臺灣博碩士論文加值系統

將轉殖葉表生菌及葉綠體技術應用於蘇力菌抗蟲基因表現與自發型轉錄終止，是本研究的重點。第一章概論蘇力菌抗蟲基因的應用。分析基因種類與轉基因作物種植比例的年相關性、總結抗蟲策略，並回顧文獻。第二~五章分別舉實例說明抗蟲基因在微生物與植物體內的應用效果，以及開發新基因之必要性。第二章實驗說明，將蘇力菌殺蟲晶體蛋白基因cry1Aa1以電穿孔法送入常見於植物葉表且無害的草生軟腐菌Erwinia herbicola中，毒害小菜蛾可達最高83.3%的死亡率，效期約2週，死亡率隨效期下降，符合用藥安全，除抗蟲效果，該菌分泌代謝物亦可有效抑制蔬菜黑腐與軟腐病害微生物之生長。第二章附錄說明本案已行專利申請。持續開發蘇力菌基因種類，掌握本土資源，是第三章之主旨。重點針對目前商品化蘇力菌製劑無法有效防治的夜蛾科Spodoptera屬害蟲，說明此類蟲害在全球的發生狀況，並發掘對其具毒害效果的新型cry1C基因。利用快速的複合式聚合酶連鎖反應配合生物檢定，尋得殺蟲致死率達36.7%的新型cry1C基因。第四、五章以實驗說明蘇力菌基因在植物細胞核與葉綠體之表現。轉入細胞核的基因是人工合成的cry1Ac基因，轉入葉綠體的是未經修飾的cry1C基因，且不含協助形成晶體的基因端。在細胞核的表現因單、雙子葉植物系統的差異，使抗蟲基因呈弱勢態，仍對番茄夜蛾幼蟲具有抑制生長的效果。在葉綠體表現強勢的cry1C基因，可產生約1%的殺蟲蛋白，對斜紋夜盜蛾幼蟲具有近100%的致死率，子代中的Cry1C殺蟲蛋白可達4%，100%的殺蟲成效由3天縮短為2天。表現cry1C基因的轉錄單元中，使用了E. coli thra終止子，為了解該型終止子是否可具自發型的轉錄終止反應，第六章實驗設計了由aadA與cry1C基因組成的dicistron，將終止子放置基因間，利用poly-T長短可控制終止效率的特性，觀察生體內轉錄終止的發生。結果顯示，自發型的轉錄終止仍可參與葉綠體基因表現的調控，並可能是由細胞核傳送至葉綠體的RNA聚合酶所使用的轉錄終止訊號之一。反轉對稱序列看似不能在葉綠體內終止轉錄，利用電腦分析20種綠色植物的葉綠體全基因序列，同時加上「反轉對稱序列形成莖環結構」與「poly-T在RNA聚合酶內形成不穩定的DNA-RNA hybrid」的兩項空間配位因素，發現葉綠體基因後方仍存有自發型轉錄終止子。測試水稻葉綠體rps18基因下游的rpsWT終止子，發現其轉錄終止的發生與RNA的建造基質，即rNTP的濃度有關，類似自發型的E. coli thra終止子在生體外的反應。葉綠體內轉錄終止子將可能依賴莖環結構形勢、其下游形成DNA-RNA hynrid的穩定度、供應rNTP的濃度等三因素決定其是否終止轉錄。

Transgenic approaches applying for pest control were accomplished by expressing Bacillus thuringiensis (Bt) endotoxins in epiphytic bacteria or plant chloroplasts. Intrinsic transcription termination for gene expression in chloroplast was also demonstrated hereby. In chapter 1, we summaried distribution of Bt gene types, the correlation between the annually increasing number of gene and the production and commercialization of transgenic Bt plants. A complete summary for transgenic strategies and the literature review were also included. Experiments in chapter 2~5 carried out the strategies regarding to Bt genes in epiphytic bacteria, in plant nuclei or in chloroplasts for their insecticidal activities. In chapter 2, the epiphytic Erwinia herbicola has been successfully transformed with a Bt toxin gene cry1Aa1 presented in plasmid pUN4. One of the transformed E. herbicola strains, Eh4, was sprayed on cabbage leaves for colonization to test its insecticidal efficacy and persistency against the diamondback moth (DBM). The mortality rates of DBM were 36.7~60% and 40~83.3% after feeding for 48 and 72 hours, respectively. A decrease of insecticidal efficacy with a concomitant increase of time interval after spraying E. herbicola was observed. The antagonistic effects of E. herbicola showed that there was a significant suppression against Xanthomonas campestris pv. campestris and Erwinia carotovora subsp. carotovora. Results in chapter 2 were used to apply for an official patent in 2002. In chapter 3, eleven in 807 Bt isolates in Taiwan were identified for their cry1 genotypes, cry1C gene content, and the related insecticidal activities. Multiplex polymerase chain reaction (mPCR) genotyping and assays for insecticidal activity or pest growth inhibition helped to explore a novel type of cry1C gene from E05-20a strain causing a mortality rate of 36.7% against the tobacco cutworm (Spodoptera litura, TCW). In chapter 4, two Bt genes, a synthetic cry1A(c) and a PCR-amplified cry1C, containing no C-terminal domain region responsible for crystallization, were delivered into nucleus and chloroplast of the tobacco plant cells, respectively. Three transgenic lines, Nt-Ac/C-1, -2, -3, were assayed for the coexistence