View source for Streptomyces

{{Short description|Genus of bacteria}}
{{Automatic taxobox
| image = Streptomyces_sp_01.png
| image_caption = Slide [[microbial culture|culture]]
| taxon = Streptomyces
| authority = Waksman and Henrici 1943 (Approved Lists 1980)
| type_species = ''[[Streptomyces albus]]''
| type_species_authority =(Rossi Doria 1891) Waksman and Henrici 1943 
| diversity_link = List of Streptomyces species
| diversity = About 550 species
| synonyms_ref = <ref name="LPSN">{{cite web| vauthors = Euzéby JP, Parte AC |url=https://lpsn.dsmz.de/genus/streptomyces |title=''Streptomyces'' |access-date=June 9, 2021 |publisher=[[List of Prokaryotic names with Standing in Nomenclature]] (LPSN)}}</ref>
| synonyms ={{collapsible list
   |title={{small|List}}
   |bullets = true
   |''Actinacidiphila'' <small>Madhaiyan et al. 2022</small>
   |''Actinopycnidium'' <small>Krassilnikov 1962 (Approved Lists 1980)</small>
   |''Actinosporangium'' <small>Krassilnikov and Yuan 1961 (Approved Lists 1980)</small>
   |''Chainia'' <small>Thirumalachar 1955 (Approved Lists 1980)</small>
   |''Elytrosporangium'' <small>Falcão de Morais ''et al''. 1966 (Approved Lists 1980)</small>
   |"''Indiella''" <small>Brumpt 1906</small>
   |"''Indiellopsis''" <small>Brumpt 1913</small>
   |''Kitasatoa'' <small>Matsumae and Hata 1968 (Approved Lists 1980)</small>
   |?"''Macrospora''" <small>Tsyganov ''et al''. 1964</small>
   |"''Microechinospora''" <small>Konev ''et al''. 1967</small>
   |''Microellobosporia'' <small>Cross, Lechevalier & Lechevalier 1963 (Approved Lists 1980)</small>
   |"''Oospora''" <small>Krüger 1904</small>{{citation needed|date=November 2021}}
   |''Streptantibioticus'' <small>Madhaiyan et al. 2022</small>
   |''Streptoverticillium'' <small>Baldacci 1958 (Approved Lists 1980)</small>
   |?"''Verticillomyces''" <small>Shinobu 1965</small>
  }}
}}

[[File:Streptomyces mycelial sheets.webp|thumb|254px|{{center|[[Mycelial sheet]]s{{hsp}}<ref>Van der Meij, A., Willemse, J., Schneijderberg, M.A., Geurts, R., Raaijmakers, J.M. and van Wezel, G.P. (2018) "Inter-and intracellular colonization of Arabidopsis roots by endophytic actinobacteria and the impact of plant hormones on their antimicrobial activity". ''Antonie van Leeuwenhoek'', '''111'''(5): 679–690. {{doi|10.1007/s10482-018-1014-z}}</ref>}}]]

'''''Streptomyces''''' is the largest [[genus]] of [[Actinomycetota]], and the [[type genus]] of the family [[Streptomycetaceae]].<ref name="Kämpfer">{{cite book | vauthors = Kämpfer P  |year=2006 |chapter=The Family Streptomycetaceae, Part I: Taxonomy |chapter-url=https://books.google.com/books?id=swciHNNWZDEC&pg=PA538 |pages=538–604 | veditors = Dworkin M, Falkow S, Rosenberg E, Schleifer KH, Stackebrandt E |title=The Prokaryotes |doi=10.1007/0-387-30743-5_22 |isbn=978-0-387-25493-7}}</ref> Over 700 species of ''Streptomyces'' [[bacteria]] have been described.<ref>{{cite web | vauthors = Euzéby JP | url=http://www.bacterio.cict.fr/s/streptomycesa.html |title=Genus Streptomyces | work=List of Prokaryotic names with Standing in Nomenclature |year=2008 |access-date=2008-09-28}}</ref><ref name=":0">{{Cite journal |last1=Nikolaidis |first1=Marios |last2=Hesketh |first2=Andrew |last3=Frangou |first3=Nikoletta |last4=Mossialos |first4=Dimitris |last5=Van de Peer |first5=Yves |last6=Oliver |first6=Stephen G. |last7=Amoutzias |first7=Grigorios D. |date=June 2023 |title=A panoramic view of the genomic landscape of the genus Streptomyces |journal=Microbial Genomics |volume=9 |issue=6 |doi=10.1099/mgen.0.001028 |doi-access=free |issn=2057-5858 |pmid=37266990|pmc=10327506 |s2cid=259025020 }}</ref><ref name=":1">{{Cite web |title=Genus: Streptomyces |url=https://www.bacterio.net/genus/streptomyces |access-date=2023-06-21 |website=www.bacterio.net |language=en}}</ref> As with the other Actinomycetota, streptomycetes are [[gram-positive]],  and have very large [[genomes]] with high [[GC-content|GC content]].<ref name=":0" /><ref name=Brock>{{cite book |veditors=Madigan M, Martinko J | title = Brock Biology of Microorganisms | edition = 11th | publisher = Prentice Hall | year = 2005 | isbn= 978-0-13-144329-7 }}{{page needed|date=October 2014}}</ref>  Found predominantly in soil and decaying vegetation, most streptomycetes produce [[spore]]s, and are noted for their distinct "earthy" odor that results from production of a volatile [[metabolite]], [[geosmin]].<ref>{{Cite book |url=https://onlinelibrary.wiley.com/doi/book/10.1002/047001590X |title=eLS |date=2001-05-30 |publisher=Wiley |isbn=978-0-470-01617-6 |editor-last=John Wiley & Sons, Ltd |edition=1 |language=en |doi=10.1002/9780470015902.a0020392.pub2}}</ref> Different strains of the same species may colonize very diverse environments.<ref name=":0" />

