Biology:Bacillus

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Bacillus, from Latin "bacillus", meaning "little staff, wand", is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum Bacillota, with 266 named species. The term is also used to describe the shape (rod) of other so-shaped bacteria; and the plural Bacilli is the name of the class of bacteria to which this genus belongs. Bacillus species can be either obligate aerobes which are dependent on oxygen, or facultative anaerobes which can survive in the absence of oxygen. Cultured Bacillus species test positive for the enzyme catalase if oxygen has been used or is present.[1]

Bacillus can reduce themselves to oval endospores and can remain in this dormant state for years. The endospore of one species from Morocco is reported to have survived being heated to 420 °C.[2] Endospore formation is usually triggered by a lack of nutrients: the bacterium divides within its cell wall, and one side then engulfs the other. They are not true spores (i.e., not an offspring).[3] Endospore formation originally defined the genus, but not all such species are closely related, and many species have been moved to other genera of the Bacillota.[4] Only one endospore is formed per cell. The spores are resistant to heat, cold, radiation, desiccation, and disinfectants. Bacillus anthracis needs oxygen to sporulate; this constraint has important consequences for epidemiology and control.[5] In vivo, B. anthracis produces a polypeptide (polyglutamic acid) capsule that kills it from phagocytosis.[5] The genera Bacillus and Clostridium constitute the family Bacillaceae. Species are identified by using morphologic and biochemical criteria.[1] Because the spores of many Bacillus species are resistant to heat, radiation, disinfectants, and desiccation, they are difficult to eliminate from medical and pharmaceutical materials and are a frequent cause of contamination. Not only are they resistant to heat, radiation, etc., but they are also resistant to chemicals such as antibiotics.[6] This resistance allows them to survive for many years and especially in a controlled environment.[6] Bacillus species are well known in the food industries as troublesome spoilage organisms.[1]

Ubiquitous in nature, Bacillus includes symbiotic (sometimes referred to as endophytes) as well as independent species. Two species are medically significant: B. anthracis causes anthrax;[7] and B. cereus causes food poisoning.[8]

Many species of Bacillus can produce copious amounts of enzymes, which are used in various industries, such as in the production of alpha amylase used in starch hydrolysis and the protease subtilisin used in detergents. B. subtilis is a valuable model for bacterial research. Some Bacillus species can synthesize and secrete lipopeptides, in particular surfactins and mycosubtilins.[9][10][11] Bacillus species are also found in marine sponges.[11] Marine sponge associated Bacillus subtilis (strains WS1A and YBS29) can synthesize several antimicrobial peptides.[11][12] These Bacillus subtilis strains can develop disease resistance in Labeo rohita.[11]

Structure

Cell wall

Bacillus subtilis (right|thumb The cell wall of Bacillus is a structure on the outside of the cell that forms the second barrier between the bacterium and the environment, and at the same time maintains the rod shape and withstands the pressure generated by the cell's turgor. The cell wall is made of teichoic and teichuronic acids. B. subtilis is the first bacterium for which the role of an actin-like cytoskeleton in cell shape determination and peptidoglycan synthesis was identified and for which the entire set of peptidoglycan-synthesizing enzymes was localized. The role of the cytoskeleton in shape generation and maintenance is important.[13]

Bacillus species are rod-shaped, endospore-forming aerobic or facultatively anaerobic, Gram-positive bacteria; in some species cultures may turn Gram-negative with age. The many species of the genus exhibit a wide range of physiologic abilities that allow them to live in every natural environment. Only one endospore is formed per cell. The spores are resistant to heat, cold, radiation, desiccation, and disinfectants.[1]

Origin of name

The genus Bacillus was named in 1835 by Christian Gottfried Ehrenberg, to contain rod-shaped (bacillus) bacteria. He had seven years earlier named the genus Bacterium. Bacillus was later amended by Ferdinand Cohn to further describe them as spore-forming, Gram-positive, aerobic or facultatively anaerobic bacteria.[14] Like other genera associated with the early history of microbiology, such as Pseudomonas and Vibrio, the 266 species of Bacillus are ubiquitous.[15] The genus has a very large ribosomal 16S diversity.[16]

Isolation and identification

Established methods for isolating Bacillus species for culture primarily involve suspension of sampled soil in distilled water, heat shock to kill off vegetative cells leaving primarily viable spores in the sample, and culturing on agar plates with further tests to confirm the identity of the cultured colonies.[17] Additionally, colonies which exhibit characteristics typical of Bacillus bacteria can be selected from a culture of an environmental sample which has been significantly diluted following heat shock or hot air drying to select potential Bacillus bacteria for testing.[18]

Cultured colonies are usually large, spreading, and irregularly shaped. Under the microscope, the Bacillus cells appear as rods, and a substantial portion of the cells usually contain oval endospores at one end, making them bulge.[19]

Characteristics of Bacillus spp.

