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Bergey's Manual of Systematic Bacteriology第1版 163 総 説 腸内細菌学雑誌 22 : 163–169，2008 「Bergey’s Manual of Systematic Bacteriology第1版」に記載された 分類とその後の分類の異同―グラム陰性菌 平山 和宏＊ ＊東京大学大学院農学生命科学研究科・獣医公衆衛生学教室 Phylogenetic Position of Gram-Negative Bacteria in Modern Molecular Taxonomy Kazuhiro HIRAYAMA* *Department of Veterinary Public Health, Graduate School of Agricultural and Life Sciences, The University of Tokyo 要 旨 1984年に出版された「Bergey’s Manual of Systematic Bacteriology」は細菌分類学における膨大な知見を包括的 にまとめたものであり，細菌分類学の分野で最も権威を認められている著書の一つである．しかし，近年の分子生物学的 な細菌学の進歩により16S rRNAをコードする遺伝子（16S rDNA）の塩基配列の解読が急速に進み，16S rDNAの塩基配 列を基とした系統分類が用いられるようになった結果，1984年のManual出版の時には近縁と考えられていた菌がまった く異なるグループに分類されることもしばしば見受けられる．「Bergey’s Manual of Systematic Bacteriology」は現在第2 版が出版されつつあるところではあるが，本稿では消化管内の菌叢として重要な種が多く含まれる偏性嫌気性グラム陰性 菌について，1984年の第1版に記載された種や属が現在の分類体系ではどのように分類されているかを解説した． Abstract Bergey’s Manual of Systematic Bacteriology is one of the most comprehensive and authoritative works in the field of bacterial taxonomy. However, rapid advances in molecular sequencing of highly conserved regions of the prokary- otic genome, most notably genes coding for the RNA of the small ribosomal subunit, have resulted in numerous taxo- nomic rearrangements and changes in nomenclature since the first edition of the Manual was published in 1984. This paper summarizes the present state of taxonomy of anaerobic Gram-negative bacteria described in the first edition of the Manual. Key words : 16S rDNA ; bacterial taxonomy ; Gram-negative bacteria ; digestive tract ているかを解説した． はじめに 偏性嫌気性グラム陰性球菌 口腔から大腸に至る消化管に生息する細菌の中で，グ ラム陰性菌，特に偏性嫌気性グラム陰性桿菌は重要な位 Bergey’s Manualには，偏性嫌気性グラム陰性球菌は 置を占めている．しかし，16S rDNAの塩基配列を基に Veillonellaceae 科（Family）に分類される属（Genus） した新しい分類体系の採用により，従来のグラム染色性 として Veillonella，Acidaminococcus，Megasphaera によるグループ分けは必ずしも現在の分類体系とは対応 の 3属が記載されており，この他に分類位置が不明な菌 しなくなっており，かつて近縁と考えられていた菌がま として Gemmiger formicilis が記載されている． ったく異なるグループに分類されることもしばしば見受 Veillonella 属，Acidaminococcus 属，Megasphaera 属 けられる．そこで，本稿では1984年に出版された は今も同じ Veillonellaceae 科に分類されているが，現 「Bergey’s Manual of Systematic Bacteriology第 1版」 在の分類体系ではグラム陽性菌のグループである （以下 Bergey’s Manualと略す）に記載された偏性嫌気 Firmicutes 門（Phylum）に入り，Clostridia 綱（Class） 性のグラム陰性菌が現在の体系ではどのように分類され Clostridiales 目（ Order）に属している（Table 1）．これら の3属にはBergey’s Manualの出版以降に新菌種の報告は あったが，再分類の対象となった種はない．Gemmiger 2008年5月9日受付 ＊ 〒113–8657 東京都文京区弥生1–1–1 1–1–1 Yayoi, Bunkyo- formicilis は現在は Proteobacteria 門Alphaproteobacteria ku, Tokyo 113–8657, Japan 綱 Rhizobiales 目 Hyphomicrobiaceae 科に分類されてい 164 腸内細菌学雑誌 22巻3号 2008 Table 1. Anaerobic Gram negative cocci Table 2. Species of Genus Bacteroides Phylum Firmicutes, Class Clostridia, Order Clostridiales, Species described in the first edition of Bergey’s Manual of Family Veillonellaceae Systematic Bacteriology Genus Veillonella B. capillosus V. atypica* B. coagulans V. caviae* B. eggerthii* V. criceti* B. fragilis* V. denticariosi B. ovatus* V. dispar* B. splanchnicus V. montpellierensis B. thetaiotaomicron* V. parvula*,** B. uniformis* V. ratti* B. ureolyticus V. rodentium* B. vulgatus* Genus Acidaminococcus Species reported after the first edition of Bergey’s Manual A. fermentans* of Systematic Bacteriology A. intestini B. acidifaciens B. barnesiae