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| DC 欄位 | 值 | 語言 |
|---|---|---|
| dc.contributor.advisor | 劉瑞芬(Ruey-Fen Liou) | |
| dc.contributor.author | Ya-Yuan Yang | en |
| dc.contributor.author | 楊雅媛 | zh_TW |
| dc.date.accessioned | 2021-06-07T17:41:09Z | - |
| dc.date.copyright | 2020-07-29 | |
| dc.date.issued | 2020 | |
| dc.date.submitted | 2020-07-16 | |
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| dc.identifier.uri | http://tdr.lib.ntu.edu.tw/jspui/handle/123456789/15491 | - |
| dc.description.abstract | OPEL為Phytophthora parasitica所分泌的質外體效應蛋白 (apoplastic effector)，序列分析顯示其具有signal peptide, thaumatin-like domain, glycine-rich domain及glycosyl hydrolase (GH) 16 domain，後者包含β-1,3-glucanase的保守性序列。以OPEL重組蛋白處理Nicotiana tabacum cv. Samsun-NN可引發明顯之壞疽斑 (necrosis)、癒傷葡聚醣 (callose) 沉積、活性氧分子 (reactive oxygen species, ROS) 累積及誘導防禦相關基因表現。此外，OPEL引發植物免疫反應的關鍵構造為GH16 domain，將其預測的酵素活性位點進行單點突變後，即大幅降低OPEL激發植物防禦反應的能力，因此OPEL很可能藉由分解植物細胞壁產生DAMP而引發免疫反應。為探討這個可能性，本研究製備OPEL重組蛋白及其酵素活性區雙點突變蛋白 (OPEL-dm)，用以處理菸草後，收集質外體液 (分別稱為AF-OPEL及AF-OPEL-dm)，防禦反應分析結果顯示AF-OPEL可引發菸草細胞壞疽、誘導防禦相關基因表現及增加對P. parasitica的抗性，但AF-OPEL-dm僅在菸草葉片引發微弱黃化。以95 oC加熱15分鐘後，AF-OPEL仍具有引發菸草細胞壞疽的活性；除此之外，AF-OPEL也較AF-OPEL-dm含有較多的還原糖。為了進一步找出AF-OPEL內引發植物免疫反應的活性物質，先後以正己烷及乙酸乙酯萃取，將AF-OPEL的成分分成低極性、中極性以及高極性等三個部份，續以the luminol-based chemiluminescence分析方法檢測各極性層萃取物引發活性氧分子累積的活性。結果顯示AF-OPEL的低極性層 (AF-OPEL/H)、中極性層 (AF-OPEL/EA) 及高極性層 (AF-OPEL-A) 萃取物均具有誘導活性氧分子累積的能力，但AF-OPEL/EA展現的活性明顯高於其他二者。進一步以高效液相層析法分析，發現三個AF-OPEL/EA特有的波峰，分別稱為peak I, peak II與peak III，其中僅peak II具有誘導活性氧分子累積的能力。這些結果顯示OPEL引發植物免疫反應之機制很可能是藉由辨識DAMPs而不是PAMPs且與其酵素活性密切相關。未來研究目標將著重於找出peak II所代表之物質，以瞭解其對於OPEL及植物間的交互作用。 | zh_TW |
| dc.description.abstract | OPEL is an elicitor protein identified in Phytophthora parasitica, which contains signal peptide in the N-terminus, followed by a thaumatin-like domain, a glycine rich domain and a GH16 domain. GH16 domain is characterized by the presence of a conserved active site for beta-1,3-glucanase. It has been shown previously that infiltration of OPEL recombinant protein into leaves of Nicotiana tabacum cv. Samsun-NN resulted in necrosis, callose deposition, ROS production as well as the induction of defense gene expression. Moreover, domain analysis indicated the elicitor activity of OPEL depends solely on the GH16 domain. Interestingly, the elicitor activity is compromised with OPEL recombinant protein harboring a single-point mutation in the predicted active site. It suggests an essential role of the enzymatic activity of OPEL in inducing plant defense responses, which might involve the generation of damage/danger-associated molecular pattern in the apoplast. To investigate this possibility, we prepared OPEL recombinant protein and