Streptomycetes are characterised by a complex [[secondary metabolism]].<ref name=Brock/> Between 5-23% (average: 12%) of the protein-coding genes of each ''Streptomyces'' species are implicated in secondary metabolism.<ref name=":0" /> Streptomycetes produce over two-thirds of the clinically useful [[antibiotic]]s of natural origin (e.g., [[neomycin]], [[streptomycin]], [[cypemycin]], [[grisemycin]], [[bottromycin]]s and [[chloramphenicol]]).<ref>{{cite book |vauthors=Kieser T, Bibb MJ, Buttner MJ, Chater KF, Hopwood DA |title=Practical Streptomyces Genetics |edition=2nd |publisher=John Innes Foundation |location=Norwich, England |year=2000 |isbn=978-0-7084-0623-6}}{{page needed|date=October 2014}}</ref><ref name="pmid24256223">{{cite journal | vauthors = Bibb MJ | title = Understanding and manipulating antibiotic production in actinomycetes | journal = [[Biochemical Society Transactions]] | volume = 41 | issue = 6 | pages = 1355–64 | date = December 2013 | pmid = 24256223 | doi = 10.1042/BST20130214 }}</ref> The antibiotic [[streptomycin]] takes its name directly from ''Streptomyces''. Streptomycetes are infrequent [[pathogens]], though infections in humans, such as [[Eumycetoma|mycetoma]], can be caused by ''[[Streptomyces somaliensis|S. somaliensis]]'' and ''[[Streptomyces sudanensis|S. sudanensis]]'', and in plants can be caused by ''[[Streptomyces caviscabies|S. caviscabies]]'', ''[[Streptomyces acidiscabies|S. acidiscabies]]'', ''[[Streptomyces turgidiscabies|S. turgidiscabies]]'' and ''[[Streptomyces scabies|S. scabies]]''.

==Taxonomy==
{{See also|List of Streptomyces species}}

''Streptomyces'' is the type genus of the family [[Streptomycetaceae]]<ref name=Anderson2001>{{cite journal | vauthors = Anderson AS, Wellington EM | title = The taxonomy of Streptomyces and related genera | journal = International Journal of Systematic and Evolutionary Microbiology | volume = 51 | issue = Pt 3 | pages = 797–814 | date = May 2001 | pmid = 11411701 | doi = 10.1099/00207713-51-3-797 | doi-access = free }}</ref> and currently covers more than 700 [[species]] with the number increasing every year.<ref name=Labeda2011>{{cite journal | vauthors = Labeda DP | title = Multilocus sequence analysis of phytopathogenic species of the genus Streptomyces | journal = International Journal of Systematic and Evolutionary Microbiology | volume = 61 | issue = Pt 10 | pages = 2525–2531 | date = October 2011 | pmid = 21112986 | doi = 10.1099/ijs.0.028514-0 | doi-access = free }}</ref><ref name=":1" /> It is estimated that the total number of ''Streptomyces'' species is close to 1600.<ref name=":0" /> [[Acidophilic]] and acid-tolerant strains that were initially classified under this genus have later been moved to ''[[Kitasatospora]]'' (1997) <ref>{{cite journal | vauthors = Zhang Z, Wang Y, Ruan J | title = A proposal to revive the genus Kitasatospora (Omura, Takahashi, Iwai, and Tanaka 1982) | journal = International Journal of Systematic Bacteriology | volume = 47 | issue = 4 | pages = 1048–54 | date = October 1997 | pmid = 9336904 | doi = 10.1099/00207713-47-4-1048 | doi-access = free }}</ref> and ''[[Streptacidiphilus]]'' (2003).<ref>{{cite journal | vauthors = Kim SB, Lonsdale J, Seong CN, Goodfellow M | title = Streptacidiphilus gen. nov., acidophilic actinomycetes with wall chemotype I and emendation of the family Streptomycetaceae (Waksman and Henrici (1943)AL) emend. Rainey et al. 1997 | journal = Antonie van Leeuwenhoek | volume = 83 | issue = 2 | pages = 107–16 | year = 2003 | pmid = 12785304 | doi = 10.1023/A:1023397724023 | s2cid = 12901116 }}</ref> Species nomenclature are usually based on their color of [[hyphae]] and [[spores]].

''[[Saccharopolyspora erythraea]]'' was formerly placed in this genus (as ''Streptomyces erythraeus'').

== Morphology ==

The genus ''Streptomyces'' includes [[aerobic organism|aerobic]], [[Gram-positive]], multicellular, filamentous bacteria that produce well-developed vegetative hyphae (between 0.5-2.0&nbsp;μm in diameter) with branches. They form a complex [[Mycelium|substrate mycelium]] that aids in scavenging organic compounds from their substrates.<ref name=Chater/> Although the mycelia and the [[Hypha|aerial hyphae]] that arise from them are amotile, mobility is achieved by dispersion of spores.<ref name=Chater/> Spore surfaces may be hairy, rugose, smooth, spiny or warty.<ref>{{cite journal | vauthors = Dietz A, Mathews J | title = Classification of Streptomyces spore surfaces into five groups | journal = Applied Microbiology | volume = 21 | issue = 3 | pages = 527–33 | date = March 1971 | pmid = 4928607 | pmc = 377216 | doi = 10.1128/AEM.21.3.527-533.1971 }}</ref> In some species, aerial hyphae consist of long, straight filaments, which bear 50 or more spores at more or less regular intervals, arranged in whorls (verticils). Each branch of a verticil produces, at its apex, an umbel, which carries from two to several chains of spherical to ellipsoidal, smooth or rugose spores.<ref name=Chater>{{cite book | vauthors = Chater K, Losick R |title= Microbial development|chapter= Morphological and physiological differentiation in ''Streptomyces''|chapter-url=http://cshmonographs.org/index.php/monographs/article/view/4367 |access-date= 2012-01-19|doi=10.1101/0.89-115|year= 1984 |volume= 16|isbn= 978-0-87969-172-1 |pages= 89–115|doi-broken-date= 2024-09-12 }}
</ref> Some strains form short chains of spores on substrate hyphae. Sclerotia-, pycnidia-, sporangia-, and synnemata-like structures are produced by some strains.