Genus Megasphaera B. caccae M. cerevisiae B. cellulosilyticus M. elsdenii*,** B. cellulosolvens M. micronuciformis** B. coprocola M. paucivorans B. coprophilus M. sueciensis B. coprosuis B. dorei Phylum Proteobacteria, Class Alphaproteobacteria, Order B. finegoldii Rhizobiales, Family Hyphomicrobiaceae B. galacturonicus Genus Gemmiger B. gallinarum G. formicilis*,** B. helcogenes B. intestinalis * : Species described in the first edition of Bergey’s Manual of B. massiliensis Systematic Bacteriology, B. nordii ** : Species found in human intestine. B. pectinophilus B. plebeius B. polypragmatus る（Table 1）． B. propionicifaciens B. pyogenes 偏性嫌気性グラム陰性桿菌 B. salanitronis B. salyersiae Bergey’s Manualでは，偏性嫌気性グラム陰性桿菌と B. stercoris* B. suis して Bacteroides，Fusobacterium，Leptotrichia， B. tectus Butyrivibrio，Succinimonas，Succinivibrio， B. xylanolyticus Anaerobiospirillum，Wolinella，Selenomonas， * : Species found in human intestine. Anaerovibrio，Pectinatus，Acetivibrio，Lachnospira の 13の属が記載され，全て Bacteroidaceae 科に分類さ れていた．しかし，現在の分類体系ではBacteroides 属 Bacteroidetes 綱 Bacteroidales 目 Bacteroidaceae 科に分 以外は Bacteroidaceae 科から離れているばかりでなく， 類されている．Bacteroides 属は消化管内菌叢を形成す Bacteroides 属と同じ Bacteroidetes 門に属しているものも る優勢な菌群であり，1984年に出版されたBergey’s ない．Fusobacterium 属と Leptotrichia 属は Fusobacteria Manualには亜種を含めて40の菌名が記載されている 門に所属している．Butyrivibrio 属，Selenomonas 属， が，分子生物学的な分類学が用いられるようになったこ Anaerovibrio 属，Pectinatus 属，Acetivibrio 属およ とにより，多くの菌が再分類の対象となった． び Lachnospira 属はグラム陽性菌のグループであ Bacteroides 属に分類される菌は B. fragilis とその近縁 る Firmicutes 門に入っている．Succinimonas 属， の種のみとすることが提唱され（17），Bergey’s Manual Succinivibrio 属，Anaerobiospirillum 属および に記載のあった Bacteroides 属の菌のうち新しい分類体 Wolinella 属は Proteobacteria 門である． 系でも Bacteroides 属に分類されるものは10菌種となっ た（Table 2）．ただし，腸内菌叢として重要であり， Bacteroides 属 B. fragilis と非常に近縁と考えられていた種のうち， Bacteroides 属は現在の分類体系では Bacteroidetes 門 B. distasonis のみは他の2 種とともにBacteroidales 平山和宏：新しい分類における嫌気性グラム陰性菌 165 目の異なる科である Porphyromonadaceae 科の からも分離されるB. furcosus は同じ Bacteroidaceae 科 Parabacteroides 属（12）に再分類された（Table 3）． の Anaerorhabdus 属に再分類されて A. furcosa となっ また，Bacteroides 属にはBergey’s Manualの出版以降 た（15）が，その他は科以上で異なる分類単位へと移 に 38菌種が報告されたが，そのうち11菌種が現在は 動した（Table 3）．口腔内には多くの偏性嫌気性グラム Bacteroides 属以外に分類されている． 陰性桿菌が生息し，重要な口腔内菌叢を形成している． これらの再分類の対象となった菌種のうち，消化管内 これらはかつて Bacteroides 属に含まれていたが，多く Table 3. Species reclassified from Genus Bacteroides to other genera of Phylum Bacteroidetes Phylum Bacteroidetes, Class Bacteroidetes, Order Bacteroidales, Family Bacteroidaceae Genus Anaerorhabdus B. furcosus* → A. furcosa** Phylum Bacteroidetes, Class Bacteroidetes, Order Bacteroidales Family Rikenellaceae Genus Alistipes B. putredinis* → A. putredinis Genus Rikenella B. microfusus* → R. microfusus Family Porphyromonadaceae Genus Parabacteroides B. distasonis* → P. distasonis** B. goldsteinii → P. goldsteinii B. merdae → P. merdae Genus Porphyromonas B. asaccharolyticus* → P. asaccharolytica B. endodontalis → P. endodontalis B. gingivalis* → P. gingivalis** B. levii* → P. levii B. macacae → P. macacae B. salivosus → P. salivosa � P. macacae Genus Tannerella B. forsythus → T. forsythensis Family Prevotellaceae Genus Prevotella B. bivius* → P. bivia** B. buccalis → P. buccalis B. buccae* → P. buccae B. corporis* → P. corporis