of cry1A(c) and cry1C genes in plant cell and showed the insecticidal activity of 100% against the TCW, and of 33% against the Helicoverpa armigera. Although the mortality rate caused by the synthetic cry1A(c) gene’s product was not significant among larvae of H. armigera, their weight increments (10.6~51.7%) were much less than those of the controlled larvae (up to 255%). It indicated that expressing cry1A(c) gene in co-transformed tobacco provides a protection effect. Chloroplast is a better location for over-expression of Bt endotoxin. We proved it in chapter 5. The cry1C gene constructed under the control of rbcL promoter and the E. coli thra terminator in a plasmid vector as shown in chapter 4 was delivered into the chloroplast alone. High expression level of the integrated cry1C gene in chloroplast was observed through Northern and Western hybridization assays (~1% in soluble proteins). The transplastomic tobacco plants are highly toxic to TCW, causing a mortality rate of 76.9% to 100% after 72 hours of feeding test. T1 progeny of the transplastomic line IC-5 produced about 4% of the Cry1C in total soluble proteins. This very toxic tobacco plant to TCW resulted in a mortality of 100% within two days. Since the plastid transgenes are not transmitted through pollen, highly expressed Cry1C in chloroplasts facilitated not only the improvement in breeding for insect-resistant plants, but also the prevention of contamination of transgenes among crop plants. In chapter 6, we focused on the problem of which the E. coli thra terminator is able to be or not to be intrinsic in chloroplast transcription in vivo. A dicistronic gene expression unit was introduced in the assay. Wild-type thra terminator (T9WT) standing between aadA and cry1C interrupted 64.3% of the dicistronic transcription, implying that intrinsic termination still play its role in the regulation of plastid gene expression. Terminator with a shorter poly-T tract (T4) had no termination function in vitro but displayed a termination efficiency of 18.1% in vivo. It indicated that a nuclear-encoded polymerase (NEP) may response to the intrinsic termination. Despite some inverted repeats (IR) did not work for transcription termination in plastid, a survey of all published plastid genomes was developed by the GeSTer algorithm to look for intrinsic terminator-like structures in the 3’-untranslated region (3’-UTR) of plastid genes. A candidate (rpsWT) was found at 3’-UTR of the rice rps18 gene in the form of an easy-to-form stem-loop structure immediately followed by an unstable DNA-RNA hybrid such as the poly-T tract. The rpsWT is composed of a GC-rich IR and an immediately followed poly-T tract (TTCTTTTTT). Its IR showed a concensus motif (TCCCGGAGN6CTCCGGG) among some higher plants. Termination of the rpsWT is determined by the concentration of rNTP, similar to that we observed previously in transcription termination of the E. coli thra or leu terminators in vitro. Results in chapter 6 prompted us to propose a model for intrinsic termination in chloroplast, which is the combination of the factor 1 (a stable stem-loop structure), factor 2 (an unstable DNA-RNA hybrid appears in time after the stem-loop starts to form), and factor 3 (an interferenced RNA chain synthesis at certain position on DNA template due to factor 1, 2, plus the insufficient concentration of the complement rNTP).

