ウイルス 152 図1 第 62 巻 第 2 号 カブモザイクウイルスの (A) カブと (B) ダイコンでのモザイク症状 的研究は早くから開始され 現在では地球規模で発生して ポティウイルス属のウイルス種では 一部は新興集団と いる分離株ゲノムの塩基配列が網羅的にまた継続的に調べ しての報告もある
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1 62 2 pp RNA 7250 Turnip mosaic virus, TuMV TuMV TEL: FAX: ohshimak@cc.saga-u.ac.jp Turnip mosaic virus TuMV 1,2) TuMV 3-7) 20
2 ウイルス 152 図1 第 62 巻 第 2 号 カブモザイクウイルスの (A) カブと (B) ダイコンでのモザイク症状 的研究は早くから開始され 現在では地球規模で発生して ポティウイルス属のウイルス種では 一部は新興集団と いる分離株ゲノムの塩基配列が網羅的にまた継続的に調べ しての報告もあるが それよりも昔から農作物に大きな被 られ 現在ではおおよそ地球上に存在するゲノム 遺伝子 害を及ぼし続けているために 世界中の多くの研究者が長 型が分かっているのではと思われる その報告の数は 我々 年研究してきたウイルスである と言っても その研究の 植物ウイルスの研究者から見て すさまじい進展を感じる 歴史は一世紀にも満たない 1920 年の後半 イギリスの しかしながら それらの研究を追いかけるように幾つかの Kenneth Smith11) がジャガイモのウイルス 後にジャガイ 植物ウイルス分離株も国規模 大陸規模 地球規模で採集 モ Y ウイルス Potato virus Y PVY と呼ばれるが こ されるようになり ゲノムの一部や全領域について解析さ のウイルスと類似したウイルス種が研究されて以来 1971 れ 分子進化の 3 大推進力 突然変異 組換え 再集合 年にそれらを Potyvirus ポティウイルス と呼ぶように に関連した研究や集団遺伝構造の研究が開始しされ始めた なった 12,13) その後ポティウイルス属となり 1989 年に ただ多くの場合 動物ウイルスに比べて植物ウイルス分離 PVY のゲノム構造が最初に明らかになったのち 14) ここ 株を保存している農業関連機関が少ないために 分離株を 20 年間くらい それらポティウイルスのゲノムの多様性 集めることにはとても苦労する 多くの場合 研究者が様々 が深く探られるようになった ポティウイルス科には 8 属 な国々に出かけて田畑を走り回り 分離株を採集して集め あるが おそらくポティウイルスは 90% のポティウイル てくるのが実情であるので ウイルスを採集するだけでも ス科のウイルス種をカバーする またポティウイルスの中 長期間の仕事となる で 最も分子進化的に解析が進んでいる我々の研究してい 現在 分子進化的研究が進展しているウイルスを挙げる る TuMV15) は 主に世界中のアブラナ科のアブラナ属や とすれば 世界の農作物に大きな被害を与えている新興ウ ダイコン属植物の野菜類などの農作物に大きな被害を与え イルスである DNA ウイルスのジェミニウイルス科ベゴモ ているウイルス種である ウイルス属のトマト黄化葉巻ウイルス Tomato yellow leaf curl virus TYLCV であろう 8) 世界中のトマトや トルコギキョウなどに大きな被害を与えており 日本でも 発生がみられる 一方 RNA ウイルスとしては 1 本鎖 RNA ウイルスでアフリカのイネに大きな被害を与えているソベモ ウイルス属の Rice yellow mottle virus RYMV 9-10) の研究 が精力的に行われている 本ウイルスは 我が国では発生 が確認されておらず アフリカに発生が限られるエンデ ミックなウイルスである そして最も研究が進んでいるウ イルスを挙げるとすれば 我々の手掛けている TuMV と 思われる 本ウイルスは ポティウイルス科ポティウイル ス属のウイルス種である カブモザイクウイルスの分子進化 TuMV は アフリカ アジア ヨーロッパ オセアニア 南北アメリカなどの温帯 亜熱帯など世界中に広く分布し ており そして最近ではアラスカでも分離された世界中ど こにでもあるパンデミックなウイルスである 16) そのよ うなウイルスの進化を調査することは 本ウイルスの空間 的や時間的な情報だけでなく 他のウイルス進化や拡散に ついても多くのヒントを与えてくれる 1921 年に本ウイ ルスがアメリカで初記載され 2,17,18) その後日本などのア ジア諸国でも発見された 当初 野菜や園芸植物の病徴に 基 づ い て 様 々 な ウ イ ル ス 名 が つ け ら れ た 例 え ば
3 153 pp 図2 カブモザイクウイルスの分子系統樹の略図 Anemone mosaic virus Cabbage A virus Cabbage black NIb 遺伝子は複製に CP 遺伝子はアブラムシの伝搬性や ring virus Daikon mosaic virus Horseradish mosaic ウイルス粒子の会合に関与している 4,7) 最近 オーバーラッ virus などがその代表でもある 本ウイルスはアブラムシ ピングした pretty interesting Potyviridae ORF PIPO P3 で非永続的に伝播し 野菜の重要病害ウイルスとしてキュ 遺伝子上の 2 の読み枠に存在 が見つかった 21) この ウリモザイクウイルスに次いで 2 番目にランクされており 1) PIPO は 主に TuMV の我々の塩基配列データを用いて解 名前からもわかるようにカブは勿論のこと ダイコン キャ 析され明らかとなったが 実は我々も事前に気が付いてお ベツ ブロッコリー カリフラワー ナタネ カラシナ り研究を進めていたのだが 残念ながら先に論文にするこ ハクサイ チンゲンサイなどの中国野菜などのアブラナ科 とができなかった 以上は TuMV だけではなくポティウ 植物などにモザイク症状 えそ症状 時には黄色斑点など イルス一般のゲノム構造であるが ポティウイルスのタイ を呈する 図 1 またアブラナ科植物以外の植物 例え プ種としては ジャガイモやタバコなどのナス科植物に感 ばユリ科やキンポウゲ科植物などにも感染し 15) ポティ 染し大きな被害を与えている PVY が挙げられる 14,22-25) ウイルスの中でも特に広い宿主域を持つことが知られ 最 動物ウイルスは各国の大学や保健機関に過去のそして現 近ではバイオ燃料を生産する植物に被害を起こすために注 在流行している分離株が大量に保存されている 一方本ウ 目されているウイルスでもある イルスも他の多くの植物ウイルスと同様に 分離株を保存 TuMV は約 720nm の粒子で そのゲノムは一本鎖プラ ス RNA で約 9,830 塩基から構成されている している大学や農業関係の機関は少なく国内外から収集す このゲ るのが困難であった それでも世界の多数の研究者の協力 ノムから大きなポリタンパク質が翻訳され 第 1 タンパク を得て 主に我が国を含めたユーラシア大陸からの分離株 19,20) 質 P1 ヘルパー成分プロテアーゼタンパク質 HC-Pro であるが 現在まで数百分離株を採集してきており これ さらに核内封入体 a- タンパク質 NIa-Pro によりプロセッ までトルコ イラン ギリシャそして中国など延距離数にし シ ン グ さ れ 最 低 10 種 類 の 成 熟 し た タ ン パ ク 質 P1 て数万キロ以上を車や徒歩で圃場を回り調査してきた 26,27) HC-Pro 第 3 タンパク質 P3 6 キロダルトン 1 タンパ 研究を始めた当初は 地域を考慮し TuMV 76 分離株を ク質 6K1 筒状封入体タンパク質 CI 6 キロダルト 選抜し ゲノムの両末端に位置する二遺伝子 即ち最も変 ン 2 タンパク質 6K2 ゲノム結合タンパク質 VPg 異が多い遺伝子と考えられている P1 遺伝子と その時点 NIa-Pro 核内封入体 b タンパク質 NIb さらに外被タ で最も研究が進展している CP 遺伝子 各遺伝子約 1,000 ンパク質 CP が産生されると考えられている それぞ 塩基 ゲノムの 20% について塩基配列を決定後 分子 れの遺伝子は様々な役割をしているが 例えば HC-Pro 遺 系統関係を描いた 15) 研究を開始した頃の 1990 年代後半 伝子は アブラムシの伝搬性やサイレンシングの抑制に の時代では 76 分離株について 一ゲノム当り 2000 塩基
4 GenBank TuMV PCR 4 Orchis Brassica basal-b; basal-brassica world-b; world-brassica Raphanus Asian-BR; Asian-Brassica/ Raphanus basal-br; basal- Brassica/Raphanus PHYLPRO 28) SISCAN 29) TuMV 40 30)
5 pp BR Brassica/Raphanus 90 30% PHYLPRO 28) SISCAN 29) RDP3 31) 70 32) TuMV 100 P1 CI C VPg 2 33) ) P1 Asian-BR / TuMV ) DNASP 36) ARLEQUIN 37) 2 TuMV basal-br 2000 TuMV 38) 2 39) basal-br basal-br basal-br basal-br 30 DNASP 36 3
6 ) TuMV TuMV 41) Orchis 42) Orchis Orchis militaris O. morio O. simian TuMV TuMV TuMV JYMV Japanese yam mosaic virus 43,44 Scallion mosaic virus ScMV 45) NYSV Narcissus yellow stripe virus 46) TuMV TuMV TuMV Eruca sativa Cameria sativa TuMV PVY Pepper mottle virus PepMoV Peru tomato mosaic virus PTMV PVY Bean common mosaic virus BCMV 6,13,47) Soybean mosaic virus SMV 6,13,48) Peanut mottle virus PMV 6,13 TuMV JYMV ScMV NYSV Narcissus late season yellows virus 49) TuMV TuMV 1 Onion yellow dwarf virus OYDV 6,13 3 OYDV Shallot yellow stripe virus Narcissus degeneration virus Sugarcane mosaic virus SCMV 6,13 Cocksfoot streak virus Johnsongrass mosaic virus Pennisetum mosaic virus Holmes BEAST Bayesian evolutionary analysis by sampling trees 50-52) 53-58) 59-61) CP coherently evolving ccp ,63) TuMV BEAST ) 8 ICTV
7 pp Brambyvirus 1 Bymovirus Polymyxa graminis Ipomovirus Macluravirus Poacevirus 1 Potyvirus 30 1 Rymovirus eriophyid 1 Tritimovirus eriophyid Brambyvirus 1 Bymovirus 6 Ipomovirus 5 Macluravirus 6 Poacevirus 2 Rymovirus 3 Tritimovirus 4 2 Rymovirus Agropysum mosaic virus Hordeum mosaic virus Ryegrass mosaic virus Rymovirus 64,65) Adrian Gibbs 66,67) 68-70) 71,72) Huy Duc Nguyen Adrian J. Gibbs John A. Walsh
