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* ''Anoxyphotobacteria''<br /><small>(Gibbons and Murray 1978) Murray 1988<ref name="Murray 1988">{{cite book |author=Murray RGE |chapter=The higher taxa, or, a place for everything..? |editor=Krieg NR, Holt JG |title=Bergey's manual of systematic bacteriology, vol. 1. |publisher=The Williams & Wilkins Co., Baltimore |year=1984 |pages=31-34 }}</ref><br />(IJSEMリストに記載 1988)<ref name="10.1099/00207713-38-2-220">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/00207713-38-2-220 |volume=38 |issue=2 |title=Validation of the Publication of New Names and New Combinations Previously Effectively Published Outside the IJSB |date=01 April 1988 |pages=220-222 }}</ref></small>
* ''Anoxyphotobacteria''<br /><small>(Gibbons and Murray 1978) Murray 1988<ref name="Murray 1988">{{cite book |author=Murray RGE |chapter=The higher taxa, or, a place for everything..? |editor=Krieg NR, Holt JG |title=Bergey's manual of systematic bacteriology, vol. 1. |publisher=The Williams & Wilkins Co., Baltimore |year=1984 |pages=31-34 }}</ref><br />(IJSEMリストに記載 1988)<ref name="10.1099/00207713-38-2-220">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/00207713-38-2-220 |volume=38 |issue=2 |title=Validation of the Publication of New Names and New Combinations Previously Effectively Published Outside the IJSB |date=01 April 1988 |pages=220-222 }}</ref></small>
* "''Candidatus'' Zetaproteobacteria"<br /><small>Emerson ''et al''. 2007<ref name="10.1371/journal.pone.0000667">{{cite journal |journal=PLoS One |date=2007 Aug 1 |volume=2 |issue=7 |page=e667 |doi=10.1371/journal.pone.0000667 |title=A novel lineage of proteobacteria involved in formation of marine Fe-oxidizing microbial mat communities |author=David Emerson, Jeremy A Rentz, Timothy G Lilburn, Richard E Davis, Henry Aldrich, Clara Chan, Craig L Moyer |PMID=17668050 |PMC=1930151 }}</ref><br />(IJSEMリストに記載 2020)<ref name="10.1099/ijsem.0.003789">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/ijsem.0.003789 |volume=70 |issue=7 |title=Lists of names of prokaryotic Candidatus taxa |author=Aharon Oren, George M. Garrity, Charles T. Parker, Maria Chuvochina and Martha E. Trujillo |date=30 June 2020 |pages=3956-4042 }}</ref></small>
* "''Candidatus'' Zetaproteobacteria"<br /><small>Emerson ''et al''. 2007<ref name="10.1371/journal.pone.0000667">{{cite journal |journal=PLoS One |date=2007 Aug 1 |volume=2 |issue=7 |page=e667 |doi=10.1371/journal.pone.0000667 |title=A novel lineage of proteobacteria involved in formation of marine Fe-oxidizing microbial mat communities |author=David Emerson, Jeremy A Rentz, Timothy G Lilburn, Richard E Davis, Henry Aldrich, Clara Chan, Craig L Moyer |PMID=17668050 |PMC=1930151 }}</ref><br />(IJSEMリストに記載 2020)<ref name="10.1099/ijsem.0.003789">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/ijsem.0.003789 |volume=70 |issue=7 |title=Lists of names of prokaryotic Candidatus taxa |author=Aharon Oren, George M. Garrity, Charles T. Parker, Maria Chuvochina and Martha E. Trujillo |date=30 June 2020 |pages=3956-4042 }}</ref></small>
* "''Candidatus'' Mariprofundia"<br /><small>corrig. Emerson ''et al''. 2007<ref name="10.1371/journal.pone.0000667 /><br />(IJSEMリストに記載 2017)<ref name="10.1099/ijsem.0.001715">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/ijsem.0.001715 |volume=67 |issue=4 |title=A plea for linguistic accuracy – also for ''Candidatus'' taxa |author=Aharon Oren |date=01 April 2017 }}</ref></small>
* "''Candidatus'' Mariprofundia"<br /><small>corrig. Emerson ''et al''. 2007<ref name="10.1371/journal.pone.0000667" /><br />(IJSEMリストに記載 2017)<ref name="10.1099/ijsem.0.001715">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/ijsem.0.001715 |volume=67 |issue=4 |title=A plea for linguistic accuracy – also for ''Candidatus'' taxa |author=Aharon Oren |date=01 April 2017 }}</ref></small>
* カウロバクター綱<br />"''Caulobacteria''"<br /><small>Cavalier-Smith 2020<ref name="10.1007/s00709-019-01442-7">{{cite journal |journal=Protoplasma |title=Multidomain ribosomal protein trees and the planctobacterial origin of neomura (eukaryotes, archaebacteria) |date=03 January 2020 |volume=257 |pages=621–753 |author=Thomas Cavalier-Smith, Ema E-Yung Chao |PMID=31900730 |PMC=7203096 |doi=10.1007/s00709-019-01442-7 |issue=3 }}</ref></small>
