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研究生：	黃麗敏 Huang, Li-Min
論文名稱：	阿拉伯芥GDSL型酯解酶/脂肪酶之全基因分析與功能特性研究 Genome-wide analysis and functional characterization of GDSL-type esterases/lipases in Arabidopsis
指導教授：	蕭介夫 Shaw, Jei-Fu 陳榮芳 Chen, Long-Fang O.
學位類別：	博士 Doctor
系所名稱：	生物科學與科技學院 - 生物科技與產業科學系 Department of Biotechnology and Bioindustry Sciences
論文出版年：	2017
畢業學年度：	106
語文別：	英文
論文頁數：	161
中文關鍵詞：	阿拉伯芥、酯解酶、脂肪酶、逆境、葡萄糖、軟腐病
外文關鍵詞：	Arabidopsis, GDSL, lipase, esterase, stress
相關次數：	點閱：113 下載：6
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GDSL型酯解酶/脂肪酶為一群具有多功能特性的水解酵素，廣泛存在微生物與植物之中，在醫藥、食品及生物功能上提供很重要的應用價值，本研究從模式植物阿拉伯芥中定義出105個 GDSL型酯解酶/脂肪酶基因（AtGELP1-105），結合演化、基因表現及蛋白結構分析進行推測此基因家族可能具有之功能特性，並利用T-DNA插入剔除或過量表現特定AtGELP基因進行生理功能的探討，從SALK種源庫獲得199株可能插入AtGELP基因的突變株，研究結果確認出123株分別插入71 個AtGELP基因，其中88株確認分別為54個AtGELP基因剔除轉殖株。其中AtGELP72具有提升植物對高濃度葡萄糖的耐受性，且AtGELP72 經調控脂肪酸分解路徑（β-oxidation pathway）的相關因子導致種子中的脂肪酸含量及組成改變。此外，AtGELP65參與細菌性軟腐菌Pectobacterium carotovorum subsp. carotovorum的抗性反應，研究結果顯示AtGELP65經調控茉莉酸/乙烯路徑相關因子ERF1與PDF1.2的表現，提升阿拉伯芥植苗抵抗細菌性軟腐病菌的感染。本論文研究結果說明了阿拉伯芥GDSL型酯解酶/脂肪酶基因家族具非常多樣性的生理功能，包含參與高葡萄糖或高鹽逆境的反應，以及參與抗微生物病菌感染之各種生理功能。

In this study, 105 GDSL-type esterase/lipase (AtGELP) genes were identified in Arabidopsis thaliana by conducting a comprehensive computational analysis. The phylogenetic analysis, combined with expression profiling and protein motif architectures, were used to predict the roles of AtGELP genes. To further investigate the physical roles of the AtGELP gene family, 88 AtGELP T-DNA knockout lines for 54 AtGELP genes from 199 putative SALK T-DNA mutants were successfully screened out. AtGELP72 has the function on tolerance to high glucose concentrations. Furthermore, it increased the expression of genes involved in fatty acid metabolism during seed germination and seedling development, such as COMATOSE (CTS), LACS6 and LACS7, PED1 and PNC1 which function in β-oxidation pathway controlling fatty acid degradation. AtGELP65 has the function on resistant to necrotropic bacterial pathogen, Pectobacterium carotovorum subsp. carotovorum (Pcc). AtGELP65 expression in leaves were induced by Pcc and jasmonic acid (JA), ethylene (ET) and salicylic acid (SA), known as crucial roles in pathogen response. AtGELP65 regulated the expression of ERF1 and PDF1.2, JA/ET-dependent signals, to make Arabidopsis plants more resistant to Pcc pathogen. Overall, the results suggest that the AtGELP genes have diverse physiological functions, such as affecting the early growth of seedlings and germination rate subjected to high concentrations of glucose and salt, or biotic stress responses.

Chinese Abstract (中文摘要)	......................	I
Abstract	...............................	II
Acknowledgements	......................	VI
Table of Contents	..........................	VII
Contents of Tables	.....................	X
Contents of Figures	.......................	XI
Abbreviations List	.........................	XIII
Chapter 1 Research Background	....................	1
1-1	GDSL-type esterases/lipases	..............	1
1-2	GDSL-type esterases/lipases in plants	....	2
1-3	GDSL-type esterases/lipases in Arabidopsis.....	3
1-4	Research Objetives	.......	4
Chapter 2 Materials and Methods	.........	6
2-1	Plant materials and growth conditions	.......	6
2-2	Bioinformatic analysis	............	8
2-3	Expression analysis of Arabidopsis GDSL-type esterase/lipase genes	...........	9
2-4	Phylogenetic analysis of GDSL-type esterases/lipases and motif search	.......	10
2-5	Isolation of genomic DNA and PCR	.......	10
2-6	Southern blot	........................	11 
2-7	RT-PCR	..................................	12
2-8	Plant transformation and GUS staining	.......	12
2-9 	Quantitative real-time PCR and Statistical analysis	........................................ 	15
2-10	Sequence and motif analysis of ATGELP72	.......	16
2-11 	Construction and transformation of AtGELP72 in a yeast expression system ............................	17
2-12	Specific activity of lipase or esterase activity plate assays and protein purification	..............	17
2-13 	Antibody preparation	.......	18
2-14 	Hormones treatment	.........	19
Chapter 3 Genome-wild analysis of GDSL-type esterases/lipases in Arabidopsis	.....	21
3-1	Introduction	...........	21
3-2	Results	..................	23
3-3	Discussion	.......................	33
3-4	Summary	.........................	36
Chapter 4 A novel GDSL-type esterase, AtGELP72, involved in fatty acid degradation and glucose tolerance	.......	38
4-1	Introduction	........	38
4-2	Results	......................	42
4-3	Discussion	..........	52
4-4	Summary	...............	55
Chapter 5 AtGELP65 (GLIP4) plays a critical role in resistant to Pectobacterium
carotovorum subsp. carotovorum	..........	56
5-1	Introduction	.........................	56
5-2	Results	.............................	58
5-3	Discussion	.......................	64
5-4	Summary	.......................	66
Chapter 6 Conclusions	.....................	68
References	................................	70
Tables	......................................	81
Figures	......................................	113
Related Paper Publications	...........	162
 
