDNA ends (3'-RACE) ……………………………………………26 2.3.2.4. Rapid amplification of 5' cDNA ends (5'-RACE) ……………………………………………27 2.3.2.5. Agarose gel electrophoresis ……………………………………………………………………29 2.3.2.6. Cloning ………………………………………………………………………………………….29 2.3.2.7. Clones identification …………………………………………………………………………...30 2.3.2.8. Nucleotide sequencing …………………………………………………………………………32 2.3.3. Nucleotide sequences alignment and analysis ………………………………………………..32 2.3.3.1. Nucleotide sequence identification …………………………………………………………..32 2.3.3.2. Sequence alignment ……………………………………………………………………………33 2.3.3.3. Promoter and cis-acting regulatory DNA elements prediction ……………………………..33 2.3.3.4. Homology test …………………………………………………………………………………..34 2.3.4. Expression analysis of candidate genes ………………………………………………………34 2.3.4.1. RT-PCR ……………………………………………………………………………………….34 2.3.4.2. In situ hybridization …………………………………………………………………………...36 2.3.4.2.1. Samples fixation and dehydration ……..………………………………………………...36 2.3.4.2.2. Paraffin embedding ……………………………………………………………………….36 2.3.4.2.3. In vitro transcription for DIG-labeled probes generation ……………………………...37 2.3.4.2.4. Section, deparaffinization and rehydration……………………………………………...38 2.3.4.2.5. Proteinase K treatment, post-fixation and acetylation of samples……..……………….39 2.3.4.2.6. Dehydration ……………………………………………………………………………….40 2.3.4.2.7. Hybridization ……………………………………………………………………………...40 2.3.4.2.8. Post-hybridization washing and RNase A treatment …………………………………...40 2.3.4.2.9. Blocking and antibody reaction ….………………………………………………………41 2.3.4.2.10. Dectection and mounting …...…………………………………………………………….42 3. Results ………………………………………………………………..43 3.1. Field observation ……………………………………………………………...43 3.2. Paraffin section ………..…...………………………………………………....45 3.3. Candidate genes cloning ………………………………………………….…..49 3.3.1. Homolog of TFL1 and CEN …………………………………………………………….……..49 3.3.2. Homolog of LFY/FLO …………………………………………………………………….……53 3.3.3. Homolog of WUS/ROA …………………………………………………………….…………..53 3.3.4. Homolog of PLE, the AG-like MADS gene ………………………………………………......55 3.4. RT-PCR results for all candidate genes in our research …………………...58 3.4.1. RT-PCR design …………………………………………………………………………………58 3.4.1.1. Primer sets design in RT-PCR ………………………………………………………........58 3.4.1.2. cDNA resources selected in RT-PCR ………………………………………………….....60 3.4.2. RT-PCR results …………………………………………………………………………….......60 3.4.3. Further confirm in RT-PCR result of ToTFL and ToPLE ………………………………......64 3.4.3.1. Restrictional fragment length analysis in ToTFL RT-PCR result …………………......64 3.4.3.2. Restrictional fragment length analysis in ToTFL RT-PCR result ……………………...65 3.5. RNA in situ hybridization results for ToFLO and ToCEN …………………67 3.5.1. Choosing of fixatives in my research: FAA was better than PGA in RNA in situ hybridization for less background …………………………………………………………....67 3.5.2. RNA in situ hybridization for ToCEN ………………………………………………………...68 3.5.3. RNA in situ hybridization for ToFLO ………………………………………………………...69 3.5.4. Summary of RNA in situ hybridization of ToCEN and ToFLO in meristems on Titanotrichum inflorescence …………………………………………………………………...70 4. Discussion ………………………………………………………….....77 4.1. Observation of inflorescence transition in Titanotrichum ………………….77 4.1.1. Stages categorization of inflorescent