S.I. Paul et al. (2021)[11] isolated and identified multiple strains of Bacillus subtilis (strains WS1A,[20] YBS29,[21] KSP163A,[22] OA122,[23] ISP161A,[24] OI6,[25] WS11,[26] KSP151E,[27] and S8,[28]) from marine sponges of the Saint Martin's Island Area of the Bay of Bengal, Bangladesh. Based on their study, colony, morphological, physiological, and biochemical characteristics of Bacillus spp. are shown in the Table below.[11]

Test type Test Characteristics
Colony characters Size Medium
Type Round
Color Whitish
Shape Convex
Morphological characters Shape Rod
Physiological characters Motility +
Growth at 6.5% NaCl +
Biochemical characters Gram's staining +
Oxidase -
Catalase +
Oxidative-Fermentative O/F
Motility +
Methyl Red +
Voges-Proskauer -
Indole -
H2S Production +/–
Urease -
Nitrate reductase +
β-Galactosidase +
Hydrolysis of Gelatin +
Aesculin +
Casein +
Tween 40 +
Tween 60 +
Tween 80 +
Acid production from Glycerol +
Galactose +
D-Glucose +
D-Fructose +
D-Mannose +
Mannitol +
N-Acetylglucosamine +
Amygdalin +
Maltose +
D-Melibiose +
D-Trehalose +
Glycogen +
D-Turanose +

Note: + = Positive, – =Negative, O= Oxidative, F= Fermentative

Phylogeny

It's been long known that the (pre-2020) definition of Bacillus is overly vague.[29]

One clade, formed by Bacillus anthracis, Bacillus cereus, Bacillus mycoides, Bacillus pseudomycoides, Bacillus thuringiensis, and Bacillus weihenstephanensis under the 2011 classification standards, should be a single species (within 97% 16S identity), but for medical reasons, they are considered separate species[39] (an issue also present for four species of Shigella and Escherichia coli).[40]

Alcaraz et al. 2010[35] 16S rRNA based LTP_10_2024[41][42][43] 120 marker proteins based GTDB 09-RS220[44][45][46]
Root
 
"pathogenic"
 

Bacillus weihenstephanensis

 
 

Bacillus cereus/thuringiensis/anthracis

 
 
 
 
 
"soil"
 

Bacillus pumilus

 
 
 

Bacillus subtilis

 
 

Bacillus licheniformis

 
 
 
 
 
 
"benthic"

Geobacillus kaustophilus

 
 
 
 
"aquatic"
 
 

Bacillus coahuilensis

 
 

Bacillus sp. m3-13

 
 
 

Bacillus sp. NRRLB-14911

 
 
 
 
"benthic"

Oceanobacillus iheyensis

 
"halophiles"
 

Bacillus halodurans

 
 

Bacillus clausii

 
 
 
 
 
 
 
Bacillus s.s.
 
 

B. capparidis Wang et al. 2017

 
 

B. gobiensis Liu et al. 2016

 
 
 
 
 
 

B. changyiensis Xiao et al. 2023

 
 

B. haynesii Dunlap et al. 2017

 
 
 
 

B. glycinifermentans Kim et al. 2015

 
 
 

B. paralicheniformis Dunlap et al. 2015

 
 
 
 

B. aerius Shivaji et al. 2006

 
 

B. licheniformis (Weigmann 1898) Chester 1901

 
 
 
 

B. sonorensis Palmisano et al. 2001

 
 

B. swezeyi Dunlap et al. 2017

 
 
 
 
 
 
 
 

B. atrophaeus Nakamura 1989

 
 
 
 

B. altitudinis Shivaji et al. 2006

 
 
 
 

B. aerophilus Shivaji et al. 2006

 
 

B. xiamenensis Lai, Liu & Shao 2014

 
 
 
 

B. safensis Satomi, La Duc & Venkateswaran 2006

 
 
 

B. australimaris Liu et al. 2016

 
 
 

B. pumilus Meyer & Gottheil 1901

 
 

B. zhangzhouensis Liu et al. 2016

 
 
 
 
 
 
 
 

B. velezensis Ruiz-Garcia et al. 2005

 
 
 
 

B. siamensis Sumpavapol et al. 2010

 
 

B. amyloliquefaciens Fukumoto 1943 ex Priest et al. 1987

 
 
 
 
 

B. vallismortis Roberts, Nakamura & Cohan 1996

 
 
 

B. mexicanus de los Santos Villalobos et al. 2023

 
 

B. stercoris (Adelskov & Patel 2017) Dunlap, Bowman & Zeigler 2020

 
 
 
 
 

B. spizizenii (Nakamura et al. 1999) Dunlap, Bowman & Zeigler 2020

 
 
 
 

B. tequilensis Gatson et al. 2006

 
 
 

B. halotolerans (Delaporte & Sasson 1967) Tindall 2017

 
 

B. mojavensis Roberts, Nakamura & Cohan 1994

 
 
 
 
 

B. nakamurai Dunlap et al. 2016

 
 
 

B. cabrialesii tritici de los Santos-Villalobos et al. 2023

 
 
 

B. cabrialesii de los Santos Villalobos et al. 2019

 
 
 

B. inaquosorum (Rooney et al. 2009) Dunlap, Bowman & Zeigler 2020

 
 

B. subtilis (Ehrenberg 1835) Cohn 1872

 
 
 
 
 
 
 
 
 
 
 
 
 
 
Bacillus s.s.
 