Contents of Tables
Table 1. Primers of AtGELPs for genomics PCR and RT-PCR	........................................	82
Table 2. Primers of AtGELPs for overexpression and promoter construction and
qRT-PCR	.............................................................................................................	88
Table 3. Annotations of Arabidopsis GDSL-type esterase/lipase family members from
TAIR database	..................................................................................................	90
Table 4. Characteristics of Arabidopsis GDSL-type esterase/lipase family members	......	94
Table 5. The putative gene structure of Arabidopsis GDSL-type esterase/lipase family
members	............................................................................................................... 	98
Table 6. 15 identified AtGELP family genes and 13 non-Arabidopsis GDSL 
orthologues or homologues	.................................................................................	103
Table 7. Characteristics of T-DNA insertion lines of AtGELP family members	...............	107
Table 8. Activity analysis of recombinant AtGELP72 enzymes from Pichia pastoris 
to the p-nitrophenyl butyrate in purified process	.................................................	110
Table 9. Characteristics of 24 plant GDSL esterases/lipases with known functions	.........	111  
Contents of Figures
Figure 1.	Example of Alignment Arabidopsis GDSL-type lipases/esterases with
AtGELP97	.......................................................................................................	114
Figure 2.	Chromosomal distribution and segmental duplication events for 
AtGELPs	..........................................................................................................	115
Figure 3.	The expression patterns for AtGELP genes	......................................................	117
Figure 4. 	An unrooted phylogenetic tree of AtGELPs and 13 non-Arabidopsis GDSL
esterase/lipase proteins	....................................................................................	119
Figure 5.	Analysis of the phylogenetic relationship and protein motif structure among
	the AtGELPs and plant homologues with known function	..............................	121
Figure 6. 	Identification of the number of T-DNA inserts for AtGELP T-DNA insetion
	mutants	.............................................................................................................	122
Figure 7. 	Analysis of AtGELP expression in AtGELP T-DNA insertion mutants	...........	125
Figure 8.	An unrooted phylogenetic tree and functional analysis of AtGELP genes
	and plant homologues with known function	....................................................	127
Figure 9.	Conditional phenotypic alterations of the AtGELP T-DNA knockout mutants
	in response to salt sress	....................................................................................	128
Figure 10.	Conditional phenotypic alterations of the AtGELP T-DNA knockout mutants
	in response to Pcc	.............................................................................................	129
Figure 11.	The genomic organization of ATGELP72 and structure for ATGELP72
	protein	..............................................................................................................	132
Figure 12.	Phylogenetic analysis of ATGELP72 protein sequence and 24 plant GDSL
 	esterases/ lipases	...............................................................................................	133
Figure 13.	Recombinant ATGELP72 proteins for lipase/esterase activity analysis	..........	134
Figure 14.	The enzyme activity assay of recombinant ATGELP72 proteins	.....................	136
Figure 15.	Protein motif structure and location of DELLA downregulated GDSL-type
	enzymes in Arabidopsis	....................................................................................	137
Figure 16.	The expression pattern of ATGELP72 in Arabidopsis	.....................................	138
Figure 17.	Ghenotype analysis of ATGELP72 transgenic plants	.......................................	140
Figure 18.	Phenotype analysis of ATGELP72 transgenic plants under salt stress	.............	142
Figure 19.	Phenotype analysis of ATGELP72 transgenic plants under osmotic stress	......	145
Figure 20.	The relative transcription levels of HXK1 in imbibed seeds and 
	post-germinated seedlings	................................................................................	148
Figure 21.	The relative transcription levels of fatty acid metabolic factors in imbibed
	and post-germinated seedlings	.........................................................................	149
Figure 22.	The complementary DNA (cDNA) sequence of AtGELP65 (GLIP4)	.............	153
Figure 23.	Multiple alignment of GLIP1-7 in Arabidopsis	................................................	154
Figure 24.	An unrooted phylogenetic tree of Arabidopsis GLIPs	.....................................	155
Figure 25.	Expression pattern of ATGELP65 in Arabidopsis	............................................	156
Figure 26.	Response of AtGELP65 transgenic plants to Pcc inoculation	..........................	157
Figure 27.	Semi-quantitative RT-PCR analysis of ATGELP65 expression after Pcc
	inoculation	.......................................................................................................	159
Figure 28.	Analysis of ATGELP65 gene expression in response to defense hormone	......	160
Figure 29.	Analysis of defense genes expression in transgenic Arabidopsis plants	..........	161

                                    

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