* カウロバクター綱<br />"''Caulobacteria''"<br /><small>Cavalier-Smith 2020<ref name="10.1007/s00709-019-01442-7">{{cite journal |journal=Protoplasma |title=Multidomain ribosomal protein trees and the planctobacterial origin of neomura (eukaryotes, archaebacteria) |date=03 January 2020 |volume=257 |pages=621–753 |author=Thomas Cavalier-Smith, Ema E-Yung Chao |PMID=31900730 |PMC=7203096 |doi=10.1007/s00709-019-01442-7 |issue=3 }}</ref></small>
* マグネトコッカス綱<br />''Magnetococcia''<br /><small>Chuvochina ''et al''. 2024<ref name="10.1093/femsle/fnad071">{{cite journal |doi=10.1093/femsle/fnad071 |title=Proposal of names for 329 higher rank taxa defined in the Genome Taxonomy Database under two prokaryotic codes |author=Maria Chuvochina, Aaron J Mussig, Pierre-Alain Chaumeil, Adam Skarshewski, Christian Rinke, Donovan H Parks, Philip Hugenholtz |journal=FEMS Microbiology Letters |volume=370 |page=fnad071 |date=21 July 2023 |PMID=37480240 |PMC=10408702 }}</ref><br />(IJSEMリストに記載 2024)<ref name="10.1099/ijsem.0.006173">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/ijsem.0.006173 |volume=74 |issue=1 |title=Validation List no. 215. Valid publication of new names and new combinations effectively published outside the IJSEM |author=Aharon Oren and Markus Göker |date=01 February 2024 |page=6173 }}</ref></small>
* マグネトコッカス綱<br />''Magnetococcia''<br /><small>Chuvochina ''et al''. 2024<ref name="10.1093/femsle/fnad071">{{cite journal |doi=10.1093/femsle/fnad071 |title=Proposal of names for 329 higher rank taxa defined in the Genome Taxonomy Database under two prokaryotic codes |author=Maria Chuvochina, Aaron J Mussig, Pierre-Alain Chaumeil, Adam Skarshewski, Christian Rinke, Donovan H Parks, Philip Hugenholtz |journal=FEMS Microbiology Letters |volume=370 |page=fnad071 |date=21 July 2023 |PMID=37480240 |PMC=10408702 }}</ref><br />(IJSEMリストに記載 2024)<ref name="10.1099/ijsem.0.006173">{{cite journal |journal=International Journal of Systematic and Evolutionary Microbiology |doi=10.1099/ijsem.0.006173 |volume=74 |issue=1 |title=Validation List no. 215. Valid publication of new names and new combinations effectively published outside the IJSEM |author=Aharon Oren and Markus Göker |date=01 February 2024 |page=6173 }}</ref></small>
* マリプロフンドゥス綱<br />"''Mariprofundia''"<br /><small>Cavalier-Smith 2020<ref name="10.1007/s00709-019-01442-7 /></small>
* マリプロフンドゥス綱<br />"''Mariprofundia''"<br /><small>Cavalier-Smith 2020<ref name="10.1007/s00709-019-01442-7" /></small>
* ゼータプロテオバクテリア綱<br />"''Zetaproteobacteria''"<br /><small>Makita ''et al''. 2017<ref name="10.1007/s00203-016-1307-4">{{cite journal |doi=10.1007/s00203-016-1307-4 |journal=Archives of Microbiology |title=''Mariprofundus micogutta'' sp. nov., a novel iron-oxidizing zetaproteobacterium isolated from a deep-sea hydrothermal field at the Bayonnaise knoll of the Izu-Ogasawara arc, and a description of ''Mariprofundales'' ord. nov. and ''Zetaproteobacteria'' classis nov. |date=20 October 2016 |volume=199 |pages=335–346 |author=Hiroko Makita, Emiko Tanaka, Satoshi Mitsunobu, Masayuki Miyazaki, Takuro Nunoura, Katsuyuki Uematsu, Yoshihiro Takaki, Shinro Nishi, Shigeru Shimamura & Ken Takai |PMID=27766355 }}</ref></small>
* ゼータプロテオバクテリア綱<br />"''Zetaproteobacteria''"<br /><small>Makita ''et al''. 2017<ref name="10.1007/s00203-016-1307-4">{{cite journal |doi=10.1007/s00203-016-1307-4 |journal=Archives of Microbiology |title=''Mariprofundus micogutta'' sp. nov., a novel iron-oxidizing zetaproteobacterium isolated from a deep-sea hydrothermal field at the Bayonnaise knoll of the Izu-Ogasawara arc, and a description of ''Mariprofundales'' ord. nov. and ''Zetaproteobacteria'' classis nov. |date=20 October 2016 |volume=199 |pages=335–346 |author=Hiroko Makita, Emiko Tanaka, Satoshi Mitsunobu, Masayuki Miyazaki, Takuro Nunoura, Katsuyuki Uematsu, Yoshihiro Takaki, Shinro Nishi, Shigeru Shimamura & Ken Takai |PMID=27766355 }}</ref></small>
|下位分類名=下位分類([[目 (分類学)|目]])(2024年6月現在)<ref name=LPSN>{{Cite web |publisher=List of Prokaryotic names with Standing in Nomenclature |author=Jean P. Euzéby, Aidan C. Parte |url=https://lpsn.dsmz.de/class/alphaproteobacteria |title=Class ''Alphaproteobacteria'' |access-date=2024-06-11 }}</ref>
|下位分類名=下位分類([[目 (分類学)|目]])(2024年6月現在)<ref name=LPSN>{{Cite web |publisher=List of Prokaryotic names with Standing in Nomenclature |author=Jean P. Euzéby, Aidan C. Parte |url=https://lpsn.dsmz.de/class/alphaproteobacteria |title=Class ''Alphaproteobacteria'' |access-date=2024-06-11 }}</ref>

2024年6月29日 (土) 00:14時点における版

アルファプロテオバクテリア綱
昆虫細胞内のボルバキア属（Wolbachia）の透過電子顕微鏡写真。
出典:Public Library of Science / Scott O'Neill
タイプ属カウロバクター属（Caulobacter）のタイプ種カウロバクター・ビブリオイデス(Caulobacter vibrioides)の顕微鏡画像。鞭毛を持つ遊走細胞と、付属器を伸ばす有柄細胞とが結合している。
分類
ドメイン : 真正細菌
Bacteria
門 : Pseudomonadota門
綱 : アルファプロテオバクテリア綱
Alphaproteobacteria
学名
Alphaproteobacteria
Garrity et al. 2006[1]
(IJSEMリストに記載 2006)[2]
タイプ属
カウロバクター属
Caulobacter
Henrici and Johnson 1935[3]
(IJSEMリストに記載 1980)[4]
修正　Abraham et al. 1999[5]
(IJSEMリストに記載 1980)[6]
シノニム
  • アルファバクテリア綱
    Alphabacteria
    Cavalier-Smith 2002[7]
    (IJSEMリストに記載 2002)[8]
  • Anoxyphotobacteria
    (Gibbons and Murray 1978) Murray 1988[9]
    (IJSEMリストに記載 1988)[10]
  • "Candidatus Zetaproteobacteria"
    Emerson et al. 2007[11]
    (IJSEMリストに記載 2020)[12]
  • "Candidatus Mariprofundia"
    corrig. Emerson et al. 2007[11]
    (IJSEMリストに記載 2017)[13]
  • カウロバクター綱
    "Caulobacteria"
    Cavalier-Smith 2020[14]
  • マグネトコッカス綱
    Magnetococcia
    Chuvochina et al. 2024[15]
    (IJSEMリストに記載 2024)[16]
  • マリプロフンドゥス綱
    "Mariprofundia"
    Cavalier-Smith 2020[14]
  • ゼータプロテオバクテリア綱
    "Zetaproteobacteria"
    Makita et al. 2017[17]
下位分類(目)(2024年6月現在)[46]

