臺灣博碩士論文加值系統

本論文主要針對莧白銹菌 (Albugo bliti) 之生物特性及其病害防治進行研究。白銹菌為卵菌綱 (Oomycetes) 、露菌目 (Peronosporales) 、白銹菌科 (Albuginaceae) 、白銹菌屬 (Albugo) 之絕對寄生菌，供試菌株分別來自白莧 (Amaranthus mangostanus) 、紅莧 (A. mangostanus forma ruber) 、鳥莧 (A. lividus) 與滿天星 (Alternanthera sessilis) 等人工接種後之罹病植株，各菌株間之游走孢子囊、游走孢子與卵孢子在形態上並無顯著差異，經比對均為莧白銹菌 (A. bliti) 。游走孢子為莧菜白銹病之主要感染源，直接發芽後從氣孔侵入，於氣孔下腔室 (substomatal chamber) 內形成膨大泡囊 (substomatal vesicles) 與管狀菌絲 (coenocytic hyphae) ，菌絲在細胞間隙生長，菌絲兩側則產生球型吸器 (global haustoria) 侵入寄主細胞內吸取養分。莧白銹菌侵入寄主植物 4 - 5 天後，於海綿葉肉組織 (spongy mesophyll) 形成游走孢子囊 (sporangia) 與游走孢子囊堆 (sori) ，同時在海綿組織與葉脈 (veins) 形成藏精器 (antheridia) 與藏卵器 (oogonia) ，藏卵器經授精後產生卵孢子 (oopores) ，一般莧白銹菌之卵孢子在菌絲侵入寄主植物 10 - 11天後形成。莧白銹菌僅感染莧菜子葉與真葉，而葉背氣孔開放率比葉表者多，乃莧白銹菌之主要侵入部位，且為游走孢子囊堆之主要分布位置。以植株、子葉或切葉進行病原性接種測試之結果顯示，莧白銹菌各菌株僅能感染原寄主植物，具寄主專一性。莧菜接種時，葉表水膜只要持續 1 小時以上，就可使莧菜被感染而發病。莧白銹菌侵入寄主 8 - 9 天後所形成之游走孢子囊成熟度最好，釋放游走孢子的比率最高，將此游走孢子囊置於蒸餾水 20 - 40 分鐘後，開始釋放游走孢子，於 4 小時後的釋放率達到高峰，適合游走孢子囊釋放游走孢子之溫度介於 12 - 28 oC，最適溫度為 20 - 24 oC，32 - 36oC抑制游走孢子之釋放，但是移置於 20 oC經 4 小時後可恢復部份游走孢子囊釋放游走孢子之能力。莧白銹菌游走孢子囊在室溫可存活 1 個月，而在土壤中則可存活 1 年左右，游走孢子囊可在潮溼土壤中釋放游走孢子感染植株。田間試驗發現莧菜白銹病之初次感染源來自土壤，在莧菜真葉形成初期之子葉與真葉被感染後，成為田間第二次感染源的主要來源，當植株生長到3-4葉期時，是第二次感染源大量產生之時。白銹病在田間之病勢進展，初期緩慢，到播種10-14天後才開始發病，爾後罹病率與罹病程度呈現急速上升趨勢。病斑在植株上之分布主要集中在由上向下數之第4-8葉位，而白銹病在田間之分布，於初期之子葉期與一葉期時屬逢機分布，隨後轉變成均勻分布。藥劑篩選結果發現銅快得寧及含有銅離子之藥劑抑制游走孢子囊釋放游走孢子之效果最佳。對病害防治的最適當施藥時機，是在子葉期僅施一次就可達防治效果。隧道式覆蓋法及應用界面活性劑十二烷基硫酸鈉 (sodium dodecyl sulfate) 也有良好的防治效果。以逢機增幅多型性 DNA 片段圖譜法及 rDNA 內轉錄區間隔定序法對所採集之莧白銹菌各菌株、甕菜白銹菌 (A. ipomoeae-aquaticae) 與馬齒莧白銹菌 (A. portulacae) 進行親緣性分析 (Phylogenetic analysis) ，兩種分析法所得之白銹菌親緣關係類似，顯示莧屬植物上之莧白銹菌菌株由兩群所組成，其中一群為白莧與紅莧菌株，另一群為鳥莧菌株，滿天星屬植物上之滿天星菌株也單獨形成一群，甕菜白銹菌與對照菌株薺菜 (Capsella bursa-pastoris) 白銹菌 (A. candida) 親緣性較近，馬齒莧白銹菌與所測試之白銹菌菌株親緣性最遠，顯示莧白銹菌為適應寄主植物而與寄主植物經長期共同演化後，使莧白銹菌各菌株間之物種演化有所變異。

The aim of this study was to investigate the biological characteristics of Albugo bliti and its disease control. Albugo is an obligate parasitic Oomycete genus classified in the family Albuginaceae of Peronosporales, and among which A. bliti causes white rust of amaranth. Isolates were collected from the artificial inoculated plants of Amaranthus mangostanus, A. mangostanus forma ruber, A. lividus, and Alternanthera sessilis, respectively. However, there is no distinguishable difference in the morphology of isolates from different hosts and all identified as A. bliti. Zoospores, the main inocula of amaranth white rust, germinated and penetrated host plant via stomata, and then formed substomatal vesicles and coenocytic hyphae in the substomatal chamber. Hyphae grew intercellularly with lateral globose haustoria absorbing the host nutrients. The fungus formed sporangia and sori in spongy mesophyll at 4 - 5 days after inoculation, and in the meanwhile formed antheridia and oogonia in spongy mesophyll and veins. Subsequently, mating between antheridia and oogonia occurred, and then oospores developed at 10 - 11 days after inoculation. Infection and symptoms only occurred on cotyledon and leaves of amaranth. As stomata open more frequently on the abaxial leaf surfaces than on the adaxial leaf surfaces, infection sites and sori were mainly distributed