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A spirochaete (/ˈspaɪroʊˌkiːt/)[1] or spirochete is a member of the phylum Spirochaetes (/-ˈkiːtiːz/),[2] which contains distinctive diderm (double-membrane) bacteria, most of which have long, helically coiled (corkscrew-shaped or spiraled, hence the name) cells.[3] Spirochaetes stain gram-negative and are chemoheterotrophic in nature, with lengths between 3 and 500 µm and diameters around 0.09 to at least 3 µm.[4] Spirochaetes are distinguished from other bacterial phyla by the location of their flagella, sometimes called axial filaments, which run lengthwise between the bacterial inner membrane and outer membrane in periplasmic space. These cause a twisting motion which allows the spirochaete to move about. When reproducing, a spirochaete will undergo asexual transverse binary fission. Most spirochaetes are free-living and anaerobic, but there are numerous exceptions. Spirochaetes bacteria are diverse in their pathogenic capacity, the ecological niches that they inhabit, as well as molecular characteristics including guanine-cytosine content and genome size.[5][6]

Contents

1 Pathogenicity 2 Taxonomy and molecular signatures 3 Phylogeny 4 Taxonomy 5 See also 6 References 7 External links

Pathogenicity[edit] Many organisms within the Spirochaetes phylum cause prevalent diseases. Pathogenic members of this phylum include the following:

Leptospira
Leptospira
species, which causes leptospirosis[7] Borrelia
Borrelia
burgdorferi, B. garinii, and B. afzelii, which cause Lyme disease Borrelia
Borrelia
recurrentis, which causes relapsing fever[8] Treponema pallidum
Treponema pallidum
subspecies which cause treponematoses such as syphilis and yaws. Brachyspira pilosicoli and Brachyspira aalborgi, which cause intestinal spirochaetosis[9]

Spirochaetes may also cause dementia and may be involved in the pathogenesis of Alzheimer's disease.[10] Salvarsan, the first partially organic synthetic antimicrobial drug in medical history, was effective against spirochaetes only and was primarily used to cure syphilis. Taxonomy and molecular signatures[edit] The class currently consists of 14 validly named genera across 4 orders and 5 families.[11][12][13] The orders Brachyspirales, Brevinematales and Leptospirales each contain a single family, Brachyspiraceae, Brevinemataceae and Leptospiraceae, respectively. The Spirochaetales
Spirochaetales
order harbours two families, Spirochaetaceae and Borreliaceae. Molecular markers in the form of conserved signature indels (CSIs) and CSPs have been found specific for each of the orders, with the exception of Brevinimetales, that provide a reliable means to demarcate these clades from one another within the diverse phylum.[12] Additional CSIs have been found exclusively shared by each family within the Spirochaetales. These molecular markers are in agreement with the observed phylogenetic tree branching of two monophyletic clades within the Spirochaetales
Spirochaetales
order.[12] CSIs have also been found that further differentiate taxonomic groups within the Borreliaceae family that further delineate evolutionary relationships that are in accordance with physical characteristics such as pathogenicity (viz. Borrelia
Borrelia
emend. Borreliella gen. nov.).[14] A conserved signature indel has also been found exclusively shared by all Spirochaetes species.[12] This CSI is a 3 amino acid insert in the flagellar basal body rod protein FlgC which is an important part of the unique endoflagellar structure shared by Spirochaetes species.[15] Given that the CSI is exclusively shared by members within this phylum, it has been postulated that it may be related to the characteristic flagellar properties observed among Spirochaetes species.[12][15] Historically, the all families belonging to the Spirochaetes phylum were assigned to a single order, the Spirochaetales.[5][6] However, the current taxonomic view is more connotative of accurate evolutionary relationships, where the distribution of conserved signature indels are indicative of shared ancestry within each respective clade for which they are specific. These synapomorphic characteristics justify the phylogenetic divisions and are a means to identify each order/family within the phylum.[12] Phylogeny[edit] The phylogeny is based on 16S rRNA-based LTP release 123 by 'The All-Species Living Tree' Project.[16]

