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{{Short description|Species of bacterium}}
{{italic title}}
{{Speciesbox
| image = Bacillus_anthracis.png
| image_caption = Photomicrograph of ''Bacillus anthracis'',<br>stained using [[fuchsin]]-[[methylene blue]] (spore stain)
| genus = Bacillus
| species = anthracis
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''B. anthracis'' measures about 3 to 5 μm long and 1 to 1.2 μm wide. The reference genome consists of a 5,227,419 bp circular chromosome and two extrachromosomal DNA [[plasmids]], pXO1 and pXO2, of 181,677 and 94,830 bp respectively,<ref>{{cite web |url=https://www.ncbi.nlm.nih.gov/genome/?term=txid1392 |title=Reference genome: ''Bacillus anthracis'' str. 'Ames Ancestor' |author=<!--Not stated--> |date=February 13, 2022 |website=NCBI Genomes |publisher=[[National Center for Biotechnology Information]] |access-date=February 28, 2022}}</ref> which are responsible for the pathogenicity. It forms a protective layer called [[endospore]] by which it can remain inactive for many years and suddenly becomes infective under suitable environmental conditions. Because of the resilience of the endospore, the bacterium is one of the most popular [[biological weapon]]s. The [[Bacterial capsule|protein capsule]] (poly-D-gamma-glutamic acid) is key to evasion of the immune response. It feeds on the heme of blood protein [[haemoglobin]] using two secretory [[siderophore]] proteins, IsdX1 and IsdX2.
[[File:Bacillus anthracis belongs to the Bacillus cereus group of strains.png|thumb|Phylogenic tree showing ''B. anthracis''
[[Image:B anthracis diagram en.png|thumb|right|Structure of ''B. anthracis'']]Untreated ''B. anthracis'' infection is usually deadly. Infection is indicated by inflammatory, black, necrotic lesions ([[eschar
==Description==
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''B. anthracis'' are [[bacillus (shape)|rod-shaped]] bacteria, approximately 3 to 5 μm long and 1 to 1.2 μm wide.<ref name="Bergeys">{{cite book |doi=10.1002/9781118960608.gbm00530 |chapter=Bacillus |title=Bergey's Manual of Systematics of Archaea and Bacteria |year=2015 |last1=Logan |first1=Niall A. |last2=Vos |first2=Paul De |pages=1–163 |isbn=978-1-118-96060-8 }}</ref> When grown in [[Microbiological culture|culture]], they tend to form long chains of bacteria. On [[agar]] plates, they form large colonies several millimeters across that are generally white or cream colored.<ref name="Bergeys" /> Most ''B. anthracis'' strains produce a [[Bacterial capsule|capsule]] that gives colonies a slimy mucus-like appearance.<ref name="Bergeys" />
It is one of few bacteria known to synthesize a weakly immunogenic and antiphagocytic [[Bacterial capsule|protein capsule]] (poly-D-gamma-glutamic acid) that disguises the vegetative bacterium from the host immune system.<ref>Choo, M. K., Sano, Y., Kim, C., Yasuda, K., Li, X. D., Lin, X., … Park, J. M. (2017). TLR sensing of bacterial spore-associated RNA triggers host immune responses with detrimental effects. ''Journal of Experimental Medicine'', ''214''(5), 1297–1311.
