木黴菌是環境中常見的生物防治劑，其作為生物防治劑的機制可能為分泌胞外蛋白質、次級代謝物及誘導植物抗病性。其中胞外蛋白質L-amino acid oxidase及次級代謝物chrysophanol已被證明具有促使甘藍生長及誘導甘藍產生抗性的相關作用。Trichoderma harzianum ETS 323分泌於細胞外的L-amino acid oxidase（Th-LAAO）對L-phenylalanine amino acid的催化能力最好，Th-LAAO具有抗細菌、拮抗植物病原真菌以及誘導甘藍產生抗性之相關活性；但是甘藍是否有相關表面受器與Th-LAAO結合導致此抗病機制及訊息傳遞目前我們仍不清楚。本研究利用親合管柱層析，Th-LAAO能與甘藍葉膜蛋白質Chlorophyll a/b binding protein結合。探討甘藍抗病下cab表現，將Th-LAAO貼附於甘藍葉表面後以灰黴菌感染，能增加cab表現比控制組增加約2.5倍；將cab轉殖於大腸桿菌表達之CAB（His6-CAB）貼附於甘藍葉表面，再以灰黴菌感染，cab表現比控制組下降約7倍；將His6-CAB與Th-LAAO以莫耳數比1:1混和置於甘藍葉面，隨後以灰黴菌感染，His6-CAB能拮抗Th-LAAO誘導甘藍抗性，使甘藍葉發病；經此拮抗作用後甘藍葉內之cab表現量則比控制組下降約10倍。上述各處理的甘藍葉發病面積比率與葉內cab表現量成反比，Th-LAAO處理發病面積為0-15%，His6-CAB處理發病面積為50-100 %，His6-CAB與Th-LAAO結合處理發病面積為70-100 %之間。再將Th-LAAO-Cy5.5處理在甘藍根部上，Th-LAAO-Cy5.5會往葉子輸送並進入葉綠體。本研究證明甘藍葉內葉綠體之膜蛋白質Chlorophyll a/b binding protein為T. harzianum ETS 323 L-amino acid oxidase結合標的，啟動甘藍抗灰黴菌能力。

T. harzianum ETS 323 is a well-known environmental biological control agent. Its mechanisms mainly contributes to production of extracellular proteins, secondary metabolites and resistance induction. Among these, L-amino acid oxidase and chrysophanol have been proven antibiotic activities and resistance induction of Brassica oleracea. The further interest would be to investigating binding proteins of the B. oleracea to ThLAAO. By using ThLAAO affiliated affinity chromatography, a 28.3 kDa ThLAAO binding membrane protein of B. oleracea was isolated and identified as Chlorophyll a/b binding protein. In the present of the ThLAAO, later infected with Botrytis cinerea, cab expression of B. oleracea leaf increase 2.5 folds, the cab of B. oleracea leaf that treated with His6-CAB later infection B. cinerea decreasing 7 folds. A mole ratio 1:1 of ThLAAO and His6-CAB mixture was applied to the B. oleracea leaf later infected with B. cinerea showing that His6-CAB can antagonize the ThLAAO resistance induction capability corresponding to decreasing cab expression 10 folds. The percentage of infection area of B. oleracea leaf were ThLAAO is 0 to 15 ％ His6-CAB is 50 to 100 ％ and ThLAAO mixture with His6-CAB is 70 to 100 ％accordingly decrease cab expression, respectively. ThLAAO-Cy5.5 treated B. oleracea root showing that ThLAAO-Cy5.5 enter root and transport to the leaves where have been proven in chloroplast in other investigation. This study showed that B. oleracea chlorophyll a/b binding protein is the binding protein of T. harzianum ETS 323 L-amino acid oxidase that induces pathogen resistance.

致謝 i
中文摘要 ii
目錄 vi
1 前言 1
2 介紹 3
2.1 真菌的介紹 3
2.2 真菌的分類 3
2.3 真菌的應用 3
2.4 木黴菌的介紹 5
2.4.1 木黴菌的分類與特性 5
2.4.2 木黴菌的生物防治機制 6
2.4.3 木黴菌的次級代謝物 6
2.5 L-amino acid oxidase（LAAO） 6
2.5.1 木黴菌胞外蛋白質—左旋胺基酸氧化酶（ThLAAO） 7
2.6 Light-harvesting Chlorophyll a/b binding protein（CAB） 8
2.7 植物抗性 9
3 研究動機與目的 11
4 材料與方法 13
4.1 化學藥品 13
4.2 菌種、植物 13
4.3 菌種培養 13
4.4 ThLAAO製備 14
4.5 純化L-amino acid oxidase（ThLAAO） 14
4.6 聚丙烯硫胺膠體電泳 Sodium dodecylsulfate Polyacrylamid Gel Electrophoresis（SDS-PAGE） 15
4.7 抽取甘藍葉子膜蛋白 15
4.8 ThLAAO與甘藍膜蛋白作用分離 16
4.9 In gel digestion 16
4.10 LC-MS MS 17
4.11 甘藍葉 Totoal RNA 萃取 18
4.12 將mRNA反轉錄成cDNA 18
4.13 Real-time PCR即時聚合酶鏈鎖反應 19
4.13.1 QRT-PCR primer設計 19
4.14 建構重組甘藍 Chlorophyll a/b binding protein質體DNA於大腸桿菌BL-21表現載體 19
4.14.1 將Chlorophyll a/b binding protein質體DNA表現於Escherichia coli菌株DH5α 19
4.14.2 將質體移至pET28a載體系統上形成標記-His6 tag重組蛋白質 21
4.14.3 IPTG誘導大腸桿菌BL21-CAB表現His6-Chlorophyll a/b binding protein, His6-CAB 22
4.15 純化His6-CAB 23
4.16 ThLAAO和His6-CAB對甘藍抗性之實驗 23
4.17 ThLAAO標記Cy5.5 23
4.19 ThLAAO與His6-CAB interaction via pull down assay 24
5 實驗結果 27
5.1 菌種生長觀察 27
5.2 T. harzianum ETS 323與deactivated R. solani的菌絲共培養 27
5.3 蛋白質ThLAAO純化結果 27
5.4 甘藍葉膜蛋白質抽取結果 28
5.5 ThLAAO標記Cy5.5 處理甘藍根 28
5.6 大量表現重組蛋白質His6-CAB於大腸桿菌 29
5.7 ThLAAO與His6-CAB 交互作用結果 29
5.8 His6-CAB拮抗ThLAAO對甘藍葉抗病誘導 30
5.9 探討甘藍抗病下ThLAAO誘導甘藍2天後感染灰黴菌，cab mRNA q-PCR結果 30
6 討論 33
7 結論 39
8 參考文獻 41
9 圖表結果 49
10 附錄 59

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