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研究生(外文):Huan-Yu Lin
論文名稱:阿拉伯芥 AtMAPR3 參與對抗灰黴病菌感染之功能性研究
論文名稱(外文):Functional studies of Arabidopsis AtMAPR3 in the defense against B. cinerea
指導教授(外文):Chien-Chih Yang
口試委員(外文):Chiu-Ping ChengMei-Chun ChengRita P.-Y. ChenYee-yung Charng
外文關鍵詞:Membrane-associated progesterone receptorBotrytis cinereareactive oxygen speciesplant defensesalicylic acid
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膜聯合孕酮受質蛋白 (MAPRs) 是一種廣泛存在於真核生物的血基質結合蛋白質,存在於動物、植物及真菌中。AtMAPR 是 MAPR 家族在阿拉伯芥中的同源蛋白,目前已知的功能有參與去除內質網壓力 (ER stress) 與參與木質素的生合成等等。本研究探討 AtMAPR3 在植物防禦上可能扮演的角色。根據微陣列資料庫的資料顯示,AtMAPR3 的表現量會被植物病原菌灰黴病菌、丁香假單胞菌等病原菌感染所誘導。其中在灰黴病菌感染後, AtMAPR3 表現量提升最高。為了瞭解 AtMAPR3 是否參與對抗灰色葡萄孢菌的防禦,本研究利用兩個 AtMAPR3 功能缺失的突變株 (msbp2-1 和 msbp2-2) 進行研究。 msbp2 突變株中被灰黴病菌感染後,感染面積比野生型植株大 1.2 倍。然而在感染的突變株植株中,病菌的生長情況與野生型植株無明顯相異。此結果暗示 msbp2 突變株的初期防禦機制受損,但是當植物防禦啟動後,突變株能夠限制感染真菌的生長。分析感染後不同時間產生的活性氧化物質 (ROS),發現在感染 48 小時後,msbp2 突變株中的 ROS 累積量比野生型多,這和呼吸爆發氧化酶 D (RBOHD) 的表現量增加相關。此外,突變株中水楊酸 (SA) 反應基因表現量也比較高。綜合上述,缺乏 AtMAPR3 似乎影響病原菌感染的早期感應階段。AtMAPR3 可能在辨識病原菌上或是早期防禦訊息傳導上扮演了某個角色,所以後續引發的防禦才會被延遲。
Membrane-associated progesterone receptors (MAPRs) are widely found in eukaryotes, including animals, plants, and fungi. The functions of MAPR in Arabidopsis (AtMAPR, also named as MSBPs) are diversified, ranging from participating in the release of ER stress to lignin biosynthesis. Here, the possible role of AtMAPR3 in plant defense is studied. Based on microarray databases, the expression of AtMAPR3 can be induced by the infection of phytopathogen Botrytis cinerea (B. cinerea), Pseudomonas syringae (P. syringae), etc.; among these factors, B. cinerea has been shown to generate the highest induction levels of AtMAPR3 expression. To investigate if AtMAPR3 is involved in plant defense against B. cinerea, two mutants defected in AtMAPR3, namely msbp2-1 and msbp2-2, were employed in this study. The lesion areas of msbp2 infected by B. cinerea were larger by 1.2-fold than that of WT. However, the fungal growth of B. cinerea was similar in the mutants and WT. This suggested that the defense mechanisms in the early stage of infection were impaired in msbp2 mutants. However, the mutants were able to limit the growth of the infected fungi after the activation of late plant defense mechanisms. The ROS accumulation triggered by the infection was detected at different time points post-inoculation of the pathogen. It was interesting to note that ROS accumulated more in msbp2 mutants after 48 hours of infection, as revealed by DAB staining. This enhanced accumulation of ROS in msbp2 mutants correlated with the enhanced expression of the respiratory burst oxidase homolog D (RBOHD) in msbp2 mutants.
Furthermore, the expression levels of salicylic acid (SA) responsive genes were also higher in mutants. Taken together, the lack of AtMAPR3 appears to affect the early stage responses for the pathogen infection. AtMAPR3 might likely play roles in pathogen sensing or early signaling, which could have resulted in the delayed triggering of the defense system in the msbp2 mutants.
