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File:Spirochete Cross Section.png
Cross section of a spirochaete cell
File:Endoflagella Components.pdf
Endoflagella Components. Legend: Fig. 1: A cross-section of a typical spirochete cell showing endoflagella located in the periplasm between the inner cytoplasmic membrane and the outer membrane. Periplasm, consisting of a gel-like matrix, provides a semi-stable medium to secure endoflagella during rotation. The axial filament, indicated in red, is composed of bundles of endoflagella.
Fig. 2: A side-view of a spirochete cell which shows two axial filaments in opposing motion. One axial filament rotates in a clockwise orientation; an adjacent axial filament rotates in a counter-clockwise orientation. Rotation of the endoflagella creates torsion and drives the corkscrew rotation of the cell.
Fig. 3: An expanded view of the cellular membranes that surround endoflagellum. Both the inner and outer membrane contain a phospholipid bi-layer, with non-polar fatty acid chains in-ward of polar phosphorus heads. Peptidoglycan, the cell wall, provides structure in bacterial microorganisms. Axial filaments are superior to the peptidoglycan.

A spirochaete (Template:IPAc-en)<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> or spirochete is a member of the phylum Spirochaetota (also called Spirochaetes<ref name="Dorlands">Template:Cite book</ref> Template:IPAc-en), which contains distinctive diderm (double-membrane) Gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or spiraled, hence the name) cells.<ref name=Sherris>Template:Cite book</ref> Spirochaetes are chemoheterotrophic in nature, with lengths between 3 and 500 μm and diameters around 0.09 to at least 3 μm.<ref>Template:Cite journal</ref>

Spirochaetes are distinguished from other bacterial phyla by the location of their flagella, called endoflagella, or periplasmic flagella, which are sometimes called axial filaments.<ref name="Nakamura">Template:Cite journal</ref><ref name="McGraw">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Endoflagella are anchored at each end (pole) of the bacterium within the periplasmic space (between the inner and outer membranes) where they project backwards to extend the length of the cell.<ref name="Brock">Template:Cite book</ref> These cause a twisting motion which allows the spirochaete to move. When reproducing, a spirochaete will undergo asexual transverse binary fission. Most spirochaetes are free-living and anaerobic, but there are numerous exceptions. Spirochaete bacteria are diverse in their pathogenic capacity and the ecological niches that they inhabit, as well as molecular characteristics including guanine-cytosine content and genome size.<ref name="Paster_2011">Template:Cite book</ref><ref name="Paster_2011b">Template:Cite book</ref>

PathogenicityEdit

Many organisms within the Spirochaetota phylum cause prevalent diseases. Pathogenic members of this phylum include the following:

Salvarsan, the first partially organic synthetic antimicrobial drug in medical history, was effective against spirochaetes and primarily used to cure syphilis. Additionally, oral spirochaetes are known to play a significant role in the pathogenesis of human periodontal disease.<ref>Template:Cite journal</ref>

Taxonomy and molecular signaturesEdit

The class currently consists of 14 validly named genera across four orders and five families.<ref name=NCBI>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="pmid23908650">Template:Cite journal</ref><ref name="Oren & Garrity ">Template:Cite journal</ref> The orders Brachyspirales, Brevinematales and Leptospirales each contain a single family, Brachyspiraceae, Brevinemataceae and Leptospiraceae, respectively. The 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.<ref name="pmid23908650"/> 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 order.<ref name="pmid23908650"/> 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 emend. Borreliella gen. nov.).<ref name=pmid24744012>Template:Cite journal</ref> However, this study has been criticized, and other studies using different approaches do not support the proposed split.<ref name="Winslow2019">Template:Cite journal</ref> The new naming system for the Lyme and relapsing fever Borrelia has not been adopted by the scientific literature.<ref name="Winslow2019"/>

A CSI has also been found exclusively shared by all Spirochaetota species.<ref name="pmid23908650"/> This CSI is a three-amino-acid insert in the flagellar basal body rod protein FlgC which is an important part of the unique endoflagellar structure shared by Spirochaetota species.<ref name=Macnab03>Template:Cite journal</ref> 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 Spirochaetota species.<ref name="pmid23908650"/><ref name=Macnab03/>

Historically, all families belonging to the Spirochaetota phylum were assigned to a single order, the Spirochaetales.<ref name="Paster_2011"/><ref name="Paster_2011b"/> However, the current taxonomic view is more connotative of accurate evolutionary relationships. The distribution of a CSI is indicative of shared ancestry within the clade for which it is specific. It thus functions as a synapomorphic characteristic, so that the distributions of different CSIs provide the means to identify different orders and families within the phylum and so justify the phylogenetic divisions.<ref name="pmid23908650"/>

PhylogenyEdit

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TaxonomyEdit

The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN)<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and National Center for Biotechnology Information (NCBI).<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

  • Phylum Spirochaetota Garrity and Holt 2021
    • Genus ?"Spirosymplokos" Guerrero et al. 1993
    • Class Leptospiria Chuvochina et al. 2024
      • Order Turneriellales Chuvochina et al. 2024
        • Family Turneriellaceae Chuvochina et al. 2024
      • Order Leptospirales Gupta et al. 2014
    • Class Brachyspiria Chuvochina et al. 2024
    • Class Brevinematia Chuvochina et al. 2024
      • Order Brevinematales Gupta et al. 2014
        • Family "Longinemataceae" Karnachuk et al. 2021
          • Genus ?"Longinema" corrig. Karnachuk et al. 2021
        • Family Brevinemataceae Paster 2012
        • Family Thermospiraceae Ben Ali Gam et al. 2023
    • Class Spirochaetia Paster 2020
      • Order Spirochaetales Buchanan 1917
      • Order "Exilispirales" Pallen, Rodriguez-R & Alikhan 2022
        • Family "Exilispiraceae" Pallen, Rodriguez-R & Alikhan 2022
      • Order Borreliales Chuvochina et al. 2024
        • Family Borreliaceae Gupta et al. 2014
          • Genus Borrelia Swellengrebel 1907 (relapsing fever Borrelia, reptile-associated Borrelia and Echidna-associated Borrelia)
          • Genus Borreliella Adeolu & Gupta 2015 (Lyme disease Borrelia)
          • Genus ?Cristispira Gross 1910
      • Order "Entomospirales" Pallen, Rodriguez-R & Alikhan 2022
        • Family "Entomospiraceae" Pallen, Rodriguez-R & Alikhan 2022
          • Genus Entomospira Grana-Miraglia et al. 2024 non Enderlein 1917
      • Order "Marispirochaetales" Pallen, Rodriguez-R & Alikhan 2022
        • Family Marispirochaetaceae Chuvochina et al. 2024
      • Order "Sediminispirochaetales" Pallen, Rodriguez-R & Alikhan 2022
        • Family Sediminispirochaetaceae Chuvochina et al. 2024
      • Order Sphaerochaetales Chuvochina et al. 2024
      • Order Treponematales Chuvochina et al. 2024

See alsoEdit

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ReferencesEdit

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External linksEdit

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