Please use this identifier to cite or link to this item: http://hdl.handle.net/11455/89353
標題: 內生菌根絲核菌的生物多樣性與菌根共生研究
Biodiversity and mycorrhizal symbiosis of endomycorrhizal Rhizoctonia fungi in mycoheterotrophic orchids
作者: Jr-Hau Jiang
蔣志豪
關鍵字: 形式屬絲核菌
菌根共生
真菌纏據
真菌-植物共生界面
植物保護
form-genus Rhizoctonia
mycorrhizal symbiosis
fungal colonization
fungal-plant interface
plant protection
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摘要: 真菌異營蘭花可分成綠色及非綠色蘭花,前者的菌根真菌多屬於形式屬絲核菌(form-genus Rhizoctonia)。藉由菌絲形態、隔膜孔構造、融合群以及核醣體DNA之內轉錄區間等鑑定方法,顯示分離自本島蘭花菌根處的內生菌根絲核菌包含Thanatephorus、Tulasnella及Ceratobasidium等真菌。Tulasnella真菌主要分離自樹蘭亞科的蘭花菌根,包含T. calospora、T. danica及二種未知的新種。Ceratobasidium真菌則分離自蘭亞科及樹蘭亞科之菌根中獲得,包含AG-A、AG-B、AG-G、AG-P及AG-R等融合群。除AG-A外,其他融合群皆是首次被發現與蘭花菌根共生相關。本研究證實同一區域內綠色蘭花之內生菌根絲核菌種類具多樣性。此外調查台灣各地金線連的菌根真菌,顯示Rhizoctonia solani AG-6廣泛地存在於台灣金線連菌根中,而AG-G、AG-P及AG-R也可在野生台灣金線連菌根中發現,顯示不同區域生長的台灣金線連也有多樣的內生菌根絲核菌。研究顯示支序群I內的AG-6、AG-R及AG-P分離株能提高台灣金線連種子44–91%的萌發率,並促進圓球體生長至階段III – VI,反之非共生種子萌發的對照組萌發率僅13%。種子活力程度可能與共生結果偏向菌根共生或寄生有關。然而支序群I – III內的所有供試菌株,皆能與台灣金線連組織培養小苗形成菌根共生並顯著增加植體鮮種,顯示金線連種子的菌根專一性與組織培養苗不同。除了AG-R分離株外,異地栽培(ex vitro)台灣金線連顯示接種支序群I內的內生菌根絲核菌均能顯著地提高植株鮮種,而該群對試管內的台灣金線連組織培養小苗僅能造成低程度的真菌纏據(p ̂_i = 0.30 – 0.47)。中高程度的真菌纏據可來自接種支序群II (p ̂_i = 0.63 – 0.82)及III (p ̂_i = 0.63 – 0.75)內的分離株,並且本研究首次嘗試以菌根解剖的序數資料證實蘭花菌根真菌的纏據程度與植株鮮種呈現負相關(r = -0.8801)。 接種內生菌根絲核菌的台灣金線連真菌根莖處之切片可區分為消化區與非消化區。以電子顯微鏡觀察真菌根莖處非消化區內的真菌-植物共生界面,顯示至少存在三種類型。類型I與類型II主要由一層源自植物的低密度電子透明層及一層高密度的電子不透明層所組成,而類型II幾乎由高密度電子層組成。而類型III的解剖特徵為首次發現,包含二層厚度不均的高密度電子不透明層及二層低密度電子透明層所組成,共生界面常可見條狀缺口或紡錘形的透化區。共生界面可以被單株抗體JIM 5標幟非酯化的同聚半乳糖醛酸聚糖,液泡化的菌絲及崩解菌絲之共生界面有相對較多的金屬金沉澱,顯示非酯化果膠物質可能與菌絲活性有關連。而單株抗體JIM 7僅能標誌植物細胞間的中膠層。所有的共生界面均可被單元抗體CCRC-M26偵測到β-(1→3) / (1→6)-葡聚醣。 此外本次研究首次以融合群為基礎探討內生菌根絲核菌的毒力與病原性,並證實內生菌根絲核菌AG-6、AG-A、AG-B、AG-G、AG-P以及Tulasnalla分離株,均表現較低的罹病程度,其相對處理效應數值為蘿蔔(p ̂_i = 0.1 – 0.61)、胡瓜(p ̂_i = 0.28 – 0.54)及小白菜(p ̂_i = 0.18 – 0.65),均不會造成10天大的受試植物表現典型猝倒病徵而死亡。而病原AG-4幾乎造成所有受試植物下胚軸崩解倒伏及子葉萎凋等病徵(p ̂_i = 0.88 – 0.96)。這些已知融合群的內生菌根絲核菌均與已發表可保護植物並且為弱毒力絲核菌的分離株為相同融合群。試驗結果顯示若預先接種內生菌根絲核菌AG-P分離株6天後再接種AG-4病原,對26天大的小白菜則具有最高的植物保護率(91% – 100%)。因此內生菌根絲核菌除能應用在蘭花的保育與生產外,亦有潛力開發成有用的生物防治製劑應用於農作物的栽培。
Orchids can be divided into non-green (myco-heterotrophic) and green (mixotrophic) orchids. Green orchids have mycorrhizal associations with the form genus Rhizoctonia. With phylogenetic analysis and anastomosis group determination, mycorrhizal fungi in our study belonged to Thanatephorus, Tulasnella and Ceratobasidium. Tulasnella fungi distributed mainly in Epidendroideae, including T. calospora, T. danica and two unknown isolates. Ceratobasidium fungi (i.e., AG-A, AG-B, AG-G, AG-P and AG-R) were isolated from terrestrial orchids. Except for AG-A, 4 AGs of Ceratobasidium were found the first occurrence in Orchidaceae. Our results suggested that green orchids in the same geographic region associate with a diversity of Rhizoctonia groups. Furthermore, Thanatephorus (AG-6) and Ceratobasidium (AG-G, P, and R) were also identified in native A. formosanus from different geographic