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Eremothecium Coryli によるダイズ子実汚斑病（新称）

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Eremothecium Coryli によるダイズ子実汚斑病（新称） 日植病報 73： 283–288（2007）4 Jpn. J. Phytopathol. 73: 283–288 (2007) Eremothecium coryli によるダイズ子実汚斑病（新称） 木村 重光 1* ABSTRACT KIMURA, S. (2007). Yeast-spot disease of soybean caused by Eremothecium coryli (Peglion) Kurtzman in Japan. Jpn. J. Phythopathol. 73: 283–288. An ascomycetous, yeast-like fungus was isolated from lesions of soybean (cv. Murasakizukin) seeds that had been sucked by pentato- mid bugs (Hemiptera: Pentatomidae) in Kyoto prefecture. Based on morphological and physiological characteristics and sequence data of the internal transcribed spacer (ITS) regions including 5.8S rDNA, these yeast-like fungi were identified as Eremothecium coryli (Peglion) Kurtzman (syn. Nematospora coryli). After healthy, immature soybean seeds (cv. Maihime and Enrei) were inoculated with the isolated fungus, symptoms of the disease were reproduced, and the fungus was reisolated from the lesions. This fungus is widely known as causing a “yeast-spot disease” pathogen of soybean seeds, but has not previously been reported in Japan. Thus, the common name yeast spot (‘Shijitsu-ohan-byo’ in Japanese) is proposed for this disease of soybean. (Received October 23, 2006; Accepted February 28, 2007) Key words: Eremothecium coryli, shijitsu-ohan-byo, yeast-spot disease, soybean, pentatomid bugs れており，その地域も，南北アメリカ，ヨーロッパ，アジア， 緒 言 アフリカ，オーストラリアと全世界的である．本菌による病 京都府特産の黒大豆（品種：新丹波黒）は，丹波，丹後地 害は，Ashby and Nowell（1926）により“Stigmatomycosis”と 域を中心に約 260 ha 栽培されている．また近年，枝豆として いう病名が与えられているが Preston and Ray（1943）および 出荷し，ブランド京野菜「紫ずきん」として販売される黒大 Lehman（1943）がダイズから N. coryli を分離したそれぞれ 豆（品種：紫ずきんおよび新丹波黒）も約 50 ha に面積を拡 の報告において“yeast-spot disease”を病名に用いており， 大してきた．子実害虫の防除は莢伸長期から殺虫剤をスケ 以降ダイズにおいては“yeast spot”が用いられている．ま ジュール散布することで行っているが，その被害は決して少 た，本菌は，Daugherty（1967）によりカメムシ類によりダ ないとはいえない．特に，カメムシ類の吸汁による子実の奇 イズへ伝搬されることが明らかにされている． 形や陥没は，商品化率を低下させ大きな損失をもたらしてい 本研究では，観察された酵母様微生物と Nematospora 属菌 る．そのような被害粒を切断すると，子実内部が汚白色に変 との関係およびカメムシ類によるダイズ被害粒症状との関 色し，周囲に壊死を伴うもの（Fig. 1A）が見られる．変色し 係を明らかにするために，カメムシ類被害粒より酵母様微生 た被害部位切片の検鏡を行うと，紡錘形でむち状の付属糸を 物を単離し，これらの菌の形態，生化学的性質，5.8S rDNA 持つ多数の胞子と，円形または楕円形で出芽を認める酵母様 を含む ITS 領域の塩基配列解析等から酵母様微生物の同定 微生物が観察された．これらは，子嚢酵母である Nematospora を行った．また，酵母様微生物の同定後，接種により分離菌 coryli Peglion の記載（曾根田，1975）に類似しているように 株のダイズ子実に対する病原性と yeast-spot disease の症状 思われた．本邦での Nematospora 属菌に関する記録は，国内 について比較検討し，得た知見を報告する． 発生未詳であるがワタさく腐敗病（日本植物病理学会編， 材料および方法 2000）とインゲンマメからの分離（瀧元，1954）があるが， ダイズからの分離例はない．