== Genomics ==
The complete [[genome]] of "''[[Streptomyces coelicolor|S. coelicolor]]'' strain A3(2)" was published in 2002.<ref name=Bentley_2002>{{cite journal | vauthors = Bentley SD, Chater KF, Cerdeño-Tárraga AM, Challis GL, Thomson NR, James KD, Harris DE, Quail MA, Kieser H, Harper D, Bateman A, Brown S, Chandra G, Chen CW, Collins M, Cronin A, Fraser A, Goble A, Hidalgo J, Hornsby T, Howarth S, Huang CH, Kieser T, Larke L, Murphy L, Oliver K, O'Neil S, Rabbinowitsch E, Rajandream MA, Rutherford K, Rutter S, Seeger K, Saunders D, Sharp S, Squares R, Squares S, Taylor K, Warren T, Wietzorrek A, Woodward J, Barrell BG, Parkhill J, Hopwood DA | display-authors = 6 | title = Complete genome sequence of the model actinomycete Streptomyces coelicolor A3(2) | journal = Nature | volume = 417 | issue = 6885 | pages = 141–7 | date = May 2002 | pmid = 12000953 | doi = 10.1038/417141a | s2cid = 4430218 | doi-access = free | bibcode = 2002Natur.417..141B }}</ref> At the time, the "''S. coelicolor''" genome was thought to contain the largest number of [[gene]]s of any [[bacterium]].<ref name=Bentley_2002/> The chromosome is 8,667,507 [[Base pair|bp]] long with a GC-content of 72.1%, and is predicted to contain 7,825 protein-encoding genes.<ref name=Bentley_2002/> In terms of taxonomy, "''S. coelicolor'' A3(2)" belongs to the species ''[[Streptomyces violaceoruber|S. violaceoruber]]'', and is not a validly described separate species; "''S. coelicolor'' A3(2)" is not to be mistaken for the actual ''[[Streptomyces coelicolor|S. coelicolor]]'' (Müller), although it is often referred to as ''S. coelicolor'' for convenience.<ref>{{cite journal | vauthors = Chater KF, Biró S, Lee KJ, Palmer T, Schrempf H | title = The complex extracellular biology of Streptomyces | journal = FEMS Microbiology Reviews | volume = 34 | issue = 2 | pages = 171–98 | date = March 2010 | pmid = 20088961 | doi = 10.1111/j.1574-6976.2009.00206.x | doi-access = free }}</ref> The transcriptome and translatome analyses of the strain A3(2) were published in 2016.<ref>{{cite journal | vauthors = Jeong Y, Kim JN, Kim MW, Bucca G, Cho S, Yoon YJ, Kim BG, Roe JH, Kim SC, Smith CP, Cho BK | display-authors = 6 | title = The dynamic transcriptional and translational landscape of the model antibiotic producer Streptomyces coelicolor A3(2) | journal = Nature Communications | volume = 7 | issue = 1 | pages = 11605 | date = June 2016 | pmid = 27251447 | pmc = 4895711 | doi = 10.1038/ncomms11605 | bibcode = 2016NatCo...711605J }}</ref>

The first complete genome sequence of ''[[Streptomyces avermitilis|S. avermitilis]]'' was completed in 2003.<ref name=Ikeda_2003>{{cite journal | vauthors = Ikeda H, Ishikawa J, Hanamoto A, Shinose M, Kikuchi H, Shiba T, Sakaki Y, Hattori M, Omura S | display-authors = 6 | title = Complete genome sequence and comparative analysis of the industrial microorganism Streptomyces avermitilis | journal = Nature Biotechnology | volume = 21 | issue = 5 | pages = 526–31 | date = May 2003 | pmid = 12692562 | doi = 10.1038/nbt820 | doi-access = free }}</ref> Each of these genomes forms a [[chromosome]] with a linear structure, unlike most bacterial genomes, which exist in the form of circular chromosomes.<ref name="Dyson2011">{{cite book| vauthors = Dyson P |title=Streptomyces: Molecular Biology and Biotechnology|url=https://books.google.com/books?id=3z9_QwFumi8C|access-date=16 January 2012|date=1 January 2011|publisher=Horizon Scientific Press|isbn=978-1-904455-77-6|page=5}}</ref> The genome sequence of ''[[Streptomyces scabiei|S. scabiei]]'', a member of the genus with the ability to cause potato scab disease, has been determined at the [[Sanger Institute|Wellcome Trust Sanger Institute]]. At 10.1 Mbp long and encoding 9,107 provisional genes, it is the largest known ''Streptomyces'' genome sequenced, probably due to the large [[pathogenicity island]].<ref name="Dyson2011"/><ref>{{cite web|url=http://www.sanger.ac.uk/Projects/S_scabies|title=Streptomyces scabies|publisher=Sanger Institute|access-date=2001-02-26}}</ref>