B. gobiensis [B. capparidis]

 
 
 
 
 

B. glycinifermentans

 
 

B. sonorensis

 
 
 
 

B. swezeyi

 
 
 

B. licheniformis

 
 
 

B. haynesii

 
 

B. paralicheniformis [B. crescens Shivani et al. 2015]

 
 
 
 
 
 
 
 
 

B. altitudinis [B. aerius; B. aerophilus]

 
 

B. xiamenensis

 
 
 
 

B. zhangzhouensis

 
 
 

B. pumilus

 
 
 

B. australimaris

 
 

B. safensis

 
 
 
 
 
 
 

B. atrophaeus

 
 
 
 

B. nakamurai

 
 
 

B. amyloliquefaciens

 
 
 

B. siamensis

 
 

B. velezensis [B. lentimorbus Dutky 1940]

 
 
 
 
 
 
 

B. halotolerans

 
 

B. mojavensis

 
 
 
 

B. tequilensis

 
 
 
 

B. vallismortis

 
 
 

"B. rugosus" Bhattacharya et al. 2020

 
 

B. spizizenii

 
 
 
 
 

B. cabrialesii

 
 
 

B. inaquosorum

 
 
 

B. stercoris

 
 

B. subtilis [Paenibacillus solisilvae Kong et al. 2020]

 
 
 
 
 
 
 
 
 
 
 
 
 
Metabacillus_D

Metabacillus arenae

 
 
Bacillus_A
 

B. rhizoplanae

 
 
 

B. manliponensis

 
 
 

B. cytotoxicus

 
 
 
 
 

B. gaemokensis

 
 
 
 

B. arachidis

 
 

B. bingmayongensis

 
 
 
 

B. pseudomycoides

 
 
 
 
 
 
 

B. clarus

 
 
 
 
 
 
 
 
 
 

B. paramycoides

 
 
 
 

B. nitratireducens

 
 

B. proteolyticus

 
 
 
 

B. hominis

 
 

B. mycoides

 
 
 
 
 

B. toyonensis

 
 
 
 
 
 

B. wiedmannii

 
 

B. mobilis

 
 
 
 
 

B. luti

 
 
 
 
 
 

B. sanguinis

 
 
 
 

B. albus

 
 
 
 
 

B. anthracis

 
 
 

B. pacificus

 
 

B. paranthracis

 
 
 
 

B. tropicus

 
 
 
 
 
 
 

B. fungorum

 
 
 
 
 
 
 

B. bombysepticus

 
 

B. cereus

 
 
 

B. thuringiensis

 
 
 
 
 
 
 
 
 
 

Ectobacillus

 

Species

Species orphaned and assigned to other genera:

