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研究生：	張氏秀貞 Trinh, Truong Thi Tu
論文名稱：	探討阿拉伯芥葉片生長發育時PAP-FIBRILLIN基因族群的表現模式及可能受賀爾蒙和非生物逆境調控途徑 Investigate the expression patterns of PAP-fibrillin family genes and their putative regulatory networks in response to developmental and environmental signals in Arabidopsis
指導教授：	李瑞花 Lee, Ruey-Hua 吳文鑾 Wu, Wen-Luan
學位類別：	碩士 Master
系所名稱：	生物科學與科技學院 - 生命科學系 Department of Life Sciences
論文出版年：	2015
畢業學年度：	103
語文別：	英文
論文頁數：	87
中文關鍵詞：	阿拉伯芥、葉綠體、葉片老化、類囊膜瓦解、PAP-fibrillin 、Plastoglobuli
外文關鍵詞：	Arabidopsis thaliana, leaf senescence, thylakoid membrane degradation, PAP-fibrillin, Plastoglobuli
相關次數：	點閱：105 下載：3
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葉片黃化時葉綠體瓦解時，類囊膜的瓦解是最早受破壞的胞器內結構，而且可以觀察到葉綠體內部大量形成plastoglobule且數目隨著瓦解程度而增加，Plastoglobule是單膜結構主要成分為半乳糖酯，並有蛋白質(如PAP-fibrillins)崁鑲在單膜表面上，PAP-fibrillins 被認為扮演穩定plastoglobule的結構型態、新陳代謝物運輸及抵抗逆境功能，為了瞭解發育及環境網絡如何調控PAP-fibrillin族群基因的表現，本研究利用阿拉伯芥為模式植物，探討14個PAP-fibrillin族群基因在葉片發育及老化不同發育時期的表現模式及是否受不同賀爾蒙及非生物逆境的調控，大部分這些基因在不同葉片發育及黃化時期有雙峰表現模式，包括早期的葉片發育及葉片開始黃化前期。目前使用的賀爾蒙及非生物逆境處理的阿拉伯芥植株的葉片，只有少數基因受部分處理方法有改變表現模式。從promoter序列分析，我們初步有分離40個引子基因可能扮演調控PAP-fibrillin族群基因表現的腳色，其中有分離出17個葉片老化相關的引子基因，針對目前分析結果，針對 PAP-fibrillin族群基因表現的調控網絡進行討論

Degradation of the thylakoid membrane is the first step for chloroplast breakdown during leaf senescence and is masked by increase number and size of plastoglobuli. Plastoglobuli consist of an outer galactolipid monolayer studded with PAP-fibrillin on the surface of plastoglobuli. PAP-fibrillin suggested playing roles in maintaining the structural stability of plastoglobuli, metabolite transport and against biotic and abiotic stresses. We are interested in dissecting the regulatory network of PAP-fibrillin genes in response to developmental and environmental signals during leaf senescence in Arabidopsis. We have analyzed expression profiles of 14 PAP-fibrillin genes during different stages of leaf development and senescence. Most of these genes were induced to high levels of expression in early leaf development and then reduced to low levels as leaf developed to maturity. These genes then again induced to high levels during onset of leaf senescence. We have also analyzed the expression profiles of these genes in response to different abiotic stresses and hormonal treatments. Promoter analysis for these family genes, we have identified 40 putative transcription factors and 17 of these were induced during senescence. The putative regulatory networks of PAP-fibrillin gene family in response to developmental and environmental signals are discussed.

