臺灣博碩士論文加值系統

在注重基因轉殖作物於環境釋放帶來效益的同時，基因轉殖作物潛在的生態風險及可能帶來的環境問題也受到相當程度的關注，而基因轉殖株之遺傳穩定及性狀是為評估首要項目。本研究材料利用異戊烯轉移酶(isopentenyltransferase, ipt)之基因轉殖青花菜(Brassica oleracea var. italica)經過4年田間性狀評估，顯示出產量性狀受環境因子的影響大過於轉殖品系間差異，基因轉殖ipt青花菜植株性狀及產量優於對照組104(CK)，與商業雜交一代品種「綠王」相當，而將花球儲存在25±2℃，轉殖系花球黃化面積達50%天數相較於非基因轉殖青花菜能延長2-3天，對於在櫥架壽命(shelf-life)方面具有實際的應用價值。
第二階段環境生態安全評估中，基因流佈(gene flow)是基因轉殖作物生態風險評估中的一個重要指標，而花粉活性與雜交稔實率是影響作物異交潛力的主要因子。基因轉殖青花菜103品系T5代為父本，與5種近緣亞種作物(青花菜雜交一代品種-綠王及同為綠王T5代104(CK)(Brassica oleracea var. italica)、白花芥藍-富悅(B. oleracea var. alboglabra)、黃花芥藍-白格林(B. oleracea var. alboglabra)、花椰菜-麗雪(B. oleracea var. botrytis) 和不同種的小白菜(B. rapa var. chinensis)、小油菜(B. rapa))進行雜交，透過分析花粉活性與雜交率對青花菜異交潛力進行評估，提供基因轉殖青花菜商業栽培隔離距離與建立共存之參考。結果顯示在一般栽培條件下，基因轉殖青花菜與其近緣亞種作物花期同步天數在2007-2008兩年間分別為3-28天及8-44天。兩種染色法及離體萌芽法分析各品種(系)親本間花粉活性，除以螢光染色法(fluorescent diacetate, FDA)檢測其間差異未達顯著水準(P＞0.05)外，以氯化三苯基四氮唑染色法(2,3,5-triphenyltetrazolium chloride, TTC)及離體萌芽法(in vitro germination)花粉活性親本間有顯著差異(P＜0.05)；經由基因轉殖青花菜103品系與其5種近緣亞種作物之人工授粉、蜜蜂授粉及田間自然雜交的稔實率調查，種內變幅各在86.0-100%、1.1-42.5% 及0-9.0%間，而屬於種間雜交之小白菜及小油菜並無發現雜交現象，本研究顯示基因轉殖異戊烯轉移酶插入對青花菜品種異交潛力的改變甚微，但仍存在著基因流佈的可能性風險。進一步探討轉殖株雜交後代植株之競爭能力，於溫室利用同為自交系基因轉殖ipt青花菜-103轉殖系與104(CK)為花粉貢獻親，分別與4種近緣亞種作物進行人工雜交，整體而言，F1後代植株性狀以103轉殖系雜交F1後代株高較104雜交F1後代為矮，其多數性狀仍在104(CK)雜交F1後代、近緣亞種及104(CK)範圍以內，性狀改變甚微，且變異程度小於104(CK)雜交後代；103轉殖系雜交F1後代花粉活性雖未高於親本，但每果莢種子數顯著高於104(CK)雜交F1後代及雜交親本，顯示基因轉殖ipt青花菜雜交近緣亞種F1後代仍具有相當的雜交優勢。
探討基因轉殖與非基因轉殖青花菜對環境微生物群落之影響中，研究發現基因轉殖ipt青花菜之土壤pH為7.2

While focusing on the benefits of releasing genetically modified crops to environment, there have been considerable degrees of concern for potential ecological risks of transgenic crops and environmental issues. Genetic stabilities of traits on the transgenic plants are the main goals for assessment. Three selfed T5 lines of ipt transformed broccoli (lines 101, 102 and 103) have been screened by selections for single copy insertion, desirable horticultural traits and the activity of ipt transgene. These 3 transgenic inbred lines were evaluated in field during 2004-2007 to determine their growth, yield and shelf-life after harvest, compared with the non-transgenic inbred line (104-CK) and the original parental variety Green King. For most of the vegetative growth parameters observed. Variability measured from year-to-year was larger than that from line-to-line. Inbreeding had no significant impact morphologically and yield potential of the broccoli lines. Head productions of the transgenic inbred lines 102 and 103 were comparable to the parental variety Green King, but were significantly higher than the non-transgenic inbred line 104, as more lines 102 and 103 plants produced heavier flower heads. When flower heads were stored at 25±2℃, the period required to become 50% floret yellowing were 7.5 and 8.5 d for the transgenic lines 102 and 103 respectively, compared with 5.6 d for the non-transgenic line 104, and 5.1 d for the parental variety Green King. This study confirmed that the ipt-transformed inbred lines of broccoli exhibited acceptable appearance and yields with enhanced shelf-life.