アルファプロテオバクテリア綱（ーこう、Alphaproteobacteria）は細菌ドメインのPseudomonadota門の綱の一つである[47]。生態は光合成細菌から植物共生菌まで非常に多様である。ミトコンドリアの祖先にあたる生物が属していたグループとしても知られ、アルファプロテオバクテリアの進化についての研究は真核生物の起源を探る上で重要である。グラム陰性であるが、細胞内寄生種はペプチドグリカンを持たず、したがってグラム染色試験での結果は陽性になることもある[48]。ベータプロテオバクテリアおよびガンマプロテオバクテリアと近縁である。

特徴

アルファプロテオバクテリア綱は非常に多様な生物群である。ある属は光栄養細菌であり、その一部はC1化合物を代謝する(例:Methylobacterium spp.)。またある属は植物共生細菌であり(例: Rhizobium spp.)、ある属は節足動物の内生菌であり(Wolbachia属)、ある属は細胞内侵襲性の感染菌である(例: Rickettsia属)。すでに絶滅したが、アルファプロテオバクテリア綱の細菌にはかつて真核生物の細胞に侵入して現在のミトコンドリアとなったプロトミトコンドリアがいた（細胞内共生説）[49] 。Rhizobium radiobacter (過去のAgrobacterium tumefaciens)は植物細胞に外来DNAを転移させるために用いられる[50] 。Pelagibacter ubiqueといった好気性の非酸素光栄養細菌はアルファプロテオバクテリア綱であり、外洋の微生物コミュニティの10%以上を構成すると考えられている海中プランクトンである。

分類

アルファプロテオバクテリアの系統関係については大まかな合意はあるものの、いくつかの目（order）については議論が続いている（例えばペラジバクター目、 リケッチア目、ホロスポラ目）[51]。他の細菌や古細菌と同様、その分類や系統関係については現在も絶えず改定が続いている[24][52]。NCBIのデータベースでは現在20の目が設定されている[53]。一方、GTDBでは35の目が設定されている[54]。分子系統解析とアルファプロテオバクテリアのゲノムに特徴的なインデルの分析[55]により、アルファプロテオバクテリア綱の分岐は細菌ドメインの中ではかなり遅く、ベータプロテオバクテリア綱とガンマプロテオバクテリア綱のみがアルファプロテオバクテリア綱より後に分岐したと推定されている[56][57]。

アルファプロテオバクテリアのうち、最も早くに分岐したのはマグネトコッカス目（Magnetococcales）であることはおおむね支持されているが、この目をアルファプロテオバクテリアに含めるかについては議論があり、独立した目（イータプロテオバクテリア[Etaproteobacteria]）の設立も提案されている[58][59]（ただし、2021年現在も認められていない）。2018年の研究では、ミトコンドリアはマグネトコッカス目を除く全てのアルファプロテオバクテリアに対して姉妹群となることが示唆された[60]。これはミトコンドリアにつながる祖先の系統分岐がアルファプロテオバクテリア綱内で非常に早く起こったことを示している。またミトコンドリアの系統が、リケッチア目と近縁ではあるものの、独立した系統群を形成することも示唆している。

系統樹

アルファプロテオバクテリア綱の分類体系は"List of Prokaryotic names with Standing in Nomenclature"(LPSN) [46]と米国立生物工学情報センター(NCBI)[61]で公開されている。下記の系統樹は呼吸複合体IのサブユニットNuoLを用いて2022年に作成されたものである[62]。

 
 

Magnetococcales（マグネトコッカス目）

 
 
 
 