The genomes of the various ''Streptomyces'' species demonstrate remarkable plasticity, via ancient single gene duplications, block duplications (mainly at the chromosomal arms) and horizontal gene transfer.<ref name=":0" /><ref>{{Cite journal |last1=McDonald |first1=Bradon R. |last2=Currie |first2=Cameron R. |date=2017-06-06 |title=Lateral Gene Transfer Dynamics in the Ancient Bacterial Genus Streptomyces |journal=mBio |volume=8 |issue=3 |pages=e00644–17 |doi=10.1128/mBio.00644-17 |issn=2150-7511 |pmc=5472806 |pmid=28588130}}</ref> The size of their chromosome varies from 5.7-12.1 Mbps (average: 8.5 Mbps), the number of chromosomally encoded proteins varies from 4983-10,112 (average: 7130), whereas their high GC content varies from 68.8-74.7% (average: 71.7%).<ref name=":0" /> The 95% soft-core proteome of the genus consists of approximately 2000-2400 proteins.<ref name=":0" /> The [[pangenome]] is open.<ref>{{Cite journal |last1=Caicedo-Montoya |first1=Carlos |last2=Manzo-Ruiz |first2=Monserrat |last3=Ríos-Estepa |first3=Rigoberto |date=2021 |title=Pan-Genome of the Genus Streptomyces and Prioritization of Biosynthetic Gene Clusters With Potential to Produce Antibiotic Compounds |journal=Frontiers in Microbiology |volume=12 |pages=677558 |doi=10.3389/fmicb.2021.677558 |issn=1664-302X |pmc=8510958 |pmid=34659136 |doi-access=free }}</ref><ref>{{Cite journal |last1=Otani |first1=Hiroshi |last2=Udwary |first2=Daniel W. |last3=Mouncey |first3=Nigel J. |date=2022-11-07 |title=Comparative and pangenomic analysis of the genus Streptomyces |journal=Scientific Reports |volume=12 |issue=1 |pages=18909 |doi=10.1038/s41598-022-21731-1 |issn=2045-2322 |pmc=9640686 |pmid=36344558|bibcode=2022NatSR..1218909O }}</ref> In addition, significant genomic plasticity is observed even between strains of the same species, where the number of accessory proteins (at the species level) ranges from 250 to more than 3000.<ref name=":0" /> Intriguingly, a correlation has been observed between the number of carbohydrate-active enzymes and secondary metabolite biosynthetic gene clusters ([[siderophore]]s, e-Polylysin and type III [[lanthipeptide]]s) that are related to competition among bacteria, in ''Streptomyces'' species.<ref name=":0" /> Streptomycetes are major biomass degraders, mainly via their carbohydrate-active enzymes.<ref>{{Cite journal |last1=Chater |first1=Keith F. |last2=Biró |first2=Sandor |last3=Lee |first3=Kye Joon |last4=Palmer |first4=Tracy |last5=Schrempf |first5=Hildgund |date=March 2010 |title=The complex extracellular biology of Streptomyces |journal=FEMS Microbiology Reviews |volume=34 |issue=2 |pages=171–198 |doi=10.1111/j.1574-6976.2009.00206.x |issn=1574-6976 |pmid=20088961|doi-access=free }}</ref> Thus, they also need to evolve an arsenal of siderophores and antimicrobial agents to suppress competition by other bacteria in these nutrient-rich environments that they create.<ref name=":0" /> Several evolutionary analyses have revealed that the majority of evolutionarily stable genomic elements are localized mainly at the central region of the chromosome, whereas the evolutionarily unstable elements tend to localize at the chromosomal arms.<ref name=":0" /><ref>{{Cite journal |last1=Lorenzi |first1=Jean-Noël |last2=Lespinet |first2=Olivier |last3=Leblond |first3=Pierre |last4=Thibessard |first4=Annabelle |date=September 2019 |title=Subtelomeres are fast-evolving regions of the Streptomyces linear chromosome |journal=Microbial Genomics |volume=7 |issue=6 |pages=000525 |doi=10.1099/mgen.0.000525 |doi-access=free |issn=2057-5858 |pmc=8627663 |pmid=33749576}}</ref><ref>{{Cite journal |last1=Tidjani |first1=Abdoul-Razak |last2=Lorenzi |first2=Jean-Noël |last3=Toussaint |first3=Maxime |last4=van Dijk |first4=Erwin |last5=Naquin |first5=Delphine |last6=Lespinet |first6=Olivier |last7=Bontemps |first7=Cyril |last8=Leblond |first8=Pierre |date=2019-09-03 |title=Massive Gene Flux Drives Genome Diversity between Sympatric Streptomyces Conspecifics |journal=mBio |volume=10 |issue=5 |pages=e01533–19 |doi=10.1128/mBio.01533-19 |issn=2150-7511 |pmc=6722414 |pmid=31481382}}</ref><ref>{{Cite journal |last1=Volff |first1=J. N. |last2=Altenbuchner |first2=J. |date=January 1998 |title=Genetic instability of the Streptomyces chromosome |url=https://pubmed.ncbi.nlm.nih.gov/9484880 |journal=Molecular Microbiology |volume=27 |issue=2 |pages=239–246 |doi=10.1046/j.1365-2958.1998.00652.x |issn=0950-382X |pmid=9484880|s2cid=20438399 }}</ref><ref>{{Cite journal |last1=Chen |first1=Carton W. |last2=Huang |first2=Chih-Hung |last3=Lee |first3=Hsuan-Hsuan |last4=Tsai |first4=Hsiu-Hui |last5=Kirby |first5=Ralph |date=October 2002 |title=Once the circle has been broken: dynamics and evolution of Streptomyces chromosomes |url=https://pubmed.ncbi.nlm.nih.gov/12350342 |journal=Trends in Genetics |volume=18 |issue=10 |pages=522–529 |doi=10.1016/s0168-9525(02)02752-x |issn=0168-9525 |pmid=12350342}}</ref> Thus, the chromosomal arms emerge as the part of the genome that is mainly responsible for rapid adaptation at both the species and strain level.<ref name=":0" />

