臺灣博碩士論文加值系統

近年來，百香果產業在台灣蓬勃發展而栽培面積也逐年增加，以區域共同防治和健康種苗方式解決病毒病害對產業的威脅後，真菌性病害在栽培上造成的危害逐漸受到注意，因此進行百香果上之病害調查和鑑定，以預防潛在病害對百香果栽培上造成之產量損失。於此對3個百香果新發生之病害進行描述，包含未知的黃暈葉斑、褐色葉斑及白粉病之病徵。黃暈葉斑之樣本採集自南投和台中，病原菌在葉面造成淺褐色或白色之壞疽斑，並於其上形成柄子殼，高濕環境下黃白色孢子堆呈長鬚狀擠出，病原菌孢子為絲狀或桿狀，具有1-3隔，藉由形態特徵鑑定其為Septoria屬。利用TEF1-α和β-tubulin基因進行親緣分析，鑑定為Septoria passifloricola，為台灣新報導病害，依病徵和病原菌種類將其命名為白星病。褐色葉斑菌株收集自台中、南投、彰化和屏東，共分離得33株菌株，依據形態特徵鑑定病原菌為Alternaria屬，為已紀錄之褐斑病，分離菌株分別屬於Alternaria屬之A. porri section或A. alternaria section兩類群的親緣種，再各別增幅Alt a1、GAPDH、ITS、TEF、RPB2、OPA10-2等基因進行親緣分析，鑑定有4個支序群的Alternaria可造成病害，包含A. alternata、A. jacinthicola及2種可能之新種。白粉病菌樣本收集自台中，孢子形態為鏈生，內不具有fibrosin bodies，發芽管呈棍棒狀，菌絲上之附著器呈乳突狀，以形態特徵初步將菌株鑑定為Golovinomyces屬，並以ITS序列特徵鑑定屬G. ambrosiae/G. circumfusus/G. spadiceus complex之一員，之後比對孢子長寬比等形態特徵鑑定為G. circumfusus。經科霍氏法則確認各菌株之病原性，證明白星病可感染葉片及果實，惟孢子發芽須97%以上的高相對溼度，此外發現S. passifloricola在液體培養中會經微循環產孢以出芽形式產生許多孢子，此類型孢子毒力與來自固體培養基的分生孢子毒力相當，而液體培養可獲得的孢子相對穩定且大量。褐斑病菌亦可感染葉片及果實，其中E clade的菌株對葉片有較高的毒力，其餘菌株僅造成細小病斑；相對於葉片，各病原菌皆可於果實造成明顯的褐斑。田間接種褐斑病菌於百香果之各發育生長期，褐斑病菌可感染各種花器組織及各生育期之果實，其中小果期感染可能造成果實畸形的病徵；白粉病可感染黃百香果的苗期植株，於子葉、新葉和莖部造成病徵，並可跟大花咸豐草進行交互感染。為提供化學藥劑之防治建議，對白星病菌和褐斑病菌進行藥劑篩選，前者以百克敏和三氟派瑞在有效成分1 ppm時即能達到100% 孢子發芽抑制；後者以三氟派瑞、亞托待克立、得克立、賽普護汰寧和克熱淨在100 ppm有效成分時能對5個clade的菌株達到80%以上的菌絲生長抑制，其中500 ppm得克立和100 ppm克熱淨可完全抑制菌絲生長，而E clade菌株對藥劑感受性相對其他4個類群則較高。此外於南投魚池遮雨設施和露天之百香果園進行病害調查，發現以遮雨設施種植之植株褐斑病、疫病較露天種植者罹病率較低。本研究針對台灣未紀錄之百香果病害進行描述、鑑定和藥劑篩選，期能提供資訊做為擬定防治策略之參考。