8 Walsh JA and Jenner CE: Turnip mosaic virus and the quest for durable resistance. Mol Plant Pathol 3: , Ohshima K: Turnip mosaic virus. In Characterization, Diagnosis & Management of Plant Viruses. G. P. Rao, P. L. Kumar, Holguin-Pena, R. J. eds., Studium Press LLC, Houston, Texas, USA, p313-30, Revers F, Le Gall O, Candresse T and Maule AJ: New advances in understanding the molecular biology of plant/potyvirus interactions. Mol Plant-Microbe Interact 12: , Riechmann JL, Lain S, Garcia JA: Highlights and prospects of potyvirus molecular biology. J Gen Virol 73: 1 16, Shukla DD, Ward CW and Brunt AA: The Potyviridae. Wallingford, UK: CABI 516 pp Shukla DD, Ward CW, Brunt AA and Berger PH: Potyviridae family. In AAB Descriptions of Plant Viruses. dpvno =366, Urcuqui-Inchima S, Haenni AL and Bernardi F: Potyvirus proteins: a wealth of functions. Virus Res 74: , Abhary M, Patil B and Fauquet C: Molecular biodiversity, taxonomy, and nomenclature of tomato yellow leaf curl-like viruses. Czosnek H, ed. In Tomato yellow leaf curl virus disease. pp , Fargette D, Pinel-Galzi A, S r m D, Lacombe S, H brard E, Traor O amd Konat G: Diversification of Rice yellow mottle virus and related viruses spans the history of agriculture from the neolithic to the present. PLoS Pathog 15: e , Fargette D, Pinel A, Rakotomalala M, Sangu E, Traor O, S r m D, Sorho F, Issaka S, H brard E, S r Y, Kanyeka Z and Konat G: Rice yellow mottle virus, an RNA plant virus, evolves as rapidly as most RNA animal viruses. J Virol 82: , Smith KM: On the composite nature of certain potato virus diseases of the mosaic group as revealed by the use of plant indicators and selective methods of transmission. Proc R Soc B;109: 251, Harrison BD, Finch JT, Gibbs AJ, Hollings M, Shepherd RJ, Valenta V and Wetter C: Sixteen groups of plant viruses. Virology 45: , King AMQ, Adams MJ, Carstens EB and Lefkowitz EJ: Virus taxonomy: Classification and nomenclature of viruses. Ninth report of the International Committee on Taxonomy of Viruses. San Diego: Elsevier/Academic Press pp, Robaglia C, Durand-Tardif M, Tronchet M, Boudazin G, Astier-Manifacier S and Casse-Delbart F: Nucleotide sequence of potato virus Y (N Strain) genomic RNA. J Gen Virol 70: , Ohshima K, Yamaguchi Y, Hirota R, Hamamoto T, Tomimura K Tan Z, Sano T, Azuhata F, Walsh JA, Fletcher J, Chen J, Gera A and Gibbs AJ: Molecular evolution of Turnip mosaic virus ; evidence of host adaptation, genetic recombination and geographical spread. J Gen Virol 83: , Robertson NL and Ianson DC: First report of Turnip mosaic virus in rhubarb in Alaska. Plant Dis 89: 430, Gardner MW and Kendrick JB: Turnip mosaic. J Agric Res 22: 123 4, Brunt AA, Crabtree