regions, suggesting that single orchid species exhibits diverse associations with Rhizoctonia fungi. Isolates of AG-6, AG-R and AG-P in Clade I increased seed germination 44–91% and promoted protocorm growth from phase III to VI compared to asymbiotic treatments (13%) and isolates in Clade II and III. Different levels of seed viability may affect the result of mycorrhizal symbiosis: a mycorrhizal interaction or a parasitic interaction. However, all isolates in Clades I to III formed fungal pelotons in tissue-cultured seedlings of A. formosanus. An analysis of the relative effect of treatment (p ̂_i) showed that the low level of colonization (p ̂_i = 0.30 – 0.47) by isolates in Clade I resulted in a significant increase in seedling growth compared to isolates in Clades II (0.63 – 0.82) and III (0.63 – 0.75) which caused moderate to high colonization in plant. There was also a negative correlation (r = -0.8801) with fresh plant weight and fungal colonization. Sections of A. formosanus mycorrhizome consisted of a non-digested zone and a digested zone. Both type I and II of symbiotic interface consisted of an electron-lucent layer and an electron-opaque layer; however, type II exhibited much thicker electron-opaque layer than type I. However, type III consisted of two electron-lucent layers and two electron-opaque layers with occasional nicks, which is first occurrence in orchid mycorrhizae. Pectin substances have been proved at symbiotic interface by using innunogld labelling for un-esterified homogalacturonan (HG). Un-esterified HG was recognized heavily at the interface of vacuolar hyphae. It seems that fungal activity is associated with the presence of pectin substances. However, methyl-esterified HG was only found in plant cell wall and middle lamella, and β-(1→3) / (1→6)-glucan was only found in fungal cell wall, especially at dolipore septum. Evaluating the virulence of endomycorrhizal Rhizoctonia fungi is important for the purposes on biological control against plant disease. All isolates, excepting the AG-R isolate, did not cause seedling death with low values of relative treatment effects for disease severity in 10-day-old radish (0.1 – 0.61), cucumber (0.28 – 0.54), and Chinese mustard (0.18 – 0.65); however, pathogenic AG-4 isolates resulted almost all the test plants died (0.88 – 0.96). AG-P isolate Cno10-3 and CalS1-2 exhibited 91% and 100% of plant protection with 0.14 and 0.20 of relative treatment effects for disease severity in 26-day-old Chinese mustard grown in soil. The results presented here are potentially useful for advancing research on the medicinal properties, production and conservation of orchid in diverse ecosystems. Moreover, endomycorrhizal Rhizoctonia may have the potential for biological control, and that isolates of AG-P successfully controlled the damping-off of Chinese mustard.
URI: http://hdl.handle.net/11455/89353
文章公開時間: 2016-12-03
Appears in Collections:植物病理學系

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