Gene flow is an important index for the second phase of environmental and ecological safety assessment.. Pollens are potential carriers for genetically modified DNA sequence to other plants. For phanerogams, pollen viability and cross-compatibility are critical factors for successful outcross hybridization. In this study, ipt transformed broccoli designated as line 103 was used to investigate potential outcross by pollen viability and pollination with non-transgenic broccolis (line 104 and Green King) and other closely related varieties, such as Chinese kales (B. oleracea var. alboglabra) ‘Fu Yue’ (white flower) and ‘Bai Ge Lin’ (green flower), cauliflower (B. oleracea var. botrytis) ‘Li Syue’, ‘Pakchoi’ (B. rapa var. chinensis), and ‘Rapa’ (B. rapa). During field trial, flowering periods of transgenic ipt broccolis and non transgenic recipients were overlapping by 3-28 days in 2007 and 8-44 days in 2008. The pollen viability was determined by triphenyl tetrazolium chloride (TTC) analysis, fluorescein diacetate (FDA) and in vitro germination, respectively. Pollen viabilities by FDA showed no significant difference (p＞0.05) for transgenic line 103, non-transgenic and other tested cultivars . However, the differences of pollen viabilities were significant (p<0.05) using TTC and in vitro germination. Outcross rate was determined by the existence of ipt gene in outcross progenies analyzed by PCR. The range of fertility rates followed by artificial, insect (bee), and nature pollination were 86-100%, 1.1-42.5% and 0-9% respectively, while no hybridization was observed for ‘Pakchoi’ and ‘Rapa’ as pollen recipients. The results showed that transgenic DNA insertion did not alter potential risk for outcross among broccoli varieties. However, there was possibility of gene transfer between transgenic and non-transgenic broccoli varieties. In order to reveal the competitive advantage of F1 progenies breed from transgenic ipt broccoli crossed with four closely related varieties, 103 transgenic line and 104(CK) inbred line were used as pollen donors and hybridized with related varieties by artificial pollination. F1 offsprings from hybridization with 103 showed larger plant height, while 104 F1 hybrid offspring showed shorter. It was also found that the seed number per pod of 103 F1 hybrid was higher than 104 (CK) hybrid F1. Results showed F1 progeny from transgenic broccoli hybrid still exhibited the phenomenon of heterosis.
Another concern of potential impact from transgenic broccoli is microbial communities in environment. According to our results, transgenic ipt broccoli had no significant effects on soil pH, and 103 transgenic line exhibited higher rhizosphere organic matter content on the day of transplantation. However, soil organic matter data showed no significant difference among all lines after 90 days. In 2010, total numbers of fungi and bacteria were evaluated, and inbred line 104(CK) had significantly higher than those of other lines. Moreover, almost all examined microbial populations showed significant difference among transgenic and non-transgenic lines during growing season, and there was no significant difference after harvest and tillage. Nevertheless, the populations of gram-negative bacteria, urea-oxidizing bacteria and cellulolytic bacteria showed difference on 105th days after transplanting in 2010. Similarity coefficient ofsoil microorganism were scored between transgenic and non-transgenic broccoli lines also examined. The result showed that commercial F1 hybrid variety ‘Green King’ had lower similarity coefficient score than the other two lines.