  • B. Symun
  • B. acidicola
  • B. acidiproducens
  • B. acidocaldarius
  • B. acidoterrestris
  • B. aeolius
  • B. agaradhaerens
  • B. agri
  • B. aidingensis
  • B. akibai
  • B. albus
  • B. alcalophlus
  • B. algicola
  • B. alginolyticus
  • B. alkalidiazotrophicus
  • B. alkalinitrilicus
  • B. alkalisediminis
  • B. alkalitelluris
  • B. alveayuensis
  • B. alvei
  • B. aminovorans[47]
  • B. amylolyticus
  • B. andreesenii
  • B. aneurinilyticus
  • B. anthracis
  • B. aquimaris
  • B. arenosi
  • B. arseniciselenatis
  • B. arsenicus
  • B. aurantiacus
  • B. arvi
  • B. aryabhattai
  • B. asahii
  • B. axarquiensis
  • B. azotofixans
  • B. azotoformans
  • B. badius
  • B. barbaricus
  • B. bataviensis
  • B. beijingensis
  • B. benzoevorans
  • B. beringensis
  • B. berkeleyi
  • B. beveridgei
  • B. bogoriensis
  • B. boroniphilus
  • B. borstelensis
  • B. brevis
  • B. butanolivorans
  • B. canaveralius
  • B. carboniphilus
  • B. cecembensis
  • B. cellulosilyticus
  • B. centrosporus
  • B. cereus
  • B. chagannorensis
  • B. chitinolyticus
  • B. chondroitinus
  • B. choshinensis
  • B. chungangensis
  • B. cibi
  • B. circulans
  • B. clarkii
  • B. clausii
  • B. coagulans
  • B. coahuilensis
  • B. cohnii
  • B. composti
  • B. curdlanolyticus
  • B. cycloheptanicus
  • B. cytotoxicus
  • B. daliensis
  • B. decisifrondis
  • B. decolorationis
  • B. deserti
  • B. dipsosauri
  • B. drentensis
  • B. edaphicus
  • B. ehimensis
  • B. eiseniae
  • B. enclensis
  • B. endophyticus
  • B. endoradicis
  • B. farraginis
  • B. fastidiosus
  • B. fengqiuensis
  • B. filobacterium rodentuim
  • B. firmus
  • B. flexus
  • B. foraminis
  • B. fordii
  • B. formosus
  • B. fortis
  • B. fumarioli
  • B. funiculus
  • B. fusiformis
  • B. gaemokensis
  • B. galactophilus
  • B. galactosidilyticus
  • B. galliciensis
  • B. gelatini
  • B. gibsonii
  • B. ginsengi
  • B. ginsengihumi
  • B. ginsengisoli
  • B. glucanolyticus
  • B. gordonae
  • B. gottheilii
  • B. graminis
  • B. halmapalus
  • B. haloalkaliphilus
  • B. halochares
  • B. halodenitrificans
  • B. halodurans
  • B. halophilus
  • B. halosaccharovorans
  • B. hemicellulosilyticus
  • B. hemicentroti
  • B. herbersteinensis
  • B. horikoshii
  • B. horneckiae
  • B. horti
  • B. huizhouensis
  • B. humi
  • B. hwajinpoensis
  • B. idriensis
  • B. indicus
  • B. infantis
  • B. infernus
  • B. insolitus
  • B. invictae
  • B. iranensis
  • B. isabeliae
  • B. isronensis
  • B. jeotgali
  • B. kaustophilus
  • B. kobensis
  • B. kochii
  • B. kokeshiiformis
  • B. koreensis
  • B. korlensis
  • B. kribbensis
  • B. krulwichiae
  • B. laevolacticus
  • B. larvae
  • B. laterosporus
  • B. lautus
  • B. lehensis
  • B. lentus
  • B. ligniniphilus
  • B. litoralis
  • B. locisalis
  • B. luciferensis
  • B. luteolus
  • B. luteus
  • B. macauensis
  • B. macerans
  • B. macquariensis
  • B. macyae
  • B. malacitensis
  • B. mannanilyticus
  • B. marisflavi
  • B. marismortui
  • B. marmarensis
  • B. massiliensis
  • B. megaterium
  • "B. mesentericus"
  • B. mesonae
  • B. methanolicus
  • B. methylotrophicus
  • B. migulanus
  • B. mucilaginosus
  • B. muralis
  • B. murimartini
  • B. mycoides
  • B. naganoensis
  • B. nanhaiensis
  • B. nanhaiisediminis
  • B. nealsonii
  • B. neidei
  • B. neizhouensis
  • B. niabensis
  • B. niacini
  • B. novalis
  • B. oceanisediminis
  • B. odysseyi
  • B. okhensis
  • B. okuhidensis
  • B. oleronius
  • B. oryzaecorticis
  • B. oshimensis
  • B. pabuli
  • B. pakistanensis
  • B. pallidus
  • B. pallidus
  • B. panacisoli
  • B. panaciterrae
  • B. pantothenticus
  • B. parabrevis
  • B. paraflexus
  • B. pasteurii
  • B. patagoniensis
  • B. peoriae
  • B. persepolensis
  • B. persicus
  • B. pervagus
  • B. plakortidis
  • B. pocheonensis
  • B. polygoni
  • B. polymyxa
  • B. popilliae
  • B. pseudalcalophilus
  • B. pseudofirmus
  • B. pseudomycoides
  • B. psychrodurans
  • B. psychrophilus
  • B. psychrosaccharolyticus
  • B. psychrotolerans
  • B. pulvifaciens
  • B. purgationiresistens
  • B. pycnus
  • B. qingdaonensis
  • B. qingshengii
  • B. reuszeri
  • B. rhizosphaerae
  • B. rigui
  • B. ruris
  • B. salarius
  • B. salexigens
  • B. saliphilus
  • B. schlegelii
  • B. sediminis
  • B. selenatarsenatis
  • B. selenitireducens
  • B. seohaeanensis
  • B. shacheensis
  • B. shackletonii
  • B. silvestris
  • B. simplex
  • B. siralis
  • B. smithii
  • B. soli
  • B. solimangrovi
  • B. solisalsi
  • B. songklensis
  • B. sphaericus
  • B. sporothermodurans
  • B. stearothermophilus
  • B. stratosphericus
  • B. subterraneus
  • B. taeanensis
  • B. thermantarcticus
  • B. thermoaerophilus
  • B. thermoamylovorans
  • B. thermocatenulatus
  • B. thermocloacae
  • B. thermocopriae
  • B. thermodenitrificans
  • B. thermoglucosidasius
  • B. thermolactis
  • B. thermoleovorans
  • B. thermophilus
  • B. thermoproteolyticus
  • B. thermoruber
  • B. thermosphaericus
  • B. thiaminolyticus
  • B. thioparans
  • B. thuringiensis
  • B. tianshenii
  • B. toyonensis
  • B. trypoxylicola
  • B. tusciae
  • B. validus
  • B. vedderi
  • B. vietnamensis
  • B. vireti
  • B. vulcani
  • B. wakoensis
  • B. xiaoxiensis
  • B. zanthoxyli
  • B. zhanjiangensis

Ecological and clinical significance

Bacillus species are ubiquitous in nature, e.g. in soil. They can occur in extreme environments such as high pH (B. alcalophilus), high temperature (B. thermophilus), and high salt concentrations (B. halodurans). They also are very commonly found as endophytes in plants where they can play a critical role in their immune system, nutrient absorption and nitrogen fixing capabilities.[48][49][50][51][52] B. thuringiensis produces a toxin that can kill insects and thus has been used as insecticide.[53] B. siamensis has antimicrobial compounds that inhibit plant pathogens, such as the fungi Rhizoctonia solani and Botrytis cinerea, and they promote plant growth by volatile emissions.[54] Some species of Bacillus are naturally competent for DNA uptake by transformation.[55]

  • Two Bacillus species are medically significant: B. anthracis, which causes anthrax; and B. cereus, which causes food poisoning, with symptoms similar to that caused by Staphylococcus.[56]
    • B. cereus produces toxins which cause two different set of symptoms:
      • emetic toxin which can cause vomiting and nausea
      • diarrhea
  • B. thuringiensis is an important insect pathogen, and is sometimes used to control insect pests.
  • B. subtilis is an important model organism. It is also a notable food spoiler, causing ropiness in bread and related food.
    • B. subtilis can also produce and secrete antibiotics.
  • Some environmental and commercial strains of B. coagulans may play a role in food spoilage of highly acidic, tomato-based products.