Rickettsiales（リケッチア目）、Holosporales（ホロスポラ目）

 
 
 

Rhodospirillales（ロドスピリルム目）

 
 
 

Sneathiellales（スネアチエラ目）

 
 
 

Emcibacterales

 
 
 
 
 

Rhodothalassiales

 
 

Iodidimonadales

 
 
 

Kordiimonadales（コーディイモナス目）

 
 
 
 

Hyphomicrobiales（ヒフォミクロビウム目）

 
 
 
 

Caulobacterales（カウロバクター目）

 
 

Sphingomonadales（スフィンゴモナス目）

 
 
 

Rhodobacterales（ロドバクター目）

 
 
 
 
 
 
 
 
 

ミトコンドリア

 
 
 

自然遺伝子形質転換

アルファプロテオバクテリア綱の自然遺伝子形質転換の過程についてはAgrobacterium tumefaciens[63] 、Methylobacterium organophilum[64] 、およびBradyrhizobium japonicum[65] で明らかになっている。形質転換はDNAをある細胞から別の細胞に移動させる過程であり、相同組換えによって供給DNAは受容ゲノムに組み込まれる。

脚注

  1. ^ Garrity GM, Bell JA, Lilburn T (2005). “Class I. Alphaproteobacteria class. nov.”. In Brenner DJ, Krieg NR, Staley JT, Garrity GM. Bergey's Manual of Systematic Bacteriology, 2nd edn, vol. 2 (The Proteobacteria), part C (The Alpha-, Beta-, Delta-, and Epsilonproteobacteria). Springer (New York). p. 1 
  2. ^ a b c “List of new names and new combinations previously effectively, but not validly, published”. International Journal of Systematic and Evolutionary Microbiology 56 (1): 1-6. (01 January 2006). doi:10.1099/ijs.0.64188-0. 
  3. ^ a b Arthur T. Henrici, Delia E. Johnson (1 July 1935). “Studies of Freshwater Bacteria: II. Stalked Bacteria, a New Order of Schizomycetes”. Journal of Bacteriology 30 (1): 61-93. doi:10.1128/jb.30.1.61-93.1935. PMC 543637. PMID 16559821. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC543637/. 
  4. ^ a b c d e V. B. D. Skerman, Vicki. McGOWAN, P. H. A. Sneath (01 January 1980). “Approved Lists of Bacterial Names”. International Journal of Systematic and Evolutionary Microbiology 30 (1): 225-420. doi:10.1099/00207713-30-1-225. 
  5. ^ Wolf-Rainer Abraham, Carsten Strömpl, Holger Meyer, Sabine Lindholst, Edward R. B. Moore, Ruprecht Christ, Marc Vancanneyt, B. J. Tindall, Antonio Bennasar, John Smit and Michael Tesar (01 July 1999). “Phylogeny and polyphasic taxonomy of Caulobacter species. Proposal of Maricaulis gen. nov. with Maricaulis maris (Poindexter) comb. nov. as the type species, and emended description of the genera Brevundimonas and Caulobacter”. International Journal of Systematic and Evolutionary Microbiology 49 (3): 1053-73. doi:10.1099/00207713-49-3-1053. PMID 10425763. 
  6. ^ Anonymous (01 October 1999). “Notification that new names and new combinations have appeared in volume 49, part 3, of the IJSB”. International Journal of Systematic and Evolutionary Microbiology 49 (4): 1327-1328. doi:10.1099/00207713-49-4-1327. 
  7. ^ T Cavalier-Smith (01 January 2002). “The neomuran origin of archaebacteria, the negibacterial root of the universal tree and bacterial megaclassification”. International Journal of Systematic and Evolutionary Microbiology 52 (1): 7-76. doi:10.1099/00207713-52-1-7. PMID 11837318. 
  8. ^ “Notification list. Notification that new names and new combinations have appeared in volume 52, part 1 of the IJSEM”. International Journal of Systematic and Evolutionary Microbiology 52: 293-295. (2002). doi:10.1099/ijs.0.02318-0. 
  9. ^ Murray RGE (1984). “The higher taxa, or, a place for everything..?”. In Krieg NR, Holt JG. Bergey's manual of systematic bacteriology, vol. 1.. The Williams & Wilkins Co., Baltimore. pp. 31-34 
  10. ^ “Validation of the Publication of New Names and New Combinations Previously Effectively Published Outside the IJSB”. International Journal of Systematic and Evolutionary Microbiology 38 (2): 220-222. (01 April 1988). doi:10.1099/00207713-38-2-220. 