on the abaxial leaf epidermis. Cross inoculations by using whole plants, cotyledons and detached-leaf method showed that infection only occurred on their original host species, manifesting the host specificity of A. bliti. Maintaining for a minimum of one hour of water film on the leaf surface was required for the infection of A. bliti. The highest amount of discharged zoospores was obtained from the sporangia at 8 - 9 days after inoculation. Zoospores would be discharged within 20-40 minutes as mature sporangia were incubated in distilled water at 24 oC, and discharged most abundantly about 4 hr after treatment. The temperatures for sporangia to discharge zoospores ranged from 12 to 28 oC, while 20 - 24 oC was optimal. The zoospore discharging was inhibited at 32 -36 oC, but resumed partially upon transferring these sporangia to 20 oC for 4 hr. Sporangia of A. bliti could survive for one month at room temperature and for one year in soil. The surviving sporangia were able to discharge zoospores in wet soil and caused infection. Salt solutions like acetic acid and copper acetate, and buffers used were found to be inhibitory to the sporangial germination of A. bliti. Disease development of white rust was studied in field. Initial inocula of infection came from soil. The infected cotyledons and diseased leaves of seedlings yielded sporangia abundantly at 3-4 leaf stage, serving as the secondary inocula for infection. Disease development of white rust was slow in the beginning. Symptom did not appear until 10 - 14 days after sowing seed, since then disease incidence and severity increased rapidly. The symptoms mainly occurred among the 4th to 8th leaf counted from the top. Diseased plant distributed randomly in the field at cotyledon and one leaf stage, and the disease was spreading to the entire field afterward. Among the several fungicides tested against sporangial germination, copper hydroxide + oxine-copper and those containing cupric ions were the most effective. The best time of fungicide application was at cotyledon stage, and only one application was adequate. Planting amaranth in plastic tunnel or application of surfactant sodium dodecyl sulfate were effective for controlling amaranth white rust in field. The phylogenetic relationship among A. bliti, A. ipomoeae-aquaticae, and A. portulacae collected in Taiwan were analyzed using random amplified polymorphic DNA method (RAPD) and rDNA internal transcribed spacers (ITS) analysis. Results of RAPD and ITS showed the same phylogenetic relationship among tested Albugo species. The genetic distance of A. bliti isolates from Amaranthus spp. could be differentiated into two groups. One group contained Abw and Abr isolates from A. mangostanus, another one contained Abl isolates from A. lividus. Albugo bliti Aba isolates from Al. sessilis formed distinctly a separate group. Albugo ipomoeae-aquaticae was close to A. candida from Capsella bursa-pastoris, while A. portulacae was related most distantly among these tested Albugo spp. The phylogenetic diversity of A. bliti isolates suggested that they have coevolved with their hosts.