Leptospirales

Leptospiraceae

Turneriella parva (Hovind-Hougen et al. 1982) Levett et al. 2005

Leptonema illini Hovind-Hougen 1983

Leptospira
Leptospira
Noguchi 1917 emend. Faine and Stallman 1982

Euspirochaetae

Brevinemataceae

Brevinema andersonii
Brevinema andersonii
Defosse et al. 1995

Brachyspiraceae

Brachyspira Hovind-Hougen et al. 1982

Spirochaetales

Exilispira thermophila Imachi et al. 2008

Spirochaetaceae

Spirochaeta Ehrenberg 1835 emend. Pikuta et al. 2009

Treponema
Treponema
Schaudinn 1905 emend Abt et al. 2013

Spirochaeta clade 2

Spirochaeta aurantia (ex Vinzent 1926) Canale-Parola 1980

Borreliaceae

Borreliella Adeolu & Gupta 2015 ( Lyme disease
Lyme disease
Borrelia)

Borrelia
Borrelia
Swellengrebel 1907 emend. Adeolu & Gupta 2014 (relapsing fever Borrelia)

Taxonomy[edit] The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN)[17] and National Center for Biotechnology Information (NCBI).[18]

Phylum Spirochaetes Garrity & Holt 2001 [Spirochaetae Cavalier-Smith 2002; Spirochaetaeota Oren et al. 2015]

Class Spirochaetae Cavalier-Smith 2002 ["Protozoobacteriales"; "Spirochaetia" Paster 2011]

Order Leptospirales Gupta et al. 2014

Family Leptospiraceae Hovind-Hougen 1979 emend. Levett et al. 2005

Genus Leptonema Hovind-Hougen 1983 Genus Leptospira
Leptospira
Noguchi 1917 emend. Faine and Stallman 1982 Genus Turneriella Levett et al. 2005

Order Brachyspirales corrig. Gupta et al. 2014

Family Brachyspiraceae Paster 2012

Genus Brachyspira Hovind-Hougen et al. 1982

Order Brevinematales Gupta et al. 2014

Family Brevinemataceae Paster 2012

Genus Brevinema Defosse et al. 1995

Order Spirochaetales
Spirochaetales
Buchanan 1917 emend. Gupta et al. 2013

Genus Exilispira Imachi et al. 2008 Family Borreliaceae Gupta et al. 2014

Genus Borreliella Adeolu & Gupta 2015 [ Borrelia
Borrelia
burgdorferi species-group] ( Lyme disease
Lyme disease
Borrelia) Genus Borrelia
Borrelia
Swellengrebel 1907 emend. Adeolu & Gupta 2014 (relapsing fever Borrelia) Genus Cristispira pectinis ♪ Gross 1910

Family Spirochaetaceae Swellengrebel 1907

Genus ? Clevelandina reticulitermitidis ♦ Bermudes et al. 1988 Genus ? Diplocalyx calotermitidis ♦ (ex Gharagozlou 1968) Bermudes et al. 1988 Genus ? Hollandina pterotermitidis ♦ (ex To et al. 1978) Bermudes et al. 1988 Genus ? Pillotina calotermitidis ♦ (ex Hollande and Gharagozlou 1967) Bermudes et al. 1988 Genus ?Spironema culicis ♠ Turk et al. 1999 Genus Spirochaeta Ehrenberg 1835 emend. Pikuta et al. 2009 ["Ehrenbergia" Gieszczykiewiez 1939 non Spreng.; incl. Sphaerochaeta Ritalahti et al. 2012 emend. Miyazaki et al. 2014; Salinispira Ben Hania et al. 2015] Genus Treponema
Treponema
Schaudinn 1905 emend Abt et al. 2013 ["Spironema" Vuillemin 1905 non Klebs 1892; "Microspironema" Stiles & Pfender 1905]

Notes: ♦ Type strain lost or not available ♪ Prokaryotes where no pure (axenic) cultures are isolated or available, i. e. not cultivated or can not be sustained in culture for more than a few serial passages ♠ Strains found at the National Center for Biotechnology Information (NCBI) but not listed in the List of Prokaryotic names with Standing in Nomenclature (LSPN) See also[edit]

Bacteriology Borrelia Flagellum Lyme disease
Lyme disease
microbiology Pinta (disease) Prokaryote Treponema
Treponema
pallidum Brevinema andersonii Syphilis Morgellons

References[edit]

