TheChlamydiota(synonymChlamydiae) are abacterialphylumandclasswhose members are remarkably diverse, includingpathogensof humans and animals,symbiontsof ubiquitousprotozoa,[4]and marine sediment forms not yet well understood.[5]All of the Chlamydiota that humans have known about for many decades are obligate intracellular bacteria; in 2020 many additional Chlamydiota were discovered in ocean-floor environments, and it is not yet known whether they all havehosts.[5]Historically it was believed that all Chlamydiota had apeptidoglycan-free cell wall, but studies in the 2010s demonstrated a detectable presence of peptidoglycan, as well as other important proteins.[6][7][8][9][10][11]
Chlamydiota | |
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Chlamydia trachomatis | |
Scientific classification | |
Domain: | Bacteria |
Superphylum: | PVC superphylum |
Phylum: | Chlamydiota Garrity & Holt 2021[3] |
Class: | Chlamydiia Horn 2016[1][2] |
Orders and families | |
Synonyms | |
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Among the Chlamydiota, all of the ones long known to science grow only by infectingeukaryotichost cells. They are as small as or smaller than manyviruses.They are ovoid in shape and stainGram-negative.They are dependent on replication inside the host cells; thus, some species are termed obligateintracellular pathogensand others are symbionts of ubiquitous protozoa. Most intracellular Chlamydiota are located in aninclusion bodyorvacuole.Outside cells, they survive only as an extracellular infectious form.
These Chlamydiota can grow only where their host cells grow, and develop according to a characteristic biphasic developmental cycle.[12][13][14]Therefore,clinically relevantChlamydiotacannot be propagatedin bacterial culture media in the clinical laboratory. They are most successfully isolated while still inside their host cells.
Of various Chlamydiota that cause human disease, the two most important species areChlamydia pneumoniae,which causes a type ofpneumonia,andChlamydia trachomatis,which causeschlamydia.Chlamydia is the most common bacterialsexually transmitted infectionin the United States, and 2.86 million chlamydia infections are reported annually.
History
editChlamydia-like disease affecting the eyes of people was first described in ancient Chinese and Egyptian manuscripts. A modern description of chlamydia-like organisms was provided by Halberstaedrrter andvon Prowazekin 1907.
Chlamydial isolates cultured in the yolk sacs of embryonating eggs were obtained from a humanpneumonitisoutbreak in the late 1920s and early 1930s, and by the mid-20th century, isolates had been obtained from dozens of vertebrate species. The term chlamydia (a cloak) appeared in the literature in 1945, although other names continued to be used, including Bedsonia, Miyagawanella, ornithosis-, TRIC-, and PLT-agents. In 1956,Chlamydia trachomatiswas first cultured byTang Fei-fan,though they were not yet recognized as bacteria.[15]
Nomenclature
editIn 1966, Chlamydiota were recognized as bacteria and the genusChlamydiawas validated.[16]The orderChlamydialeswas created by Storz and Page in 1971. The classChlamydiiawas recently validly published.[17][18][19]Between 1989 and 1999, new families, genera, and species were recognized. The phylum Chlamydiae was established inBergey's Manual of Systematic Bacteriology.[20]By 2006, genetic data for over 350 chlamydial lineages had been reported.[21]Discovery of ocean-floor forms reported in 2020 involves newclades.[5]In 2022 the phylum was renamed Chlamydiota.[3]
Taxonomy and molecular signatures
editThe Chlamydiota currently contain eight validly named genera, and 14 genera.[22]The phylum presently consist of two orders (Chlamydiales, Parachlamydiales) and nine families within a single class (Chlamydiia).[17][18]Only four of these families are validly named (Chlamydiaceae,Parachlamydiaceae,Simkaniaceae,Waddliaceae)[23][24]while five are described as families (Clavichlamydiaceae,Criblamydiaceae,Parilichlamydiaceae,Piscichlamydiaceae,andRhabdochlamydiaceae).[25][26][27]
The Chlamydiales order as recently described contains the families Chlamydiaceae, and theClavichlamydiaceae,while the new Parachlamydiales order harbors the remaining seven families.[17]This proposal is supported by the observation of two distinct phylogenetic clades that warrant taxonomic ranks above the family level. Molecular signatures in the form ofconserved indels(CSIs) and proteins (CSPs) have been found to be uniquely shared by each separate order, providing a means of distinguishing each clade from the other and supporting the view of shared ancestry of the families within each order.[17][28]The distinctness of the two orders is also supported by the fact that no CSIs were found among any other combination of families.