海外では，熱帯，亜熱帯植物種 病原菌の分離 2005 年 11 月に採取したダイズ（品種：紫 子の重要病原菌として，ワタ，ダイズ等 11 属で報告がなさ ずきん）の子実を 70％エタノールに数秒，続いて 1％次亜塩 1 京都府農業総合研究所（〒 621-0806 京都府亀岡市余部町和久成 9） Kyoto Prefectual Agricultural Research Institute, 9 Wakunari, Amarubecho, Kameoka, 621-0806 Kyoto, Japan * Corresponding author (E-mail:[email protected]) 本研究で明らかにした塩基配列は，GenBank/EmBL/DDBJ DNA データベースに Accession Number AB271231 および AB271232 とし て登録した． 284 日本植物病理学会報 第 73 巻 第 4 号 平成 19 年 11 月 素酸ナトリウム溶液に 1 分間浸漬し表面殺菌した．滅菌水で （SANYO 社製）を用いて，94°C 3 分間熱変成させた後，35 サ 十分洗浄した後，臍を中心にして長径と直交する方向に切断 イクル（熱変成 94°C，1 分間，アニーリング 52°C，1 分間， し，子葉断面に汚白色で黒点の散在する病徴のある子実 12 伸長反応 72°C，2 分間）行い，続いて 72°C で 7 分間の伸長 粒から菌の分離を行った．病変部から火炎滅菌したカミソリ 反応を行った．5.8S を含む ITS 領域の PCR プライマーには， で 5mm程度の切片を切り取り，ストレプトマイシン硫酸塩 White ら（1990）のユニバーサルプライマー ITS5 の 5' 末端 （Wako 社製）および，シクロヘキシミド（Sigma 社製）をそ から 5 塩基目の G を C に変えたものをフォワードプライマー れぞれ 100 ppm 添加したブドウ糖加用ジャガイモ煎汁寒天 （5'GGAACTAAAAGTCGTAACAAGG3'）に，リバースプライ （Difco 社製）（以下，PDA ＋ SC と略記）平板培地に置床し マーには ITS4（5'TCCTCCGCTTATTGATATGC3'）（White et て，24°C（暗黒下）で 7 日間の培養により分離株を得た．得 al., 1990）を用いた．得られた PCR 産物は，MicroconTM-100 られた菌株は，PDA 平板培地上で単コロニー分離を数回繰り （TaKaRa 社製）により精製し，PCR に用いた各プライマーと 返し，以下の実験に供した． DTCS Quick Start kit（BECKMAN COULTER 社製）を用いた 形態観察および生化学的性質の調査 分離菌株は，PDA 平 ダイレクトシーケンス法によって塩基配列を決定した．サイ 板培地および 10％麦芽寒天平板培地に画線後，24°C（暗黒 クルシーケンス反応は，GeneAmpR PCR System 9700（Applied 下）で 14 日間および 1ヶ月間培養し，コロニーの性状，栄養 Biosystem 社製）を用いて標準のプロトコルに従って行い， 細胞の形態の観察を行った．菌糸および分芽胞子の形成は， CEQTM 2000XL DNA Analysis System（BECKMAN COULTER コーンミール寒天平板培地上で Dalmau plate 培養（Dalmau, 社製）によってデータを回収した．塩基配列の解析は， 1929）を 24°C（暗黒下）で 7 日間行い観察した．また，5％ Genetix-win ver. 6.0.2（ソフトウェア開発社製）を用いて行っ 麦汁液体培地への接種培養を 24°C（暗黒下）で 3 日間行い， た．解析した配列データを基に，日本 DNA データバンク 皮膜の形成等液体培地での生育形態の観察を行った．分離菌 （DDBJ）が提供している相同性検索メニュー（BLAST）によ 株の生化学的性質は，API 20C AUX（BIOMERIEUX 社製）を り相同性検索を行い，類縁菌の 5.8S を含む ITS 領域の塩基 用い炭素源の同化性を，また，2％ブドウ糖，0.05％MgSO4・ 配列を DDBJ の配列検索メニュー（SRS）で取得した（Table 7H2O，0.1％KH2PO4 を含む基礎培地を用い，オキサノグラフ 1）．また，分離菌株の 5.8S を含む ITS 領域の塩基配列は，ア 法によって硝酸塩の同化性を試験（後藤，1975）した． クセッション番号によって Table 1 に示した．これらの配列 PCR 増幅および塩基配列の解析 分離菌株のうち NC1 と データにより近隣結合法による系統解析を行った． NC2 を PDA 斜面培地で 24°C（暗黒下），7 日間培養後，滅菌 接種試験 前述の試験を行い分離菌の同定を行った後，分 水 100 µl に懸濁し，懸濁液を遠心分離（15,000 × g，5 分間） 離菌株の病原性の確認およびその病徴を再現するため，次に した．上清を除去して菌体を収集し，Gen とるくん（TaKaRa より接種試験を行った．2006 年 4 月 22 日播種のダイズ（品 社製）を用いて，添付されたプロトコルのとおり抽出した 種：エンレイ）および 5 月 23 日播種の黒ダイズ（品種：舞 DNA を鋳型 DNA として PCR 法により 5.8S を含む ITS1 と 姫）を，それぞれ，1 プランター（23 × 72 × 23 cm）に 5 株 ITS2 領域を増幅した．50 µl の PCR 反応液には 1.0 µM の各 ずつ植え，ガラス室内で慣行に準じて管理した．エンレイの プライマーと1.25 ユニットの TaKaRa EX TaqTM（TaKaRa 社 開花盛期は 6 月 15 日，舞姫は 6 月 21 日であった．