^ spirochaete - Oxford Dictionaries ^ Elsevier, Dorland's Illustrated Medical Dictionary, Elsevier.  ^ Ryan KJ; Ray CG (editors) (2004). Sherris Medical Microbiology (4th ed.). McGraw Hill. ISBN 0-8385-8529-9. CS1 maint: Extra text: authors list (link) ^ Margulis, L.; Ashen, J. B.; Solé, M.; Guerrero, R. (1993-08-01). "Composite, large spirochetes from microbial mats: spirochete structure review" (PDF). Proceedings of the National Academy of Sciences. 90 (15): 6966–6970. doi:10.1073/pnas.90.15.6966. ISSN 0027-8424. PMC 47056 . PMID 8346204.  ^ a b Paster BJ (2011) Phylum XV. Sprochaetes Garrity and Holt. In: Bergey’s Manual of Systematic Bacteriology, pp. 471. Eds D.J. Brenner, N.R.Krieg, G.M.Garrity, and J.T. Staley Springer-: New York. ^ a b Paster BJ (2011) Family I. Sprochaetes Swellengrebel 1907, 581AL. In: Bergey’s Manual of Systematic Bacteriology, pp. 473-531. Eds D.J. Brenner, N.R.Krieg, G.M.Garrity, and J.T. Staley Springer-: New York. ^ McBride A, Athanazio D, Reis M, Ko A (2005). "Leptospirosis". Current Opinion in Infectious Diseases. 18 (5): 376–86. doi:10.1097/01.qco.0000178824.05715.2c. PMID 16148523.  ^ Schwan T (1996). "Ticks and Borrelia: model systems for investigating pathogen-arthropod interactions". Infect Agents Dis. 5 (3): 167–81. PMID 8805079.  ^ Amat Villegas I, Borobio Aguilar E, Beloqui Perez R, de Llano Varela P, Oquiñena Legaz S, Martínez-Peñuela Virseda JM (January 2004). "[Colonic spirochetes: an infrequent cause of adult diarrhea]". Gastroenterol Hepatol (in Spanish). 27 (1): 21–3. PMID 14718105.  ^ Miklossy, Judith (2011-08-04). " Alzheimer's disease
Alzheimer's disease
- a neurospirochetosis. Analysis of the evidence following Koch's and Hill's criteria". Journal of Neuroinflammation. 8: 90. doi:10.1186/1742-2094-8-90. ISSN 1742-2094. PMC 3171359 . PMID 21816039.  ^ Sayers; et al. "TherSpirochaetia". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 2016-10-25.  ^ a b c d e f Gupta RS, Mahmood S, Adeolu M (2013). "A phylogenomic and molecular signature based approach for characterization of the phylum Spirochaetes and its major clades: proposal for a taxonomic revision of the phylum". Front Microbiol. 4 (217). doi:10.3389/fmicb.2013.00217. PMC 3726837 . PMID 23908650.  ^ Oren A, Garrity GM (2014). "List of new names and new combinations previously effectively, but not validly, published" (PDF). Int. J. Syst. Evol. Microbiol. 64: 693–696. doi:10.1099/ijs.0.062521-0.  ^ Adeolu M, Gupta RS (2014). "A phylogenomic and molecular marker based proposal for the division of the genus Borrelia
Borrelia
into two genera: the emended genus Borrelia
Borrelia
containing only the members of the relapsing fever Borrelia, and the genus Borreliella gen. nov. containing the members of the Lyme disease
Lyme disease
Borrelia
Borrelia
(Borrelia burgdorferi sensu lato complex)". Antonie Van Leeuwenhoek. 105 (6): 1049–1072. doi:10.1007/s10482-014-0164-x. PMID 24744012.  ^ a b Macnab RM (2003). "How bacteria assemble flagella". Annu Rev Microbiol. 57: 77–100. doi:10.1146/annurev.micro.57.030502.090832. PMID 12730325.  ^ 'The All-Species Living Tree' Project."16S rRNA-based LTP release 123 (full tree)" (PDF). Silva Comprehensive Ribosomal RNA Database. Retrieved 2013-03-20.  ^ J.P. Euzéby. "Spirochaetes". List of Prokaryotic names with Standing in Nomenclature (LPSN). Retrieved 2013-03-20.  ^ Sayers; et al. "Spirochaetes". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 2013-03-20. 

External links[edit]

Introduction to the Spirochetes University of California Museum of Paleontology (UCMP)

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Prokaryotes: Bacteria
Bacteria
classification (phyla and orders)

Domain Archaea Bacteria Eukaryota (Supergroup Plant Hacrobia Heterokont Alveolata Rhizaria Excavata Amoebozoa Opisthokonta

Animal Fungi)

G-/ OM

Terra-/ Glidobacteria (BV1)

Eobacteria

Deinococcus–Thermus

Deinococcales Thermales

Chloroflexi

Anaerolineales Caldilineales Chloroflexales Herpetosiphonales Dehalococcoidales Ktedonobacterales Thermogemmatisporales Thermomicrobiales Sphaerobacterales

other glidobacteria

Thermodesulfobacteria thermophiles

Aquificae Thermotogae

Cyanobacteria

Proteobacteria (BV2)