Industrial significance

A portion of the Bacillus thuringiensis genome was incorporated into corn[57] and cotton[58] crops. The resulting plants are resistant to some insect pests.[59]

Bacillus subtilis (natto) is the key microbial participant in the ongoing production of the soya-based traditional natto fermentation, and some Bacillus species are on the Food and Drug Administration's GRAS (generally regarded as safe) list.[60]

The capacity of selected Bacillus strains to produce and secrete large quantities (20–25 g/L) of extracellular enzymes has placed them among the most important industrial enzyme producers. The ability of different species to ferment in the acid, neutral, and alkaline pH ranges, combined with the presence of thermophiles in the genus, has led to the development of a variety of new commercial enzyme products with the desired temperature, pH activity, and stability properties to address a variety of specific applications. Classical mutation and (or) selection techniques, together with advanced cloning and protein engineering strategies, have been exploited to develop these products.{{citation needed|date=February 2023} Efforts to produce and secrete high yields of foreign recombinant proteins in Bacillus hosts initially appeared to be hampered by the degradation of the products by the host proteases. Recent studies have revealed that the slow folding of heterologous proteins at the membrane-cell wall interface of Gram-positive bacteria renders them vulnerable to attack by wall-associated proteases.{{citation needed|date=February 2023} isulphide oxidoreductases in B. subtilis may be beneficial in the secretion of disulphide-bond-containing proteins. Such developments from our understanding of the complex protein translocation machinery of Gram-positive bacteria should allow the resolution of current secretion challenges and make Bacillus species preeminent hosts for heterologous protein production.[citation needed] Bacillus strains have also been developed and engineered as industrial producers of nucleotides, the vitamin riboflavin, the flavor agent ribose, and the supplement poly-gamma-glutamic acid. With the recent characterization of the genome of B. subtilis 168 and of some related strains, Bacillus species are poised to become the preferred hosts for the production of many new and improved products as we move through the genomic and proteomic era.[61]

Use as model organism

Colonies of the model species Bacillus subtilis on an agar plate

Bacillus subtilis is one of the best understood prokaryotes, in terms of molecular and cellular biology. Its superb genetic amenability and relatively large size have provided the powerful tools required to investigate a bacterium from all possible aspects. Recent improvements in fluorescent microscopy techniques have provided novel insight into the dynamic structure of a single cell organism. Research on B. subtilis has been at the forefront of bacterial molecular biology and cytology, and the organism is a model for differentiation, gene/protein regulation, and cell cycle events in bacteria.[62]

