臺灣博碩士論文加值系統

富亮胺酸重複類受體激酶 (LRR-RLKs) 是植物細胞膜上最大的一群RLKs，但大部分成員之功能尚未知。為了探討RLKs基因在熱逆境反應 (HSR) 的角色，本研究檢測九個阿拉伯芥之RLK的突變株在熱逆境 (HS) 下之性狀，並且進一步分析一個受熱誘導表現的RLK基因STRUBBELIG-RECEPTOR-FAMILY 6 (SRF6) 的功能。SRF6 (At1g53730) 編碼一個LRR-RLK，經由先前的分析發現SRF6可能參與在HSR當中。SRF6為一個座落在細胞膜上的蛋白質，利用β-glucuronidase (GUS) 染色分析，發現SRF6廣泛分佈在植物體各時期的組織中。本篇利用細菌表現SRF6的激酶區段並進行活性分析，結果未顯示出其磷酸化活性。兩個不同程度減低SRF6表現量的T-DNA突變株在後天耐熱性 (SAT) 的檢測結果顯示，其存活率高於阿拉伯芥野生型，而大量表現SRF6的轉殖株之存活率則較低。另外，本篇在熱反應的表型觀察中發現，SRF6可能在植物熱逆境下的根生長恢復扮演重要的功能，且srf6植物之內質網 (ER) 逆境反應基因bZIP28和bZIP60的表現量顯著增加。本研究指出一個在植物根組織中非典型熱反應路徑的存在，且與SRF6以及ER逆境反應相關。

Leucine-Rich Repeat Receptor-like kinases (LRR-RLKs) comprise the largest family of plant RLKs, but most of them are still functionally unknown. To explore the role of RLKs in heat stress response (HSR), we characterized 9 RLK-mutant lines of Arabidopsis under HS condition and focused on functional analysis of a heat-responsive gene: STRUBBELIG-RECEPTOR- FAMILY 6 (SRF6). SRF6 (At1g53730) encodes a putative LRR-RLK and was suggested to play a role in the HSR pathway. Subcellular localization and tissue-specific expression analysis demonstrated that SRF6 is a membrane-bound protein and ubiquitously expressed in plants. This study concluded that the cytoplasmic kinase domain (CKD) of SRF6 is catalytically inactive. The srf6 mutants is less sensitive than wild type when challenged with a short-term acquired thermotolerance (SAT) test. Besides, analysis of the thermo-responsive root elongation phenotypes in srf6 highlighted that SRF6 plays a role in the control of root growth recovery from the HS. Analysis of the endodermis reticulum (ER)-stress-responsive genes showed that the expression levels of bZIP28 and bZIP60 were higher in srf6. The preliminary study indicates that the existence of a non-canonical HSR pathway involving SRF6 and ER-stress response in root growth.

摘要………………………………………………….…………….…….....1
ABSTRACT…………………………………………...……………….…..2
LIST OF TABLES………………………………………………………….3
LIST OF FIGURES……………………………………………...….……...4
ABBREVIATIONS………………………………………………………...6
INTRODUCTION………………………………………………………….7
Heat stress/shock (HS)………………………………………………………………......8
Heat shock/stress response and heat-mediated regulatory pathways…........9
Receptor-like kinases (RLKs)…………………………………………….……………..13
STRUBBELIG-RECEPTOR FAMILY…………………...…………………………....17
Objectives…………………………………………………...…………………………....20
MATERIALS AND METHODS……………………………………….....22
Plants materials and growth conditions…………………..…………………………....22
DNA/RNA Preparation, cDNA Synthesis and qRT-PCR………………...…………...22
Construction of transgenic SRF6-overexpression lines generation………..…….…...23
Protoplast Preparation and Transient Expression Assay……………………....……..24
Subcellular Localization…………………………………………………………………25
GUS reporter activity staining and cleared tissue for observation……………….…..25
Thermotolerance test……………………………………………………….……………26
In vitro kinase assay……………………………………………………………..….…....27
Coomassie Brilliant Blue and Pro-Q diamond staining……………………………….27
Purification of membrane protein…………………………………………..……….….28
Immunoprecipitation and Western blotting…………………………………..…….....29
Ponceau S staining………………………………………………………………...……..29
Statistical analysis…………………………………………………………..……………30
RESULTS………………………………………………………...……….31
Expression of SRF6 is regulated by HSFA1s and HSFA7b in response to heat
shock…………………………………………….…………………………….…….31
Phenotyping of RLKs- mutant plants by quantitation of the hypocotyl and root
elongation in response to heat stress (HS)………………….…………………….32
SRF6 localized to plasma membrane……………………………...……………………34
Tissue-specific expression of SRF6…………………………………...…………………34
SRF6 encodes an atypical RLK………………………………………………................35
SRF6 is a negative regulator in short-term acquired thermotolerance……………....36
Expression of SRF6 in srf6-2 mutant was not responsive to heat shock………..……37
Overexpression of SRF6 restored the mutated phenotype of srf6……………………37
SRF6 does not play a role in hypocotyl elongation control in response to heat
shock…………………………………………………………………………….......38
SRF6 may play a role in root control in response to heat shock………………..…….38
Expression of the abiotic stress-responsive genes in srf6 mutants in response to heat
shock…………………………………………………………………………….…..39
Phosphorylation of SRF6 was undetectable in response to heat shock………………40
Expression of SRF7 was not responsive to heat shock but higher in SRF6-KO
mutants……………………………………………………………………………...41
DISCUSSION……………………………………………………...……..42
SRF6 is a downstream regulator of HSFA1s and HSFA7b in specific
thermoresponsive mechanism required for proper thermotolerance……….….42
The function of SRF6 in root elongation may relate to ER-stress regulators…….….44
SRF6 is an atypical RLK and may not be phosphorylated by HS……………………45
The expression of SRF7 is not heat-inducible but affected by SRF6 transcriptional
level………………………………………………………………………………….49
TABLES…………………………………………………………………..51
FIGURES………………………………………………...……………….54
SUPPLEMENTARY FIGURES…………………………………………..79
REFERENCES……………………………………………………………81
APPENDIX………………………………………………….……………88

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