OPEL mutant (OPEL-dm) with mutations at two key residues in the putative active site of the GH16 domain, infiltrated them individually into N. tabacum leaves, and then collected apoplastic fluid from the treated leaves, named AF-OPEL and AF-OPEL-dm, respectively. When infiltrated into tobacco leaves, AF-OPEL caused necrosis as well as induced defense gene expression and resistance against P. parasitica. In contrast, the activity of AF-OPEL-dm was significantly compromised. After heat treatment at 95 oC for 15 mins, AF-OPEL still retained most of its necrosis-inducing activity. Moreover, AF-OPEL contained more reducing sugars than AF-OPEL-dm. To identify the PTI-inducing substances, AF-OPEL was extracted by using n-hexane (H) and then ethyl acetate (EA) as the solvent, resulting in the collection of three fractions: AF-OPEL/H (low polar), AF-OPEL/EA (medium polar), and AF-OPEL/A (high polar). All three fractions induced ROS production as shown by the luminol-based chemiluminescence assay. However, the activity of AF-OPEL/EA was much higher than the other two.Further purification by high performance liquid chromatography (HPLC) identified three peaks (known as peak I, peak II, and peak III), which were found only in AF-OPEL/EA but not in the control experiment.Notably, only peak II induced ROS production. Based on these results, the mechanism for OPEL to induce PTI is through recognition of DAMP rather than PAMP and related to its putative site. In the future, we hope to identify the peak II-represented substance to understand the interaction between OPEL and plants. | en |
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| dc.description.tableofcontents | 致謝………………………………………………………………………………………i 中文摘要………………………………………………………………………………..iii 英文摘要………………………………………………………………………………...v 壹、 前言………………………………………………………………………..1 1. 植物防禦反應…………………………………………………………...1 1.1 MAMPs簡介…………………………………………………………….2 1.2 DAMPs簡介……………………………………………………………..4 1.3 PTI反應…………………………………………………………………....6 2. 細胞壁分解酵素在植物防禦反應扮演的角色……………………7 3. 疫病菌簡介……………………………………………………………….7 4. 植物對疫病菌之防禦反應…………………………………………….9 4.1 質外體效應蛋白………………………………………………………...9 4.2 細胞質效應蛋白………………………………………………………...11 5. 疫病菌外泌蛋白OPEL於植物防禦反應的角色………………….12 5.1 醣苷水解酶家族16 (GH16 family) 的特性……………………..12 5.2 OPEL同源性基因於植物免疫反應的角色………………………..13 6. 植物質外體液 (apoplastic fluid, AF) 的組成分……………….14 7. 研究動機…………………………………………………………………..14 貳、 材料與方法………………………………………………………………..15 1. 供試植株及菌種來源……………………………………………………15 1.1 植物材料及生長條件…………………………………………………...15 1.2 表現重組蛋白之大腸桿菌菌株來源及保存………………………..15 2. 以大腸桿菌表現OPEL重組蛋白……………………………………...15 2.1 OPEL重組蛋白表現及純化…………………………………………….15 2.2 凝膠過濾法 (Gel filtration)…………………………………………...16 3. 