See also

References

  1. ↑ 1.0 1.1 1.2 1.3 "Bacillus". Barron's Medical Microbiology (4th ed.). Univ of Texas Medical Branch. 1996. ISBN 978-0-9631172-1-2. https://www.ncbi.nlm.nih.gov/books/NBK7699/. Retrieved 2019-11-14. 
  2. ↑ "Life from the ashes: survival of dry bacterial spores after very high temperature exposure". Extremophiles: Life Under Extreme Conditions 22 (5): 751–759. September 2018. doi:10.1007/s00792-018-1035-6. PMID 29869718. 
  3. ↑ "Bacterial Endospores". Cornell University College of Agriculture and Life Sciences, Department of Microbiology. https://cals.cornell.edu/microbiology/research/active-research-labs/angert-lab/epulopiscium/bacterial-endospores. 
  4. ↑ Brock Biology of Microorganisms (11th ed.). Prentice Hall. 2005. ISBN 978-0-13-144329-7. 
  5. ↑ 5.0 5.1 "Bacillus". Medical Microbiology (4th ed.). Galveston (TX): University of Texas Medical Branch at Galveston. 1996. ISBN 978-0-9631172-1-2. https://www.ncbi.nlm.nih.gov/books/NBK7699/. Retrieved 2024-03-18. 
  6. ↑ 6.0 6.1 "Bacillus spore germination: Knowns, unknowns and what we need to learn". Cellular Signalling 74. October 2020. doi:10.1016/j.cellsig.2020.109729. PMID 32721540. 
  7. ↑ "Bacillus anthracis". Journal of Clinical Pathology 56 (3): 182–187. March 2003. doi:10.1136/jcp.56.3.182. PMID 12610093. 
  8. ↑ "Bacillus cereus food poisoning and its toxins". Journal of Food Protection 68 (3): 636–648. March 2005. doi:10.4315/0362-028X-68.3.636. PMID 15771198. 
  9. ↑ "Biocontrol traits of Bacillus licheniformis GL174, a culturable endophyte of Vitis vinifera cv. Glera". BMC Microbiology 18 (1). October 2018. doi:10.1186/s12866-018-1306-5. PMID 30326838. 
  10. ↑ "Characterization of lipopeptides produced by Bacillus licheniformis using liquid chromatography with accurate tandem mass spectrometry". Rapid Communications in Mass Spectrometry 30 (20): 2237–2252. October 2016. doi:10.1002/rcm.7705. PMID 27487987. Bibcode: 2016RCMS...30.2237F. 
  11. ↑ 11.0 11.1 11.2 11.3 11.4 11.5 "Identification of marine sponge-associated bacteria of the Saint Martin's island of the Bay of Bengal emphasizing on the prevention of motile Aeromonas septicemia in Labeo rohita". Aquaculture 545. 2021-12-15. doi:10.1016/j.aquaculture.2021.737156. Bibcode: 2021Aquac.54537156P. 
  12. ↑ "Whole-Genome Sequence of Bacillus subtilis WS1A, a Promising Fish Probiotic Strain Isolated from Marine Sponge of the Bay of Bengal". Microbiology Resource Announcements 9 (39): e00641–20. September 2020. doi:10.1128/MRA.00641-20. PMID 32972930. 
  13. ↑ "The bacterial cytoskeleton". Microbiology and Molecular Biology Reviews 70 (3): 729–754. September 2006. doi:10.1128/MMBR.00017-06. PMID 16959967. 
  14. ↑ "Untersuchungen über Bakterien." (in DE). Beiträge zur Biologie der Pflanzen 2 (1): 127–224. 1872. 
  15. ↑ Bacillus entry in LPSN [Euzéby, J.P. (1997). "List of Bacterial Names with Standing in Nomenclature: a folder available on the Internet". Int J Syst Bacteriol (Microbiology Society) 47 (2): 590–2. doi:10.1099/00207713-47-2-590. ISSN 0020-7713. PMID 9103655. https://ijs.microbiologyresearch.org/content/journal/ijsem/10.1099/00207713-47-2-590. Retrieved 2019-02-23. ]
  16. ↑ "Diversity of 16S rRNA genes within individual prokaryotic genomes". Applied and Environmental Microbiology 76 (12): 3886–3897. June 2010. doi:10.1128/AEM.02953-09. PMID 20418441. Bibcode: 2010ApEnM..76.3886P. 
  17. ↑ "Selective Process for Efficient Isolation of Soil Bacillus spp". Applied and Environmental Microbiology 53 (6): 1263–1266. June 1987. doi:10.1128/aem.53.6.1263-1266.1987. PMID 16347359. Bibcode: 1987ApEnM..53.1263T. 
  18. ↑ "Isolation and characterization of Bacillus sp. strain BC01 from soil displaying potent antagonistic activity against plant and fish pathogenic fungi and bacteria". Journal, Genetic Engineering & Biotechnology 16 (2): 387–392. December 2018. doi:10.1016/j.jgeb.2018.01.005. PMID 30733751. 
  19. ↑ (in en) The Students Reference Guide to Bacteria. Lulu.com. 2011-04-06. ISBN 978-1-257-42416-0. https://books.google.com/books?id=MpBpAwAAQBAJ&dq=Bacillus+cells+appear+as+rods%2C+and+a+substantial+portion+of+the+cells+usually+contain+oval+endospores+at+one+end%2C&pg=PA35. 
  20. ↑ "Bacillus subtilis strain WS1A". GenBank (U.S. National Library of Medicine). 19 May 2019. https://www.ncbi.nlm.nih.gov/nuccore/MK910097.1/. Retrieved 5 September 2021. 
  21. ↑ "Bacillus subtilis strain YBS29". GenBank (U.S. National Library of Medicine). 19 June 2020. https://www.ncbi.nlm.nih.gov/nuccore/MT605348.1/. Retrieved 5 September 2021. 
  22. ↑ "Bacillus subtilis strain KSP163A". GenBank (U.S. National Library of Medicine). 19 May 2019. https://www.ncbi.nlm.nih.gov/nuccore/MK910103.1/. Retrieved 5 September 2021. 
