本論文目的在證明木黴菌ETS-323分泌之ThLAAO會與葉肉細胞之Lhcb1結合，進而激活植物防禦機制以誘導抗性對抗病原菌感染。我們使用Bimolecular Fluorescence Complementation (BiFC)技術可觀察到在阿拉伯芥葉肉細胞表現之LAAO-NYFP會在葉綠體內與Lhcb1-CYFP結合。我們使用Fluorescence Resonance Energy Transfer (FRET)技術可觀察到在阿拉伯芥葉肉細胞表現之LAAO-GFP會在葉綠體內與Lhcb1-OFP結合。由此可知當ThLAAO表現在葉肉細胞時會進入葉綠體而與類囊體上之Lhcb1結合。我們接著用Agrobacterium-mediated transformation建構ETS-323之ThLAAO基因缺失株(deletion mutants)。我們共得到8株轉形株，並命名為ETS-△LAAO1及ETS-△LAAO3－ETS-△LAAO8，其中#2轉形株之ThLAAO基因並沒缺失，故命名為ETS-2。我們發現ETS-△LAAO1－ETS-△LAAO8培養在培養基時會導致其無法產生孢子，故推測ThLAAO可幫助木黴菌在缺氧環境的土壤中產生孢子。此外、ETS-△LAAO1刺激植物生長的效果與ETS-323相似而不受ThLAAO基因缺失影響，顯見ThLAAO與木黴菌刺激植物生長的能力無關。在這ThLAAO基因缺失株中ETS-△LAAO1拮抗病原真菌Rhizoctonia solani之能力顯著降低。我們將ETS-323與ETS-△LAAO1孢子注入土壤以感染阿拉伯芥根部，然後在其葉子接種Botrytis cinerea，觀察到ETS-△LAAO1誘導阿拉伯芥抗性能力下降，推測ThLAAO可能為Trichoderma誘導植物抗性的一個重要分子。此外、我們發現當將LAAO基因轉殖入阿拉伯芥Col-0 (35S:PR1b-LAAO)時則抗病能力增強，但是若將LAAO基因轉殖入阿拉伯芥Lhcb1及Lhcb2的反義股突變株(antisense mutants) Col-0:asLhcb2-12時罹病程度與Col-0及Col-0:asLhcb2-12相似，顯示ThLAAO是經由與Lhcb1結合而誘導抗性。總而言之、木黴菌ETS-323分泌之ThLAAO會進入葉肉細胞之葉綠體內而與Lhcb1結合，進而激活植物防禦機制以誘導抗性對抗病原菌感染。

The purpose of this paper is to prove that ThLAAO secreted by Trichoderma ETS-323 will bind to Lhcb1 in mesophyll cells, thereby activating the plant defense mechanism to induce resistance against pathogen infection. Using Bimolecular Fluorescence Complementation (BiFC) technology, we observed that LAAO-NYFP expressed in Arabidopsis mesophyll cells would bind to Lhcb1-CYFP in chloroplasts. We then used Fluorescence Resonance Energy Transfer (FRET) technology and observed that LAAO-GFP expressed in Arabidopsis mesophyll cells would enter into chloroplasts and bind to Lhcb1-OFP on thylakoids. Therefore, ThLAAO expressed in mesophyll cells will enter thylakoids to bind to Lhcb1. We then constructed the ThLAAO gene deletion mutants of ETS-323 by Agrobacterium-mediated transformation and obtained 8 transformants. Among these transformants, the ThLAAO gene of the #2 transformant was not deleted. We named these transformants ETS-△LAAO1, ETS-2 and ETS-△LAAO3－ETS-△LAAO8. We found that ETS-△LAAO1－ETS-△LAAO8 could not produce spores when cultured in plates, so it was speculated that ThLAAO could help Trichoderma to produce spores in hypoxic soil environment. In addition, the effect of ETS-△LAAO1on stimulating plant growth was similar to that of ETS-323 and was not affected by the deletion of ThLAAO gene. Therefore, ThLAAO obviously had no effect on the ability of Trichoderma to stimulate plant growth. Among the ThLAAO deletion mutants, the ability of ETS-△LAAO1 to antagonize the pathogenic fungus Rhizoctonia solani was significantly reduced. We then spread the spores of ETS-323 and ETS-△LAAO1 into the soil to infect the roots of Arabidopsis seedling and then inoculated Botrytis cinerea on the leaves. We observed that the ability of ETS-△LAAO1 to induce resistance of Arabidopsis thaliana is decreased comparing to that of ETS-323. In addition, we found that when the LAAO gene was transferred into Arabidopsis Col-0 (35S:PR1b-LAAO), it enhances the disease resistance aginst the infection of Botrytis cinerea. However, if the LAAO gene was transferred into the antisense mutants (Col-0:asLhcb2-12) of Arabidopsis, the degree of disease is similar to that of Col-0 and Col-0:asLhcb2-12, indicating that ThLAAO induces resistance through binding to Lhcb1. Overall, ThLAAO secreted by Trichoderma ETS-323 will enter the chloroplasts of mesophyll cells and bind to Lhcb1, thereby activating plant defense mechanisms to induce resistance against pathogen infection.

