台灣蝴蝶蘭之低溫逆境反應和PaCBF1和PaICE1表現的分析

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研究生：	彭柏欣 Peng, Po-Hsin
論文名稱：	台灣蝴蝶蘭之低溫逆境反應和PaCBF1和PaICE1表現的分析 Cold response in Phalaenopsis aphrodite and characterization of PaCBF1 and PaICE1
指導教授：	林彩雲
口試委員:	曾夢蛟黃鎮剛楊明德楊長賢
學位類別：	博士 Doctor
系所名稱：	生命科學暨醫學院 - 生物資訊與結構生物研究所 Institute of Bioinformatics and Structural Biology
論文出版年：	2014
畢業學年度：	102
語文別：	英文
論文頁數：	91
中文關鍵詞：	低溫逆境、臺灣白花蝴蝶蘭
外文關鍵詞：	C-repeat binding factor, dyhydrin, inducer of CBF expression, MYC motif, Phalaenopsis aphrodite
相關次數：	點閱：2 下載：0
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蝴蝶蘭為一種冬天開花的蘭科植物，廣泛栽培於亞洲許多熱帶國家。由於蝴蝶蘭是台灣主要經濟花卉之一，故在低溫等逆境下蝴蝶蘭生理及基因調控具有進一步研究之重要性。這篇論文主要探討臺灣白花蝴蝶蘭 (P. aphrodite) 對低溫及其他非生物逆境的反應，由我們的實驗結果得知臺灣白花蝴蝶蘭對低溫敏感，尤其在植株開花期，4°C處理的生長期植株比開花期植株葉片細胞膜穩定且光合作用效率較高。在分子層面，低溫誘導C-repeat binding factor1 (PaCBF1) 基因及可能受其調控的下游基因dehydrin1 (PaDHN1) mRNAs累積，但inducer of CBF expression1 (PaICE1) 的mRNA則持續表現。PaICE1可活化PaCBF1啟動子及存在該區域的MYC motifs，推測PaICE1可能藉由MYC motifs調控PaCBF1；PaCBF1在低溫下可活化PaDHN1啟動子，顯示PaCBF1可能參與低溫調控PaDHN1的表現。在未受低溫刺激的環境，大量表現PaCBF1基因的轉殖阿拉伯芥相較於野生型具有較高的逆境基因COR6.6及RD29a表現量，且在低溫處理後維持較好的細胞膜完整性。在此篇論文中我們提供證據說明在蘭科植物低溫逆境經由PaICE1蛋白調控PaCBF1基因表現，並誘導DHN等cold-regulated (COR) 基因表現達到降低低溫傷害，就我們所知這也是第一篇從蘭科植物選殖並分析CBF、DHN及ICE基因的論文。

Phalaenopsis is a winter-blooming orchid genus commonly cultivated in tropical Asian countries. Because orchid is one of the most economically important flower crops in Taiwan, it is crucial to understand its response to cold and other abiotic stresses. The present study focused on gene regulation of P. aphrodite in response to abiotic stress, mainly cold. Our results demonstrate that P. aphrodite is sensitive to low temperatures, especially in its reproductive stage. We found that after exposure to 4°C, plants in the vegetative stage maintained better membrane integrity and photosynthetic capacity than in the flowering stage. At the molecular level, C-repeat binding factor1 (PaCBF1) and its putative target gene dehydrin1 (PaDHN1) mRNAs were induced by cold, whereas inducer of CBF expression1 (PaICE1) mRNAs were constitutively expressed. PaICE1 transactivated MYC motifs in the PaCBF1 promoter, indicating that up-regulation of PaCBF1 may be mediated by the binding of PaICE1 to MYC motifs. PaCBF1 transactivated PaDHN1 at cold treatment, suggesting PaCBF1 was involved in the cold-regulation of PaDHN1. Overexpression of PaCBF1 in transgenic Arabidopsis induced AtCOR6.6 and RD29a without cold stimulus and maintain better membrane integrity after cold stress. Herein, we present evidence that cold induction of PaCBF1 transcripts in P. aphrodite may be transactivated by PaICE1 and consequently protect plants from cold damage through up-regulation of cold-regulated (COR) genes, such as DHN. To our knowledge, this study is the first report of isolation and characterization of CBF, DHN and ICE genes in the Orchidaceae family.