  23. ↑ "Bacillus subtilis strain OA122". GenBank (U.S. National Library of Medicine). 20 June 2020. https://www.ncbi.nlm.nih.gov/nuccore/MT611945.1/. Retrieved 5 September 2021. 
  24. ↑ "Bacillus subtilis strain ISP161A". GenBank (U.S. National Library of Medicine). 20 June 2020. https://www.ncbi.nlm.nih.gov/nuccore/MT611943.1/. Retrieved 5 September 2021. 
  25. ↑ "Bacillus subtilis strain OI6". GenBank (U.S. National Library of Medicine). 19 June 2020. https://www.ncbi.nlm.nih.gov/nuccore/MT605347.1/. 
  26. ↑ "Bacillus subtilis strain WS11". GenBank (U.S. National Library of Medicine). 19 May 2019. https://www.ncbi.nlm.nih.gov/nuccore/MK910101.1/. 
  27. ↑ "Bacillus subtilis strain KSP151E". GenBank (U.S. National Library of Medicine). 19 June 2020. https://www.ncbi.nlm.nih.gov/nuccore/MT605346.1/. 
  28. ↑ "Bacillus subtilis strain S8". GenBank (U.S. National Library of Medicine). 20 June 2020. https://www.ncbi.nlm.nih.gov/nuccore/MT611946.1/. Retrieved 5 September 2021. 
  29. ↑ 29.0 29.1 Ash, Carol; Farrow, J.A.E.; Wallbanks, Sally; Collins, M.D. (28 June 2008). "Phylogenetic heterogeneity of the genus Bacillus revealed by comparative analysis of small-subunit-ribosomal RNA sequences". Letters in Applied Microbiology 13 (4): 202–206. doi:10.1111/j.1472-765x.1991.tb00608.x. ISSN 0266-8254. http://dx.doi.org/10.1111/j.1472-765x.1991.tb00608.x. Retrieved 13 November 2024. 
  30. ↑ "Phylogenetic relationships between Bacillus species and related genera inferred from comparison of 3' end 16S rDNA and 5' end 16S-23S ITS nucleotide sequences". International Journal of Systematic and Evolutionary Microbiology 53 (Pt 3): 695–704. May 2003. doi:10.1099/Ijs.0.02346-0. PMID 12807189. 
  31. ↑ "The All-Species Living Tree project: a 16S rRNA-based phylogenetic tree of all sequenced type strains". Systematic and Applied Microbiology 31 (4): 241–250. September 2008. doi:10.1016/j.syapm.2008.07.001. PMID 18692976. Bibcode: 2008SyApM..31..241Y. https://tede.ufrrj.br/jspui/handle/jspui/5297. Retrieved 2023-01-03. 
  32. ↑ "Update of the All-Species Living Tree Project based on 16S and 23S rRNA sequence analyses". Systematic and Applied Microbiology 33 (6): 291–299. October 2010. doi:10.1016/j.syapm.2010.08.001. PMID 20817437. Bibcode: 2010SyApM..33..291Y. 
  33. ↑ "Release LTPs104 of the all-species living tree". Systematic and Applied Microbiology 34 (3): 169–70. May 2011. doi:10.1016/j.syapm.2011.03.001. PMID 21497273. Bibcode: 2011SyApM..34..169M. http://www.arb-silva.de/fileadmin/silva_databases/living_tree/LTP_release_104/LTPs104_SSU_tree.pdf. 
  34. ↑ "Comparative Genomics of the Genus Lactobacillus Reveals Robust Phylogroups That Provide the Basis for Reclassification". Applied and Environmental Microbiology 84 (17). September 2018. doi:10.1128/AEM.00993-18. PMID 29915113. Bibcode: 2018ApEnM..84E.993S. 
  35. ↑ 35.0 35.1 "Understanding the evolutionary relationships and major traits of Bacillus through comparative genomics". BMC Genomics 11 (1). May 2010. doi:10.1186/1471-2164-11-332. 1471216411332. PMID 20504335. Bibcode: 2010BMCG...11..332A. 
  36. ↑ 36.0 36.1 Gupta, Radhey S.; Patel, Sudip; Saini, Navneet; Chen, Shu (1 November 2020). "Robust demarcation of 17 distinct Bacillus species clades, proposed as novel Bacillaceae genera, by phylogenomics and comparative genomic analyses: description of Robertmurraya kyonggiensis sp. nov. and proposal for an emended genus Bacillus limiting it only to the members of the subtilis and cereus clades of species" (in en). International Journal of Systematic and Evolutionary Microbiology 70 (11): 5753–5798. doi:10.1099/ijsem.0.004475. ISSN 1466-5026. PMID 33112222. 
  37. ↑ 37.0 37.1 Patel, Sudip; Gupta, Radhey S. (1 January 2020). "A phylogenomic and comparative genomic framework for resolving the polyphyly of the genus Bacillus: Proposal for six new genera of Bacillus species, Peribacillus gen. nov., Cytobacillus gen. nov., Mesobacillus gen. nov., Neobacillus gen. nov., Metabacillus gen. nov. and Alkalihalobacillus gen. nov." (in en). International Journal of Systematic and Evolutionary Microbiology 70 (1): 406–438. doi:10.1099/ijsem.0.003775. ISSN 1466-5026. PMID 31617837. 
  38. ↑ 38.0 38.1 "A Comparative Analysis of the Core Proteomes within and among the Bacillus subtilis and Bacillus cereus Evolutionary Groups Reveals the Patterns of Lineage- and Species-Specific Adaptations". Microorganisms 10 (9): 1720. August 2022. doi:10.3390/microorganisms10091720. PMID 36144322. 
  39. ↑ "Chapter 2: Genomics of Bacillus species". Genomics of Foodborne Bacterial Pathogens. Food Microbiology and Food Safety. 29. New York, NY: Springer. December 2010. pp. 29–53 (34–35). doi:10.1007/978-1-4419-7686-4_2. ISBN 978-1-4419-7686-4. 
  40. ↑ "Family I. Enterobacteriaceae Rahn 1937, Nom. fam. cons. Opin. 15, Jud. Com. 1958, 73; Ewing, Farmer, and Brenner 1980, 674; Judicial Commission 1981, 104.". Bergey's Manual of Systematic Bacteriology. 1 (first ed.). Baltimore: The Williams & Wilkins Co. 1984. pp. 408–420. 
