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Development of marker-free transgenic plants with resistance to Tomato leaf curl Taiwan virus and Tomato spotted wilt virus
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由粉蝨傳播的雙生病毒與由薊馬所傳播的番茄萎凋病毒是目前極為重要的兩群植物病毒。利用基因工程技術藉由基因轉殖的方式為目前有效的抗病毒的策略之一。然而在轉殖的過程中，通常都會使用抗生素抗性基因或殺草劑抗性基因作為篩選用的標誌基因以區分其中少數具有轉基因的植物。但是站在生態與食品安全的角度考量，對於使用此類抗性基因的安全疑慮越來越受大眾矚目，因此本論文的主旨即是建構可穩定移除標誌基因的轉殖系統，並利用此系統生產可抗雙生病毒與番茄萎凋病毒的無篩選標誌基因之轉基因植物。本研究所分析的雙生病毒是主要危害台灣番茄生長的台灣番茄捲葉病毒，首先，為了得知基因片段中何者可提供轉基因植物對雙生病毒表現較高的抗性，因此將台灣番茄捲葉病毒全基因體分割為數個基因片段後，再分別轉殖至圓葉菸草中進行抗性分析，發現其中四個構築可提供轉基因植物較高的抗性分別為：IRC1(基因間區域與C1開放讀碼區5′ 端之序列)，C2 (部分C2開放讀碼區之序列)，C2C3 (C2與C3開放讀碼區重疊區域之序列) 以及Rep2 (C1開放讀碼區之3′ 端序列)；並藉偵測轉基因植物中所累積的siRNA確認其抗病機制是經由基因沉寂所誘發。除此之外，番茄斑萎病毒是屬於Tospovirus屬而且擁有最廣泛寄主範圍的植物病毒之一，故此研究利用其N基因片段與台灣番茄捲葉病毒的部分C2開放讀碼區連結在一起作為一個嵌合轉基因，再將此構築轉殖入圓葉菸草與番茄中，以期能發展具有多重抗性之轉基因植物。經抗性分析後發現轉基因植物以農桿菌接種台灣番茄捲葉病毒後不會產生病徵，而機械接種番茄斑萎病毒後亦呈現高度抗性的現象，進一步分析抗病的轉基因植物則可偵測到siRNA的累積，證明此抗性是經由基因沉寂的機制所提供。同時也證實藉由連結不同基因體組成的病毒基因片段的策略可提供轉基因植物對DNA與RNA病毒具有抗性。與此同時，本論文亦以共轉殖法發展出三種生產無篩選標誌基因轉基因植物的轉殖系統，包括 (1) pGANP-CP1/pBin19：單一菌株內兩個獨立的質體各自攜帶有目標基因或標誌基因的T-DNA；(2) pGA2T-CP1：同一質體中同時帶有目標基因及標誌基因的T-DNA；(3) 為了發展可適用更多植物種類的載體，尤其是對kanamycin敏感度較低的作物，因此構築了pGA2TNH：單載體攜帶兩組T-DNA而於其中一組T-DNA中帶有兩種不同的選擇性標誌基因。經此三種系統再生之轉基因菸草的共轉型效率皆很相近，約為於50%。至於兩組T-DNA於子代的分離現象亦符合預期，於單一標誌基因重複數的轉基因植物中，移除標誌基因的比例在雙載體系統為24.1%，單載體系統可達17.5%~18.6%。結果顯示這些系統確實可行且同樣都能有效率地移除選擇性標誌基因，進一步可提供簡便及實用性兼具的工具應用至生產無篩選標誌基因的轉基因植物。因此，本論文將先前研究結果所提及之可提供轉基因植物對台灣番茄捲葉病毒產生較高抗性的基因片段，包括IRC1、C2、C2C3、Rep2並與番茄斑萎病毒之部分N基因片段連結在一起後，構築至二位元載體pGA2TNH藉此生產具有病毒抗性的無篩選標誌基因轉基因菸草。並利用農桿菌接種法篩選出對台灣番茄捲葉病毒具有抗性的轉基因親本植物，且經由自交的方式於其子代中獲得無篩選標誌基因轉基因抗病菸草。總而言之，本論文的結果提供了可應用在田間防治雙生病毒與番萎凋病毒的轉基因策略。而所研發的生產無選擇性標誌基因轉基因植物的轉殖系統，也將有助於提升一般大眾對轉基因作物的接受度。|
Abstract Whitefly-transmitted geminiviruses (Geminiviridae) and thrips-borne tospoviruses (Bunyaviridae) are two groups of extremely important plant viruses. Current transgenic approach based on genetic engineering technology provides an efficient strategy to breed plants to resist viral infection. However, public concerns about the use of antibiotic- and herbicide-resistance genes for the selection of transgenic plants during the transformation process have increased tremendously. Therefore, the objectives of this study were to develop the reliable transformation system that could remove selectable markers while generating transgenic plants that would resist the infection of geminiviruses and tospoviruses. Firstly, Tomato leaf curl Taiwan virus (ToLCTWV), a predominant tomato-infecting geminivirus in Taiwan, was subjected to investigate which viral gene fragments can confer high resistance to geminiviruses in transgenic plants. Individual transgenic constructs covering the entire ToLCTWV genome was transformed into Nicotiana benthamiana plants. Four constructs including IRC1 (intergenic region flanked with 5' end of C1), C2 (partial C2 ORF), C2C3 (overlapping region of C2 and C3 ORFs) and Rep2 (3' end of the C1 ORF) of high resistance