Like ''[[Bordetella pertussis]]'', it forms a [[calmodulin]]-dependent [[adenylate cyclase]] exotoxin{{further explanation needed|date=June 2024}} known as [[Anthrax toxin|anthrax edema factor]], along with [[Anthrax lethal factor endopeptidase|anthrax lethal factor]]. It bears close [[genotypic]] and [[phenotypic]] resemblance to ''[[Bacillus cereus]]'' and ''[[Bacillus thuringiensis]]''. All three species share cellular dimensions and [[morphology (biology)|morphology]]. All form oval [[spore]]s located centrally in an unswollen [[sporangium]]. ''B. anthracis'' endospores, in particular, are highly resilient, surviving extremes of temperature, low-nutrient environments, and harsh chemical treatment over decades or centuries.{{cn|date=March 2023}}
The endospore is a dehydrated cell with thick walls and additional layers that form inside the cell membrane. It can remain inactive for many years, but if it comes into a favorable environment, it begins to grow again. It initially develops inside the rod-shaped form. Features such as the location within the rod, the size and shape of the endospore, and whether or not it causes the wall of the rod to bulge out are characteristic of particular species of ''Bacillus''. Depending upon the species, the endospores are round, oval, or occasionally cylindrical. They are highly [[wikt:refractile|refractile]] and contain [[dipicolinic acid]]. Electron micrograph sections show they have a thin outer endospore coat, a thick [[spore cortex]], and an inner [[spore membrane]] surrounding the endospore contents. The endospores resist heat, drying, and many disinfectants (including 95% ethanol).<ref>Bergey's Manual of Systematic Bacteriology, vol. 2, p. 1105, 1986, Sneath, P.H.A.; Mair, N.S.; Sharpe, M.E.; Holt, J.G. (eds.); Williams & Wilkins, Baltimore, Maryland, USA</ref> Because of these attributes, ''B. anthracis'' endospores are extraordinarily well-suited to use (in powdered and aerosol form) as [[biological weapon]]s. Such weaponization has been accomplished in the past by at least five state bioweapons programs—those of the [[United Kingdom]], [[Imperial Japan|Japan]], the [[United States]], [[Russia]], and [[Iraq]]—and has been attempted by several others.<ref>Zilinskas, Raymond A. (1999), "Iraq's Biological Warfare Program: The Past as Future?", Chapter 8 in: [[Joshua Lederberg|Lederberg, Joshua]] (editor), ''Biological Weapons: Limiting the Threat'' (1999), [[The MIT Press]], pp 137-158.</ref>
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==Strains==
The 89 known strains of ''B. anthracis'' include:
* [[Sterne strain]] (34F2; aka the "Weybridge strain"), used by [[Max Sterne]] in his 1930s vaccines
* [[Vollum strain]], formerly weaponized by the US, UK, and Iraq; isolated from a cow in [[Oxfordshire]], UK, in 1935
** Vollum M-36, virulent British research strain; passed through macaques 36 times
** Vollum 1B, weaponized by the US and UK in the 1940s-60s
** Vollum-14578, used in UK bio-weapons trials which severely contaminated [[Gruinard Island]] in 1942
** V770-NP1-R, the avirulent, nonencapsulated strain used in the ''[[BioThrax]]'' vaccine
* Anthrax 836, highly virulent strain weaponized by the USSR; discovered in [[Kirov, Kirov Oblast|Kirov]] in 1953
* [[Ames strain]], isolated from a cow in [[Texas]] in 1981; famously used in [[AMERITHRAX]] letter attacks (2001)
** Ames Ancestor
** Ames Florida
* H9401, isolated from human patient in Korea; used in investigational anthrax vaccines<ref name="Chun" />
==Evolution==
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''B. anthracis'' belongs to the ''B. cereus'' group consisting of the strains: ''B. cereus'', ''B. anthracis'', ''B. thuringiensis'', ''[[Bacillus mycoides|B. mycoides]]'', and ''B. pseudomycoides''. The first three strains are pathogenic or opportunistic to insects or mammals, while the last three are not considered pathogenic. The strains of this group are genetically and phenotypically heterogeneous overall, but some of the strains are more closely related and phylogenetically intermixed at the chromosome level. The ''B. cereus'' group generally exhibits complex genomes and most carry varying numbers of plasmids.<ref name="Kolstø" />