Transformation of a cytokinin limiting enzyme adenosine ipt gene enhanced stress tolerance by delaying the onset of flower head wilting which prolongs the shelf-life of broccolis. Assess the effectiveness of transgenic ipt gene on improving broccoli stress tolerance and its commercialization value, transgenic broccoli was sampled by lyophilizing the 2nd leaf from top. Overall, transgenic line 103 expressed higher ipt transcripts levels than those of 102, and as a result line 103 withstood unfavorable environmental stresses better than 102 in general. Interestingly, the ipt expression levels varied over time as observed in those experiments for drought and heat treatments. Drought stress can decrease the expression levels of ipt gene but the effect of heat stress on ipt gene expression was temperature and duration specific, therefore it was inclusive. The introduction of ipt gene to transgenic broccoli line 103 enhanced the ability of water retention in leaves while ethylene production was reduced compared with transgenic line 102 or other non-transgenic lines whenever broccolis were subjected to the treatments of flood, drought, or heat stress.

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中文摘要 I
英文摘要 IV
誌謝 VII
誌謝後記 IX
表目錄 XV
圖目錄 XVII
縮字表 XIX
附錄目錄 XXIII
第一章 前言 1
第二章 前人研究 6
第三章 異戊烯轉移酶(ipt)基因轉殖青花菜之遺傳性狀評估 29
一、前言 29
二、材料與方法 29
(一) 試驗材料 29
(二) 田間試驗 30
(三) 植株性狀 30
(四) 花球性狀 31
(五) 花球櫥架壽命 31
(六) 統計分析 31
三、結果 32
(一) 2004-2007年基因轉殖與非基因轉殖青花菜園藝性狀
調查 32
(二) 2004-2007年基因轉殖與非基因轉殖青花菜花球性狀
調查 32
(三) 花球櫥架壽命調查 33
四、討論 33
第四章 異戊烯轉移酶(ipt)青花菜花粉活性與其近緣種雜交之可
能性 46
一、前言 46
二、材料與方法 47
(一) 試驗材料 47
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(二) 花粉活性與性狀測定 47
1. 花粉活性 47
2. 花粉性狀 48
3. 果莢結實率(pod setting) 49
(三) 與近緣亞種之雜交試驗 49
1. 人工雜交 49
2. 蜜蜂進行近緣種雜交 49
3. 田間自然試驗 49
(四)基因轉殖植體檢測分析 50
(五) 統計分析 51
三、結果 51
(一) 花粉的活性測定 51
(二) 花粉性狀調查 51
(三) 基因轉殖青花菜與近緣種作物之花期重疊觀察 52
(四) 人工雜交、蜜蜂授粉及田間之自然雜交試驗 52
四、討論 53
第五章 異戊烯轉移酶(ipt)基因轉殖青花菜雜交後之性狀探討 66
一、前言 66
二、材料與方法 67
(一) 試驗材料 67
(二) 雜交F1後代植株生育性狀調查 67
(三) 雜交F1後代花粉活性及性狀檢測 68
