臺灣博碩士論文加值系統

水稻白葉枯病是由白葉枯病菌Xanthomonas oryzae pv. oryzae (Xoo)所感染的系統性病害，白葉枯病可導致水稻產量減少20 %至50 %，目前大多使用農藥防治，但對環境造成影響，且防治效果有限。前人研究顯示，在植株內堆疊多個不同抗性機制的抗性基因能抵抗更多生理小種，達到廣幅抗性。台南11號 (TN11) 具有高產、抗倒伏、抗褐飛蝨等優良特性，是目前台灣種植面積最廣的品種，但其對白葉枯病不具有抗性。本研究以國際水稻研究所 (International Rice Research Institute, IRRI) 育成帶有五個抗白葉枯病基因 (Xa4+xa5+Xa7+xa13+Xa21) 的秈稻品種IRBB66作為抗病基因提供親 (donor parent)，與稉稻TN11進行雜交和回交，導入白葉枯病抗性基因，再以分子標誌輔助選拔法 (marker- assisted selection，MAS) 選出帶有五個抗性基因的植株後，自交純化，並進行外表型選拔，希望選出抗白葉枯病且農藝性狀良好之植株，以育成優良抗病品種。本試驗使用BC1及BC2兩個回交族群，不同回交世代在外表性狀上會有更多的選擇，從BC1F2與BC2F2世代始，以與抗病基因連鎖之分子標誌篩選帶有四個 (xa5+Xa7+xa13+Xa21) 或五個抗病基因的植株，於植株最高分蘗期接種病原菌進行抗病檢定，透過農藝性狀篩選後，選拔優良植株於下一期作進行抗病基因和外表性狀純化，並繼續篩選。至2014Ⅱ期作，選育出的18個抗病品系已達BC1F5及BC2F6世代，種植至苗栗改良場、霧峰農試所與北溝農場並接種Xoof2、Xf89-b及Xe2三種病原菌，檢定結果呈抗性反應，且植株外觀、米質及產量表現良好，抗褐飛蝨又抗倒伏，未來可推廣成為抗白葉枯病的新品種，亦可做為良好的雜交親本。本試驗亦利用禾本科網站中，比較圖譜的資訊進行圖譜交叉比對，並經由親本間多型性檢定，搜尋到位在抗病基因上和與抗病基因連鎖距離更近之SSR分子標誌，不僅可節省成本及時間的耗費，更有利於育種效率的提升。

摘要 i
Abstract ii
目錄 iii
表目錄 v
圖目錄 vi
附錄 viii
壹、前言 1
貳、前人研究 3
一、台灣水稻主要栽培品種之演變及品種遺傳多樣性之變遷 3
二、秈稉雜交品種之優勢 5
三、水稻白葉枯病 6
四、台灣及全世界目前遭遇之白葉枯病相關問題 8
五、水稻白葉枯病抗性基因 9
(一) Xa4 9
(二) xa5 10
(三) Xa7 11
(四) xa13 11
(五) Xa21 12
六、白葉枯病抗性品系IRBB近同源品系之建立 13
七、分子輔助育種及分子輔助回交育種 14
八、基因堆疊 16
參、材料與方法 19
一、輪迴親、供給親及抗病檢定對照品種 19
二、水稻種植與栽培管理 19
三、水稻宿根栽培流程 20
四、水稻雜交 20
五、育種流程 21
六、白葉枯病接種檢定 22
七、菌流測試 23
八、農藝性狀調查 23
九、優良品系之食味品質、米粒理化特性分析 23
十、水稻大量基因組DNA萃取 25
十一、水稻小量基因組DNA萃取 26
十二、聚合酶連鎖反應 (polymerase chain reaction, PCR) 26
十三、聚丙烯醯胺膠體製作 27
十四、聚丙烯醯胺膠體分析 27
十五、洋菜膠體製作 27
十六、洋菜膠體電泳 28
十七、電泳資料分析 28
十八、白葉枯病抗性基因連鎖分子標誌之開發 28
肆、結果 29
一、白葉枯病抗性基因連鎖分子標誌之開發 29
(一) Xa4 29
(二) xa5 30
(三) Xa7 30
(四) xa13 31
(五) Xa21 31
二、白葉枯病抗性基因植株基因型分析與白葉枯病抗性檢定 32
(一) BC1 33
(二) BC2 34
三、以病斑長度篩選抗性植株 37
四、農藝性狀調查 38
(一) 感性對照品種TN1、IR24、親本TN11與IRBB66，接種不同病原菌，是否影響產量之分析 39
(二) 接種不同白葉枯病菌系對抗病品系與親本TN11在產量之影響 39
五、米質分析 41
(一) 優良抗病品系與親本TN11之比較 41
(二) TN11接種對米質之影響 42
六、秈稉稻雜交之雜交優勢及不同世代之選拔效率 42
伍、討論 45
一、MAS在堆疊育種中的優點 45
二、白葉枯病抗性基因連鎖分子標誌之開發 46
三、優良抗病品系之白葉枯病抗性檢定分析 47
四、農藝性狀調查與米質分析 49
(二) 優良抗病品系於外表性狀之同質化分析 49
(二) 白葉枯病對產量及米質的影響 50
五、秈稉稻雜交之雜交優勢及不同世代之選拔效率 53
陸、結論 55
柒、參考文獻 57