== Biotechnology ==
[[Biotechnology]] researchers have used ''Streptomyces'' species for [[heterologous expression]] of proteins. Traditionally, ''[[Escherichia coli]]'' was the species of choice to express [[Eukaryote|eukaryotic]] genes, since it was well understood and easy to work with.<ref name=Brawner_1991>{{cite journal | vauthors = Brawner M, Poste G, Rosenberg M, Westpheling J | title = Streptomyces: a host for heterologous gene expression | journal = Current Opinion in Biotechnology | volume = 2 | issue = 5 | pages = 674–81 | date = October 1991 | pmid = 1367716 | doi = 10.1016/0958-1669(91)90033-2 }}</ref><ref name=Payne_1990>{{cite journal | vauthors = Payne GF, DelaCruz N, Coppella SJ | title = Improved production of heterologous protein from Streptomyces lividans | journal = Applied Microbiology and Biotechnology | volume = 33 | issue = 4 | pages = 395–400 | date = July 1990 | pmid = 1369282 | doi = 10.1007/BF00176653 | s2cid = 19287805 }}</ref> Expression of eukaryotic proteins in ''E. coli'' may be problematic. Sometimes, proteins do not fold properly, which may lead to insolubility, deposition in [[inclusion bodies]], and loss of bioactivity of the product.<ref name=Binnie_1997>{{cite journal | vauthors = Binnie C, Cossar JD, Stewart DI | title = Heterologous biopharmaceutical protein expression in Streptomyces | journal = Trends in Biotechnology | volume = 15 | issue = 8 | pages = 315–20 | date = August 1997 | pmid = 9263479 | doi = 10.1016/S0167-7799(97)01062-7 }}</ref> Though ''E. coli'' strains have secretion mechanisms, these are of low efficiency and result in secretion into the [[periplasmic space]], whereas secretion by a Gram-positive bacterium such as a ''Streptomyces'' species results in secretion directly into the extracellular medium. In addition, ''Streptomyces''  species have more efficient secretion mechanisms than ''E.coli''. The properties of the secretion system is an advantage for industrial production of heterologously expressed protein because it simplifies subsequent purification steps and may increase yield. These properties among others make ''Streptomyces'' spp. an attractive alternative to other bacteria such as ''E. coli'' and ''[[Bacillus subtilis]]''.<ref name="Binnie_1997"/> In addition, the inherently high genomic instability suggests that the various Streptomycetes genomes may be amenable to extensive genome reduction for the construction of synthetic minimal genomes with industrial applications.<ref name=":0" />

== Plant pathogenic bacteria ==
Several species belonging to this genus have been found to be pathogenic to plants:<ref name=Labeda2011/>
# ''[[Streptomyces scabiei|S. scabiei]]''
# ''[[Streptomyces acidiscabies|S. acidiscabies]]''
# ''[[Streptomyces europaeiscabiei|S. europaeiscabiei]]''
# ''[[Streptomyces luridiscabiei|S. luridiscabiei]]''
# ''[[Streptomyces niveiscabiei|S. niveiscabiei]]''
# ''[[Streptomyces puniciscabiei|S. puniciscabiei]]''
# ''[[Streptomyces reticuliscabiei|S. reticuliscabiei]]''
# ''[[Streptomyces stelliscabiei|S. stelliscabiei]]''
# ''[[Streptomyces turgidiscabies|S. turgidiscabies]]'' (scab disease in [[potato]]es)
# ''[[Streptomyces ipomoeae|S. ipomoeae]]'' (soft rot disease in [[sweet potato]]es)
# ''[[Streptomyces brasiliscabiei|S. brasiliscabiei]]'' (first species identified in Brazil)<ref>{{cite journal |last1=Corrêa |first1=Daniele Bussioli Alves |last2=do Amaral |first2=Danilo Trabuco |last3=da Silva |first3=Márcio José |last4=Destéfano |first4=Suzete Aparecida Lanza |title=Streptomyces brasiliscabiei, a new species causing potato scab in south Brazil |journal=Antonie van Leeuwenhoek |date=July 2021 |volume=114 |issue=7 |pages=913–931 |doi=10.1007/s10482-021-01566-y|pmid=33881637 }}</ref>
# ''[[Streptomyces hilarionis|S. hilarionis]]'' and ''[[Streptomyces hayashii|S. hayashii]]'' (new species identified in Brazil)<ref>{{cite journal |last1=Vitor |first1=Lucas |last2=Amaral |first2=Danilo Trabuco |last3=Corrêa |first3=Daniele Bussioli Alves |last4=Ferreira-Tonin |first4=Mariana |last5=Lucon |first5=Emanuel Torres |last6=Appy |first6=Mariana Pereira |last7=Tomaseto |first7=Alex Augusto |last8=Destéfano |first8=Suzete Aparecida Lanza |title=Streptomyces hilarionis sp. nov. and Streptomyces hayashii sp. nov., two new strains associated with potato scab in Brazil |journal=International Journal of Systematic and Evolutionary Microbiology |date=15 June 2023 |volume=73 |issue=6 |doi=10.1099/ijsem.0.005916|pmid=37319004 }}</ref>

== Medicine ==
''Streptomyces'' is the largest [[antibiotic]]-producing genus, producing antibacterial, [[antifungal drug|antifungal]], and antiparasitic drugs, and also a wide range of other [[Biological activity|bioactive]] compounds, such as [[Immunosuppressive drug|immunosuppressants]].<ref name="pmid11702082">{{cite journal | vauthors = Watve MG, Tickoo R, Jog MM, Bhole BD | title = How many antibiotics are produced by the genus Streptomyces? | journal = Archives of Microbiology | volume = 176 | issue = 5 | pages = 386–90 | date = November 2001 | pmid = 11702082 | doi = 10.1007/s002030100345 | s2cid = 603765 }}</ref> Almost all of the bioactive compounds produced by ''Streptomyces'' are initiated during the time coinciding with the aerial hyphal formation from the substrate mycelium.<ref name=Chater/>

=== Antifungals ===
{{See also|Polyene antimycotic}}
Streptomycetes produce numerous antifungal compounds of medicinal importance, including  [[nystatin]] (from ''[[Streptomyces noursei|S. noursei]]''), [[amphotericin B]] (from ''[[Streptomyces nodosus|S. nodosus]]''),<ref name="pmid22975171">{{cite journal | vauthors = Procópio RE, Silva IR, Martins MK, Azevedo JL, Araújo JM | title = Antibiotics produced by Streptomyces | journal = The Brazilian Journal of Infectious Diseases | volume = 16 | issue = 5 | pages = 466–71 | year = 2012 | pmid = 22975171 | doi = 10.1016/j.bjid.2012.08.014 | doi-access = free }}</ref> and [[natamycin]] (from ''[[Streptomyces natalensis|S. natalensis]]'').