K, Dallwitz MJ, Gibbs AJ, Watson L and Zurcher EJ: Plant Viruses Online: Descriptions and Lists from the VIDE Database. Ver. 20 th, URL 19 Nicolas O and Lalibert JF: The complete nucleotide sequence of turnip mosaic potyvirus RNA. J Gen Virol 73: , Ohshima K, Tanaka M and Sako N: The complete nucleotide sequence of Turnip mosaic virus RNA Japanese strain. Arch Virol 141: , Chung BY, Miller WA, Atkins JF and Firth AE: An overlapping essential gene in the Potyviridae. Proc Natl Acad Sci U S A 105: , Ohshima K, Sako K, Hiraishi C, Nakagawa A, Matsuo K, Ogawa T, Shikata E and Sako N: Potato Tuber Necrotic Ringspot Disease Occurring in Japan: Its Association with Potato virus Y Necrotic Strain. Plant Dis 10: , Ogawa T, Tomitaka Y, Nakagawa A and Ohshima K: Genetic structure of a population of Potato virus Y inducing potato tuber necrotic ringspot disease in Japan; comparison with North American and European populations. Virus Res 131, , Ogawa T, Nakagawa A, Hataya T and Ohshima K: The genetic structure of populations of Potato virus Y in Japan; based on the analysis of 20 full genomic sequences. J Phytopath 160, , Ward CW, Weiller G, Shukla DD and Gibbs AJ: Molecular systematics of the Potyviridae, the largest plant virus family. In Molecular Basis of Virus Evolution. Gibbs AJ, Calisher CH, Garc a-arenal H, editors. Cambridge: Cambridge University Press. pp , Korkmaz S, Tomitaka Y, Onder S and Ohshima K: Occurrence and molecular characterization of Turkish isolates of Turnip mosaic virus. Plant Pathol 57: , Farzadfar S, Tomitaka Y, Ikematsu M, Golnaraghi A, Pourrahim R and Ohshima K: Molecular characterisation of Turnip mosaic virus isolates from Brassicaceae weeds. Eur J Pl Pathol 124: 45-55, Weiller GF: Phylogenetic profiles: a graphical method for detecting genetic recombinations in homologous sequences. Mol Biol Evol 15: , Gibbs MJ, Armstrong JS and Gibbs AJ: Sister-scanning: a Monte Carlo procedure for assessing signals in recombinant sequences. Bioinformatics 16: , Tomimura K, Gibbs AJ, Jenner CE, Walsh JA and
9 pp Ohshima K: The phylogeny of Turnip mosaic virus; comparisons of 38 genomic sequences reveal a Eurasian origin and a recent 'emergence' in east Asia. Mol Ecol 12: , Martin DP, Lemey P, Lott M, Moulton V, Posada D and Lefeuvre P: RDP3: a flexible and fast computer program for analyzing recombination. Bioinformatics 26: , Tan Z, Wada Y, Chen J and Ohshima K: Inter- and intralineage recombinants are common in natural populations of Turnip mosaic virus. J Gen Virol 85: , Ohshima K, Tomitaka Y, Wood JT, Minematsu Y, Kajiyama H, Tomimura K and Gibbs AJ: Patterns of recombination in Turnip mosaic virus genomic sequences indicate hotspots of recombination. J Gen