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Spirochaete
Phylum of bacteria
This article is about the bacterial phylum. For spiral-shaped bacteria in general, see spiral bacteria.
Spirochaetes
Treponema pallidum, a spirochaete which causes syphilis
Scientific classification
Domain:
Bacteria
Kingdom:
Pseudomonadati
Phylum:
Spirochaetota Garrity and Holt 2021[3]
Class:
Spirochaetia Paster 2020[1]: 471–563 [2]
Orders
Brachyspirales
Brevinematales
Leptospirales
Spirochaetales
"Treponematales"
Synonyms
Spirochaetota:
"Protozoobacteriales" Prévot 1958
"Spirochaetae" Cavalier-Smith 2002
"Spirochaetaeota" Oren et al. 2015
"Spirochaetes" Garrity and Holt 2001
"Spirochaetota" Whitman et al. 2018
Spirochaetia:
Spirochaetes Cavalier-Smith 2002
"Spirochaetia" Cavalier-Smith 2020
"Leptospiria" Cavalier-Smith 2020
Cross section of a spirochaete cellEndoflagella 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 (/ˈspaɪroʊˌkiːt/)[4] or spirochete is a member of the phylum Spirochaetota (also called Spirochaetes[5]/ˌspaɪroʊˈkiːtiːz/), which contains distinctive diderm (double-membrane) Gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or spiraled, hence the name) cells.[6] Spirochaetes are chemoheterotrophic in nature, with lengths between 3 and 500 μm and diameters around 0.09 to at least 3 μm.[7]
Spirochaetes are distinguished from other bacterial phyla by the location of their flagella, called endoflagella, or periplasmic flagella, which are sometimes called axial filaments.[8][9] 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.[10] 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.[11][12]
Pathogenicity
[edit]
Many organisms within the Spirochaetota phylum cause prevalent diseases. Pathogenic members of this phylum include the following:
Leptospira species, which causes leptospirosis[13]
Borrelia burgdorferi, B. mayonii, B. bissettiae, B. garinii, B. afzelii, B. spielmanii, B. lusitaniae, which cause Lyme disease.[14]
Borrelia recurrentis, which causes relapsing fever[15]
Treponema pallidum subspecies which cause treponematoses such as syphilis and yaws.
Brachyspira pilosicoli and Brachyspira aalborgi, which cause intestinal spirochaetosis[16]
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.[17]
Taxonomy and molecular signatures
[edit]
The class currently consists of 14 validly named genera across four orders and five families.[18][19][20] 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.[19] 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.[19] 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.).[21] However, this study has been criticized, and other studies using different approaches do not support the proposed split.[22] The new naming system for the Lyme and relapsing fever Borrelia has not been adopted by the scientific literature.[22]
A CSI has also been found exclusively shared by all Spirochaetota species.[19] 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.[23] 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.[19][23]
Historically, all families belonging to the Spirochaetota phylum were assigned to a single order, the Spirochaetales.[11][12] 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.[19]
Phylogeny
[edit]
See also: Bacterial taxonomy
16S rRNA based LTP_10_2024[24][25][26]
120 marker proteins based GTDB 09-RS220[27][28][29]
Brevinematia
Brevinematales
Brevinemataceae
Brevinema