=== Antibacterials ===
Members of the genus ''Streptomyces'' are the source for numerous antibacterial pharmaceutical agents; among the most important of these are:
* [[Chloramphenicol]] (from ''[[Streptomyces venezuelae|S. venezuelae]]'')<ref>{{cite journal | vauthors = Akagawa H, Okanishi M, Umezawa H | title = A plasmid involved in chloramphenicol production in Streptomyces venezuelae: evidence from genetic mapping | journal = Journal of General Microbiology | volume = 90 | issue = 2 | pages = 336–46 | date = October 1975 | pmid = 1194895 | doi = 10.1099/00221287-90-2-336 | doi-access = free }}</ref>
* [[Daptomycin]] (from ''[[Streptomyces roseosporus|S. roseosporus]]'')<ref>{{cite journal | vauthors = Miao V, Coëffet-LeGal MF, Brian P, Brost R, Penn J, Whiting A, Martin S, Ford R, Parr I, Bouchard M, Silva CJ, Wrigley SK, Baltz RH | display-authors = 6 | title = Daptomycin biosynthesis in Streptomyces roseosporus: cloning and analysis of the gene cluster and revision of peptide stereochemistry | journal = Microbiology | volume = 151 | issue = Pt 5 | pages = 1507–1523 | date = May 2005 | pmid = 15870461 | doi = 10.1099/mic.0.27757-0 | doi-access = free }}</ref>
* [[Fosfomycin]] (from ''[[Streptomyces fradiae|S. fradiae]]'')<ref name="pmid17113999">{{cite journal | vauthors = Woodyer RD, Shao Z, Thomas PM, Kelleher NL, Blodgett JA, Metcalf WW, van der Donk WA, Zhao H | display-authors = 6 | title = Heterologous production of fosfomycin and identification of the minimal biosynthetic gene cluster | journal = Chemistry & Biology | volume = 13 | issue = 11 | pages = 1171–82 | date = November 2006 | pmid = 17113999 | doi = 10.1016/j.chembiol.2006.09.007 | doi-access =  }}</ref>
* [[Lincomycin]] (from ''[[Streptomyces lincolnensis|S. lincolnensis]]'')<ref>{{cite journal | vauthors = Peschke U, Schmidt H, Zhang HZ, Piepersberg W | title = Molecular characterization of the lincomycin-production gene cluster of Streptomyces lincolnensis 78-11 | journal = Molecular Microbiology | volume = 16 | issue = 6 | pages = 1137–56 | date = June 1995 | pmid = 8577249 | doi = 10.1111/j.1365-2958.1995.tb02338.x | s2cid = 45162659 }}</ref>
* [[Neomycin]] (from ''S. fradiae'')<ref>{{cite journal | vauthors = Dulmage HT | title = The production of neomycin by Streptomyces fradiae in synthetic media | journal = Applied Microbiology | volume = 1 | issue = 2 | pages = 103–6 | date = March 1953 | pmid = 13031516 | pmc = 1056872 | doi = 10.1128/AEM.1.2.103-106.1953 }}</ref>
* [[Nourseothricin]] {{citation needed|date= August 2015}}
* [[Puromycin]] (from ''[[Streptomyces alboniger|S. alboniger]]'')<ref>{{cite journal | vauthors = Sankaran L, Pogell BM | title = Biosynthesis of puromycin in Streptomyces alboniger: regulation and properties of O-demethylpuromycin O-methyltransferase | journal = Antimicrobial Agents and Chemotherapy | volume = 8 | issue = 6 | pages = 721–32 | date = December 1975 | pmid = 1211926 | pmc = 429454 | doi = 10.1128/AAC.8.6.721 }}</ref>
* [[Streptomycin]] (from ''[[Streptomyces griseus|S. griseus]]'')<ref>{{cite journal | vauthors = Distler J, Ebert A, Mansouri K, Pissowotzki K, Stockmann M, Piepersberg W | title = Gene cluster for streptomycin biosynthesis in Streptomyces griseus: nucleotide sequence of three genes and analysis of transcriptional activity | journal = Nucleic Acids Research | volume = 15 | issue = 19 | pages = 8041–56 | date = October 1987 | pmid = 3118332 | pmc = 306325 | doi = 10.1093/nar/15.19.8041 }}</ref>
* [[Tetracycline]] (from ''[[Streptomyces rimosus|S. rimosus]]'' and ''[[Streptomyces aureofaciens|S. aureofaciens]]'')<ref name="NelsonGreenwald2001">{{cite book| vauthors = Nelson M, Greenwald RA, Hillen W |title=Tetracyclines in biology, chemistry and medicine|url=https://books.google.com/books?id=kHNW4tFhZD4C&pg=PA8|access-date=17 January 2012|year=2001|publisher=Birkhäuser|isbn=978-3-7643-6282-9|pages=8–}}</ref>
* [[Oleandomycin]] (from ''[[Streptomyces antibioticus|S. antibioticus]]'')<ref>{{cite web|title=What are Streptomycetes?|url=http://home.hiroshima-u.ac.jp/mbiotech/hosenkin_lab/Strepto-E.html | archive-url = https://web.archive.org/web/20160304052228/https://home.hiroshima-u.ac.jp/mbiotech/hosenkin_lab/Strepto-E.html | archive-date = 4 March 2016 |website=Hosenkin Lab; Hiroshima-University|access-date=10 August 2015}}</ref><ref>{{cite journal | vauthors = Swan DG, Rodríguez AM, Vilches C, Méndez C, Salas JA | title = Characterisation of a Streptomyces antibioticus gene encoding a type I polyketide synthase which has an unusual coding sequence | journal = Molecular & General Genetics | volume = 242 | issue = 3 | pages = 358–62 | date = February 1994 | pmid = 8107683 | doi = 10.1007/BF00280426 | s2cid = 2195072 }}</ref><ref>{{cite web|title= Finto: MeSH: Streptomyces antibioticus|url=http://finto.fi/mesh/en/page/D013303|website=finto: Finnish Thesaurus and Ontology Service|access-date=10 August 2015}}</ref>
* [[Tunicamycin]] (from ''S. torulosus'')<ref>{{cite journal| vauthors = Atta HM |title=Biochemical studies on antibiotic production from Streptomyces sp.: Taxonomy, fermentation, isolation and biological properties|journal=Journal of Saudi Chemical Society|date=January 2015|volume=19|issue=1|pages=12–22|doi=10.1016/j.jscs.2011.12.011|doi-access=free}}</ref>
* Mycangimycin (from ''Streptomyces sp. SPB74'' and ''[[Streptomyces antibioticus|S. antibioticus]]'')<ref>{{cite journal | vauthors = Oh DC, Scott JJ, Currie CR, Clardy J | title = Mycangimycin, a polyene peroxide from a mutualist Streptomyces sp | journal = Organic Letters | volume = 11 | issue = 3 | pages = 633–6 | date = February 2009 | pmid = 19125624 | pmc = 2640424 | doi = 10.1021/ol802709x }}</ref>
* [[Boromycin]] (from ''[[Streptomyces antibioticus|S. antibioticus]]'')<ref>{{cite journal| vauthors = Chen TS, Chang CJ, Floss HG |title=Biosynthesis of boromycin|journal=The Journal of Organic Chemistry |date=June 1981 |volume=46 |issue=13 |pages=2661–2665 |doi=10.1021/jo00326a010 }}</ref>
* [[Bambermycin]] (from ''[[Streptomyces bambergiensis|S. bambergiensis]]'' and ''[[Streptomyces ghanaensis|S. ghanaensis]]'', the active compound being [[Moenomycin Family Antibiotics|moenomycins A and C]])<ref>{{cite web | publisher = National Center for Biotechnology Information. | work = PubChem Compound Database | title = CID=53385491 | url =  https://pubchem.ncbi.nlm.nih.gov/compound/53385491 | access-date = 8 March 2017 }}</ref>
* [[Vulgamycin]]<ref>{{Cite journal |last1=Babczinski |first1=Peter |last2=Dorgerloh |first2=Michael |last3=Löbberding |first3=Antonius |last4=Santel |first4=Hans-Joachim |last5=Schmidt |first5=Robert R. |last6=Schmitt |first6=Peter |last7=Wünsche |first7=Christian |date=1991 |title=Herbicidal activity and mode of action of vulgamycin |url=https://onlinelibrary.wiley.com/doi/10.1002/ps.2780330406 |journal=Pesticide Science |language=en |volume=33 |issue=4 |pages=439–446 |doi=10.1002/ps.2780330406}}</ref>