''B. cereus'' is a soil-dwelling bacterium which can colonize the gut of invertebrates as a symbiont<ref>{{cite journal |last1=Jensen |first1=G. B. |last2=Hansen |first2=B. M. |last3=Eilenberg |first3=J. |last4=Mahillon |first4=J. |title=The hidden lifestyles of Bacillus cereus and relatives: The hidden lifestyles of B. cereus and relatives |journal=Environmental Microbiology |date=18 July 2003 |volume=5 |issue=8 |pages=631–640 |doi=10.1046/j.1462-2920.2003.00461.x |pmid=12871230 |doi-access=free }}</ref> and is a frequent cause of food poisoning<ref>{{cite journal |last1=Drobniewski |first1=F A |title=Bacillus cereus and related species |journal=Clinical Microbiology Reviews |date=October 1993 |volume=6 |issue=4 |pages=324–338 |doi=10.1128/cmr.6.4.324 |pmid=8269390 |pmc=358292 }}</ref> It produces an emetic toxin, enterotoxins, and other virulence factors.<ref>{{cite journal |last1=Stenfors Arnesen |first1=Lotte P. |last2=Fagerlund |first2=Annette |last3=Granum |first3=Per Einar |title=From soil to gut: Bacillus cereus and its food poisoning toxins |journal=FEMS Microbiology Reviews |date=July 2008 |volume=32 |issue=4 |pages=579–606 |doi=10.1111/j.1574-6976.2008.00112.x |pmid=18422617 |doi-access=free }}</ref> The enterotoxins and virulence factors are encoded on the chromosome, while the emetic toxin is encoded on a 270-kb plasmid, pCER270.<ref name="Kolstø" />
''B. thuringiensis'' is an microrganism pathogen and is characterized by production of [[Parasporal body|parasporal crystals]] of insecticidal toxins Cry and Cyt.<ref>{{cite journal |last1=Schnepf |first1=E. |last2=Crickmore |first2=N. |last3=Van Rie |first3=J. |last4=Lereclus |first4=D. |last5=Baum |first5=J. |last6=Feitelson |first6=J. |last7=Zeigler |first7=D. R. |last8=Dean |first8=D. H. |title=Bacillus thuringiensis and Its Pesticidal Crystal Proteins |journal=Microbiology and Molecular Biology Reviews |date=1 September 1998 |volume=62 |issue=3 |pages=775–806 |doi=10.1128/MMBR.62.3.775-806.1998 |pmid=9729609 |pmc=98934 }}</ref> The genes encoding these proteins are commonly located on plasmids which can be lost from the organism, making it indistinguishable from ''B. cereus''.<ref name="Kolstø" />
===Pseudogene===
''PlcR'' is a global transcriptional regulator which controls most of the secreted virulence factors in ''B. cereus'' and ''B. thuringiensis''. It is chromosomally encoded and is ubiquitous throughout the cell.<ref>{{cite journal |last1=Agaisse |first1=Herve |last2=Gominet |first2=Myriam |last3=Okstad |first3=Ole Andreas |last4=Kolsto |first4=Anne-Brit |last5=Lereclus |first5=Didier |title=PlcR is a pleiotropic regulator of extracellular virulence factor gene expression in Bacillus thuringiensis |journal=Molecular Microbiology |date=June 1999 |volume=32 |issue=5 |pages=1043–1053 |doi=10.1046/j.1365-2958.1999.01419.x |pmid=10361306 |doi-access=
The ''plcR'' gene is part of a two-gene operon with ''papR''.<ref>{{cite journal |last1=Økstad |first1=Ole A. |last2=Gominet |first2=Myriam |last3=Purnelle |first3=Bénédicte |last4=Rose |first4=Matthias |last5=Lereclus |first5=Didier |last6=Kolstø |first6=Anne-Brit |title=Sequence analysis of three Bacillus cereus loci carrying PlcR-regulated genes encoding degradative enzymes and enterotoxin |journal=Microbiology |date=1 November 1999 |volume=145 |issue=11 |pages=3129–3138 |doi=10.1099/00221287-145-11-3129 |pmid=10589720 |doi-access=free }}</ref><ref name="Slamti">{{cite journal |last1=Slamti |first1=L. |last2=Lereclus |first2=D |title=A cell-cell signaling peptide activates the PlcR virulence regulon in bacteria of the Bacillus cereus group |journal=The EMBO Journal |date=2 September 2002 |volume=21 |issue=17 |pages=4550–4559 |doi=10.1093/emboj/cdf450 |pmid=12198157 |pmc=126190 }}</ref> The ''papR'' gene encodes a small protein which is secreted from the cell and then reimported as a processed heptapeptide forming a quorum-sensing system.<ref name="Slamti" /><ref>{{cite journal |last1=Bouillaut |first1=L. |last2=Perchat |first2=S. |last3=Arold |first3=S. |last4=Zorrilla |first4=S. |last5=Slamti |first5=L. |last6=Henry |first6=C. |last7=Gohar |first7=M. |last8=Declerck |first8=N. |last9=Lereclus |first9=D. |title=Molecular basis for group-specific activation of the virulence regulator PlcR by PapR heptapeptides |journal=Nucleic Acids Research |date=June 2008 |volume=36 |issue=11 |pages=3791–3801 |doi=10.1093/nar/gkn149 |pmid=18492723 |pmc=2441798 }}</ref> The lack of PlcR in ''B. anthracis'' is a principle characteristic differentiating it from other members of the ''B. cereus'' group. While ''B. cereus'' and ''B. thuringiensis'' depend on the ''plcR'' gene for expression of their virulence factors, ''B. anthracis'' relies on the pXO1 and pXO2 plasmids for its virulence.<ref name="Kolstø" /> [[Bacillus cereus biovar anthracis|''Bacillus cereus'' biovar ''anthracis'']], i.e. ''B. cereus'' with the two plasmids, is also capable of causing anthrax.