(四) 雜交F1後代花球性狀 68
(五) 統計分析 69
三、結果 69
(一) 雜交F1後代植株生育性狀調查 69
(二) 雜交F1後代花粉活性測定、果莢結實率及種子數 70
(三) 雜交F1後代花粉性狀 70
(四) 雜交F1後代花球生理變化 71
1. 乙烯生成量 71
2. 色差 71
四、討論 72
第六章 異戊烯轉移酶(ipt)基因轉殖青花菜微生物相之研究 88
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一、前言 88
二、材料與方法 89
(一) 試驗材料與採樣 89
(二) 土壤質地分析 89
1. 土壤pH值測定 90
2. 土壤有機質測定 90
(三) 葉表微生物相 90
(四) 土壤微生物相 91
1. 總真菌數 91
2. 總細菌數 91
3. 尿素氧化菌 91
4. 溶磷菌 91
5. 蛋白質分解菌 91
6. 纖維分解菌 91
7. 格蘭氏陰性菌 91
8. 放線菌 91
(五) 以分生技術分析土壤微生物相 92
1. 微生物群落之去氧核糖核酸萃取 92
2. DNA濃度測定 92
3. 進行PCR反應(放大16S rDNA與18S rDNA) 92
4. 以DGGE電泳分離PCR產物 93
5. 電泳圖譜多型性分析 93
(六) 統計分析 94
三、結果 94
(一) 基因轉殖ipt青花菜對土壤質地的影響 94
(二) 基因轉殖ipt青花菜對葉表微生物數量消長之影響 94
(三) 基因轉殖ipt青花菜對土壤微生物數量消長之影響 95
(四) 以分生技術分析土壤微生物相似性 97
四、討論 97
第七章 異戊烯轉移酶(ipt)基因轉殖青花菜野化能力之評估 118
一、前言 118
二、材料與方法 119
(一) 試驗材料 119
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(二) 乾旱試驗 120
1. 植株葉片萎凋率 120
2. 乙烯釋放量之測定 120
3. 脯氨酸累積含量之測定 121
4. 葉片相對含水量(related water content, RWC) 121
5. Real-time PCR定量 122
(三) 高溫試驗 124
(四) 淹水試驗 124
1. 葉片萎凋率(%) 124
2. 葉片黃化率(%) 124
3. 乙烯釋放量之測定 125
4. 根系生長 125
(五) 統計分析 125
三、結果 125
(一) 乾旱試驗 125
(二) 高溫試驗 126
(三) 淹水試驗 127
四、討論 128
(一) 基因轉殖ipt青花菜在乾旱及高溫逆境下之ipt基因表
現 128
(二) 基因轉殖ipt青花菜在乾旱、高溫及淹水逆境之乙烯
生成量影響 130
(三) 基因轉殖ipt青花菜在乾旱及高溫逆境之脯胺酸累積
影響 132
(四) 基因轉殖ipt青花菜在淹水逆境之不定根生成 134
第八章 綜合討論 162
中文參考文獻 177
英文參考文獻 182
附錄4-1. 蕓薹屬植物由3個初級物種因雜交及染色體數加倍進化為後成物種之三角形的圖解(宇氏三角) 215
附錄4-2. 十字花科蕓薹屬作物之相關物種 216
附錄6-1. 農業試驗所基因轉殖隔離田植前土壤質地分析 217
附錄6-2. 土壤有機質含量測定 218
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附錄6-3. 變性梯度膠體電泳之凝膠配方 220
簡歷 221
攻讀碩、博士學位期間公開發表的論文 224

表目錄
頁次
表2-1.
基因轉殖作物分類情況
27
表2-2.
基因轉殖作物核准隔離田間試驗申請案
28
表3-1.
2004-2007年田間基因轉殖ipt青花菜與非基因轉殖青花菜植株性狀之綜合變方分析
39
表3-2.
2004-2007年田間基因轉殖ipt青花菜與非基因轉殖青花菜之植株性狀
40
表3-3.
2004-2007年間經LSD測驗顯示基因轉殖ipt青花菜與非基因轉殖青花菜間之植株性狀顯著性差異總整理
41
表3-4.
2004-2007年田間基因轉殖ipt青花菜與非基因轉殖青花菜花球性狀之綜合變方分析
42
表3-5.
2004-2007年田間基因轉殖ipt青花菜與非基因轉殖青花菜之花球性狀
43
表3-6.
2004-2007年間經LSD測驗顯示基因轉殖ipt青花菜與非基因轉殖青花菜間之花球產量性狀顯著性差異總整理
44
表3-7.
2004-2007年田間基因轉殖ipt青花與非基因轉殖青花菜花球採收後之櫥架壽命
45
表4-1.
基因轉殖ipt基因青花菜(103品系)與其近緣亞種作物的花粉活性、果莢結實率及種子數之比較
60
表4-2.