In recent years, passion fruit becomes a popular crop and its plantation area is increasing in Taiwan. The threat of viral diseases to the industry was overcome by the conduction of areawide pest management with healthy seedlings. The threat of fungal diseases was increasingly noticed. Diseases of passion fruit were investigated and identified to prevent the potential threats on production. Here, we characterized three new diseases in detail: yellow halo leaf spots, brown spots, and powdery mildew. The pathogens of the yellow halo leaf spots were isolated from Nantou and Taichung. The fungal pathogens formed pycnidia on light brown lesions or white leaf spots. The yellowish white conidial heap was extruded in long tendrils from pycnidia in high humid condition. Conidia were filiform to cylindrical with 1-3 septa. The pathogens were identified as a species of Septoria based on morphological characteristics. Phylogenetic analysis identified the pathogens as Septoria passifloricola, a first recored fungus in Taiwan, based on sequences of TEF1-α and β-tubulin genes. According to the symptoms and pathogens, the new disease was named Septoria blotch. In the study of brown spots, 33 isolates were collected from Nantou, Taichung, Changhua and Pingtung. According to the morphological characteristics, these isolates were grouped to either the A. porri section or A. alternaria section of Alternaria genus. The molecular identification was performed by amplification of Alt a1, GAPDH, ITS, TEF, RPB2 and OPA10-2 genes for multi-genes phylogenetic tree analysis. The molecular analysis clustered pathogenic isolates into 4 clades, including A. alternata, A. jacinthicola and 2 inconclusive groups which possibly belonged to new species. The powdery mildew specimens were collected fromTaichung. The conidia of powdery mildew were catenescent, and did not have fibrosin bodies. The germinated tubes were club-shaped and the appressoria on the hyphae were nipple-shaped. The strains were identified as Golovinomyces genus by morphological characteristics. Molecular identification based on ITS sequences concluded that the strains were of G. ambrosiae/G. circumfusus/G. spadiceus complex. Later, the strains were identified as G. circumfusus according to the ratio of conidia length and width and other morphological characteristics. The pathogenicity of causal agents of the three new diseases were proved by the Koch's postulates. Septoria blotch could infect leaves and fruits, but the spores germinated required a high relative humidity of 97% or more. In addition, we found that S. passifloricola could produce a number of budding conidia through microcycle conidiation in liquid culture. The virulence of budding conidia was comparable to that of conidia from the solid medium. Notably, the sporulation in liquid culture was relatively stable and large in amount. The brown spot pathogen could also infect leaves and fruits. The strains of E clade had high virulence to the leaves while the other strains only cause small lesions. Compared with lesions on leaves, more obvious brown spots on fruits were caused by each clade of pathogens. Field inoculation with brown spot pathogens on different growth stages of passion fruit, indicating that the pathogens were able to infect allorgans of flower and stages of developing fruits. The infection on small developing fruit stage might cause fruit malformation. Powdery mildew infected cotyledons, true leaves and stems of seedlings of yellow passion fruit. Cross-infection with G. circumfusus between Bidens pilosa var. radiata and P. edulis f. flavicarpa wassuccessfully performed. In order to recommend chemical control strategies, fungicide assays on pathogens of brown spots and Septoria blotch were carried out in this study. Fungicides such as pyraclostrobin, fluopyram + trifloxystrobin were effective to inhibit 100% spore germination of Septoria blotch isolates at 100 ppm. Fungicides such as Fluopyram + Trifloxystrobin, Azoxystrobin + Difenoconazole, Tebuconazole, Cyprodinil + Fludioxonil and Iminoctadine triacetate were effective to inhibit 80% mycelial growth of 5 clades isolates of Alternaria brown spot disease. Isolates from E clade were more sensitive to tested fungicides. Moreover, a disease investigation was conducted in the shading and open field passion fruit orchards in Nantou Yuchi, and it was found that the shading facility effectively reduced the diseases incidence of Alternaria brown spot and Phytophthora blight. This study described unrecorded passion fruit diseases in Taiwan, identified pathogens, and screened fungicides in order to provide disease information for developing control strategies.

中文摘要 i
Abstract iii
表次索引 viii
圖次索引 ix
前言 1
前人研究 3
一、台灣百香果產業現況 3
二、台灣百香果種植之品種 4
三、台灣與國外紀錄之百香果病害 6
百香果白星病 (Septoria blotch) 8
百香果褐斑病 (Alternaria brown spot) 9
百香果白粉病 (Powdery mildew) 11
材料與方法 12
一、供試菌株之來源、分離和保存 12
二、病原菌之接種 12
(1) 白星病 (Septoria blotch) 菌之病原性測試 12
(2) 褐斑病 (Alternaria brown spot) 菌之病原性測試及各生育期果實之抗感病測試 13
(3) 百香果、咸豐草白粉病之病原性測試 14
三、病原菌形態鑑定 14
四、供試菌株之genomic DNA萃取 15
五、基因片段之核甘酸序列增幅與定序 16
六、基因片段之核甘酸序列親緣演化分析 16
七、溫度對病原菌株孢子發芽 (菌絲生長) 之影響 17
八、相對溼度對白星病分離菌株孢子發芽之影響 18
九、白星病菌株於液態培養之產孢量與病原性測試 18
十、病原菌對殺菌劑之感受性 19
十一、病害調查 20
十二、統計分析 21
結果 22
(一) 白星病 22
(1) 菌株蒐集、孢子及菌落形態特徵 22
(2) 分子鑑定 22
(3) 病原性測試 23
(4) 溫度、相對溼度對病原菌孢子發芽之影響 23
(5) 液態培養之產孢量、病原性及葉片接種部位比較 23
(6) 殺菌劑對Septoria passifloricola孢子發芽之影響 24
(二) 褐斑病 25
(1) 菌株蒐集、菌絲形態及分子鑑定 25
(2) 葉部及果實病原性測試 26
(3) 溫度與分離菌株菌絲生長關係 27
(4) 百香果生育期感病性試驗 27
(5) 分離菌株之菌絲對殺菌劑感受性 28
(三) 百香果苗期白粉病 29
(1) 病原菌造成之病徵及分子鑑定 29
(2) 形態鑑定 30
(3) 百香果及咸豐草白粉病之病原性 31
(四) 田間設施及灌溉系統對病害發生之影響 31
(1) 設施及灌溉方式對葉及未成熟青果的病害發生與影響 31
(2) 設施及灌溉方式對成熟落果的病害發生影響 32
討論 33
參考文獻 42
附錄 85

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