目 錄
中文摘要 1
英文摘要 3
前 言 5
引用文獻 15
第一章莧白銹菌病之分類、病原性測試與寄主專一性 24
摘要 24
前言 24
材料與方法 25
供試菌株來源、保存與繁殖 25
寄主植物之繁殖 26
莧科白銹菌菌株鑑定 26
接種源之製備 26
病原性測定 26
結果 27
白銹菌菌株鑑定 27
莧白銹病之病徵發展 27
病原性測定 28
討論 28
引用文獻 30
英文摘要 32
圖表 33
第二章莧白銹菌侵染過程及生活史 40
摘要 40
前言 40
材料與方法 41
莧白銹菌侵入部位與影響因子 41
1.侵入部位 41
(1)根、莖、葉組織 41
(2)葉表和葉背 42
2.植物氣孔開關與莧白銹菌侵入之關係 42
3.植物氣孔開關之影響因子 43
(1)光照與黑暗 43
(2)葉面水膜 43
莧白銹菌在寄主體內之侵染過程 43
莧白銹菌生活史 44
結果 44
莧白銹菌侵入部位與影響因子 44
1.侵入部位 44
(1)根、莖、葉組織 44
(2)葉表和葉背 45
2.植物氣孔開關與莧白銹菌侵入之關係 45
3.植物氣孔開關之影響因子 45
(1)光照與黑暗 45
(2)葉面水膜 46
莧白銹菌在寄主體內之侵染過程過程 46
莧白銹菌生活史 46
討論 47
引用文獻 50
英文摘要 53
圖表 54
第三章莧白銹菌游走孢子囊釋放游走孢子之影響因子與存活 62
摘要 62
前言 63
材料與方法 64
游走孢子囊成熟所需時間 64
游走孢子囊釋放游走孢子之時間 65
游走孢子囊釋放游走孢子之影響因子 65
1.溫度 65
(1)發芽適溫 65
(2)高溫作用 65
2.葉背水膜持續時間 66
3.醋酸與醋酸銅等溶液 66
4.pH值 66
5.蔗糖 67
游走孢子囊之存活 67
1.乾燥之影響 67
2.游走孢子囊於土壤中之存活 68
3.土壤萃出液對游走孢子囊釋放游走孢子之影響 68
4.混合游走孢子囊之土壤懸浮液與病害發生之關係 68
統計分析 69
結果 69
游走孢子囊成熟所需時間 69
游走孢子囊釋放游走孢子之時間 70
游走孢子囊釋放游走孢子之影響因子 70
1.溫度 70
(1)發芽溫度 70
(2)高溫作用 71
2.葉背水膜持續時間 71
3.醋酸與醋酸銅等溶液 72
4.pH值 72
5.蔗糖 72
游走孢子囊之存活 73
1.乾燥之影響 73
2.游走孢子囊於土壤中之存活 73
3.土壤萃出液對游走孢子囊釋放游走孢子之影響 73
4.混合游走孢子囊之土壤懸浮液與病害發生之關係 74
討論 74
引用文獻 78
英文摘要 80
圖表 81
第四章莧菜白銹病之病勢進展 94
摘要 94
前言 94
材料與方法 95
供試莧菜品種與莧菜白銹病試驗區之規劃 95
大氣溫度與相對溼度之測定 95
莧菜生長與白銹病病勢進展之關係 96
莧菜連作田土壤中之莧菜白銹病初次感染源調查 96
莧菜白銹病第二次感染源密度調查 97
莧菜白銹病於植株上之分布與田間之空間分布 97
莧菜白銹病田間調查之最適取樣數分析 98
結果 98
大氣溫度與相對溼度對病害發生之分析 98
莧菜生長與白銹病病勢進展之關係 98
莧菜白銹病之病勢進展 99
莧菜連作田土壤中之莧菜白銹病初次感染源調查 100
莧菜白銹病第二次感染源密度調查 100
莧菜白銹病於植株上之分布與田間之空間分布 101
莧菜白銹病田間調查之最適取樣數分析 101
討論 101
引用文獻 104
英文摘要 107
圖表 108
第五章莧菜白銹病之防治 114
摘要 114
前言 114
材料與方法 115
室內莧白銹菌之藥劑篩選 115
田間不同殺菌劑防治莧菜白銹病之效果 116
田間莧菜白銹病之用藥時機試驗 116
隧道式覆蓋法防治莧菜白銹病之效果 116
界面活性劑十二烷基硫酸鈉 (sodium dodecyl sulfate) 防治莧菜白銹病之效果 117
結果 117
室內莧白銹菌之藥劑篩選 117
田間不同殺菌劑防治莧菜白銹病之效果 118
田間莧菜白銹病之用藥時機試驗 118
隧道式覆蓋法防治莧菜白銹病之效果 118
界面活性劑十二烷基硫酸鈉 (sodium dodecyl sulfate) 防治莧菜白銹病之效果 118
討論 119
引用文獻 121
英文摘要 124
圖表 125
第六章莧白銹菌之親緣性分析 129
摘要 129
前言 129
材料與方法 131
供試之白銹菌菌株來源 131
莧白銹菌與比對之白銹菌菌株之DNA純化與保存 131
以RAPD分析莧白銹菌之親緣性 132
以ITS分析莧白銹菌之親緣性 133
結果 134
以RAPD分析莧白銹菌之親緣性 134
以ITS分析莧白銹菌之親緣性 135
討論 135
引用文獻 136
英文摘要 138
圖表 139
附錄 149
討 論 153
引用文獻 158

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