Virol 88: , Tomimura K, Špak J, Katis N, Jenner CE, Walsh JA, Gibbs, AJ and Ohshima K: Comparisons of the genetic structure of populations of Turnip mosaic virus in West and East Eurasia. Virology 330: , Tomitaka Y and Ohshima K: A phylogeographic study of the Turnip mosaic virus population in east Asia reveals an emergent lineage in Japan. Mol Ecol 15: , Librado P and Rozas J: DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics 25: , Excoffier L and Lischer HEL: Arlequin suite ver 3.5: A new series of programs to perform population genetics analyses under Linux and Windows. Mol Ecol Res 10: 564-7, Wang HY, Liu JL, Gao R, Chen J, Shao YH and Li XD: Complete genomic sequence analyses of Turnip mosaic virus basal-br isolates from China. Virus Genes 38: 421 8, Tomitaka Y, Yamashita T and Ohshima K: The genetic structure of populations of Turnip mosaic virus in Kyushu and central Honshu, Japan. J Gen Plant Pathol 73: , Nguyen HD, Tran HTN and Ohshima K: Genetic variation of the Turnip mosaic virus population of Vietnam; a case study of founder, regional and local influences. Virus Res in press, Nguyen HD, Tomitaka Y, Ho SYW, Duch ne S, Vetten H-J, Lesemann D, Walsh JA, Gibbs AJ and Ohshima K: Turnip mosaic potyvirus probably first spread to Eurasian brassica crops from wild orchids about 1000 years ago. submitted, Lesemann DE and Vetten HJ: The occurrence of tobacco rattle and Turnip mosaic viruses in Orchis ssp., and of an unidentified potyvirus in Cypripedium calceolus. Acta Hort 164: 45-54, Fuji S and Nakamae H: Complete nucleotide sequence of the genomic RNA of a Japanese yam mosaic virus, a new potyvirus in Japan. Arch Virol 144: , Fuji S and Nakamae H: Complete nucleotide sequence of the genomic RNA of a mild strain of Japanese yam mosaic potyvirus. Arch Virol 145: , Chen J, Zheng HY, Chen JP and Adams MJ: Characterisation of a potyvirus and a potexvirus from Chinese scallion. Arch Virol 147: , Chen J, Chen JP, Langeveld SA, Derks AFLM and Adams MJ: Molecular characterization of carla- and potyviruses from Narcissus in China. J Phytopathol 151: 26-9, Gibbs AJ, Trueman JWH and Gibbs MJ: The bean common mosaic virus lineage of potyviruses; where did it arise and when? Arch Virol 153: , Seo JK, Ohshima K, Lee HG, So M, Choi HS, Lee SH, Sohn SH and Kim, KH: Molecular variability and genetic structure of the population of Soybean mosaic virus based on the analysis of complete genome sequences. Virology 393: , Lin SQ, Shen JG, Gao FL, Cai W, Huang Z, Xie LY and Wu ZJ: Complete genome sequence of narcissus late season yellows virus infecting Chinese narcissus in China. Arch Virol 157:1821-4, Drummond AJ and Rambault A: BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evol