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==Taxonomy and discovery== In 1902, ''B. thuringiensis'' was first discovered in [[silkworm]]s by Japanese sericultural engineer {{nihongo|Ishiwatari Shigetane|石渡 繁胤}}. He named it ''B. sotto'',<ref>{{cite book|title=New Innovative Pesticides|url=https://books.google.com/books?id=ZR3uH7O713QC&pg=PA61|year=1977|publisher=EPA|page=61|quote= In 1915 the bacterium was re-examined and named ''Bacillus sotto''. [...] At about the same time, Beriner was isolating the organism}}</ref> using the Japanese word {{nihongo||卒倒|sottō|'collapse'}}, here referring to bacillary paralysis.<ref>{{cite book|title=Natural Enemies in the Pacific Area: Biological Control|url=https://books.google.com/books?id=ynweAQAAMAAJ|year=1967|publisher=Fukuoka Entomological Society|page=99|quote= "Sotto" in Japanese means "sudden collapse" or "fainting", and "sotto" of ''Bacillus thuringiensis'' var. ''sotto'' derives its name from the "sotto" disease.}}</ref> In 1911, German microbiologist [[Ernst Berliner]] rediscovered it when he isolated it as the cause of a disease called ''{{lang|de|[[Schlaffsucht]]}}'' in [[Ephestia kuehniella|flour moth]] caterpillars in [[Thuringia]] (hence the [[specific name (zoology)|specific name]] ''thuringiensis'', "Thuringian").<ref>{{cite book | vauthors = Reardon RC, Dubois NR, McLane W |title=''Bacillus thuringiensis'' for managing gypsy moth: a review |url= https://archive.org/details/CAT10861975 |year=1994 | work = USDA Forest Service | publisher = United States Department of Agriculture |quote= Mediterranean flour moths, ''Ephestia'' (=''Anagasta'') ''kuehniella'' (Zeller), that were found in stored grain in [[Thuringia]]}}</ref> ''B. sotto'' would later be reassigned as ''B. thuringiensis'' var. ''sotto''.<ref>{{cite book| vauthors =Steinhaus E |title=Insect Pathology: An Advanced Treatise|url=https://books.google.com/books?id=qXy7rpkMJdwC&pg=PA32|year=2012|publisher=Elsevier|isbn=978-0-323-14317-2|page=32 | quote= ''Bacillus sotto'' {{small|Ishiwata}} [→] Taxonomic reassignment: ''Bacillus thuringiensis'' var. ''sotto'' {{small|Ishiwata}}. [Heimpel and Angus, 1960]}}</ref> In 1976, Robert A. Zakharyan reported the presence of a plasmid in a strain of ''B. thuringiensis'' and suggested the plasmid's involvement in endospore and crystal formation.<ref>{{cite journal | vauthors = Zakharyan RA, Israelyan YK, Agabalyan AS, Tatevosyan PE, Akopyan S, Afrikyan EK |year=1979 |title=Plasmid DNA from Bacillus thuringiensis |journal=Microbiologiya |volume=48 |issue=2 |pages=226–229 |issn=0026-3656}}</ref><ref>{{cite book | veditors = Cheng TC |year=1984 |title=Pathogens of invertebrates: application in biological control and transmission mechanisms |isbn=978-0-306-41700-9 |page=[https://archive.org/details/pathogensofinver07chen/page/159 159] |url-access=registration |url=https://archive.org/details/pathogensofinver07chen/page/159 }}</ref> ''B. thuringiensis'' is closely related to ''[[Bacillus cereus|B. cereus]]'', a soil bacterium, and ''[[Bacillus anthracis|B. anthracis]]'', the cause of [[Anthrax disease|anthrax]]; the three organisms differ mainly in their [[plasmid]]s.<ref>{{cite book | vauthors = Økstad OA, Kolstø A | title=Genomics of Foodborne Bacterial Pathogens | chapter=Genomics of Bacillus Species | author2-link=Anne-Brit Kolstø | veditors = Wiedmann M, Zhang W | pages= 29–53 | publisher = Springer Science+Business Media, LLC | year = 2011 | doi = 10.1007/978-1-4419-7686-4_2 | isbn= 978-1-4419-7685-7 }}</ref>{{rp|34–35}} Like other members of the genus, all three are capable of producing [[endospore]]s.<ref name=Brock/> <!-- Tubulin was long thought to be specific to [[eukaryote]]s. More recently, however, several [[Prokaryote|prokaryotic]] proteins have been shown to be related to tubulin.<ref name="pmid9628483">{{cite journal | vauthors = Nogales E, Downing KH, Amos LA, Löwe J | title = Tubulin and FtsZ form a distinct family of GTPases | journal = Nature Structural Biology | volume = 5 | issue = 6 | pages = 451–8 | date = June 1998 | pmid = 9628483 | doi = 10.1038/nsb0698-451 | s2cid = 5945125 }}</ref><ref name="Jenkins_2002">{{cite journal | vauthors = Jenkins C, Samudrala R, Anderson I, Hedlund BP, Petroni G, Michailova N, Pinel N, Overbeek R, Rosati G, Staley JT | title = Genes for the cytoskeletal protein tubulin in the bacterial genus Prosthecobacter | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 99 | issue = 26 | pages = 17049–54 | date = December 2002 | pmid = 12486237 | pmc = 139267 | doi = 