Thermospiraceae
Thermospira
Spirochaetia
Leptospirales
Leptospiraceae
Turneriella
Leptonema
Leptospira
"Exilispirales"
"Exilispiraceae"
Exilispira
Brachyspirales
Brachyspiraceae
Brachyspira
Spirochaetales
Sphaerochaetaceae
Pleomorphochaeta
Bullifex
Sphaerochaeta
Rectinemataceae
Rectinema
Breznakiellaceae
Gracilinema
Zuelzera
Breznakiella
Treponema primitia
Helmutkoenigia
Leadbettera
Treponemataceae
Brucepastera
Teretinema
Treponema
Treponema species-group 2
Marispirochaetaceae
Marispirochaeta
Sediminispirochaetaceae
Sediminispirochaeta
Spirochaeta thermophila
Salinispiraceae
Spirochaeta lutea
Salinispira
Spirochaetaceae
Spirochaeta
Alkalispirochaetaceae
Spirochaeta halophila
Alkalispirochaeta
Spirochaeta aurantia
Spirochaetaceae 2
Spirochaeta cellobiosiphila
Oceanispirochaeta
Thiospirochaeta
Spirochaeta species-group 2
"Entomospiraceae"
Entomospira
Borreliaceae
Borrelia
Borreliella
"Leptospirae"
Leptospiria
Turneriellales
Turneriellaceae
Turneriella
Leptospirales
Leptonemataceae
Leptonema
Leptospiraceae
Leptospira
"Euspirochaetae"
Brevinematia
Brevinematales
Brevinemataceae
Brevinema
Thermospiraceae
Thermospira
Brachyspiria
Brachyspirales
Brachyspiraceae
Brachyspira
"Exilispiria"
"Exilispirales"
"Exilispiraceae"
Exilispira
Spirochaetia
Borreliales
Borreliaceae
Borrelia
Borreliella
"Entomospirales"
"Entomospiraceae"
Entomospira
DSM‑17781
DSM‑17781
Spirochaeta cellobiosiphila
Spirochaetales_E
"Thiospirochaetaceae"
Thiospirochaeta
DSM‑2461
Spirochaeta species-group 2
Spirochaetaceae_B
Oceanispirochaeta
Spirochaetales_A
Spirochaetaceae_A
Spirochaeta thermophila
"Salinispirales"
"Salinispiraceae"
Spirochaeta lutea
Salinispira
DSM‑8902
Spirochaeta africana
"Alkalispirochaetaceae"
Alkalispirochaeta
"Marispirochaetales"
Marispirochaetaceae
Marispirochaeta
"Sediminispirochaetales"
Sediminispirochaetaceae
Sediminispirochaeta
Sphaerochaetales
Sphaerochaetaceae
Parasphaerochaeta
Sphaerochaeta
Bullifex
"Ca. Ornithospirochaeta"
Treponematales
Rectinemataceae
Rectinema
Breznakiellaceae
Gracilinema
Breznakiella
Leadbettera
Treponema primitia
Treponemataceae
"Ca. Gallitreponema"
Treponema species-group 2
Brucepastera
Teretinema
"Ca. Avitreponema"
Treponema
Taxonomy
[edit]
The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN)[30] and National Center for Biotechnology Information (NCBI).[31]
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
Genus Turneriella Levett et al. 2005
Order Leptospirales Gupta et al. 2014
Family Leptonemataceae Chuvochina et al. 2024
Genus Leptonema Hovind-Hougen 1983
Family Leptospiraceae Hovind-Hougen 1979
Genus Leptospira Noguchi 1917
Class Brachyspiria Chuvochina et al. 2024
Order Brachyspirales corrig. Gupta et al. 2014
Family Brachyspiraceae Paster 2012
Genus ?"Ca. Maribrachyspira" Matsuyama et al. 2017
Genus Brachyspira Hovind-Hougen et al. 1982 non Foliella non Pfeiffer 1855
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
Genus Brevinema Defosse et al. 1995
Family Thermospiraceae Ben Ali Gam et al. 2023
Genus Thermospira Ben Ali Gam et al. 2023
Class Spirochaetia Paster 2020
Order Spirochaetales Buchanan 1917
Family "Pillotinaceae" Margulis & Hinkle 1992
Genus ?Pillotina Hollande and Gharagozlou 1967 ex Bermudes et al. 1988
Family Spirochaetaceae Swellengrebel 1907 ["Thiospirochaetaceae" Pallen, Rodriguez-R & Alikhan 2022]
Genus ?"Ca. Allospironema" Paster & Dewhirst 2000
Genus ?"Canaleparolina" Wier, Ashen & Margulis 2000