中文總摘要 ix
英文總摘要 xi
第一章：蘇力菌毒素基因轉殖與作物蟲害防治總論(文獻回顧) 1
緒言 2
基因種類之開發與蟲害之防治 3
Bt抗蟲基因之微生物製劑與作物蟲害防治 5
Bt抗蟲基因轉殖作物與蟲害防治 8
抗蟲轉殖作物對環境安全之衝擊 11
結語 13
引用文獻 13
圖1-1 Bt毒素基因種類的分布及基因數目的增加與轉殖植物推廣
時間的相關性 22
圖1-2 目前Bt毒素基因轉殖之抗蟲策略 24
第二章：含蘇力菌毒素基因cry1Aa1之轉形葉表生菌Erwinia herbicola
的殺蟲效力與拮抗植物病原菌之分析 27
中文摘要 28
英文摘要 29
緒言 31
材料與方法 32
菌種、基因與質體來源 32
生菌製劑之製備與施用 32
殺蟲效力之生物檢定 33
抑制植物病原菌生長之測試 33
結果 33
葉表上的E. herbicola轉形菌株之殺蟲效力 33
E. herbicola可抑制植物病原菌之生長 34
討論 34
引用文獻 37
表2-1 噴灑於甘藍葉面之基因轉形與未轉形的Erwinia herbicola
對小菜蛾之殺蟲效力 39
圖2-1 含蘇力菌毒素基因cry1Aa1之質體pUN4 40
圖2-2 甘藍葉面經噴灑轉形或未轉形之葉表生菌後受到小菜蛾危
害的情形 41
圖2-3 轉形與未轉形之葉表生菌Erwinia herbicola分別對斑點病
、黑腐病與軟腐病病原菌之拮抗效果 43
第二章附錄：申請專利說明書 44
第三章：利用基因型鑑定策略發現有效殺蟲活性之台灣本土新型
蘇力菌cry1C基因(短訊) 51
中文摘要 52
英文摘要 58
引用文獻 60
表3-1 不同來源下含有cry1C基因之台灣本土蘇力菌分離株的
cry1基因組成與其對小菜蛾的殺蟲活性 62
表3-2 含有cry1C基因之台灣本土蘇力菌分離株對斜紋夜盜幼蟲
之殺蟲活性 63
圖3-1 利用複合式聚合酶連鎖反應進行cry1基因型之鑑定與cry1C
基因之確認 64
圖3-2 含Cry1C蛋白之各菌株培養液對斜紋夜盜蛾幼蟲生長的抑制
效應 66
第四章：合成型蘇力菌毒素基因cry1A(c)與野生型基因cry1C之共同
轉殖及其在菸草中殺蟲效力之表現差異 69
中文摘要 70
英文摘要 71
緒言 73
材料與方法 75
轉殖載體來源與修飾 75
基因來源與轉殖質體構築 75
粒子轟擊法與共同轉殖菸草之再生 77
微載體(microcarrier)之製備與DNA包覆(coating) 77
粒子轟擊法 77
共同轉殖菸草之再生 77
共同轉殖菸草之鑑定 77
植物總量DNA之抽取 77
PCR檢定 78
南方墨漬法(Southern blotting) 78
殺蟲活性之生物檢定法 78
結果 79
人工合成cry1A(c)基因之修飾 79
cry1C基因之分離與類型鑑定 80
共同轉殖菸草之鑑定 80
cry1A(c)與cry1C基因分別造成之殺蟲效果 81
討論 83
引用文獻 86
表4-1 含雙蘇力菌毒素基因cry1A(c)與cry1C之共同轉殖菸草
對斜紋夜盜蛾幼蟲之殺蟲效力 92
表4-2 含雙蘇力菌毒素基因cry1A(c)與cry1C之共同轉殖菸草
對番茄夜蛾(玉米穗蟲)幼蟲之殺蟲效力 93
圖4-1 轉殖載體之修飾與質體pCAM-Ac、pN-IC101之構築 94
圖4-2 人工合成的cry1A(c)基因與野生型基因密碼序列之比較 97
圖4-3 經定序之cry1C基因密碼與對應之胺基酸序列 100
圖4-4 共同轉殖菸草之轉基因鑑定 102
第五章：蘇力菌野生型毒素基因cry1C在菸草葉綠體中之表現對
斜紋夜盜蛾幼蟲具高效殺蟲力 105
中文摘要 106
英文摘要 107
緒言 109
材料與方法 111
基因與質體構築 111
葉綠體轉殖與植物再生 112
核酸墨漬法 113
蛋白質免疫偵檢與定量 114
昆蟲生物檢定 115
結果 115
葉綠體表現載體構築與轉殖 115
含蘇力菌基因之轉葉綠體植物選拔與鑑定 116