Molecular signatureshave also been found that are exclusive for the family Chlamydiaceae.[17][28]The Chlamydiaceae originally consisted of one genus,Chlamydia,but in 1999 was split into two genera,ChlamydophilaandChlamydia.The genera have since 2015 been reunited where species belonging to the genus Chlamydophila have been reclassified as Chlamydia species.[29][30]
However, CSIs and CSPs have been found specifically for Chlamydophila species, supporting their distinctness from Chlamydia, perhaps warranting additional consideration of two separate groupings within the family.[17][28]CSIs and CSPs have also been found that are exclusively shared by all Chlamydia that are further indicative of a lineage independent from Chlamydophila, supporting a means to distinguish Chlamydia species from neighbouring Chlamydophila members.
Phylogenetics
editThe Chlamydiota form a unique bacterial evolutionary group that separated from other bacteria about a billion years ago, and can be distinguished by the presence of several CSIs and CSPs.[17][28][31][14]The species from this group can be distinguished from all other bacteria by the presence of conservedindelsin a number of proteins and by large numbers of signature proteins that are uniquely present in different Chlamydiae species.[32][33]
Reports have varied as to whether the Chlamydiota are related to thePlanctomycetotaorSpirochaetota.[34][35]Genome sequencing, however, indicates that 11% of the genes inProtochlamydia amoebophilaUWE25 and 4% in the Chlamydiaceae are most similar tochloroplast,plant,andcyanobacterialgenes.[14]Cavalier-Smithhas postulated that the Chlamydiota fall into the clade Planctobacteria in the larger cladeGracilicutes.However, phylogeny and shared presence of CSIs in proteins that are lineage-specific indicate that theVerrucomicrobiotaare the closest free-living relatives of these parasitic organisms.[36]Comparison ofribosomal RNAgenes has provided aphylogenyof known strains within Chlamydiota.[21]
Human pathogens and diagnostics
editThree species of Chlamydiota that commonly infect humans are described:
- Chlamydia trachomatis,which causes the eye-diseasetrachomaand the sexually transmitted infectionchlamydia
- Chlamydophila pneumoniae,which causes a form ofpneumonia
- Chlamydophila psittaci,which causespsittacosis
The unique physiological status of the Chlamydiota including their biphasic lifecycle and obligation to replicate within a eukaryotic host has enabled the use of DNA analysis for chlamydial diagnostics.[37]Horizontal transferof genes is evident and complicates this area of research. In one extreme example, two genes encoding histone-like H1 proteins of eukaryotic origin have been found in the prokaryotic genome of C. trachomatis, an obligateintracellular pathogen.
Phylogeny
edit16S rRNA basedLTP_08_2023[38][39][40] | 120 marker proteins basedGTDB08-RS214[41][42][43] | |||||||||||||||||||||||||||||||||||||||
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Taxonomy
editThe currently accepted taxonomy is based on theList of Prokaryotic names with Standing in Nomenclature(LPSN)[44]andNational Center for Biotechnology Information(NCBI)[45]
- "Similichlamydiales"Pallen, Rodriguez-R & Alikhan 2022[Hat2]
- Family "Piscichlamydiaceae"Horn 2010
- Family "Parilichlamydiaceae"Stride et al. 2013[ "Similichlamydiaceae"Pallen, Rodriguez-R & Alikhan 2022]
- OrderChlamydialesStorz & Page 1971
- Family "Actinochlamydiaceae"Steigen et al. 2013
- Family "Criblamydiaceae"Thomas, Casson & Greub 2006
- FamilyChlamydiaceaeRake 1957[ "Clavichlamydiaceae"Horn 2011]
- FamilyParachlamydiaceaeEverett, Bush & Andersen 1999
- FamilyRhabdochlamydiaceaeCorsaro et al. 2009
- FamilySimkaniaceaeEverett, Bush & Andersen 1999
- FamilyWaddliaceaeRurangirwa et al. 1999
See also
editReferences
edit- ^Horn M. (2010). "Class I.Chlamydiiaclass. nov. ". In Krieg NR, Staley JT, Brown DR, Hedlund BP, Paster BJ, Ward NL, Ludwig W, Whitman WB. (eds.).Bergey's Manual of Systematic Bacteriology.Vol. 4 (2nd ed.). New York, NY: Springer. p. 844.doi:10.1007/978-0-387-68572-4.ISBN978-0-387-95042-6.