基因轉殖ipt青花菜與其近緣種作物之花粉性狀比較
61
表4-3.
2007及2008年基因轉殖ipt基因青花菜(103品系)及其近緣種作物之重疊花期
62
表4-4.
基因轉殖ipt基因青花菜與其近緣種作物經由人工與蜜蜂授粉之雜交情形
63
表4-5.
2007及2008年田間基因轉殖ipt青花菜與非基因轉殖青花菜花球性狀之變方分析
64
表4-6.
轉基因青花菜與其近緣種作物經田間自然授粉之雜交情形
65
表5-1.
分別利用基因轉殖ipt青花菜及非基因轉殖青花菜當花粉貢獻親與近緣種作物雜交之F1後代植株性狀變異係數(%)
76
表5-2.
分別利用基因轉殖ipt青花菜與非基因轉殖青花菜當花粉貢獻親與近緣種作物雜交之F1後代花粉活性、果莢結實率與種子數之比較
81
表5-3.
分別利用基因轉殖ipt青花菜與非基因轉殖青花菜當花粉貢獻親與近緣種作物雜交之F1後代花粉性狀比較
82
表5-4.
103雜交近緣種F1後代之花球從溫室收穫後貯藏於25±2℃之乙烯生成量影響
84
表5-5.
103雜交近緣種F1後代之花球從溫室收穫後貯藏於25±2℃之L值
85
表5-6.
103雜交近緣種F1後代之花球從溫室收穫後貯藏於25±2℃之a值
86
表5-7.
103雜交近緣種F1後代之花球從溫室收穫後貯藏於25±2℃之b值
87
表6-1.
基因轉殖ipt青花菜與非基因轉殖青花菜種植期間之土壤pH值
104
表6-2.
基因轉殖ipt青花菜與基因非轉殖青花菜種植期間之土壤有機質比較
105
表6-3.
2009年及2010年基因轉殖ipt青花菜與非基因轉殖青花菜表土細菌族群之相似性係數比較
117
表7-1.
基因轉殖ipt青花菜對乾旱處理後第4、5、6天之萎凋率及分別覆水24小時之植株回復率
138
表7-2.
乾旱對基因轉殖ipt青花菜乙烯生成量之影響
139
表7-3.
乾旱對基因轉殖ipt青花菜脯胺酸累積之影響
141
表7-4.
乾旱對基因轉殖ipt青花菜葉片相對含水量之影響
143
表7-5.
乾旱對轉殖青花菜ipt基因表現量之影響
145
表7-6.
熱處理對基因轉殖ipt青花菜乙烯生成量之影響
148
表7-7.
熱處理對基因轉殖ipt青花菜脯胺酸累積之影響
150
表7-8.
熱處理對基因轉殖ipt青花菜相對含水量之影響
152
表7-9.
熱處理後對基因轉殖青花菜ipt基因表現量之影響
154
表7-10.
淹水對基因轉殖ipt青花菜萎凋率之影響
157
表7-11.
淹水對基因轉殖ipt青花菜黃花之影響
158
表7-12.
淹水對基因轉殖ipt青花菜乙烯釋放量之影響
159
表8-1.
基因轉殖ipt青花菜與非基因轉殖青花菜間之生物安全評估項目結果總整理
171
表8-2.
基因轉殖ipt青花菜之生物安全評估的方法總整理
173

圖目錄
頁次
圖1-1.
基因轉殖ipt青花菜pSG766A載體構築
4
圖1-2
執行基因轉殖ipt青花菜評估項目之流程大綱
5
圖3-1.
2004-2007年期間基因轉殖ipt青花菜、非基因轉殖青104 (CK)與GK商業品種帶柄花球重等級百分比分布
36
圖3-2.
2004-2007年期間基因轉殖ipt青花菜、非基因轉殖青104 (CK)與GK商業品種不帶柄花球重等級百分比分布
37
圖3-3.