CONTENTS
ABSTRACT ...... I
中文摘要 ..... II
ACKNOWLEDGEMENTS ..... III
ABBREVIATIONS ...... IV
CONTENTS ...... V
1. INTRODUCTION ...... 1
1.1. Structure and function of chloroplast .... 1
1.1.1. Structure of chloroplast .... 1
1.1.2. Composition of thylakoid membrane ... 2
1.1.3. Structural function of thylakoid membrane .... 3
1.2. Chloroplast degradation during leaf senescence ... 4
1.2.1. Cellular view .... 5
1.2.2. Degradation of thylakoid membrane .... 6
1.3. Formation of plastioglobules is a hallmark of chloroplast degradation .. 7
1.3.1. Structure and composition of plastioglobules .... 8
1.3.2. Function of plastioglobuli ..... 9
1.3.3. PAPs/fibrillins ..... 10
1.4. Regulation and signaling during leaf senescence in response to developmental and environmental signals ..... 10
1.4.1. Senescence-associated genes .... 10
1.4.2. Hormonal regulation of leaf senescence ... 12
1.5. Research aim ...... 12
2. MATERIALS AND METHODS .... 13
2.1. Growth and maintenance of Arabidopsis plant .... 13
2.2. Abiotic and hormone treatments .... 14
2.3. Total RNA extraction and quantitation .... 14
2.4. Non-denaturing RNA and DNA agarose gel electrophoresis .. 15
2.5. Total chlorophyll extraction and quantitation .... 16
2.6. Total protein extraction and quantitation ... 16
2.7. Retrieval of PAP-fibrillin cDNA, gDNA, promoter and amino acid sequences and phylogenetic analysis .... 17
2.8. Promoter analysis .... 17
2.9. Semi-Quantitative Reverse Transcription-PCR (RT-PCR) .. 17
3. RESULTS ..... 18
3.1. DNA and amino acid sequence analysis ... 18
3.2. Protein and chlorophyll levels during different stages of leaf growth and senescence...... 19
3.3. Expression profiles of PAPs/fibrillins genes during different stages of leaf growth and senescence. ..... 19
3.4. Expression profiles of putative transcription factors for PAP-fibrillin genes during different stages of leaf growth and senescence... 20
3.5. Expression of PAP-fibrillin genes in response to hormonal and abiotic stress signals . 21
4. DISCUSSION ..... 21
5. CONCLUSION ...... 26
LIST OF TABLES AND FIGURES ..... 28
Table 1. Hormone and abiotic treatments for Arabidopsis thaliana ... 29
Table 2. Primers used for studying expression levels of 14 PAP-fibrillin genes by RT-PCR...... 30
Table 3. Primers used for studying expression levels of 40 putative transcription factors
having corresponding binding sites on promoters of PAP-fibrillin genes by RT-PCR. 31
Table 4. PAP-fibrillin family genes in Arabidopsis. ... 33
Table 5. Putative transcription factors for PAP-fibrillin family genes ... 34
Table 6. Summary of PAP/fibrillin genes expression during early stage of leaf development
and/or onset of leaf senescence .... 36
Table 7. Summary of PAP/fibrillin genes expression patterms modulated by hormone and
abiotic stress treatments .... 37
Figure 1. Structures of 14 PAP-fibrillin genes in Arabidopsis thaliana. .. 38
Figure 2. Conserved motifs in PAP-fibrillin family proteins in Arabidopsis. .. 39
Figure 3. Amino acid sequence alignment of 14 Arabidopsis PAPs/fibrillin proteins. .. 40
Figure 4. Phylogenetic analysis of Arabidopsis PAP-fibrillin proteins. ... 41
Figure 5. Phylogenetic analysis of the PAP-fibrillin family proteins in different plants .. 42
Figure 6. Different stages of rosette leaf growth and senescence. .. 43
Figure 7. Total protein contents in different stages of rosette leaf growth and senescence. . 44
Figure 8. Total chlorophyll contents in different stages of rosette leaf growth and
senescence. ..... 45
Figure 9. Expression patterns of PAP-fibrillin genes detected by RT-PCR. . 46
The expression patterns are divided into 4 categories during different stages of leaf growth and senescence ...... 46
Figure 10. Expression patterns of 40 transcription factors (TFs) detected by RT-PCR. These TFs having putative binding sites on the promoters of PAP-fibrillin genes. .. 51
Figure 11. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of IAA (100 M) treatments. (b) Total RNA electrophoresis (1g/lane); (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR .... 52
Figure 12. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of BA (100M) treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 53
Figure 13. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of GA (50M) treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes by RT-PCR. ..... 54
Figure 14. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of SA (100M) treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 55
Figure 15. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of MeJA (100M) treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR.... 56
Figure 16. (a) The visible phenotype of non-treated (C) and treated plants after 0-9 days of ABA (100M) treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes by RT- PCR. .... 57
Figure 17. (a) The visible phenotype of non-treated and treated plants after 0-9 days of 10M Propionyl brassinolide (BR) treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. .. 58
Figure 18. (a) The visible phenotype of non-treated and treated plants after 0-7 days of 100mM H2O2 treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 59
Figure 19. (a) The visible phenotype of non-treated and treated plants after 0-14 days of 150mM NaCl treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 60
Figure 20. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of 150mM Sorbitol treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR.... 61
Figure 21. (a) The visible phenotype of non-treated (C) and treated plants after 0-7 days of Hypoxia treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 62
Figure 22. (a) The visible phenotype of non-treated (C) and treated plants after 0-10 days of drought treatment; (b) Total RNA electrophoresis 1g/lane; (c) Pot’s weigh after dehydration for five-repeat (column) and mean (line); (d) Expression patterns of PAP-fibrillin genes by RT-PCR. ..... 63
Figure 23. (a) The visible phenotype of non-treated (C) and treated plants after 0-14 days of dark treatment ; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 64
Figure 24. (a) The visible phenotype of non-treated and treated plants after 0-14 days of high light treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 65
Figure 25. (a) The visible phenotype of non-treated (C) and treated plants after 0-14 days of cold treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 66
Figure 26. (a) The visible phenotype of non-treated (C) and treated plants after 0-3 days of heat treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 67
Figure 27. (a) The visible phenotype of non-treated (C) and treated plants after 0-10 days of wounding treatment; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 68
Figure 28. (a) The visible phenotype of non-treated and treated plants after 0-24 hours of UV-B treatment/0-24h; (b) Total RNA electrophoresis 1g/lane; (c) Expression patterns of PAP-fibrillin genes detected by RT-PCR. ... 69
REFERENCES ..... 70
APPENDIXES ...... 81
Appendix 1. Expression profiles in 14-PAP-fibrillin genes for triplicate .. 82
Appendix 2. Expression profile for 40-putative PAP-fibrillin transcription factors for triplicate ...... 83
                                    

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