- ^Oren A, Garrity GM. (2016)."Validation list no. 170. List of new names and new combinations previously effectively, but not validly, published".Int J Syst Evol Microbiol.66(7): 2463–2466.doi:10.1099/ijsem.0.001149.PMID27530111.
- ^abOren 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.PMID34694987.S2CID239887308.
- ^Sixt BS, Siegl A, Müller C, Watzka M, Wultsch A, Tziotis D, et al. (2013)."Metabolic features of Protochlamydia amoebophila elementary bodies—a link between activity and infectivity in Chlamydiae".PLOS Pathogens.9(8): e1003553.doi:10.1371/journal.ppat.1003553.PMC3738481.PMID23950718.
- ^abcDharamshi JE, Tamarit D, Eme L, Stairs CW, Martijn J, Homa F, et al. (March 2020)."Marine Sediments Illuminate Chlamydiae Diversity and Evolution".Current Biology.30(6): 1032–1048.e7.Bibcode:2020CBio...30E1032D.doi:10.1016/j.cub.2020.02.016.PMID32142706.S2CID212423997.
- ^Pilhofer M, Aistleitner K, Biboy J, Gray J, Kuru E, Hall E, et al. (2013-12-02)."Discovery of chlamydial peptidoglycan reveals bacteria with murein sacculi but without FtsZ".Nature Communications.4(1): 2856.Bibcode:2013NatCo...4.2856P.doi:10.1038/ncomms3856.PMC3847603.PMID24292151.
- ^Jacquier N, Viollier PH, Greub G (March 2015)."The role of peptidoglycan in chlamydial cell division: towards resolving the chlamydial anomaly".FEMS Microbiology Reviews.39(2): 262–275.doi:10.1093/femsre/fuv001.PMID25670734.
- ^Malhotra M, Sood S, Mukherjee A, Muralidhar S, Bala M (September 2013)."Genital Chlamydia trachomatis: an update".The Indian Journal of Medical Research.138(3): 303–316.PMC3818592.PMID24135174.
- ^Liechti GW, Kuru E, Hall E, Kalinda A, Brun YV, VanNieuwenhze M, Maurelli AT (February 2014)."A new metabolic cell-wall labelling method reveals peptidoglycan in Chlamydia trachomatis".Nature.506(7489): 507–510.Bibcode:2014Natur.506..507L.doi:10.1038/nature12892.PMC3997218.PMID24336210.
- ^Liechti G, Kuru E, Packiam M, Hsu YP, Tekkam S, Hall E, et al. (May 2016)."Pathogenic Chlamydia Lack a Classical Sacculus but Synthesize a Narrow, Mid-cell Peptidoglycan Ring, Regulated by MreB, for Cell Division".PLOS Pathogens.12(5): e1005590.doi:10.1371/journal.ppat.1005590.PMC4856321.PMID27144308.
- ^Gupta RS (August 2011)."Origin of diderm (Gram-negative) bacteria: antibiotic selection pressure rather than endosymbiosis likely led to the evolution of bacterial cells with two membranes".Antonie van Leeuwenhoek.100(2): 171–182.doi:10.1007/s10482-011-9616-8.PMC3133647.PMID21717204.
- ^Horn M (2008). "Chlamydiae as symbionts in eukaryotes".Annual Review of Microbiology.62:113–131.doi:10.1146/annurev.micro.62.081307.162818.PMID18473699.S2CID13405815.
- ^Abdelrahman YM, Belland RJ (November 2005)."The chlamydial developmental cycle".FEMS Microbiology Reviews.29(5): 949–959.doi:10.1016/j.femsre.2005.03.002.PMID16043254.
- ^abcHorn M, Collingro A, Schmitz-Esser S, Beier CL, Purkhold U, Fartmann B, et al. (April 2004)."Illuminating the evolutionary history of Chlamydiae".Science.304(5671): 728–730.Bibcode:2004Sci...304..728H.doi:10.1126/science.1096330.PMID15073324.S2CID39036549.
- ^Philip S. Brachman and Elias Abrutyn (2009-07-23).Bacterial Infections of Humans: Epidemiology and Control.ISBN9780387098425.
- ^Moulder JW (1966). "The relation of the psittacosis group (Chlamydiae) to bacteria and viruses".Annual Review of Microbiology.20:107–130.doi:10.1146/annurev.mi.20.100166.000543.PMID5330228.