[[Clavulanic acid]] (from ''[[Streptomyces clavuligerus|S. clavuligerus]]'') is a drug used in combination with some antibiotics (like [[amoxicillin]]) to block and/or weaken some bacterial-resistance mechanisms by irreversible beta-lactamase inhibition.
Novel antiinfectives currently being developed include [[Guadinomine]] (from ''Streptomyces'' sp. K01-0509),<ref name="Holmes2012">{{cite journal | vauthors = Holmes TC, May AE, Zaleta-Rivera K, Ruby JG, Skewes-Cox P, Fischbach MA, DeRisi JL, Iwatsuki M, Ōmura S, Khosla C | display-authors = 6 | title = Molecular insights into the biosynthesis of guadinomine: a type III secretion system inhibitor | journal = Journal of the American Chemical Society | volume = 134 | issue = 42 | pages = 17797–806 | date = October 2012 | pmid = 23030602 | pmc = 3483642 | doi = 10.1021/ja308622d }}</ref> a compound that blocks the [[Type III secretion system]] of Gram-negative bacteria.

=== Antiparasitic drugs ===
''[[Streptomyces avermitilis|S. avermitilis]]'' is responsible for the production of one of the most widely employed drugs against nematode and arthropod infestations, [[avermectin]],<ref name="Martin-et-al-2017">{{cite journal | last1=Martín | first1=Juan F | last2=Rodríguez-García | first2=Antonio | last3=Liras | first3=Paloma | title=The master regulator PhoP coordinates phosphate and nitrogen metabolism, respiration, cell differentiation and antibiotic biosynthesis: comparison in ''Streptomyces coelicolor'' and ''Streptomyces avermitilis'' | journal=[[The Journal of Antibiotics]] | publisher=[[Japan Antibiotics Research Association]] ([[Nature Portfolio]]) | volume=70 | issue=5 | date=2017-03-15 | issn=0021-8820 | doi=10.1038/ja.2017.19 | pages=534–541| pmid=28293039 | s2cid=1881648 | doi-access=free }}</ref> and thus its derivatives including [[ivermectin]].

