臺灣博碩士論文加值系統

本論文以香蕉品種北蕉 (Musa spp., cv. ‘Pei Chiao’, Cavendish，AAA group) 雄花序之果手原體為試驗之初始材料，進行香蕉癒傷組織誘導方式之改進及農桿菌媒介法之轉殖，並以萎縮病毒反義複製酶基因之 Raja 品種轉殖香蕉進行香蕉萎縮病毒之抗病接種試驗。
於香蕉癒傷組織之誘導上，以 picloram、dicamba、TDZ、以及 flurprimidol 四種不同生長調節劑與洋菜或水晶洋菜做為凝膠劑的相互組合試驗，結果以培養基含有 picloram 之組合能誘導多數的金黃色癒傷組織，而培養基含有 TDZ 之組合則不會產生金黃色的癒傷組織，但於 1 mg/l 之 TDZ 濃度下，能顯著降低培植體的褐化率。不同濃度之 picloram 以 1 mg/l 誘導癒傷組織的效果最佳；而以 picloram、2,4-D、IAA 與 NAA 三種植物生長素之組合，則以 picloram 與 2,4-D 之組合對癒傷組織誘導之評估最佳。
經測試影響農桿菌媒介法對香蕉果手原體進行基因轉殖效率之各因子，包括使用之藥劑及菌液濃度、所須的共培養時期，及後續的篩選時期等，結果顯示較佳條件為將果手原體先進行為期十二天的前培養，之後以農桿菌品系 LBA4404，於其濃度為 OD600 = 1.0 時，在 200 µM 之乙醯丁香酮存在之情況下，進行為期二天的共培養，並於共培養處理後第十四天開始以 50 mg/l 之 G418 進行為期一個月之篩選，培植體於篩選處理後移至不含任何抗生素之培養基進行培養。而以超音波震盪對果手原體進行創傷處理，並配合農桿菌進行轉殖，雖然轉殖率均以超音波處理二秒為最高，但與未超音波處理之對照組相較，則超音波之創傷處理為非必要的轉殖處理。
於抗病接種試驗方面，乃使用轉殖香蕉萎縮病毒反義複製酶基因之 Raja 品種香蕉進行病毒接種，於受試之二十一個轉殖系中，已確定有六株轉殖系對香蕉萎縮病毒基因型Ⅰ具有抗性，而對香蕉萎縮病毒基因型Ⅰ不具抗性之轉殖系，對病毒的反應可分為快速發病及延遲發病，而延遲發病者之病徵則遲至隔年春季才表現。轉殖系於接種香蕉萎縮病毒基因型Ⅱ之後第三個月進行病徵觀察，只有二株轉殖系具較對照組明顯之病徵。進一步將轉殖系進行香蕉萎縮病毒不同基型因之複合感染試驗，受試之原抗萎縮病毒基因型Ⅰ之三株轉殖系植株，於接種基因型Ⅱ之後，有二株出現萎縮病徵，經聚合酶連鎖反應偵測受試轉殖株體內之病毒，呈現正反應，顯示其對萎縮病毒為耐病性；而另一株則完全沒有病徵表現，以聚合酶連鎖反應偵測病毒於受試轉殖株體內之含量，亦呈現負反應，顯示該植株對萎縮病毒具有完全的抗性，故轉殖香蕉萎縮病毒反義複製酶基因於香蕉為可行之抗病方式。

The development of a regeneration system from banana (Musa spp. cv. ‘Pei Chiao’, Cavendish, AAA group) hand primordia on male flowers is described. The most important factors affecting embryogenic callus initiation were the source of explant and the composition of the culture medium. The closer the hand primordia is situated to the inflorescence apical meristem, the stronger is the regenerative capability. Improved callus initiation was obtained on culture medium supplemented with 1 mg/l picloram and 4 mg/l 2,4-D. Somatic embryogenesis was observed on callus cultures subcultured consecutively to a culture medium containing 1 mg/l picloram, 4 mg/l 2,4-D, 1 mg/l IAA, and 1 mg/l NAA. Somatic embryo germination and plantlet development was obtained using established protocols.
A protocol for the production of transgenic banana (Musa spp. cv. ‘Pei Chiao’, Cavendish, AAA group) was developed via Agrobacterium-mediated genetic transformation of hand primordia. Two disarmed Agrobacterium tumefaciens strains, A 281 and LBA 4404, both carrying the binary plasmid pBIRC1 with the nptⅡ gene were evaluated as vector systems. A number of parameters were tested with respect to maximizing transformation efficiency. While wounding was inhibitory, the pre-culture (12 days), acetosyringone treatment (200 μM), bacterial growth phase (optical density; OD600 = 1.0), co-cultivation period (2 days) had positive effects on transformation. Following co-cultivation, hand primordia were placed on multiplication medium with cefotaxime for 2 weeks then transfered onto the same medium and stressed with geneticin (50 mg/l) for 1 month. Further selection occurred in the medium at an elevated geneticin level (100 mg/l). A number of geneticin-resistant microadventitious buds were multiplied in vitro.