- ^abcdefgGupta RS, Naushad S, Chokshi C, Griffiths E, Adeolu M (September 2015). "A phylogenomic and molecular markers based analysis of the phylum Chlamydiae: Proposal to divide the class Chlamydiia into two orders, Chlamydiales and Parachlamydiales ord. nov., and emended description of the class Chlamydiia".Antonie van Leeuwenhoek.108(3): 765–781.doi:10.1007/s10482-015-0532-1.PMID26179278.S2CID17099157.
- ^abOren A, Garrity GM (July 2016)."List of new names and new combinations previously effectively, but not validly, published".International Journal of Systematic and Evolutionary Microbiology.66(7): 2463–2466.doi:10.1099/ijsem.0.001149.PMID27530111.
- ^Storz J, Page LA (1971)."Taxonomy of the Chlamydiae: reasons for classifying organisms of the genus Chlamydia, family Chlamydiaceae, in a separate order, Chlamydiales ord. nov".International Journal of Systematic Bacteriology.21(4): 332–334.doi:10.1099/00207713-21-4-332.
- ^Garrity GM, Boone DR (2001).Bergey's Manual of Systematic Bacteriology Volume 1: The Archaea and the Deeply Branching and Phototrophic Bacteria(2nd ed.). Springer.ISBN978-0-387-98771-2.
- ^abEverett KD, Thao M, Horn M, Dyszynski GE, Baumann P (July 2005)."Novel chlamydiae in whiteflies and scale insects: endosymbionts 'Candidatus Fritschea bemisiae' strain Falk and 'Candidatus Fritschea eriococci' strain Elm".International Journal of Systematic and Evolutionary Microbiology.55(Pt 4): 1581–1587.doi:10.1099/ijs.0.63454-0.PMID16014485.
- ^Sayers; et al."Chlamydiia".National Center for Biotechnology Information(NCBI) taxonomy database.Retrieved2016-10-24.
- ^Everett KD, Bush RM, Andersen AA (April 1999)."Emended description of the order Chlamydiales, proposal of Parachlamydiaceae fam. nov. and Simkaniaceae fam. nov., each containing one monotypic genus, revised taxonomy of the family Chlamydiaceae, including a new genus and five new species, and standards for the identification of organisms".International Journal of Systematic Bacteriology.49(Pt 2): 415–440.doi:10.1099/00207713-49-2-415.PMID10319462.
- ^Rurangirwa FR, Dilbeck PM, Crawford TB, McGuire TC, McElwain TF (April 1999)."Analysis of the 16S rRNA gene of micro-organism WSU 86-1044 from an aborted bovine foetus reveals that it is a member of the order Chlamydiales: proposal of Waddliaceae fam. nov., Waddlia chondrophila gen. nov., sp. nov".International Journal of Systematic Bacteriology.49(Pt 2): 577–581.doi:10.1099/00207713-49-2-577.PMID10319478.
- ^Thomas V, Casson N, Greub G (December 2006). "Criblamydia sequanensis, a new intracellular Chlamydiales isolated from Seine river water using amoebal co-culture".Environmental Microbiology.8(12): 2125–2135.Bibcode:2006EnvMi...8.2125T.doi:10.1111/j.1462-2920.2006.01094.x.PMID17107554.S2CID31211875.
- ^Stride MC, Polkinghorne A, Miller TL, Groff JM, Lapatra SE, Nowak BF (March 2013)."Molecular characterization of" Candidatus Parilichlamydia carangidicola, "a novel Chlamydia-like epitheliocystis agent in yellowtail kingfish, Seriola lalandi (Valenciennes), and the proposal of a new family," Candidatus Parilichlamydiaceae "fam. nov. (order Chlamydiales)".Applied and Environmental Microbiology.79(5): 1590–1597.Bibcode:2013ApEnM..79.1590S.doi:10.1128/AEM.02899-12.PMC3591964.PMID23275507.
- ^Kuo C-C, Horn M, Stephens RS (2011) Order I. Chlamydiales. In: Bergey's Manual of Systematic Bacteriology, vol. 4, 2nd ed. pp. 844-845. Eds Krieg N, Staley J, Brown D, Hedlund B, Paster B, Ward N, Ludwig W, Whitman W. Springer-: New York.