Table of Contents
                                                       Page
中文摘要及關鍵字 ......................................... i
Abstract and Keywords ................................. ii
謝誌 ................................................... iii
Abbreviations ......................................... iv
Table of Contents ..................................... vi
List of Tables ........................................ viii
List of Figures ....................................... ix
List of Appendix Figures .............................. xi
1. Introduction ....................................... 1
1.1 The Phalaenopsis aphrodite plants ................. 1
1.2 Low-temperature effects on P. aphrodite ........... 2
1.3 Cold-stress regulated genes ....................... 3
1.4 Thesis objectives and accomplishments ............. 5
2. Materials and Methods .............................. 6
2.1 Plant materials and stress treatments ............. 6
2.2 Electrolyte leakage measurement and photosynthesis measurement ........................................... 7
2.3 RNA isolation, cDNA synthesis and semi-quantitative/quantitative RT-PCR ...................... 9
2.4 Genomic DNA extraction ............................ 11
2.5 Gene isolation and bioinformatic analysis ......... 12
2.6 PEG-mediated protoplast transformation ............ 14
2.7 Quantification of RLUC and GUS activity in transformants                   ....................................................... 15
2.8 DNA blotting ...................................... 17
3. Results ............................................ 18
3.1 Exposure to 4°C decreased cell membrane stability and photosynthetic efficiency in P. aphrodite plants ...... 18
3.2 Isolation and characterization of PaCBF1, PaICE1 and PaDHN1 genes from P. aphrodite ........................ 19
3.3 Cold stress significantly increased transcript level of PaCBF1 and PaDHN1 but not PaICE1 ...................... 24
3.4 The PaCBF1 and its promoter was activated by abiotic stresses and PaICE1, and PaCBF1 was able to transactivate PaDHN1 with cold treatment ............................ 26
3.5 Overexpression of PaCBF1 regulated AtCOR genes in transgenic Arabidopsis plants ......................... 28
4. Discussion ......................................... 29
5. References ......................................... 34

List of Tables
                                                      Page
Table 1 List of primers used in this study ........... 43
Table 2 The websites of on-line software programs used in this thesis .......................................... 47
Table 3 Statistical analysis of Fv/Fm ratio versus time in cold stressed P. aphrodite plants .................... 48
Table 4 Sequence of deduced PaCBF1 protein was compared with other Phalaenopsis aphrodite homologues in the Orchidstra database and some AP2/ERF proteins that have been functionally-studied (Fig 5) ......................... 49
Table 5 Sequence similarity analysis of PaICE1 compared with the homologues in Fig 8 .............................. 52
Table 6 Consensus sequences for transcription factors binding to the PaDHN1 promoter predicted by PLACE .... 54
Table 7 Sequence similarity analysis of PaDHN1 compared with the homologues in Fig 9 .............................. 56
Table 8    Consensus sequences for transcription factors binding to the PaCBF1 promoter predicted by PLACE .... 58
 
List of Figures
                                                        Page
Fig. 1 Symptoms of low temperature damage to P. aphrodite plants ................................................. 59
Fig. 2 Effects of cold stress on cell membrane stability and photosynthesis of P. aphrodite ......................... 60
Fig. 3 The nucleotide sequence and deduced amino acid sequence of PaCBF1 cDNA with 5-UTR and partial promoter.61
Fig. 4 DNA blot analysis of PaCBF1 ..................... 62
Fig. 5 Sequence alignment and phylogenetic analysis of PaCBF1 and AP2/ERF proteins from different species ..... 64
Fig. 6 The nucleotide sequence and deduced amino acid sequence of PaICE1 ..................................... 70
Fig. 7 Nucleotide sequence alignment of PaICE1 and PATC134808 ............................................. 71
Fig. 8 Sequence alignment and phylogenetic analysis of PaICE1 protein and MYC homologous from different species.73
Fig. 9 The nucleotide sequence and deduced amino acid sequence of PaDHN1, and multiple sequence alignment of PaDHN1 and its homologous from other species ........... 76
Fig. 10 Subcellular localization of the PaCBF1 and PaICE1 proteins ............................................... 79
Fig. 11 Transcript levels of PaCBF1, PaDHN1 and PaICE1 in leaves of P. aphrodite in response to 4°C at different times ........................................................ 80
Fig. 12 Transcript levels of PaCBF1, PaDHN1 and PaICE1 in different tissues of P. aphrodite in response to cold stress ........................................................ 81
Fig. 13 Expression of PaCBF1, PaDHN1 and PaICE1 in leaves of P. aphrodite in response to various abiotic stresses ... 82
Fig. 14 Quantitative analysis of PaCBF1 promoter activity in response to various abiotic stresses ................... 83
Fig. 15    Quantitative analysis of PaICE1-mediated transactivation of PaCBF1 promoter and MYC box and PaCBF1-mediated transactivation of PaDHN1 promoter ............ 85
Fig. 16    Expression patterns of stress responsive genes and freezing tolerance analysis in wild type (WT, Col-0) and 35S::PaCBF1 transgenic Arabidopsis plants (OE).......... 86
 
Appendix Figures
                                                       Page
Fig. A1 35S::PaCBF1-EGFP .............................. 87
Fig. A2. 35S::PaICE1-EGFP ............................. 88
Fig. A3. 35S::PaICE1 .................................. 89
Fig. A4. PaCBF1 promoter::GUS (p1311) ................. 90
Fig. A5. PaDHN1 promoter::GUS (p1474) ................. 91

                                

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