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===Pathogenesis===
''B. anthracis'' possesses an antiphagocytic capsule essential for full virulence. The organism also
produces three plasmid-coded exotoxins: edema factor, a calmodulin-dependent adenylate cyclase that causes elevation of intracellular cAMP and is responsible for the severe edema usually seen in ''B. anthracis'' infections, lethal toxin which is responsible for causing tissue necrosis, and protective antigen, so named because of its use in producing protective anthrax vaccines, which mediates cell entry of edema factor and lethal toxin.{{cn|date=November 2023}}
===Manifestations in human disease===
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Four forms of human anthrax disease are recognized based on their [[portal of entry]].
* Cutaneous, the most common form (95%), causes a localized, inflammatory, black, necrotic lesion ([[eschar]]). Most often the sore will appear on the face, neck, arms, or hands. Development can occur within 1–7 days after exposure.
* Inhalation, a rare but highly fatal form, is characterized by flu-like symptoms, chest discomfort, diaphoresis, and body aches.<ref name="Symptoms"/> Development occurs usually a week after exposure, but can take up to two months.
* Gastrointestinal, a rare but also fatal (causes death to 25%) type, results from ingestion of spores. Symptoms include: fever and chills, swelling of neck, painful swallowing, hoarseness, nausea and vomiting (especially bloody vomiting), diarrhea, flushing and red eyes, and swelling of abdomen.<ref name="Symptoms"/> Symptoms can develop within 1–7 days
* Injection, symptoms are similar to those of cutaneous anthrax, but injection anthrax can spread throughout the body faster and can be harder to recognize and treat compared to cutaneous anthrax.<ref name="Symptoms"/> Symptoms include, fever, chills, a group of small bumps or blisters that may itch, appearing where the
===Prevention and treatment===
A number of [[anthrax vaccines]] have been developed for preventive use in livestock and humans. [[Anthrax Vaccine Adsorbed|Anthrax vaccine adsorbed]] (AVA) may protect against cutaneous and inhalation anthrax. However, this vaccine is only used for at-risk adults before exposure to anthrax and has not been approved for use after exposure.<ref>{{Cite web|url=https://www.cdc.gov/anthrax/prevention/index.html|title=How to Prevent Anthrax | CDC|date=December 14, 2020|website=www.cdc.gov}}</ref> Infections with ''B. anthracis'' can be treated with [[Beta-lactam|β-lactam]] [[antibiotic]]s such as [[penicillin]], and others which are active against Gram-positive bacteria.<ref>{{cite journal |last1=Barnes |first1=J. M. |title=Penicillin and B. anthracis |journal=The Journal of Pathology and Bacteriology |date=January 1947 |volume=59 |issue=1–2 |pages=113–125 |doi=10.1002/path.1700590113 |pmid=20266354 }}</ref> Penicillin-resistant ''B. anthracis'' can be treated with [[fluoroquinolones]] such as [[ciprofloxacin]] or tetracycline antibiotics such as [[doxycycline]].{{cn|date=November 2023}}
== Laboratory research ==
Components of [[tea]], such as [[polyphenols in tea|polyphenol]]s, have the ability to inhibit the activity both of ''B. anthracis'' and its toxin considerably; spores, however, are not affected. The addition of milk to the tea completely inhibits its antibacterial activity against anthrax.<ref>{{cite journal |last1=Baillie |first1=Les |last2=Gallagher |first2=Theresa |title=A cup of tea is the answer to everything – including the threat of bio-terrorism |journal=Microbiologist |volume=9 |issue=1 |date=March 2008 |pages=34–37 |url=https://issuu.com/societyforappliedmicrobiology/docs/micro_march08}}
* {{cite press release |date=12 March 2008 |title=Is a cup of tea really the answer to everything -- even anthrax? |website=EurekAlert! |url=https://www.eurekalert.org/pub_releases/2008-03/cu-iac031208.php}}</ref> Activity against the ''B. anthracis'' in the [[laboratory]] does not prove that drinking tea affects the course of an infection, since it is unknown how these polyphenols are absorbed and distributed within the body. ''B. anthracis'' can be cultured on PLET agar, a selective and differential media designed to select specifically for ''B. anthracis''.