植物抗病反應分析………………………………………………………..16 3.1 農桿菌注入法 (Agroinfiltration)…………………………………....16 3.2 質外體液之收集……………………………………………………….......17 3.3 癒傷葡聚醣堆積分析 (callose deposition assay)……………...17 3.4 活性氧分子染色……………………………………………………….......18 3.5 以化學冷光法 (chemiluminescence) 檢測活性氧分子………..18 3.6 防禦反應相關基因表現分析…………………………………………....19 3.6.1 純化植物total RNA………………………………………………...........19 3.6.2 製備cDNA………………………………………………………................20 3.6.3 即時定量聚合酶連鎖反應分析…………………………………..........20 3.7 疫病菌接種…………………………………………………………….......20 4. 植物質外體液之分析…………………………………………………….21 4.1 以DNS法檢測還原糖含量……………………………………………...21 4.2 植物質外體液之萃取………………………………………………….....21 4.3 高效液相層析法 (High Performance Chromatography, HPLC)……….22 5. 統計分析方法………………………………………………………………...22 參、 結果……………………………………………………………………………….23 1. 自表現OPEL之Nicotiana tabacum cv. Samsun-NN葉片所收集之質外 體液可導致菸草葉片細胞死亡………………………………………….......23 2. OPEL全長度蛋白及突變蛋白之表現及純化………………………………23 3. 突變酵素活性區降低OPEL重組蛋白在菸草葉片引發細胞死亡的活性...24 4. 突變酵素活性區降低OPEL重組蛋白在菸草引發活性氧分子累積的活 性……………………………………………………………………………...24 5. 突變酵素活性區降低OPEL重組蛋白在菸草引發癒傷葡聚醣沉積的能 力……………………………………………………………………………...25 6. 自處理OPEL重組蛋白之菸草葉片所收集的質外體液可引發菸草葉片 死亡…………………………………………………………………………...25 7. 自處理OPEL重組蛋白之菸草葉片所收集之質外體液可誘導植物防禦 反應基因表現………………………………………………………………...26 8. 經處理OPEL所收集之質外體液可提升菸草對疫病菌之抗性…………...26 9. 自處理OPEL重組蛋白之菸草葉片所收集之質外體液含有較多還原糖...27 10. 經處理OPEL所收集之質外體液可引發活性氧分子累積………………...28 11. 經處理MES及OPEL後所收集之質外體液於中極性層具有成分差異….29 12. OPEL處理後的質外體液經高效液相層析分離後的物質具有誘導活性 氧分子累積的能力…………………………………………………………...29 13. 經處理MES及OPEL後所收集之質外體液於高極性層具有成分差異….30 肆、 討論……………………………………………………………………………….31 1. OPEL雙點突變蛋白仍保留部分elicitor活性……………………………...31 2. OPEL處理後的質外體液引發植物PTI反應………………………………32 3. OPEL引發植物PTI反應的物質具有耐熱性………………………………33 4. OPEL處理後的質外體液經萃取後於各極性層均存在引發活性氧分子 累積的活性…………………………………………………………………...34 5. OPEL如何引發植物防禦反應………………………………………………35 6. 結語…………………………………………………………………………...36 伍、引用文獻………………………………………………………………………….37 陸、附表……………………………………………………………………………….55 柒、附圖……………………………………………………………………………….56 | |
| dc.language.iso | zh-TW | |
| dc.title | 探討疫病菌質外體效應蛋白OPEL如何引發植物免疫反應 | zh_TW |
| dc.title | Investigation of how OPEL, an apoplastic effector from Phytophthora parasitica, elicits plant immune responses | en |
| dc.type | Thesis | |
| dc.date.schoolyear | 108-2 | |
| dc.description.degree | 碩士 | |
| dc.contributor.oralexamcommittee | 李宗徽(Tzong-Huei Lee),林乃君(Nai-Chun Lin),吳志航(Chih-Hang Wu) | |
| dc.subject.keyword | 細胞壁分解酵素,OPEL,植物免疫反應,疫病菌,醣苷水解酶, | zh_TW |
| dc.subject.keyword | cell wall degrading enzymes,OPEL,plant immunity,Phytophthora parasitica,glycosyl hydrolase, | en |
| dc.relation.page | 70 | |
| dc.identifier.doi | 10.6342/NTU202001537 | |
| dc.rights.note | 未授權 | |
| dc.date.accepted | 2020-07-16 | |
| dc.contributor.author-college | 生物資源暨農學院 | zh_TW |
| dc.contributor.author-dept | 植物病理與微生物學研究所 | zh_TW |
| 顯示於系所單位： | 植物病理與微生物學系 | |
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