Alpha

Caulobacterales Kiloniellales Kordiimonadales Magnetococcales Parvularculales Rhizobiales Rhodobacterales Rhodospirillales Rickettsiales Sneathiellales Sphingomonadales

Beta

Burkholderiales Hydrogenophilales Methylophilales Neisseriales Nitrosomonadales Procabacteriales Rhodocyclales

Gamma

Acidithiobacillales Aeromonadales Alteromonadales Cardiobacteriales Chromatiales Enterobacteriales Legionellales Methylococcales Oceanospirillales Orbales Pasteurellales Pseudomonadales Salinisphaerales Thiotrichales Vibrionales Xanthomonadales

Delta

Bdellovibrionales Desulfarculales Desulfobacterales Desulfovibrionales Desulfurellales Desulfuromonadales Myxococcales Syntrophobacterales Syntrophorhabdales

Epsilon

Campylobacterales Nautiliales

Zeta

Mariprofundales

BV4

Spirochaetes

Spirochaetes

Sphingobacteria (FCB group)

Fibrobacteres Chlorobi

Chlorobiales Ignavibacteriales

Bacteroidetes

Bacteroidales Cytophagales Flavobacteriales Sphingobacteriales

Planctobacteria/ (PVC group)

Chlamydiae Lentisphaerae

Lentisphaerales Oligosphaerales Victivallales

Planctomycetes

Phycisphaerales Planctomycetales

Verrucomicrobia

Puniceicoccales Opitutales Chthoniobacterales Verrucomicrobiales

"Poribacteria"

Other GN

Acidobacteria

Acidobacteriales Acanthopleuribacterales Holophagales Solibacterales

Armatimonadetes

Armatimonadales Chthonomonadales Fimbriimonadales

Caldiserica Chrysiogenetes Deferribacteres Dictyoglomi Elusimicrobia Fusobacteria Gemmatimonadetes Nitrospirae Synergistetes

G+/ no OM

Firmicutes (BV3)

Bacilli

Bacillales Lactobacillales

Clostridia

Clostridiales Halanaerobiales Thermoanaerobacteriales Natranaerobiales

Erysipelotrichia

Erysipelotrichiales

Thermolithobacteria

Thermolithobacterales

Tenericutes/ Mollicutes

Mycoplasmatales Entomoplasmatales Anaeroplasmatales Acholeplasmatales Haloplasmatales

Negativicutes

Selenomonadales

Actinobacteria (BV5)

Actinobacteria

Actinomycetales Bifidobacteriales

Acidimicrobiia

Acidimicrobiales

Coriobacteriidae

Coriobacteriales

Nitriliruptoria

Euzebyales Nitriliruptorales

Rubrobacteria

Gaiellales Rubrobacterales Thermoleophilales Solirubrobacterales

Incertae sedis

†Archaeosphaeroides †Eobacterium †Leptotrichites

Source: Bergey's Manual (2001–2012). Alternative views: Wikispecies.

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Infectious diseases Bacterial diseases: BV4 non-proteobacterial G- (primarily A00–A79, 001–041, 080–109)

Spirochaete

Spirochaetaceae

Treponema

Treponema
Treponema
pallidum

Syphilis/bejel Yaws

Treponema
Treponema
carateum (Pinta) Treponema
Treponema
denticola

Borrelia

Borrelia
Borrelia
burgdorferi/ Borrelia
Borrelia
afzelii

Lyme disease Erythema chronicum migrans Neuroborreliosis

Borrelia recurrentis (Louse borne relapsing fever) Borrelia
Borrelia
hermsii/ Borrelia
Borrelia
duttoni/ Borrelia
Borrelia
parkeri (Tick borne relapsing fever)

Leptospiraceae

Leptospira

Leptospira
Leptospira
interrogans (Leptospirosis)

Chlamydiaceae

Chlamydophila

Chlamydophila psittaci (Psittacosis) Chlamydophila pneumoniae

Chlamydia

Chlamydia trachomatis

Chlamydia Lymphogranuloma venereum Trachoma

Bacteroidetes

Bacteroides fragilis Tannerella forsythia Capnocytophaga canimorsus Porphyromonas gingivalis Prevotella intermedia

Fusobacteria

Fusobacterium necrophorum (Lemierre's syndrome) Fusobacterium nucleatum Fusobacterium polymorphum

Streptobacillus moniliformis (Rat-bite fever/Haverhill fever)

Taxon identifiers

Wd: Q126

.