Banana plants transformed with the banana bunchy top virus (BBTV) antisense replicase gene were generated and twenty-one independently transformed plant lines were analyzed for resistance to BBTV. Three different responses were obtained. Some of the transgenic plants showed a pre-established, complete, and highly resistant phenotype since no viral symptoms were observed, and no virus detected. Some of the transgenic plants showed no viral symptoms before winter, but showed a delayed viral symptom in the next spring. The remaining plants from these plants showed a rapid disease symptom. Furthermore, the transformants resistant to BBTV genotypeⅠwere secondary challenged with BBTV genotypeⅡ. Transformants exhibited two different responses. One transgenic plant still showed complete, highly resistant phenotype since no viral symptoms were observed, and no virus detected. The other two transformants showed disease symptom after 2 months of inoculation.

內 容 目 次
中 文 摘 要
Abstract
前 言
前 人 研 究
一、 香蕉組織培養之再生系統
二、 香蕉萎縮病
(一)、 香蕉萎縮病防治上的限制因子
1、 交叉保護
2、 化學藥劑防治
3、 生物防治
(二)、 香蕉萎縮病毒
三、 轉殖植物的抗病機制
(一)、 Coat protein-mediated protection
(二)、 Satellite RNA-mediated resistance
(三)、 Replicase-mediated resistance
四、 基因默化 (gene silencing) 之應用
五、 過敏性反應 (Hypersensitive-response，HR)
材 料 與 方 法
一、 試驗材料
(一)、 植物材料
(二)、 聚合酶連鎖反應使用之引子對
二、 香蕉組織培養之再生系統
(一)、 生長調節劑及凝膠劑之處理
(二)、 TDZ處理
(三)、 Picloram 處理
(四)、 TDZ 與auxin之組合處理
(五)、 Picloram 與auxin 之組合處理
三、 影響農桿菌媒介法進行香蕉果手原體基因轉殖效率因子之分析
(一)、 農桿菌轉殖法
(二)、 旋轉處理對果手原體誘導不定芽之影響
(三)、 果手原體對 geneticin (G418) 之天然抗性
(四)、 農桿菌感染果手原體之最適濃度
(五)、 乙醯丁香酮 (Acetosyringone) 對轉殖效率之影響
(六)、 農桿菌品系對轉殖效率之影響
(七)、 共培養天數對果手原體轉殖效果之影響
(八)、 前培養日數 (preincubation) 對轉殖效率之影響
(九)、 不同共培養時期處理之培植體
四、 農桿菌配合超音波震盪轉殖法 (Sonication-assisted Agrobacterium - mediated transformation) 對香蕉果手原體轉殖效率之影響
五、 香蕉萎縮病毒反義複製酶基因之轉殖植株之抗病接種
(一)、 植物材料
(二)、 病毒接種源
(三)、 蚜蟲之獲毒
(四)、 病毒接種條件
(五)、 接種株之採樣部位
(六)、 香蕉萎縮病毒汁液之製備
(七)、 DAS-ELISA 步驟
(八)、 病毒核酸之製備
(九)、 聚合酶連鎖反應
(十)、 轉殖香蕉對萎縮病毒基因型Ⅰ之接種測試
(十一)、 轉殖香蕉對萎縮病毒基因型Ⅱ之接種測試
(十二)、 轉殖香蕉對萎縮病毒基因型Ⅰ及基因型Ⅱ之複合接種測試