  11. ^ a b David Emerson, Jeremy A Rentz, Timothy G Lilburn, Richard E Davis, Henry Aldrich, Clara Chan, Craig L Moyer (2007 Aug 1). “A novel lineage of proteobacteria involved in formation of marine Fe-oxidizing microbial mat communities”. PLoS One 2 (7): e667. doi:10.1371/journal.pone.0000667. PMC 1930151. PMID 17668050. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1930151/. 
  12. ^ Aharon Oren, George M. Garrity, Charles T. Parker, Maria Chuvochina and Martha E. Trujillo (30 June 2020). “Lists of names of prokaryotic Candidatus taxa”. International Journal of Systematic and Evolutionary Microbiology 70 (7): 3956-4042. doi:10.1099/ijsem.0.003789. 
  13. ^ Aharon Oren (01 April 2017). “A plea for linguistic accuracy – also for Candidatus taxa”. International Journal of Systematic and Evolutionary Microbiology 67 (4). doi:10.1099/ijsem.0.001715. 
  14. ^ a b Thomas Cavalier-Smith, Ema E-Yung Chao (03 January 2020). “Multidomain ribosomal protein trees and the planctobacterial origin of neomura (eukaryotes, archaebacteria)”. Protoplasma 257 (3): 621–753. doi:10.1007/s00709-019-01442-7. PMC 7203096. PMID 31900730. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7203096/. 
  15. ^ Maria Chuvochina, Aaron J Mussig, Pierre-Alain Chaumeil, Adam Skarshewski, Christian Rinke, Donovan H Parks, Philip Hugenholtz (21 July 2023). “Proposal of names for 329 higher rank taxa defined in the Genome Taxonomy Database under two prokaryotic codes”. FEMS Microbiology Letters 370: fnad071. doi:10.1093/femsle/fnad071. PMC 10408702. PMID 37480240. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10408702/. 
  16. ^ a b Aharon Oren and Markus Göker (01 February 2024). “Validation List no. 215. Valid publication of new names and new combinations effectively published outside the IJSEM”. International Journal of Systematic and Evolutionary Microbiology 74 (1): 6173. doi:10.1099/ijsem.0.006173. 
  17. ^ Hiroko Makita, Emiko Tanaka, Satoshi Mitsunobu, Masayuki Miyazaki, Takuro Nunoura, Katsuyuki Uematsu, Yoshihiro Takaki, Shinro Nishi, Shigeru Shimamura & Ken Takai (20 October 2016). “Mariprofundus micogutta sp. nov., a novel iron-oxidizing zetaproteobacterium isolated from a deep-sea hydrothermal field at the Bayonnaise knoll of the Izu-Ogasawara arc, and a description of Mariprofundales ord. nov. and Zetaproteobacteria classis nov.”. Archives of Microbiology 199: 335–346. doi:10.1007/s00203-016-1307-4. PMID 27766355. 
  18. ^ a b Takao Iino, Moriya Ohkuma, Yoichi Kamagata and Seigo Amachi (01 December 2016). “Iodidimonas muriae gen. nov., sp. nov., an aerobic iodide-oxidizing bacterium isolated from brine of a natural gas and iodine recovery facility, and proposals of Iodidimonadaceae fam. nov., Iodidimonadales ord. nov., Emcibacteraceae fam. nov. and Emcibacterales ord. nov.”. International Journal of Systematic and Evolutionary Microbiology 66 (12): 5016-5022. doi:10.1099/ijsem.0.001462. PMID 27566239. 
  19. ^ a b “Notification that new names of prokaryotes, new combinations and new taxonomic opinions have appeared in volume 66, part 12, of the IJSEM”. International Journal of Systematic and Evolutionary Microbiology 67 (3): 525-528. (01 March 2017). doi:10.1099/ijsem.0.001834. 
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