表目錄
表一、各期作性狀調查項目總覽 70
表二、2012Ⅱ BC2F2使用與白葉枯病抗性基因連鎖之SSR分子標誌 71
表三、2013ⅠBC2F3及BC1F2使用與白葉枯病抗性基因連鎖之SSR分子標誌 72
表四、用於篩選2013ⅠBC2F3及BC1F2的多重聚合酶反應之引子組及黏合溫度 73
表五、2013Ⅱ親本TN11、IRBB66及選育出優良之BC2F4植株性狀資料 74
表六、2013Ⅱ親本TN11、IRBB66及選育出優良之BC1F3植株性狀資料 75
表七、2014Ⅰ親本TN11及23株優良抗病品系(BC1F4、BC2F5)植株性狀資料 76
表八、2014Ⅰ抗性對照品種TN1、IR24及親本TN11、IRBB66接種病原菌後 78
表九、2014Ⅱ18個優良抗病品系與親本TN11之產量相關性狀 79
表十、優良抗性品系與親本TN11之農藝性狀 80
表十一、親本TN11和優良抗病品系2014年Ⅰ、Ⅱ期作性狀比較 81
表十二、優良抗病品系和親本TN11之2014年Ⅰ、Ⅱ期作米質分析(委託嘉義分所分析) 82
表十三、優良抗病品系和親本TN11之2014年Ⅱ期作米質分析(委託台中改良場分析) 83
表十四、2014Ⅱ期作TN11接種不同病原菌及未接種病原菌之米質分析(委託嘉義分所分析) 84
表十五、2014Ⅱ期作TN11接種不同病原菌及未接種病原菌之米質分析(委託台中改良場分析) 85
表十六、親本TN11、IRBB66與F2、BC1F2及BC2F2性狀變異域比較 86