=== Other ===
[[File:Saptomycin D and E Structures.svg|thumb|Saptomycins D and E]]
Less commonly, streptomycetes produce compounds used in other medical treatments: [[migrastatin]] (from [[Streptomyces platensis|''S. platensis'']]) and [[bleomycin]] (from [[Streptomyces verticillus|''S. verticillus'']]) are [[antineoplastic]] (anticancer) drugs; [[boromycin]] (from ''[[Streptomyces antibioticus|S. antibioticus]]'') exhibits antiviral activity against the HIV-1 strain of HIV, as well as antibacterial activity. [[Staurosporine]] (from ''[[Streptomyces staurosporeus|S. staurosporeus]]'') also has a range of activities from antifungal to antineoplastic (via the inhibition of [[protein kinase]]s).

''[[Streptomyces hygroscopicus|S. hygroscopicus]]'' and ''[[Streptomyces viridochromogenes|S. viridochromogenes]]'' produce the natural herbicide [[bialaphos]].

Saptomycins are [[chemical compound]]s isolated from ''Streptomyces''.<ref>{{Cite journal|last1=Abe|first1=N.|last2=Nakakita|first2=Y.|last3=Nakamura|first3=T.|last4=Enoki|first4=N.|last5=Uchida|first5=H.|last6=Munekata|first6=M.|year=1993|title=Novel antitumor antibiotics, saptomycins. I. Taxonomy of the producing organism, fermentation, HPLC analysis and biological activities|journal=The Journal of Antibiotics|volume=46|issue=10|pages=1530–5|doi=10.7164/antibiotics.46.1530|pmid=8244880|doi-access=free}}</ref>

== Symbiosis ==
''[[Sirex]]'' wasps cannot perform all of their own [[cellulolytic]] functions and so some ''Streptomyces'' do so in [[symbiosis]] with the wasps.<ref name="Li-et-al-2021">{{cite journal | last1=Li | first1=Hongjie | last2=Young | first2=Soleil E. | last3=Poulsen | first3=Michael | last4=Currie | first4=Cameron R. | title=Symbiont-Mediated Digestion of Plant Biomass in Fungus-Farming Insects | journal=[[Annual Review of Entomology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=66 | issue=1 | date=2021-01-07 | issn=0066-4170 | doi=10.1146/annurev-ento-040920-061140 | pages=297–316| pmid=32926791 | osti=1764729 | s2cid=221724225 }}</ref> Book ''et al.'' have investigated several of these symbioses.<ref name="Li-et-al-2021" /> Book ''et al.'', 2014 and Book ''et al.'', 2016 identify several lytic isolates.<ref name="Li-et-al-2021" /> The 2016 study isolates [[Streptomyces sp. Amel2xE9|''Streptomyces'' sp. Amel2xE9]] and [[Streptomyces sp. LamerLS-31b|''Streptomyces'' sp. LamerLS-31b]] and finds that they are equal in activity to the previously identified [[Streptomyces sp. SirexAA-E|''Streptomyces'' sp. SirexAA-E]].<ref name="Li-et-al-2021" />

== See also ==
* [[Antimycin A]] – Chemical compound produced by ''Streptomyces'' used as a piscicide
* {{Annotated link|Geosmin}}
* [[Streptomyces isolates|''Streptomyces'' isolates]]
* [[List of bacterial orders]]
* [[List of bacteria genera]]
* [[raiA-hairpin RNA motif]]

== References ==
{{Reflist|30em}}

== Further reading ==
{{Refbegin}}
* {{cite book | vauthors = Baumberg S | title = Genetics and Product Formation in Streptomyces | publisher = Kluwer Academic | year = 1991 | isbn= 978-0-306-43885-1 }}
* {{cite book | vauthors = Gunsalus IC | title = Bacteria: Antibiotic-producing Streptomyces | publisher = Academic Press | year = 1986 | isbn= 978-0-12-307209-2 | author-link = Irwin Gunsalus }}
* {{cite book | vauthors = Hopwood DA | title = Streptomyces in Nature and Medicine: The Antibiotic Makers | publisher = Oxford University Press | year = 2007 | isbn= 978-0-19-515066-7 }}
* {{cite book | veditors = Dyson P | title = Streptomyces: Molecular Biology and Biotechnology | publisher = [[Caister Academic Press]] | year = 2011 | isbn= 978-1-904455-77-6 }}
{{Refend}}

== External links ==
* {{cite web | url = http://www.micron.ac.uk/organisms/sco.html | title = Current research on ''Streptomyces coelicolor'' | work = Norwich Research Park | date = 3 January 2018 }}
* {{cite web | url = http://www.openwetware.org/wiki/Streptomyces | title = Some current ''Streptomyces'' Research & Methods / Protocols / Resources | work = www.openwetware.org }}
* {{cite web | url = http://avermitilis.ls.kitasato-u.ac.jp/ | title = ''S. avermitilis'' genome homepage | work = Kitasato Institute for Life Sciences }}
* {{cite web | url = http://www.sanger.ac.uk/Projects/S_coelicolor/ | title = ''S. coelicolor'' A3(2) genome homepage | work = Sanger Institute }}
* {{cite web | url = http://streptomyces.org.uk/ | title = Streptomyces.org.uk homepage | work = John Innes Centre }}
* {{cite web | url = http://strepdb.streptomyces.org.uk/ | title = StrepDB - the ''Streptomyces'' genomes annotation browser }}
* {{cite web | url = http://www.genomesonline.org/search.cgi?colcol=all&goldstamp=ALL&gen_type=ALL&org_name1=genus&gensp=Streptomyces&org_domain=ALL&org_status=ALL&size2=ALL&org_size=Kb&gen_gc=ALL&phylogeny2=ALL&gen_institution=ALL&gen_funding=ALL&gen_data=ALL&cont=ALL&gen_country=ALL&gen_pheno=ALL&gen_eco=ALL&gen_disease=ALL&gen_relevance=ALL&gen_avail=ALL&selection=submit+search | title = Streptomyces Genome Projects | work = Genomes OnLine Database }}

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[[Category:Streptomyces| ]]
[[Category:Bacteria genera]]