基因轉殖ipt青花與非基因轉殖青花菜花球採收當天與室溫25±2℃貯藏6天後之黃化情形
38
圖4-1.
基因轉殖ipt青花菜(103品系)及其近緣種作物在農業試驗所隔離網室進行蜜蜂授粉試驗
57
圖4-2.
基因轉殖ipt基因青花菜(103品系)及其近緣種在農業試驗所隔離田之種植配置
58
圖4-3.
基因轉殖ipt青花菜 (103品系) 與‘綠王’在田間自然雜交之後代的PCR檢測
59
圖5-1.
分別利用基因轉殖ipt青花菜及非基因轉殖青花菜當花粉貢獻親與芥藍-富悅(白花)作物雜交之F1後代性狀比較
77
圖5-2.
分別利用基因轉殖ipt青花菜及非基因轉殖青花菜當花粉貢獻親與芥藍-白格林(黃花)作物雜交之F1後代性狀比較
78
圖5-3.
分別利用基因轉殖ipt青花菜及非基因轉殖青花菜當花粉貢獻親與花椰菜-麗雪作物雜交之F1後代性狀比較
79
圖5-4.
分別利用基因轉殖ipt青花菜及非基因轉殖青花菜當花粉貢獻親與青花菜-綠王作物雜交之F1後代性狀比較
80
圖5-5.
雜交F1後代之花球黃化情形
83
圖6-1.
基因轉殖ipt與非基因轉殖青花菜在2009及2010年葉表微生物量(總真菌數及總細菌數)
106
圖6-2.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤真菌總數
107
圖6-3.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤總細菌數
108
圖6-4.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤總尿素氧化菌數
109
圖6-5.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤總溶磷菌數
頁次
110
圖6-6.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤總蛋白質分解菌菌數
111
圖6-7.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤總纖維分解菌菌數
112
圖6-8.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤革蘭氏陰性菌菌數
113
圖6-9.
基因轉殖ipt青花菜與非基因轉殖青花菜之土壤放線菌菌數
114
圖6-10.
利用DGGE分析2009年及2010年基因轉殖ipt青花菜與非基因轉殖青花菜之土壤16S rDNA核苷酸片段
115
圖6-11.
利用DGGE分析2009年及2010年基因轉殖ipt青花菜與非基因轉殖青花菜之土壤18S rDNA核苷酸片段
116
圖7-1.
乾旱對基因轉殖ipt青花菜植株生長之影響
137
圖7-2.
乾旱後對基因轉殖ipt青花菜之乙烯相對量影響
140
圖7-3.
乾旱後對基因轉殖ipt青花菜之脯胺酸相對量影響
142
圖7-4.
乾旱後對基因轉殖ipt青花菜相對葉片含水量之影響
144
圖7-5.
35/30℃對基因轉殖ipt青花菜植株生長之影響
146
圖7-6.
40/35℃對基因轉殖ipt青花菜植株生長之影響
147
圖7-7.
熱處理後對基因轉殖ipt青花菜相對乙烯量之影響
149
圖7-8.
熱處理對基因轉殖ipt青花菜脯胺酸相對含量之影響
151
圖7-9.
熱處理對基因轉殖ipt青花菜相對葉片含水量之影響
153
圖7-10.
熱處理對基因轉殖青花菜ipt基因相對表現量之影響
155
圖7-11.
不同淹水時間對基因轉殖ipt青花菜植株生長之影響
156
圖7-12.
淹水後第12天對基因轉殖ipt青花菜根系生長之影響
160
圖7-13.
基因轉殖ipt青花菜淹水後第25天不定根之生成
161

附錄目錄
頁次
附錄4-1.
蕓薹屬植物由3個初級物種因雜交及染色體數加倍進化為後成物種之三角形的圖解(宇氏三角)
215
附錄4-2.
十字花科蕓薹屬作物之相關物種
216
附錄6-1.
農業試驗所基因轉殖隔離田植前土壤質地分析
217
附錄6-2.
土壤有機質含量測定
218
附錄6-3.
變性梯度膠體電泳之凝膠配方
220

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