圖目錄
圖一、白葉枯病抗性基因導入台南11號育成新抗病優良品種之流程 87
圖二、與抗性基因Xa4和xa5連鎖之分子標誌於親本TN11、IRBB66及BC1F3子代之篩選 88
圖三、與抗性基因Xa7連鎖之分子標誌，於親本TN11、IRBB66及BC1F3子代之篩選 89
圖四、與抗性基因xa13連鎖之分子標誌，於親本TN11、IRBB66及BC1F3子代之篩選 90
圖五、與抗性基因Xa21連鎖之分標誌，於親本TN11、IRBB66及BC1F3子代之篩選 91
圖六、感性對照TN1、IR24、親本TN11、IRBB66及TN11/IRBB66 BC1F2帶有五個及四個抗性基因植株接種Xoof2之病斑長度 92
圖七、感性對照TN1、IR24、親本TN11、IRBB66及 TN11/IRBB66 BC1F3帶有五個及四個抗性基因植株接種Xoof2之病斑長度 93
圖八、感性對照TN1、IR24、親本TN11、IRBB66及TN11/IRBB66 BC2F2帶有四個抗性基因植株接種Xf89-b之病斑長度 94
圖九、感性對照TN1、IR24、親本TN11、IRBB66及TN11/IRBB66 BC2F3帶有四個抗性基因植株接種Xoof2之病斑長度 95
圖十、感性對照TN1、IR24、親本TN11、IRBB66及TN11/IRBB66 BC2F4帶有四個抗性基因植株接種Xoof2之病斑長度 96
圖十一、2013Ⅱ親本TN11、IRBB66與帶有四個抗性基因之TN11/IRBB66 BC2F3及帶有五個抗性基因之TN11/IRBB66 BC1F2宿根植株接種Xoof2及Xf89-b之病斑長度 97
圖十二、2013Ⅱ期作TN11/IRBB66 BC1F2和BC2F3宿根栽培植株，接種白葉枯病生理小種Xoof2及Xf89-b之病斑長度照片 98
圖十三、感性對照TN1、IR24、親本TN11、IRBB66及TN11/IRBB66 BC2F5帶有五個及四個抗性基因植株接種Xoof2之病斑長度 99
圖十四、感性對照TN1、IR24、親本TN11、IRBB66及TN11/IRBB66 BC2F5植株(A-R)接種白葉枯病生理小種Xoof2之病斑長度照片 100
圖十五、感性對照TN1、IR24、親本TN11、IRBB66及23株優良抗病品系接種白葉枯病生理小種Xoof2、Xf89-b及Xe2之病斑長度 101
圖十六、感性對照TN1、IR24、親本TN11、IRBB66及優良抗性品系A-W接種白葉枯病生理小種Xoof2、X89-b及Xe2之病斑長度照片 102
圖十七、植於農試所之感性對照TN1、IR24、親本TN11、IRBB66及種優良抗性品系K-R接種白葉枯病生理小種Xoof2之病斑長度 103
圖十八、種植於苗栗改良場之感性對照TN1、IR24、親本TN11、IRBB66及優良抗性品系K-R接種白葉枯病生理小種Xoof2及Xe2之病斑長度 104
圖十九、感性對照TN1、IR24、親本TN11、IRBB66及優良抗性品系A-W以病斑長度進行群集分析 105
圖二十、2014Ⅰ期作，23優良抗病品系A-W與親本TN11植株外觀、穀粒及病斑長度照片 106
圖二十一、2014Ⅰ期作，23個優良抗病品系A-W與親本TN11株高和分蘗數之變異係數 107
圖二十二、2014Ⅱ期作，優良抗病品系與親本TN11、IRBB66株高和分蘗數之變異係數 108
圖二十三、2014Ⅱ期作，優良抗病品系B-T與親本TN11植株外觀照片 109
圖二十四、優良抗病品系與親本TN11穀粒外觀照片 110
圖二十五、TN11/IRBB66 F2子代外表型變異照片 111
圖二十六、TN11、品系M及U (BC1F5)之植株外觀、穀粒外觀及寬葉照 112
圖二十七、感性對照品種TN1、IR24接種病原菌Xoof2、Xf89-b、Xe2及未接種之植株外觀照片 113
圖二十八、親本TN11、IRBB66接種病原菌Xoof2、Xf89-b、Xe2及未接種之植株外觀照片 114

附錄
附錄一、田間及溫室內白葉枯病抗性測量等級依據 115
附錄二、米質分析簡易流程圖 116
附錄三、感性對照TN1、IR24、IR64、親本TN11、IRBB66之菌流測試 117
附錄四、2013Ⅱ期作田間飛蝨類危害情形 118
附錄五、2014Ⅱ期作品系G及K田間二化螟蟲危害情形 119

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