Biol 7: 214, doi: / Drummond AJ, Ho SYW, Phillips MJ and Rambaut A: Relaxed phylogenetics and dating with confidence. PLoS Biol 4: e88, Drummond AJ, Pybus OG, Rambaut A, Forsberg R and Rodrigo AG: Measurably evolving populations. Trends Ecol Evol 18: 481-8, Twiddy SS, Holmes EC and Rambaut A: Inferring the rate and time-scale of dengue virus evolution. Mol Biol Evol 20:122-9, Shackelton LA and Holmes EC: Phylogenetic evidence for the rapid evolution of human B19 erythrovirus. J Virol 80: , Davis PL, Holmes EC, Larrous F, Van der Poel WH, Tj rneh j K, Alonso WJ and Bourhy H: Phylogeography, population dynamics, and molecular evolution of European bat lyssaviruses. J Virol 79: , Firth C, Kitchen A, Shapiro B, Suchard MA, Holmes EC and Rambaut A. Using time-structured data to estimate evolutionary rates of double-stranded DNA viruses. Mol Biol Evol 27: , Sall AA, Faye O, Diallo M, Firth C, Kitchen A and Holmes EC: Yellow fever virus exhibits slower evolutionary dynamics than dengue virus. J Virol 84: , Holmes EC and Grenfell BT: Discovering the phylodynamics of RNA viruses. PLoS Comput Biol 5:e , Pag n I, Firth C and Holmes EC: Phylogenetic analysis reveal rapid evolutionary dynamics in the plant RNA virus genus Tobamovirus. J Mol Evol 71: , Pag n I and Holmes EC: Long-term evolution of the Luteoviridae: time scale and mode of virus speciation. J Virol 84: , Duffy S and Holmes EC: Phylogenetic evidence for rapid rates of molecular evolution in the single-stranded DNA begomovirus Tomato yellow leaf curl virus. J
10 pp Virol 82: , Gibbs AJ, Ohshima K, Phillips MJ and Gibbs MJ: The prehistory of potyviruses: their initial radiation was during the dawn of agriculture. PLoS One 3: e2523, Gibbs AJ, Fargette D, Garc a-arenal F and Gibbs MJ: Time the emerging dimension of plant virus studies. J Gen Virol 91: 13-22, Sugimoto C, Kitamura T, Guo J, Al-Ahdal MN, Shchelkunov SN, Otova B, Ondrejka P, Chollet JY, El- Safi S, Ettayebi M, Gr senguet G, Kocag z T, Chaiyarasamee S, Thant KZ, Thein S, Moe K, Kobayashi N, Taguchi F and Yogo Y: Typing of urinary JC virus DNA offers a novel means of tracing human migrations. Proc Natl Acad Sci U S A 94: , Yogo Y, Zhong S, Suzuki M, Shibuya A and Kitamura T: Occurrence of the European subgroup of subtype I BK polyomavirus in Japanese-Americans suggests transmission outside the family. J Virol 81: , Gibbs AJ and Ohshima K: Potyviruses in the digital age. Annu Rev Phytopathol 48: , Gibbs AJ, Gibbs M, Ohshima K, Garcia-Arenal F: More about plant virus evolution; past, present and future. In: Domingo, E., Parrish, C.R., Holland, J.J. (Eds.), Ori- gin and Evolution of Viruses., 2nd ed. Elsevier Academic Press, London, UK pp , Suehiro