B. denticola* → P. denticola B. disiens* → P. disiens** B. heparinolyticus → P. heparinolytica B. intermedius* → P. intermedia** B. loescheii* → P. loescheii B. melaninogenicus* → P. melaninogenica** B. oralis* → P. oralis B. oris* → P. oris** B. oulorum → P. oulorum B. ruminicola subsp. brevis* → P. ruminicola subsp. brevis → P. brevis B. ruminicola subsp. ruminicola* → P. ruminicola subsp. ruminicola → P. ruminicola B. veroralis* → P. veroralis B. zoogleoformans* → P. zoogleoformans Phylum Bacteroidetes Class Flavobacteria, Order Flavobacteriales, Family Flavobacteriaceae Genus Capnocytophaga B. ochraceus → C. ochracea * : Species described in the first edition of Bergey’s Manual of Systematic Bacteriology, ** : Species found in human intestine, → : basonym, � : heterotypic synonym. 166 腸内細菌学雑誌 22巻3号 2008 が Porphyromonadaceae 科 Porphyromonas 属（8, 16, 載されているが，Megamonas 属および Mitsuokella 属 19）と Prevotellaceae 科 Prevotella 属（18）に再分類さ は現在ではさらにBacteroides 属から離れ，いずれもグラ れた．なお，B. ruminicola の 2つの亜種はPrevotella ム陽性菌のグループであるFirmicutes 門の Clostridia 属に移ったが，その後独立した種としてP. brevis， 綱 Clostridiales 目 Veillonellaceae 科に分類されている． P. ruminicola となった．B. salivosus は Bergey’s また，同じ Veillonellaceae 科には B. pneumosintes が Manualの出版以降に報告された種であるが， Dialister pneumosintes として（10），Clostridiales 目の Porphyromonas 属に移って P. salivosa となった後 Peptostreptococcaceae 科には B. praeacutus がTissierella P. macacae に含まれることとなった．糞便や病巣から分 praeacuta として再分類された（3）．B. succinogenes も 離される B. putredinis は Bacteroidales 目 Rikenellaceae 腸管内に生息する菌であるが，独立した Fibrobacteres 科の Alistipes 属に再分類された（11）．同じ Rikenellaceae 門に再分類され，現在は Fibrobacteres 門 Fibrobacteres 科の Rikenella 属には B. microfusus が再分類された 綱 Fibrobacterales 目 Fibrobacteraceae 科の Fibrobacter （4）． succinogenes subsp. succinogenes（9）となった．こ 門から異なる単位に再分類された種も少なくない の他，B. termitidis は Sebaldella 属（2）に，B. amy- （Table 4）．腸内からしばしば分離される B. hypermegas lophilus は Ruminobacter 属（20）に ，B. nodosus は と B. multiacidus は，Bergey’s Manualでもすでに原稿 Dichelobacter 属（5）に，B. gracilis は Campylobacter 執筆後出版までの間にそれぞれBacteroides 属から独立 属（22）にそれぞれ再分類された（Table 4）． した属として承認され，Megamonas hypermegale（13） および Mitsuokella multacida（14）となったことが記 Table 4. Species reclassified from Genus Bacteroides to other phyla Phylum Fibrobacteres, Class Fibrobacteres, Order Fibrobacterales, Family Fibrobacteraceae Genus Fibrobacter B. succinogenes* → F. succinogenes subsp. succinogenes** Phylum Firmicutes, Class Clostridia, Order Clostridiales Family Veillonellaceae Genus Megamonas B. hypermegas* → M. hypermegale** Genus Mitsuokella B. multiacidus* → M. multacida** Genus Dialister B. pneumosintes* → D. pneumosintes Family Peptostreptococcaceae Genus Tissierella B. praeacutus* → T. praeacuta Phylum Fusobacteria, Class Fusobacteria, Order Fusobacteriales, Family Fusobacteriaceae Genus Sebaldella B. termitidis* → S. termitidis Phylum Proteobacteria Class Epsilonproteobacteria, Order Campylobacterales, Family Campylobacteraceae Genus Campylobacter B. gracilis* → C. gracilis Class Gammaproteobacteria Order Cardiobacteriales, Family Cardiobacteriaceae Genus Dichelobacter B. nodosus* → D. nodosus Order Acidithiobacillales, Family Succinivibrionaceae Genus Ruminobacter B. amylophilus* → R. amylophilus * : Species described
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