Genus ?Clevelandina Bermudes et al. 1988
Genus ?Diplocalyx Gharagozlou 1968 ex Bermudes et al. 1988 non Richard 1850 non Presl 1845
Genus ?"Candidatus Haliotispira" Sharma et al. 2024
Genus ?Hollandina To et al. 1978 ex Bermudes et al. 1988 non Haynes 1956
Genus ?"Mobilifilum" Margulis et al. 1990
Genus Oceanispirochaeta Subhash & Lee 2017b
Genus ?Spirochaeta Ehrenberg 1835 em. Pikuta et al. 2009 non Turczaninow 1851
Genus ?"Candidatus Thalassospirochaeta" Pragya et al. 2024
Genus Thiospirochaeta Dubinina et al. 2020
Family Alkalispirochaetceae Chuvochina et al. 2024
Genus Alkalispirochaeta Sravanthi et al. 2016
Family Salinispiraceae Chuvochina et al. 2024
Genus Salinispira Ben Hania et al. 2015
Order "Exilispirales" Pallen, Rodriguez-R & Alikhan 2022
Family "Exilispiraceae" Pallen, Rodriguez-R & Alikhan 2022
Genus Exilispira Imachi et al. 2008
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
Genus Marispirochaeta Shivani et al. 2017
Order "Sediminispirochaetales" Pallen, Rodriguez-R & Alikhan 2022
Family Sediminispirochaetaceae Chuvochina et al. 2024
Genus Sediminispirochaeta Shivani et al. 2016
Order Sphaerochaetales Chuvochina et al. 2024
Family Sphaerochaetaceae Hördt et al. 2020
Genus Bullifex Wylensek et al. 2021 [incl. "Ca. Aphodenecus" Gilroy et al. 2021]
Genus "Ca. Ornithospirochaeta" Gilroy et al. 2021
Genus Parasphaerochaeta Bidzhieva et al. 2020
Genus "Ca. Physcosoma" Gilroy et al. 2022
Genus Pleomorphochaeta Arroua et al. 2016
Genus Sphaerochaeta Ritalahti et al. 2012
Order Treponematales Chuvochina et al. 2024
Family Rectinemataceae Brune et al. 2022
Genus Rectinema Koelschbach et al. 2017
Family Breznakiellaceae Brune et al. 2022
Genus Breznakiella Song et al. 2022
Genus Gracilinema Brune et al. 2022
Genus Helmutkoenigia Brune et al. 2022
Genus Leadbettera Brune et al. 2022
Genus Zuelzera Brune et al. 2022
Family Treponemataceae Robinson 1948
Genus "Ca. Avitreponema" Gilroy et al. 2021
Genus Brucepastera Song et al. 2023
Genus "Ca. Gallitreponema" Gilroy et al. 2021
Genus Teretinema Song et al. 2023
Genus Treponema Schaudinn 1905 em. Abt et al. 2013
See also
[edit]
List of bacteria genera
List of bacterial orders
Bacteriology
Borrelia
Brevinema andersonii
Flagellum
Lyme disease microbiology
Pinta (disease)
Prokaryote
Syphilis
Treponema pallidum
Yaws
References
[edit]
^Paster BJ (2010). "Class I. Spirochaetia class. nov.". In Krieg NR, Staley JT, Brown DR, Hedlund BP, Paster BJ (eds.). Bergey's Manual of Systematic Bacteriology. Vol. 4—The Bacteroidetes, Spirochaetes, Tenericutes (Mollicutes), Acidobacteria, Fibrobacteres, Fusobacteria, Dictyoglomi, Gemmatimonadetes, Lentisphaerae, Verrucomicrobia, Chlamydiae, and Planctomycetes (2nd ed.). New York, NY: Springer. doi:10.1007/978-0-387-68572-4. ISBN 978-0-387-95042-6.
^Oren A, Garrity GM (2020). "Validation list no. 195. List of new names and new combinations previously effectively, but not validly, published". Int J Syst Evol Microbiol. 70 (9): 4844–4847. doi:10.1099/ijsem.0.004366. PMID 32993851. S2CID 222147003.
^Oren A, Garrity GM (2021). "Valid publication of the names of forty-two phyla of prokaryotes". Int J Syst Evol Microbiol. 71 (10): 5056. doi:10.1099/ijsem.0.005056. PMID 34694987. S2CID 239887308.
^"SPIROCHAETE | Meaning & Definition for UK English | Lexico.com". Lexico Dictionaries | English. Archived from the original on 27 January 2021.