蘇力菌基因在葉綠體內之表現 117
殺蟲活性檢定 118
子代發芽試驗與其殺蟲活性檢定 119
討論 120
蘇力菌基因與細菌終止子可在葉綠體內發揮功能 120
插入葉綠體基因組的蘇力菌基因可表現高量蛋白質 120
葉綠體內表現蘇力菌毒素可達到高劑量抗蟲策略與防止基因流佈 121
引用文獻 123
Table 5-1 Insecticidal efficacy of the cry1C gene-transformed
transplastomic tobaccos against the larvae of tobacco
cutworm (Spodoptera litura) 127
Fig. 5-1 Construction of plasmid expression vectors 128
Fig. 5-2 DNA gel blotting assay of transplastomic tobaccos 130
Fig. 5-3 RNA and immuno blotting assays for cry1C gene
expression in transplastomic tobaccos 132
Fig. 5-4 Insecticidal activity assay of transplastomic tobaccos
and T1 seed germination test against antibiotic 134
第六章：自發型終止子可在高等植物葉綠體內終止轉錄(初稿) 137
英文摘要 138
緒言 140
材料與方法 142
質體構築 142
葉綠體轉殖與植物再生 143
PCR與南方墨漬法 144
北方墨漬法與定量分析 144
葉綠體基因組中自發轉錄終止子之電腦預測 145
推測終止子之選殖 145
生體外轉錄 146
結果 146
E. coli自發轉錄終止子可在葉綠體中終止轉錄 146
如何預測葉綠體中的轉錄終止子? 148
rpsWT的結構與功能相當接近自發終止子 149
連續的poly-T可加強rpsWT的轉錄終止功能 150
討論 151
葉綠體mRNA 3’端形成中仍存在的調節因素 151
葉綠體內自發轉錄終止子的結構限制 153
葉綠體轉錄的建造基質可能參與指揮自發轉錄終止 154
引用文獻 155
Table 6-1 Quantity of the plastid genes and the palindromes
predicted by the GeSTer program 161
Fig. 6-1 Identification of transgenic plants and in vivo termination
by E. coli thra terminators in tobacco chloroplasts 162
Fig. 6-2 Compilation of plastid rps18, rpl20, and their intergenic
regions (3’-UTR), derived from phylogenic relationships
of Viridiplantae 164
Fig. 6-3 A factor-independent terminator-like IR found in the 3’-UTR
of the rice plastid rps18 gene and its relatives from other
green plants 166
Fig. 6-4 Termination analyses of the rpsWT and rpsMT in vitro 168
Fig. 6-5 A proposed model for IR-dependent mRNA maturation
in chloroplasts 170
總結與前瞻 171

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