  41. ↑ "The LTP". https://imedea.uib-csic.es/mmg/ltp/#LTP. 
  42. ↑ "LTP_all tree in newick format". https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_all_10_2024.ntree. 
  43. ↑ "LTP_10_2024 Release Notes". https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_10_2024_release_notes.pdf. 
  44. ↑ "GTDB release 09-RS220". https://gtdb.ecogenomic.org/about#4%7C. 
  45. ↑ "bac120_r220.sp_labels". https://data.gtdb.ecogenomic.org/releases/release220/220.0/auxillary_files/bac120_r220.sp_labels.tree. 
  46. ↑ "Taxon History". https://gtdb.ecogenomic.org/taxon_history/. 
  47. ↑ "Nucleotide sequence of the sspE genes coding for gamma-type small, acid-soluble spore proteins from the round-spore-forming bacteria Bacillus aminovorans, Sporosarcina halophila and S. ureae". Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression 1396 (2): 148–152. March 1998. doi:10.1016/S0167-4781(97)00204-2. PMID 9540829. 
  48. ↑ "Isolation and identification of nitrogen-fixing bacilli from plant rhizospheres in Beijing region". Journal of Applied Microbiology 99 (5): 1271–1281. 2005. doi:10.1111/j.1365-2672.2005.02738.x. PMID 16238759. 
  49. ↑ "Isolation and identification of N2-fixing strains of Bacillus in rice rhizosphere of the Yangtze River Valley" (in Chinese). Wei Sheng Wu Xue Bao = Acta Microbiologica Sinica (Chinese Academy of Sciences) 38 (6): 480–483. Dec 1998. PMID 12548929. https://europepmc.org/article/MED/12548929. Retrieved 2022-01-30. 
  50. ↑ "Epiphytic and endophytic bacteria that promote growth of ethnomedicinal plants in the subtropical forests of Meghalaya, India". Revista de Biología Tropical 62 (4): 1295–1308. 2014. doi:10.15517/rbt.v62i4.12138. PMID 25720168. 
  51. ↑ "Nitrogen-fixing bacteria and Oxalis – evidence for a vertically inherited bacterial symbiosis". BMC Plant Biology 19 (1). 2019. doi:10.1186/s12870-019-2049-7. PMID 31646970. Bibcode: 2019BMCPB..19..441J. 
  52. ↑ "Inoculation of zinc solubilizing Bacillus aryabhattai strains for improved growth, mobilization and biofortification of zinc in soybean and wheat cultivated in Vertisols of central India". Applied Soil Ecology 73: 87–96. 2014. doi:10.1016/j.apsoil.2013.08.009. ISSN 0929-1393. Bibcode: 2014AppSE..73...87R. 
  53. ↑ Microbiology: An Evolving Science (2nd ed.). Norton. 2011. 
  54. ↑ "Draft genome sequence of the plant growth-promoting bacterium Bacillus siamensis KCTC 13613T". Journal of Bacteriology 194 (15): 4148–4149. August 2012. doi:10.1128/JB.00805-12. PMID 22815459. 
  55. ↑ "Draft Genome Sequence of the Naturally Competent Bacillus simplex Strain WY10". Genome Announcements 5 (46): e01295–17. November 2017. doi:10.1128/genomeA.01295-17. PMID 29146837. 
  56. ↑ Sherris Medical Microbiology (4th ed.). McGraw Hill. 2004. ISBN 978-0-8385-8529-0. 
  57. ↑ "History of Bt". University of California San Diego. http://www.bt.ucsd.edu/bt_history.html. 
  58. ↑ James, Clive (1996). "Global Review of the Field Testing and Commercialization of Transgenic Plants: 1986 to 1995". The International Service for the Acquisition of Agri-biotech Applications. https://www.isaaa.org/resources/publications/briefs/01/download/isaaa-brief-01-1996.pdf. 
  59. ↑ "Bt Corn: Health and the Environment – 0.707". Colorado State University Extension Office. 2013. http://extension.colostate.edu/docs/pubs/crops/00707.pdf. 
  60. ↑ Schallmey, Marcus; Singh, Ajay; Ward, Owen P (2004-01-01). "Developments in the use of Bacillus species for industrial production". Canadian Journal of Microbiology 50 (1): 1–17. doi:10.1139/w03-076. ISSN 0008-4166. PMID 15052317. https://cdnsciencepub.com/doi/10.1139/w03-076. Retrieved 2025-03-08. 
  61. ↑ "Developments in the use of Bacillus species for industrial production". Canadian Journal of Microbiology 50 (1): 1–17. January 2004. doi:10.1139/w03-076. PMID 15052317. 
  62. ↑ Bacillus: Cellular and Molecular Biology (2nd ed.). Caister Academic Press. 2012. [1]. ISBN 978-1-904455-97-4. http://www.horizonpress.com/bacillus. Retrieved 2011-12-13. 
  63. ↑ "Molecular identification of rRNA group 3 bacilli (Ash, Farrow, Wallbanks and Collins) using a PCR probe test. Proposal for the creation of a new genus Paenibacillus". Antonie van Leeuwenhoek 64 (3–4): 253–260. 1994. doi:10.1007/BF00873085. PMID 8085788. 
  64. ↑ "Virgibacillus: a new genus to accommodate Bacillus pantothenticus (Proom and Knight 1950). Emended description of Virgibacillus pantothenticus". International Journal of Systematic Bacteriology 48 (1): 99–106. January 1998. doi:10.1099/00207713-48-1-99. 
  • Bacillus genomes and related information at PATRIC , a Bioinformatics Resource Center funded by NIAID

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