接種は 製），2.5 mM の各 dNTP および 2mMの Mg2+ を含む Ex taq Daugherty（1967）の方法によった．滅菌ピペットを用い莢 reaction buffer を加えた．PCR 反応は，DNA AMPLIFIER 上に酵母懸濁液を 1 滴落とし，アルコールランプで火炎滅菌 Table 1. Fungi used in this study and Genbank/EMBL/DDBJ accession numbers Accession numbers Species Isolate Locality Origin ITS and 5.8S rDNA Eremothecium coryli NC1 (=NBRC 102356) Kyoto, Japan Black soybean AB271231 〃 NC2 (=NBRC 102357) 〃〃AB271232 〃 NRRL Y-12970a) Italy Hazel nut AY046217 E. ashbyi NRRL Y-1363b) Unknown Unknown AY046220 E. gossypii NRRL Y-1056b) Unknown Unknown AY046216 E. sinecaudum NRRL Y-17231a) Canada Oriental mustard seed AY046218 E. cymbalariae NRRL Y-17582b) Iran Brachynema garmari AY046219 Aureobasidium pullulans Orange U. S. A Pinus sylvestris AF013229 a) Type strain b) Authentic strain. There is no known type strain. Jpn. J. Phytopathol. 73（4）． November, 2007 285 した昆虫針，微針（志賀昆蟲社製，以下微針）で懸濁液上か lactose，raffinose，melezitose，D-xylose，L-arabinose，N-acetyl- ら 2 ～ 3 回突き刺した．供試した酵母懸濁液は，分離菌株 D-glucosamine，glycerol，α-methyl-D-glucoside，inositol，2-keto- NC1 を Clarke and Wilde（1970）の Nutrient broth overlay 培地 D-gluconate の 14 炭素源の同化性が Eremothecium coryli (Peg- （8.8 g/L Nutrient broth（Difco 社製），5.6 g/L yeast extract，11 g/ lion) Kurtzman の記載（do Carmo-Sausa, 1970; de Hoog et al., L dextrose）100 ml に接種し，110 rpm，24°C（暗黒下）の条 2000）に一致した．これらの同化性は記載のない炭素源の結 件で 7 日間振とう培養した．接種には，2.13 × 1011 CFU/ml 果とともに Table 2 に示した．また，硝酸塩は利用しなかった． の酵母懸濁液（胞子を含む）を用いた．舞姫では，開花 23 以上の形態的特徴および生化学的性質について，分離 12 日後に，エンレイでは，開花 31 日後に接種を行い，その時 株の菌株間に差異を認めなかった． のダイズの生育は Fehr and Caviness（1977）による分類では 分離菌株の塩基配列の相同性 R6 であった．対照は，無菌 Nutrient broth 培地を滴下し，ア 分離菌株 NC1 と NC2 は，5.8S を含む ITS 領域の塩基配列 ルコールランプで火炎滅菌した微針での付傷のみ行ったも 解析の結果，580 塩基に対して 2 塩基の置換による相違が見 のおよび無処理とした． られたが，残り 578 塩基は相同であった．また，近隣結合法 による系統解析では，NC1 および NC2 は同一のクラスター 結 果 を形成し（Fig. 2），E. coryli に近縁であることが示唆された． 病原菌の分離と形態観察 NC1 および NC2 と E. coryli の 5.8S を含む ITS 領域の塩基配 PDA ＋ SC 平板培地に置床した子実切片からは，透白色で 列は 98.1％の相同性を示した． 盛り上がる糊状のコロニーが形成されたが，しだいに白色を 分離菌株の病原性 増し光沢ある酵母様のコロニーとなった．PDA 平板培地で単 接種 7 日後の舞姫は，子実への接種部が赤褐色に変色し， コロニー分離を繰り返し行い，12 株を分離した． 接種部を中心にして周辺部が水浸状に変色していた．しか PDA 平板培地上で 7 日間培養（24°C，暗黒下） し，接種部の切断面では赤褐色の変色は接種部に限られた． 生じたコロニーは白色～乳白色，軟質，滑らかでわずかに 接種 14 日及び 21 日後では，表面上の接種部赤褐変域は変わ 盛り上がり，外縁は細かい菌糸状であった．いずれの菌株も， らなかったが，周辺部の変色域は拡大していた．切断面は， 円筒形またはわずかに変形した子嚢：60.0 ～ 95.0 × 12.5 ～ 種皮部に赤褐色域が拡大していたが，子葉には変色はなかっ 20.0 µm（平均 75.5 × 16.1 µ m ）を形成し，紡錘形でむち状 た．切片を顕微鏡下で観察し，種皮柔組織と子葉表面の境界 の付属糸をもつ子嚢胞子：55.0 ～ 100.0 × 2.0 ～ 3.0 µ m（平均 に多数の酵母体および子嚢を確認した．登熟し収穫した舞姫 75.5 × 2.5 µ m）が確認された（Fig. 1B）．