Table of Contents 1
Abbreviation list 5
中文摘要 7
Abstract 8
Chapter 1 Introduction 10
1.1. Plant pathogen classification and plant defense responses 10
1.2. The role of phytohormones in plant defense against B. cinerea 11
1.2.1. Jasmonates (JA) and ethylene (ET) in plant defense 11
1.2.2. Salicylic acid (SA) in plant defense 13
1.3. The role of reactive oxygen species (ROS) in plant defense 14
1.3.1. RBOHs and their role in the defense 15
1.3.2. PRXs and their role in the defense 16
1.3.3. The ROS scavenging systems and their role in the defense 17
1.3.4. The function of ROS in plant defense 18
1.4. Recent Studies on MAPR in Arabidopsis 19
1.5. Aims of the research 20
Chapter 2 Materials and Methods 22
2.1. Plant materials 22
2.1.1 Arabidopsis wild type (WT) 22
2.1.2 msbp2 mutants (from ABRC stock) 22
2.2 Chemicals 22
2.3 Instrument 22
2.4 Methods 23
2.4.1. Extraction of genomic DNA from Arabidopsis leaves 23
2.4.2. Agarose gel electrophoresis 23
2.4.3. Extraction of total RNA from Arabidopsis 24
2.4.4. DNase treatment 25
2.4.5. Reverse transcription 26
2.4.6. Identification of T-DNA insertion mutant 26
2.4.7. Quantitative PCR, qPCR 27
2.4.8. Fungal culture and disease assays 27
2.4.9. Trypan blue staining of Arabidopsis leaves 28
2.4.10. DAB staining of Arabidopsis leaves 28
Chapter 3 Results 29
3.1. The AtMAPR3/MSBP2 transcript level was induced by infection of B. cinerea 29
3.2. Isolation of atmapr3/msbp2 mutants 30
3.3. Mutation of AtMAPR3/MSBP2 enhanced susceptibility to B. cinerea without affecting fungal growth 31
3.4. Mutation of AtMAPR3/MSBP2 altered area of necrotic cell death and ROS accumulation in infected leaves 33
3.5. The expression of ROS generating enzymes was altered in msbp2 mutants 34
3.6. Autophagy machinery in msbp2 mutants seemed to be disrupted 35
3.7. SA-dependent defenses might promote necrotic cell death in msbp2 mutants 35
Chapter 4 Discussion 37
4.1. AtMAPR3/MSBP2 may be involved in pathogen-induced ROS accumulation 37
4.2. AtMAPR3/MSBP2 may be involved in the early stage of plant response to B. cinerea infection 39
4.3. AtMAPR3/MSBP2 may play a role in necrotic cell death under biotic stress based on SA signaling and ROS accumulation 41
4.4. Conclusion 42
Chapter 6 References 44
Figures and Tables 52
Fig 1. Expression analysis of AtMAPR3/MSBP2 in WT and msbp2 mutants after infection with B. cinerea. 52
Fig 2. Genotyping and AtMAPR3/MSBP2 expression analysis of the WT and msbp2 mutants. 54
Fig 3. The susceptibility to B. cinerea of the msbp2 mutants increased. 56
Fig 4. B. cinerea growth was quantified in WT and msbp2 mutants. 57
Fig 5. Trypan blue staining was used to detect cell death in WT and msbp2 mutants after different time points of infection. 58
Fig 6. Accumulation of H2O2 in the leaves of WT and msbp2 mutants infected with B. cinerea 60
Fig 7. The expression of ROS generating enzyme after infection in WT and msbp2 mutants. 62
Fig 8. The expression of the autophagy-related gene after infection in WT and msbp2 mutants. 64
Fig 9. Expression of the responsive genes for SA-dependent defense after infection in WT and msbp2 mutants. 66
Fig 10. The schematic diagram of the role of AtMAPR3/MSBP2 in plant defense 67
Table 1. Primer pairs used for the qRT-PCR analysis 68
Table 2. Primer pairs for other usages 70
Appendix 71
論文口試問答集及討論建議 88
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