^Dorland's Illustrated Medical Dictionary. Elsevier.
^Ryan KJ, Ray CG, eds. (2004). Sherris Medical Microbiology (4th ed.). McGraw Hill. ISBN 978-0-8385-8529-0.
^Margulis L, Ashen JB, Solé M, Guerrero R (August 1993). "Composite, large spirochetes from microbial mats: spirochete structure review". Proceedings of the National Academy of Sciences of the United States of America. 90 (15): 6966–6970. Bibcode:1993PNAS...90.6966M. doi:10.1073/pnas.90.15.6966. PMC 47056. PMID 8346204.
^Nakamura S (April 2020). "Spirochete Flagella and Motility". Biomolecules. 10 (4): 550. doi:10.3390/biom10040550. PMC 7225975. PMID 32260454.
^Carroll KC, Hobden JA, Miller S (2019). "Spirochetes and Other Spiral Microorganisms". Jawetz, Melnick, & Adelberg’s Medical Microbiology. McGraw-Hill Education. Retrieved 9 May 2021.
^Madigan MT (2019). Brock biology of microorganisms (Fifteenth, Global ed.). NY, NY: Pearson. p. 519. ISBN 9781292235103.
^ abPaster BJ (2011). "Phylum XV. Spirochaetes Garrity and Holt.". In Brenner DJ, Krieg NR, Garrity GM, Staley JT (eds.). Bergey's Manual of Systematic Bacteriology. New York: Springer. p. 471.
^ abPaster BJ (2011). "Family I. Sprochaetes Swellengrebel 1907, 581AL.". In Brenner DJ, Krieg NR, Garrity GM, Staley JT (eds.). Bergey's Manual of Systematic Bacteriology. New York: Springer. pp. 473–531.
^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. S2CID 220576544.
^Wolcott KA, Margos G, Fingerle V, Becker NS (September 2021). "Host association of Borrelia burgdorferi sensu lato: A review". Ticks and Tick-Borne Diseases. 12 (5): 101766. doi:10.1016/j.ttbdis.2021.101766. PMID 34161868.
^Schwan TG (June 1996). "Ticks and Borrelia: model systems for investigating pathogen-arthropod interactions". Infectious Agents and Disease. 5 (3): 167–181. 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. doi:10.1016/s0210-5705(03)70440-3. PMID 14718105.
^Schoch CL, Ciufo S, Domrachev M, Hotton CL, Kannan S, Khovanskaya R, et al. "TherSpirochaetia". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 25 October 2016.
^ abcdefGupta 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): 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 (3): 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 into two genera: the emended genus Borrelia containing only the members of the relapsing fever Borrelia, and the genus Borreliella gen. nov. containing the members of the Lyme disease Borrelia (Borrelia burgdorferi sensu lato complex)". Antonie van Leeuwenhoek. 105 (6): 1049–1072. doi:10.1007/s10482-014-0164-x. PMID 24744012.
^ abWinslow C, Coburn J (2019). "Recent discoveries and advancements in research on the Lyme disease spirochete Borrelia burgdorferi". F1000Research. 8: 763. doi:10.12688/f1000research.18379.1. PMC 6545822. PMID 31214329.
^"LTP_all tree in newick format". Retrieved 10 December 2024.
^"LTP_10_2024 Release Notes" (PDF). Retrieved 10 December 2024.
^"GTDB release 09-RS220". Genome Taxonomy Database. Retrieved 10 May 2024.
^"bac120_r220.sp_labels". Genome Taxonomy Database. Retrieved 10 May 2024.
^"Taxon History". Genome Taxonomy Database. Retrieved 10 May 2024.
^Euzéby JP. "Spirochaetes". List of Prokaryotic names with Standing in Nomenclature (LPSN). Retrieved 20 July 2018.
^Schoch CL, Ciufo S, Domrachev M, Hotton CL, Kannan S, Khovanskaya R, et al. "Spirochaetes". National Center for Biotechnology Information (NCBI) taxonomy database. Retrieved 20 July 2018.
External links
[edit]
Introduction to the Spirochetes University of California Museum of Paleontology (UCMP)