transition in Titanotrichum …………………………..77 4.1.2. Floral / bulbiliferous meristems are gradually changed during inflorescence transition ………………………………………………………………………………………..77 4.2. Candidate genes cloning and its expression ………………………………...80 4.2.1. Meristem identity gene ………………………………………………………………………...80 4.2.1.1. ToTFL and ToCEN ………………………………………………………………………...80 4.2.1.1.1. Duplication of TFL1/CEN-like genes in eudicots ……………………………………….80 4.2.1.1.2. ToTFL shows a novel residual substitution in conserve region ………………………...80 4.2.1.1.3. ToCEN shares similar putative promoter region with CEN …………………………....81 4.2.1.1.4. ToTFL was expressed in all tissues we sampled ………………………………………...82 4.2.1.1.5. Change of ToCEN expression matched with transition of inflorescence, indicating ToCEN is associated with floral reversion in Titanotrichum ……………………………82 4.2.1.2. ToFLO ……………………………………………………………………………………...84 4.2.1.2.1. Transcripts levels of ToFLO are not absolutely link to its development fate of inflorescence ……………………………………………………………………………….84 4.2.2. Floral meristem determinacy-related candidate genes ……………………………………...87 4.2.2.1. ToROA ……………………………………………………………………………………...87 4.2.2.2. ToPLE ……………………………………………………………………………………...88 5. Future aspect ………………………………………………………...90 Reference ………………………………………………………………..92 Appendices ……………………………………………………………..110 Appendix 1. The synthetic flowering signaling pathway in Arabidopsis …………110 Appendix 2. Gene sequence obtained in this study ………………………………..111 Appx. 2.1. Sequence of ToTFL (CDS and 3’-UTR) ……………………………………………111 Appx. 2.2. Sequence of ToCEN ………………………………………………………………….112 Appx. 2.2.1. Sequence of ToCEN (promoter and 5’-UTR, aligned with CEN) ………...112 Appx. 2.2.2. Sequence of ToCEN (CDS and 3’-UTR) …………………………………...113 Appx. 2.3. Sequence of ToFLO (CDS and 3’-UTR) ……………………………………………114 Appx. 2.4. Sequence of ToROA (CDS and 3’-UTR) …………………………………………....115 Appx. 2.5. Sequence of ToPLE …………………………………………………………………..116 Appx. 2.5.1. Sequence of ToPLE (promoter and 5’-UTR) ………………………………116 Appx. 2.5.2. Sequence of ToPLE (CDS and 3’-UTR) …………………………………....117 Appx. 2.5.3. Sequence of ToPLE (5’-RACE result) ……………………………………...118 Appendix 3. Accession numbers of genes used in homology test …………………119 Appendix 4. Fractional repeat of ToCEN and ToFLO RT-PCR ………………….121 Appendix 5. Buffers used in in situ hybridization ………………………………...122 | |
dc.language.iso | en | |
dc.title | 俄氏草花序轉換中花序特性與成花定性基因之研究 | zh_TW |
dc.title | Study of Inflorescence Identity and Floral Determinacy Genes on Inflorescence Transition in Titanotrichum | en |
dc.type | Thesis | |
dc.date.schoolyear | 98-1 | |
dc.description.degree | 碩士 | |
dc.contributor.oralexamcommittee | 黃玲瓏(Ling-Long Kuo-Huang),陳仁治(Jen-Chih Chen),陳虹樺(Hong-Hwa Chen),楊長賢(Chang-Hsien Yang) | |
dc.subject.keyword | 俄氏草,成花反轉,假性胎生,珠芽,TERMINAL FLOWER1 / CENTRORADIALIS,LEAFY / FLORICAULA, | zh_TW |
dc.subject.keyword | Titanotrichum oldhamii,floral reversion,pseudovivipary,bulbil,TERMINAL FLOWER1 / CENTRORADIALIS,LEAFY / FLORICAULA, | en |
dc.relation.page | 122 | |
dc.rights.note | 有償授權 | |
dc.date.accepted | 2010-01-11 | |
dc.contributor.author-college | 生命科學院 | zh_TW |
dc.contributor.author-dept | 生態學與演化生物學研究所 | zh_TW |
顯示於系所單位: | 生態學與演化生物學研究所 |
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