===Recent research===
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The H9401 strain isolated in the Republic of Korea was sequenced using [[454 Life Sciences|454]] GS-FLX technology and analyzed using several bioinformatics tools to align, annotate, and compare H9401 to other ''B. anthracis'' strains. The sequencing coverage level suggests a molecular ratio of pXO1:pXO2:chromosome as 3:2:1 which is identical to the Ames Florida and Ames Ancestor strains. H9401 has 99.679% sequence homology with Ames Ancestor with an [[amino acid]] sequence homology of 99.870%. H9401 has a circular chromosome (5,218,947 bp with 5,480 predicted [[Open reading frame|ORF]]s), the pXO1 plasmid (181,700 bp with 202 predicted ORFs), and the pXO2 plasmid (94,824 bp with 110 predicted ORFs).<ref name="Chun" /> As compared to the Ames Ancestor chromosome above, the H9401 chromosome is about 8.5 kb smaller. Due to the high pathogenecity and sequence similarity to the Ames Ancestor, H9401 will be used as a reference for testing the efficacy of candidate anthrax vaccines by the Republic of Korea.<ref name="Chun" />
Since the genome of B. anthracis was sequenced, alternative ways to battle this disease are being endeavored. Bacteria have developed several strategies to evade recognition by the immune system. The predominant mechanism for avoiding detection, employed by all bacteria is molecular camouflage. Slight modifications in the outer layer that render the bacteria practically invisible to lysozymes.<ref>{{cite journal |last1=Callewaert |first1=Lien |last2=Michiels |first2=Chris W. |title=Lysozymes in the animal kingdom |journal=Journal of Biosciences |date=March 2010 |volume=35 |issue=1 |pages=127–160 |doi=10.1007/s12038-010-0015-5 |pmid=20413917 |s2cid=21198203 }}</ref> Three of these modifications have been identified and characterized. These include (1) N-glycosylation of N-acetyl-muramic acid, (2) O-acetylation of N-acetylmuramic acid and (3) N-deacetylation of N-acetyl-glucosamine. Research during the last few years has focused on inhibiting such modifications.<ref>{{cite book |last1=Balomenou |first1=Stavroula |last2=Arnaouteli |first2=Sofia |last3=Koutsioulis |first3=Dimitris |last4=Fadouloglou |first4=Vassiliki E. |last5=Bouriotis |first5=Vassilis |chapter=Polysaccharide Deacetylases: New Antibacterial Drug Targets |pages=68–130 |chapter-url=https://books.google.com/books?id=_ANVDgAAQBAJ&pg=PA68 |editor1-last=Choudhary |editor1-first=M. Iqbal |title=Frontiers in Anti-Infective Drug Discovery |date=2015 |publisher=Bentham Science Publishers |isbn=978-1-68108-082-6 }}</ref> As a result the enzymatic mechanism of polysaccharide de-acetylases is being investigated, that catalyze the removal of an acetyl group from N-acetyl-glucosamine and N-acetyl-muramic acid, components of the peptidoglycan layer.{{cn|date=November 2023}}
== Host interactions ==
As with most other pathogenic bacteria, ''B. anthracis'' must acquire iron to grow and proliferate in its host environment. The most readily available iron sources for pathogenic bacteria are the [[heme]] groups used by the host in the transport of oxygen. To scavenge heme from host [[hemoglobin]] and [[myoglobin]], ''B. anthracis'' uses two secretory [[siderophore]] proteins, IsdX1 and IsdX2. These proteins can separate heme from hemoglobin, allowing surface proteins of ''B. anthracis'' to transport it into the cell.<ref>{{cite journal |last1=Maresso |first1=Anthony W. |last2=Garufi |first2=Gabriella |last3=Schneewind |first3=Olaf |title=Bacillus anthracis Secretes Proteins That Mediate Heme Acquisition from Hemoglobin |journal=PLOS Pathogens |date=22 August 2008 |volume=4 |issue=8 |pages=e1000132 |doi=10.1371/journal.ppat.1000132 |pmid=18725935 |pmc=2515342 |doi-access=free }}</ref>
''B. anthracis'' must evade the immune system to establish a successful infection. ''B. anthracis'' spores are immediately phagocytosed by macrophages and dendritic cells once they enter the host. The [[Dendritic cell#:~:text=Dendritic cells (DCs) are antigen, and the adaptive immune systems.|dendritic cells]] can control the infection through effective intracellular elimination, but the [[macrophage]]s can transport the bacteria directly inside the host by crossing a thin layer of epithelial or endothelial cells to reach the circulatory system.<ref>Hu, H., & Leppla, S. H. (2009). Anthrax Toxin Uptake by Primary Immune Cells as Determined with a Lethal Factor-β-Lactamase Fusion Protein. ''PLoS ONE'', ''4''(11), 1–6.