(十三)、 轉殖香蕉內含香蕉條紋病毒之測試
結 果
一、 利用果手原體誘導癒傷組織之研究
(一)、 生長調節劑及凝膠劑對誘導癒傷組織發生率之影響
(二)、 TDZ濃度對誘導癒傷組織發生率的影響
(三)、 TDZ 與不同生長調節劑之相互組合對誘導癒傷組織之影響
(四)、 Picloram 濃度對誘導癒傷組織發生率的影響
(五)、 Picloram 與不同生長調節劑相互組合之培養基對誘導癒傷組織之影響
二、 農桿菌媒介法進行香蕉果手原體基因轉殖效率因子之分析
(一)、 旋轉處理對誘導不定芽之影響
(二)、 果手原體對 geneticin (G418) 之天然抗性
(三)、 農桿菌接種果手原體之最適濃度
(四)、 乙醯丁香酮 (acetosyringone) 對轉殖效率之影響
(五)、 農桿菌品系對轉殖效率之影響
(六)、 共培養天數對果手原體轉殖效果之影響
(七)、 前培養日數對轉殖效率之影響
(八)、 不同共培養時期處理於前培養之培植體
(九)、 農桿菌配合超音波震盪轉殖法
三、 香蕉萎縮病毒反義複製酶基因之轉殖株系之抗病接種
(一)、 轉殖香蕉對香蕉萎縮病毒基因型Ⅰ之接種測試
1、 病徵觀察
2、 ELISA 檢測
3、 PCR 檢測
(二)、 轉殖香蕉對香蕉萎縮病毒基因型Ⅱ之接種測試
1、 病徵觀察
2、 ELISA 檢測
(三)、 轉殖香蕉對香蕉萎縮病毒基因型Ⅰ及基因型Ⅱ之複合接種測試1、 病徵觀察
2、 ELISA 檢測
3、 PCR 檢測
(四)、 轉殖香蕉之香蕉條紋病毒潛伏檢測
1、 病徵觀察
2、 PCR 檢測
討 論
一、 香蕉組織培養之再生系統
二、 香蕉萎縮病毒反義複製酶基因之轉殖
三、 香蕉萎縮病毒反義複製酶基因之轉殖植株之抗病接種
參 考 文 獻
附錄一、 香蕉幼雄花序培養所使用之培養基配方
附錄二、 香蕉癒傷組織使用之 YC 培養基與不同生長調節劑組合之配方
附錄三、 香蕉轉殖參試之農桿菌品系及所含質體
圖一、 香蕉幼雄花序衍生之癒傷組織類型
圖二、 香蕉幼雄花序培養之間接體胚發生
圖三、 果手原體對 G418 之天然抗性
圖四、 農桿菌媒介法轉殖香蕉果手原體之流程圖
圖五、 擬轉殖植株之形態
圖六、 感染香蕉萎縮病毒基因型Ⅰ之轉殖植株其病徵表現比較
圖七、 接種香蕉萎縮病毒基因型Ⅰ之未轉殖對照植株 (A) 及轉殖植株 (B) 其葉片萎縮及葉脈透明化之病徵表現比較
圖八、 接種香蕉萎縮病毒基因型Ⅰ之未轉殖對照植株 (A) 及轉殖植株 (B) 其葉柄之深綠色條斑病徵表現比較
圖九、 接種香蕉萎縮病毒基因型Ⅰ之未轉殖對照植株 (A) 及轉殖植株 (B) 之葉片比較
圖十、 感染香蕉萎縮病毒基因型Ⅰ之轉殖植株其病毒系統之聚合酶連鎖反應之分子鑑別
圖十一、 經不同基因型香蕉萎縮病毒複合感染之轉殖植株表現型
圖十二、 感染香蕉萎縮病毒之轉殖植株其不同病毒基因型複合感染之聚合酶連鎖反應之分子鑑別。
圖十三、 轉殖植株其香蕉條紋病毒之聚合酶連鎖反應之分子鑑別。
表一、 生長素及凝膠劑對誘導癒傷組織發生率之影響
表二、 TDZ對香蕉癒傷組織發生率之影響
表三、 TDZ 與不同生長素組合之培養基對果手原體誘導癒傷組織之影響表四、 Picloram對誘導香蕉癒傷組織發生率之影響
表五、 Picloram 與不同生長素組合之培養基對果手原體誘導癒傷組織之影響
表六、 旋轉處理對香蕉果手原體誘導不定芽之影響
表七、 香蕉果手原體對 G418 之天然抗性
表八、 不同濃度之 G418 對培植體的篩選效率
表九、 農桿菌接種濃度對香蕉果手原體轉殖效率之影響
表十、 Acetosyringone 對香蕉轉殖處理不定芽發生之影響
表十一、 農桿菌品系對香蕉轉殖效率之影響
表十二、 農桿菌與香蕉果手原體之共培養天數對轉殖效率之影響
表十三、 果手原體之前培養日數對轉殖效率之影響
表十四、 不同共培養時期處理之培植體對轉殖效率之影響
表十五、 農桿菌轉殖於超音波處理之果手原體
表十六、 農桿菌與香蕉果手原體共同進行超音波處理之轉殖率
表十七、 轉殖植株接種香蕉萎縮病毒基因型Ⅰ之症狀一覽表
表十八、 轉殖植株接種香蕉萎縮病毒基因型Ⅰ之 ELISA 檢測一覽表
表十九、 轉殖植株接種香蕉萎縮病毒基因型Ⅰ之 PCR 一覽表
表二十、 轉殖植株接種香蕉萎縮病毒基因型Ⅱ之症狀一覽表
表二十一、 轉殖植株接種香蕉萎縮病毒基因型Ⅱ之 ELISA 檢測一覽表
表二十二、 不同基因型香蕉萎縮病毒複合感染之轉殖植株經聚合酶連鎖反應及 ELISA 檢測結果
表二十三、 偵測轉殖植株內含香蕉條紋病毒一覽表

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