中文摘要：I
英文摘要：III
目錄：V
一、前言(背景及重要性簡介(Background and significance)) 1
1.1 木黴菌 (Trichoderma spp.) 1
1.1.1簡介木黴菌 1
1.1.2 木黴菌的生物防治機制 1
1.1.4 Trichoderma spp.產生誘導植物抗性的蛋白質及化合物 3
1.2 T. harzianum L-amino acid oxidase (ThLAAO) 3
1.3 植物抗病 4
1.4 chlorophyll a/b/ binding protein (Lhcb) 5
二、實驗室以往的ThLAAO研究結果 7
三、目的 9
四、實驗流程 11
五、材料與實驗方法： 13
5.1 真菌材料 13
5.2 植物材料 13
5.3農桿菌品系 13
5.4大腸桿菌品系 13
5.4 使用的引子與載體 13
5.5 真菌培養 13
5.6 煙草及阿拉伯芥種植方式 14
5.6質體DNA抽取 15
5.7建構BiFC所需要的農桿菌載體 15
5.7.1 PCR擴增出LAAO、Lhcb1、Epl-1、NYFP、CYFP DNA 15
5.7.2 Overlapping PCR 16
5.7.3 建構質體pGEM-LAAO-NYFP、pGEM- Epl-1-NYFP、pGEM- Lhcb1-CYFP 17
5.7.4 TA cloning 17
5.7.5以電穿孔(electroporation)方式進行大腸桿菌之轉形作用 17
5.7.6 PCR screening確認轉型株 18
5.7.7 質體DNA抽取、限制酶SpeⅠ及BstEⅠⅠ切割、回收DNA及ligation製備pCAMBIA-LAAO-NYFP、pCAMBIA-Lhcb1-CYFP及pCAMBIA-Epl1-NYFP 18
5.7.8 電穿孔到DH5α、PCR screening、抽質體、酵素試切確認 19
5.7.9 抽質體、電穿孔到LBA4404 19
5.8 建構FRET所需要的農桿菌載體 19
5.9 以BiFC及FRET檢測ThLAAO與Lhcb1結合 21
5.10 共軛焦顯微鏡觀察 21
5.11建構木徴菌ThLAAO基因缺失突變株 22
5.11.1木黴菌genomic DNA抽取 22
5.11.2 DNA 聚合酶連鎖反應檢測確認5’UTR及3’UTR引子最適合溫度 22
5.11.3 選殖ThLAAO之5’UTR及3’UTR於pGEM T-Easy載體 23
5.11.4 選殖ThLAAO之5’UTR及3’UTR於載體pPK2 24
5.11.5農桿菌法轉形木黴菌 25
5.11.6進行mitotic stability測試及single spore isolation 26
5.12 檢測Parafilm封PDA plates對ETS-323與 ETS-ΔLAAO1－8產生孢子的影響 26
5.13 ETS-ΔLAAO缺失株對立枯絲核菌拮抗作用試驗 27
5.14 檢測ETS-ΔLAAO1缺失株對阿拉伯芥生長的影響 27
5.15 檢測LAAO及Lhcb1對誘導植物抗病能力 27
5.15.1 檢測木黴菌ETS-ΔLAAO1誘導阿拉伯芥Col-0野生株抗病能力 27
5.15.2 檢測阿拉伯芥Col-0:asLhcb2-12 (35S:PR1b-LAAO)轉型株抗病能力 28
5.16 統計分析 28
六、結果 29
6.1 探討ThLAAO是否會與葉綠體之Lhcb1結合 29
6.2 建構Trichoderma ThLAAO基因缺失突變株 31
6.2.1檢視 ETS-△LAAO抑制病原菌生長能力是否減弱 31
6.2.2 探討ThLAAO基因缺失對Trichoderma孢子形成的影響 31
6.2.3 探討ThLAAO缺失是否對Trichoderma刺激植物生長有影響 32
6.3 探討ThLAAO是否為Trichoderma 誘導抗性的一個重要分子 32
6.4 探討Lhcb1的存在是否為ThLAAO誘導抗性的必要條件 33
七、討論 35
7.1 ThLAAO會在葉綠體內與Lhcb1結合 35
7.2 ThLAAO可協助木黴菌在缺氧氣環境(hypoxia)之土壤中產生孢子 36
7.3 ThLAAO為木黴菌ETS-323誘導抗性的一個重要分子 37
7.4 Lhcb1為ThLAAO誘導阿拉伯芥抗性的必要分子 39
八、總結 41
九、參考文獻 43
十、圖表 51

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