10.1073/pnas.012516899 | bibcode = 2002PNAS...9917049J }}</ref><ref name="Yutin_2012">{{cite journal | vauthors = Yutin N, Koonin EV | title = Archaeal origin of tubulin | journal = Biology Direct | volume = 7 | pages = 10 | date = March 2012 | pmid = 22458654 | pmc = 3349469 | doi = 10.1186/1745-6150-7-10 }}</ref><ref name="Larsen_2007">{{cite journal | vauthors = Larsen RA, Cusumano C, Fujioka A, Lim-Fong G, Patterson P, Pogliano J | title = Treadmilling of a prokaryotic tubulin-like protein, TubZ, required for plasmid stability in Bacillus thuringiensis | journal = Genes & Development | volume = 21 | issue = 11 | pages = 1340–52 | date = June 2007 | pmid = 17510284 | pmc = 1877747 | doi = 10.1101/gad.1546107 }}</ref> --> ===Species group placement=== ''B. thuringiensis'' is placed in the ''Bacillus cereus'' group which is variously defined as: seven closely related species: ''B. cereus'' ''sensu stricto'' (''[[Bacillus cereus|B. cereus]]''), ''[[Bacillus anthracis|B. anthracis]]'', ''B. thuringiensis'', ''[[Bacillus mycoides|B. mycoides]]'', ''[[Bacillus pseudomycoides|B. pseudomycoides]]'', and ''[[Bacillus cytotoxicus|B. cytotoxicus]]'';<ref>{{cite journal | vauthors = Guinebretière MH, Auger S, Galleron N, Contzen M, De Sarrau B, De Buyser ML, Lamberet G, Fagerlund A, Granum PE, Lereclus D, De Vos P, Nguyen-The C, Sorokin A | display-authors = 6 | title = Bacillus cytotoxicus sp. nov. is a novel thermotolerant species of the Bacillus cereus Group occasionally associated with food poisoning | journal = International Journal of Systematic and Evolutionary Microbiology | volume = 63 | issue = Pt 1 | pages = 31–40 | date = January 2013 | pmid = 22328607 | doi = 10.1099/ijs.0.030627-0 | s2cid = 2407509 }}</ref> or as six species in a ''Bacillus cereus'' sensu lato: ''[[Bacillus weihenstephanensis|B. weihenstephanensis]]'', ''B. mycoides'', ''B. pseudomycoides'', ''B. cereus'', ''B. thuringiensis'', and ''B. anthracis''. Within this grouping ''B.t.'' is more closely related to ''B.ce.'' It is more distantly related to ''B.w.'', ''B.m.'', ''B.p.'', and ''B.cy.''<ref name="Kolsto-et-al-2009">{{cite journal | vauthors = Kolstø AB, Tourasse NJ, Økstad OA | title = What sets Bacillus anthracis apart from other Bacillus species? | journal = Annual Review of Microbiology | volume = 63 | issue = 1 | pages = 451–476 | year = 2009 | pmid = 19514852 | doi = 10.1146/annurev.micro.091208.073255 | publisher = [[Annual Reviews (publisher)|Annual Reviews]] | author1-link = Anne-Brit Kolstø }}</ref> ===Subspecies=== There are several dozen recognized subspecies of ''B. thuringiensis''. Subspecies commonly used as insecticides include ''B. thuringiensis'' [[Bacillus thuringiensis kurstaki|subspecies ''kurstaki'']] (Btk), subspecies ''[[Bacillus thuringiensis israelensis|israelensis]]'' (Bti) and {{visible anchor|aizawa|text=subspecies ''aizawai''}} (Bta).<ref>{{Cite web|last=US EPA|first=OCSPP|date=2016-07-05|title=Bti for Mosquito Control|url=https://www.epa.gov/mosquitocontrol/bti-mosquito-control|access-date=2021-05-10|website=US EPA|language=en}}</ref><ref>{{cite web|title=Information on ''Bacillus thuringiensis'' subspecies ''kurstaki'' (Btk) Excerpts from a Forestry Technical Manual produced by Valent BioSciences, manufacturers of Foray® and DiPel®, two formulations of commercially produced ''Bacillus thuringiensis'' var. ''kurstaki'' (Btk)|url=https://www.fs.usda.gov/Internet/FSE_DOCUMENTS/fsbdev7_015300.pdf|website=Fs.usda.gov|access-date=2022-04-09}}</ref><ref>{{cite web|title=Commonly Asked Questions About Btk (''Bacillus thuringiensis'' var. ''kurstaki'') | vauthors = Ellis JA | work = Department of Entomology, Purdue University |url= https://www2.illinois.gov/sites/agr/Insects/Pests/Documents/GMquestions%20on%20%20Btk.pdf |archive-url=https://ghostarchive.org/archive/20221009/https://www2.illinois.gov/sites/agr/Insects/Pests/Documents/GMquestions%20on%20%20Btk.pdf |archive-date=2022-10-09 |url-status=live |access-date=2022-04-09}}</ref><ref>{{cite web|title=''Bacillus thuringiensis aizawai'' strain NB200 (006494) Fact sheet|url=https://www3.epa.gov/pesticides/chem_search/reg_actions/registration/fs_PC-006494_10-Jun-05.pdf |archive-url=https://web.archive.org/web/20170120233800/https://www3.epa.gov/pesticides/chem_search/reg_actions/registration/fs_PC-006494_10-Jun-05.pdf |archive-date=2017-01-20 |url-status=live|website=3.epa.gov|access-date=2022-04-09}}</ref> Some Bti lineages are clonal.<ref name="Kolsto-et-al-2009" />
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