N, Natsuaki T, Watanabe T and Okuda S: An important determinant of the ability of Turnip mosaic virus to infect Brassica spp. and/or Raphanus sativus is in its P3 protein. J Gen Virol 85: , Tan Z, Gibbs AJ, Tomitaka Y, Sanchez F, Ponz F and Ohshima K: Mutations in turnip mosaic virus genomes that have adapted to Raphanus sativus. J Gen Virol 86: , Ohshima K, Akaishi S, Kajiyama H, Koga R and Gibbs AJ: Evolutionary trajectory of turnip mosaic virus populations adapting to a new host. J Gen Virol 91: , Jenner CE, Tomimura K, Ohshima K, Hughes SL and Walsh JA: Mutations in Turnip mosaic virus P3 and cylindrical inclusion proteins are separately required to overcome two Brassica napus resistance genes. Virology 300: 50-9, Jenner CE, Wang X, Tomimura K, Ohshima K, Ponz F and Walsh JA: The dual role of the potyvirus P3 protein of Turnip mosaic virus as a symptom and avirulence determinant in brassicas. Mol Plant-Microbe Interact 16: , Plant potyvirus evolution: the survey of the genetic structure of populations Kazusato OHSHIMA Laboratory of Plant Virology, Department of Applied Biological Sciences, Faculty of Agriculture, Saga University, 1-banchi, Honjo-machi, Saga , Japan ohshimak@cc.saga-u.ac.jp The Potyvirus is the largest genus of the largest family of plant RNA viruses, the Potyviridae. The potyviruses infect not only dicotyledonous but also monocotyledonous plants. The potyvirus phylogeny shows that the genus probably originated from a virus of monocotyledonous plants and that it first diverged approximately 7250 years ago in Southwest Eurasia or North Africa. Turnip mosaic virus (TuMV) belongs to the genus Potyvirus and infects a wide range of plant species, most from the family Brassicaceae. TuMV is most studied a potyvirus species for molecular evolution and the genetic structure of populations. The use of computer programs for better understanding of the evolution and the genetic structures of populations of potyviruses and TuMV are illustrated.
TuMV 720 nm 1 RNA 9,830 1 P1 HC Pro a NIa Pro 10 P1 HC Pro 3 P36 1 6K1 CI 6 2 6K2VPgNIa Pro b NIb CP HC Pro NIb CP TuMV Y OGAWA et al.,
21 1980 2000 DNA I Molecular Evolution and Ecology of Turnip mosaic virus. By Kazusato OHSHIMA DNA TYLCV1 RNA Rice yellow mottle virus RYMV 1 RNA Turnip mosaic virus ; TuMV TuMV TuMV TuMV II 1 TuMV OHSHIMA
Turnip mosaic virus, TuMV 40 BEAST 20 TuMV 3,30 Cauliflower mosaic virus, CaMV Cucumber mosaic virus, CMV TuMV RNA CaMV DNA CMV RNA ,3
65 2 pp.229-238 2015 7. 3 3 1. 175 1840 Edward Hitchcock Charles Darwin 1866 Ernst Haeckel 100 1969 Adrian Gibbs 19 840-8502 1 TEL: 0952-28-8730 FAX: 0952-28-8709 E-mail: ohshimak@cc.saga-u.ac.jp 2000
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