==Sampling==
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== Historical background ==
[[Image:Bacillus anthracis - CapD protein crystal structure.jpg|160px|thumb|CapD protein crystal structure of ''B. anthracis'']]
French physician [[Casimir Davaine]] (
Throughout the 19th century, Anthrax was an infection that involved several very important medical developments. The first vaccine containing live organisms was Louis Pasteur's veterinary anthrax vaccine.<ref>{{cite journal |last1=Sternbach |first1=George |title=The history of anthrax |journal=The Journal of Emergency Medicine |date=May 2003 |volume=24 |issue=4 |pages=463–467 |doi=10.1016/s0736-4679(03)00079-9 |pmid=12745053 }}</ref>
{{clear}}
== References ==
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==Further reading==
* {{cite journal |last1=Abakar |first1=Mahamat H. |last2=Mahamat |first2=Hassan H. |title=Properties and Antibiotic Susceptibility of ''Bacillus anthracis'' Isolates from Humans, Cattle and Tabanids, and Evaluation of Tabanid as Mechanical Vector of Anthrax in the Republic of Chad |journal=Jordan Journal of Biological Sciences |volume=5 |issue=3 |date=September 2012 |pages=203–208 |s2cid=36932865 |url=http://jjbs.hu.edu.jo/files/v5n3/Paper_Number_9.pdf }}
* {{cite journal |last1=Edmonds |first1=Jason |last2=Lindquist |first2=H. D. Alan |last3=Sabol |first3=Jonathan |last4=Martinez |first4=Kenneth |last5=Shadomy |first5=Sean |last6=Cymet |first6=Tyler |last7=Emanuel |first7=Peter |title=Multigeneration Cross-Contamination of Mail with Bacillus anthracis Spores |journal=PLOS ONE |date=28 April 2016 |volume=11 |issue=4 |pages=e0152225 |doi=10.1371/journal.pone.0152225 |pmid=27123934 |pmc=4849716 |bibcode=2016PLoSO..1152225E |doi-access=free }}
* {{cite journal |last1=Sekhavati |first1=Mohammad |last2=Tadayon |first2=Keyvan |last3=Ghaderi |first3=Rainak |last4=Banihashemi |first4=Reza |last5=Jabbari |first5=Ahmad Reza |last6=Shokri |first6=Gholamreza |last7=Karimnasab |first7=Nasim |title='In-house' production of DNA size marker from a vaccinal Bacillus anthracis strain |journal=Iranian Journal of Microbiology |date=2015 |volume=7 |issue=1 |pages=45–49 |pmid=26644873 |pmc=4670467 }}
* {{cite journal |last1=Roy |first1=P. Roy |last2=Rashid |first2=M. M. |last3=Ferdoush |first3=M. J. |last4=Dipti |first4=M. |last5=Chowdury |first5=M. G. A. |last6=Mostofa |first6=M. G. |last7=Roy |first7=S. K. |last8=Khan |first8=Mahna |last9=Hossain |first9=M. M. |title=Biochemical and immunological characterization of anthrax spore vaccine in goat |journal=Bangladesh Journal of Veterinary Medicine |date=2013 |volume=11 |issue=2 |pages=151–157 |doi=10.3329/bjvm.v11i2.19140 |doi-access=free }}
* {{cite journal |last1=Kusar |first1=D. |last2=Pate |first2=M. |last3=Hubad |first3=B. |last4=Avbersek |first4=J. |last5=Logar |first5=K. |last6=Lapanje |first6=A. |last7=Zrimec |first7=A. |last8=Ocepek |first8=M. |title=Detection of Bacillus anthracis in the air, soil and animal tissue |journal=Acta Veterinaria |date=2012 |volume=62 |issue=1 |pages=77–89 |doi=10.2298/AVB1201077K |url=http://www.doiserbia.nb.rs/ft.aspx?id=0567-83151201077K |doi-access=free }}
== External links ==
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