子嚢胞子は，子嚢 の子実表面は一部くぼみ，接種後水浸状に変色を認めた部分 内にある場合は，付属糸のない鋭角な側を外側に向け，2 束 は，種皮が黒色とならず着色不良となった．ほとんどの子実 で整列，通常 4 ～ 8 個が格納されていた（Fig. 1C）．酵母体 は，接種部位側の子葉の生育が停止し，扁平となるか，完全 は円形または卵形：5.0 × 2.5 ～ 25.0 × 25.0 µ m（平均 4.3 × に生育が停止し，しわやくぼみを伴った変形粒であった．切 10.6 µ m）で，多極性出芽による増殖であった（Fig. 1D）． 断面には種皮下柔組織の茶褐変を認めたが，子葉部には病徴 10％麦芽寒天培地での画線培養（24°C，1ヶ月間） を認めなかった．接種 38 日後のエンレイでは，種皮に褐色 生じたコロニーは乳白色で，光沢があり，糊状または膜状 から黒褐色の変色部が拡がり（Fig. 1F），その断面では接種 でわずかに隆起し，粗いひだが形成され，外縁には菌糸が形 部の種皮柔組織も褐変していた（Fig. 1G）．接種 50 日後に収 成された． 穫したエンレイにおいてもしわや陥没を伴い，舞姫と同様に コーンミール寒天平板培地上での Dalmau plate 培養（24°C， 正常な粒が少なかった．微針での刺傷のみの場合には，刺傷 7 日間） 部の種皮がわずかに変色した（Fig. 1H）が，変色部は拡大す 若干の隔膜を持つ真性菌糸が豊富に形成され，卵形または ることはなく，刺傷部に沿っての変色もなかった（Fig. 1I）． 円筒形の分芽胞子が確認された（Fig. 1E）． また，有傷接種のように微針刺傷側の子葉の生育が阻害され 5％麦汁液体培地への接種培養（24°C，3 日間） ることはなかった．いずれの処理も行わなかった場合では種 やや懸濁し白色の沈渣を生じたが，皮膜は生じなかった． 皮，子葉とも変色等は無かった（Fig. 1J, K）．NC1 株は接種 酵母体は単生または連鎖状となり，卵形であった． 後，ダイズが登熟し収穫するまで，種皮柔組織に沿って広が 生化学的性質 りをみせたが，子葉組織内部への侵入は見られず，病徴の拡 API 20C AUX（BIOMERIEUX 社製）を用いて供試した 20 大は極めて緩慢であった．これらは，Lehman（1943）の記 種の炭素源のうち D-glucose，maltose，cellobiose，trehalose， 載に一致した．有傷接種したいずれの子実からも接種菌が再 286 日本植物病理学会報 第 73 巻 第 4 号 平成 19 年 11 月 Fig. 1. Symptoms and causal yeast-like fungi of soybean “yeast-spot” disease A. Cross section of a soybean damaged by a pentatomide bug. B. Ascospores of Eremothecium coryli, NC1, viewed with phase-contrast (pc) microscopy after 14 days incubation on PDA. (scale bar: 10 µm) C. Ascus of E. coryli, NC1, pc micrograph, after 14 days incubation on PDA. (scale bar: 10 µm) D. Globose and budding yeast cell of E. coryli, NC1, pc micrograph, after 14 days incubation on PDA. (scale bar: 10 µm) E. True hyphae with few septa and ovid or cylindrical blastconidia of E. coryli, NC1, pc micrograph, after 14 days incubation on corn meal agar. (scale bar: 10 µm) F. Yeast-spot lesions on soybean (cv. Enrei), after inoculation with yeast suspension of E. coryli, NC1. (Arrows: inoculation points) G. Cross section through yeast-spot symptoms on soybean (cv. Enrei), after inoculation with yeast suspension of E. coryli, NC1. (Arrow: inoculation point) H. Whole soybean seed pierced with sterile needles only. (Arrows: pierced points) I. Cross section of a soybean pierced with sterile needles only. (Arrow: pierced point) J. Whole soybean seed without any treatment. K. Cross section of a soybean without any treatment. L. Yeast-spot disease symptom with necrotic area on the cotyledons. (Arrow: necrotic area) M. Yeast-spot symptom with chalky-white area in the cotyledon. (Arrow: chalky-white area) Jpn.
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