for ToLCTWV have been observed. The detection of siRNA in transgenic plants confirmed that the mechanism of resistance was via gene silencing. Moreover, the middle half of the N gene of Tomato spotted wilt virus (TSWV), which is the type member of Tospovirus, was fused with the partial C2 ORF of ToLCTWV as the chimeric transgene and transformed into N. benthamiana and tomato to develop transgenic plants with multiple viral resistance. The transgenic plants remained symptomless post agro infected with ToLCTWV and exhibited high resistance to TSWV. The detectable siRNAs demonstrated that the resistance was mediated by gene silencing mechanism. The results also explained that linking multiple gene fragments of two viruses with different genomic organization was an effective strategy to engineer plants against both DNA and RNA viruses. Meanwhile, we developed three strategies of co-transformation to generate marker-free transgenic plants; they were (1) pGANP-CP1/pBin19, which comprises two individual plasmids carrying T-DNA of the target and marker genes separately; (2) pGA2T-CP1, which consists of one plasmid carrying two T-DNAs for the target and marker genes; and (3) pGA2TNH, which contains two T-DNAs in one plasmid in which one T-DNA carries the bi-selectable marker which can be used for more plant species especially those with low sensitivity to kanamycin. The co-transformation frequencies of the R0 transgenic N. benthamiana plants for both selection marker and target gene were similar. The co-transformation frequencies of three vector systems were about 50%. Segregation of transgene and selectable marker gene was revealed in the progeny of some co-transformed lines. The highest production ratio of marker-free transgenic plants was 24.1% in two plasmids system, followed by 18.6% in one plasmid system and 17.5% in bi-selectable marker system. We demonstrated that these strategies were feasible and efficient to eliminate the marker genes, and can provide a practical and simple tool for generating marker-free transgenic plants. Therefore, previously mentioned gene fragments that confer high resistance to ToLCTWV including IRC1, C2, C2C3 and Rep2 were linked together and fused with the middle half of the N gene from TSWV to make a chimeric transgene to be constructed into the binary vector, pGA2TNH, for the generation of viral resistance marker-free transgenic N. benthamiana. The transgenic R0 plants resistant to ToLCTWV were obtained and the marker-free resistant progeny plants were segregated by self-pollination. Overall, the results showed in this study have important implications for field deployment of transgenic strategies to control geminivirus and tospovirus. Moreover, the plant transformation systems that can generate marker-free plants would certainly boost the public acceptance of transgenic crops.
|Appears in Collections:||植物病理學系|
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