- ^abcdGriffiths E, Ventresca MS, Gupta RS (January 2006)."BLAST screening of chlamydial genomes to identify signature proteins that are unique for the Chlamydiales, Chlamydiaceae, Chlamydophila and Chlamydia groups of species".BMC Genomics.7:14.doi:10.1186/1471-2164-7-14.PMC1403754.PMID16436211.
- ^Sachse K, Bavoil PM, Kaltenboeck B, Stephens RS, Kuo CC, Rosselló-Móra R, Horn M (March 2015). "Emendation of the family Chlamydiaceae: proposal of a single genus, Chlamydia, to include all currently recognized species".Systematic and Applied Microbiology.38(2): 99–103.Bibcode:2015SyApM..38...99S.doi:10.1016/j.syapm.2014.12.004.hdl:10261/123714.PMID25618261.
- ^Oren A, Garrity GM (2015). "List of new names and new combinations previously effectively, but not validly, published".Int J Syst Evol Microbiol.65(7): 2017–2025.doi:10.1099/ijs.0.000317.PMID28056215.
- ^Greub G, Raoult D (September 2003)."History of the ADP/ATP-translocase-encoding gene, a parasitism gene transferred from a Chlamydiales ancestor to plants 1 billion years ago".Applied and Environmental Microbiology.69(9): 5530–5535.Bibcode:2003ApEnM..69.5530G.doi:10.1128/AEM.69.9.5530-5535.2003.PMC194985.PMID12957942.
- ^Griffiths E, Petrich AK, Gupta RS (August 2005)."Conserved indels in essential proteins that are distinctive characteristics of Chlamydiales and provide novel means for their identification".Microbiology.151(Pt 8): 2647–2657.doi:10.1099/mic.0.28057-0.PMID16079343.
- ^Gupta RS, Griffiths E (December 2006). "Chlamydiae-specific proteins and indels: novel tools for studies".Trends in Microbiology.14(12): 527–535.doi:10.1016/j.tim.2006.10.002.PMID17049238.
- ^Ward NL, Rainey FA, Hedlund BP, Staley JT, Ludwig W, Stackebrandt E (November 2000)."Comparative phylogenetic analyses of members of the order Planctomycetales and the division Verrucomicrobia: 23S rRNA gene sequence analysis supports the 16S rRNA gene sequence-derived phylogeny".International Journal of Systematic and Evolutionary Microbiology.50(Pt 6): 1965–1972.doi:10.1099/00207713-50-6-1965.PMID11155969.
- ^Teeling H, Lombardot T, Bauer M, Ludwig W, Glöckner FO (May 2004)."Evaluation of the phylogenetic position of the planctomycete 'Rhodopirellula baltica' SH 1 by means of concatenated ribosomal protein sequences, DNA-directed RNA polymerase subunit sequences and whole genome trees".International Journal of Systematic and Evolutionary Microbiology.54(Pt 3): 791–801.doi:10.1099/ijs.0.02913-0.PMID15143026.
- ^Griffiths E, Gupta RS (August 2007)."Phylogeny and shared conserved inserts in proteins provide evidence that Verrucomicrobia are the closest known free-living relatives of chlamydiae".Microbiology.153(Pt 8): 2648–2654.doi:10.1099/mic.0.2007/009118-0.PMID17660429.S2CID2094762.
- ^Corsaro D, Greub G (April 2006)."Pathogenic potential of novel Chlamydiae and diagnostic approaches to infections due to these obligate intracellular bacteria".Clinical Microbiology Reviews.19(2): 283–297.doi:10.1128/CMR.19.2.283-297.2006.PMC1471994.PMID16614250.
- ^"The LTP".Retrieved20 November2023.
- ^"LTP_all tree in newick format".Retrieved20 November2023.
- ^"LTP_08_2023 Release Notes"(PDF).Retrieved20 November2023.
- ^"GTDB release 08-RS214".Genome Taxonomy Database.Retrieved10 May2023.
- ^"bac120_r214.sp_label".Genome Taxonomy Database.Retrieved10 May2023.
- ^"Taxon History".Genome Taxonomy Database.Retrieved10 May2023.
- ^J.P. Euzéby."Chlamydiota".List of Prokaryotic names with Standing in Nomenclature(LPSN).Retrieved2022-09-09.
- ^Sayers; et al."Chlamydiae".National Center for Biotechnology Information(NCBI) taxonomy database.Retrieved2022-09-09.
External links
edit- Ward M."chlamydiae".University of Southampton.
The comprehensive reference and education wiki on Chlamydia and the Chlamydiales
- Chlamydia Overview