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Bacillus

Bacillus
Bacillus subtilis, stained
Scientific classification Edit this classification
Domain: Bacteria
Phylum: Bacillota
Class: Bacilli
Order: Caryophanales
Family: Bacillaceae
Genus: Bacillus
Cohn
Species

See text

Bacillus (Latin "stick") 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 (Gram stain)

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]

Bacillus phylogenetics
Phylogeny of the genus Bacillus according to Alcaraz et al. 2010[35]

Species

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.[42][43][44][45][46] B. thuringiensis produces a toxin that can kill insects and thus has been used as insecticide.[47] 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.[48] Some species of Bacillus are naturally competent for DNA uptake by transformation.[49]

  • 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.[50]
    • 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

Many Bacillus species are able to secrete large quantities of enzymes. Bacillus amyloliquefaciens is the source of a natural antibiotic protein barnase (a ribonuclease), alpha amylase used in starch hydrolysis, the protease subtilisin used with detergents, and the BamH1 restriction enzyme used in DNA research.[citation needed]

A portion of the Bacillus thuringiensis genome was incorporated into corn[51] and cotton[52] crops. The resulting plants are resistant to some insect pests.[53]

Bacillus subtilis (natto) is the key microbial participant in the ongoing production of the soya-based traditional natto fermentation, and some Bacillus species like Bacillus cereus are on the Food and Drug Administration's GRAS (generally regarded as safe) list.[54]

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.[citation needed] 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]

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.[citation needed] 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] In addition, the presence of thiol-disulphide 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.[55]

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.[56]

See also

References

  1. ^ a b c d Turnbull PC (1996). "Bacillus". In Baron S, et al. (eds.). Barron's Medical Microbiology (4th ed.). Univ of Texas Medical Branch. ISBN 978-0-9631172-1-2. PMID 21413260.
  2. ^ Beladjal L, Gheysens T, Clegg JS, Amar M, Mertens J (September 2018). "Life from the ashes: survival of dry bacterial spores after very high temperature exposure". Extremophiles: Life Under Extreme Conditions. 22 (5): 751–759. doi:10.1007/s00792-018-1035-6. PMID 29869718. S2CID 46935396.
  3. ^ "Bacterial Endospores". Cornell University College of Agriculture and Life Sciences, Department of Microbiology. Retrieved 21 October 2018.
  4. ^ Madigan M, Martinko J, eds. (2005). Brock Biology of Microorganisms (11th ed.). Prentice Hall. ISBN 978-0-13-144329-7.
  5. ^ a b Turnbull PC (1996). "Bacillus". In Baron S (ed.). Medical Microbiology (4th ed.). Galveston (TX): University of Texas Medical Branch at Galveston. ISBN 978-0-9631172-1-2. PMID 21413260. Retrieved 2024-03-18.
  6. ^ a b Christie G, Setlow P (October 2020). "Bacillus spore germination: Knowns, unknowns and what we need to learn". Cellular Signalling. 74: 109729. doi:10.1016/j.cellsig.2020.109729. PMID 32721540.
  7. ^ Spencer RC (March 2003). "Bacillus anthracis". Journal of Clinical Pathology. 56 (3): 182–187. doi:10.1136/jcp.56.3.182. PMC 1769905. PMID 12610093.
  8. ^ Schoeni JL, Wong AC (March 2005). "Bacillus cereus food poisoning and its toxins". Journal of Food Protection. 68 (3): 636–648. doi:10.4315/0362-028X-68.3.636. PMID 15771198.
  9. ^ Nigris S, Baldan E, Tondello A, Zanella F, Vitulo N, Favaro G, et al. (October 2018). "Biocontrol traits of Bacillus licheniformis GL174, a culturable endophyte of Vitis vinifera cv. Glera". BMC Microbiology. 18 (1): 133. doi:10.1186/s12866-018-1306-5. PMC 6192205. PMID 30326838.
  10. ^ Favaro G, Bogialli S, Di Gangi IM, Nigris S, Baldan E, Squartini A, et al. (October 2016). "Characterization of lipopeptides produced by Bacillus licheniformis using liquid chromatography with accurate tandem mass spectrometry". Rapid Communications in Mass Spectrometry. 30 (20): 2237–2252. Bibcode:2016RCMS...30.2237F. doi:10.1002/rcm.7705. PMID 27487987.
  11. ^ a b c d e f Paul SI, Rahman MM, Salam MA, Khan MA, Islam MT (2021-12-15). "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: 737156. Bibcode:2021Aquac.54537156P. doi:10.1016/j.aquaculture.2021.737156.
  12. ^ Rahman MM, Paul SI, Akter T, Tay AC, Foysal MJ, Islam MT (September 2020). "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. doi:10.1128/MRA.00641-20. PMC 7516141. PMID 32972930.
  13. ^ Shih YL, Rothfield L (September 2006). "The bacterial cytoskeleton". Microbiology and Molecular Biology Reviews. 70 (3): 729–754. doi:10.1128/MMBR.00017-06. PMC 1594594. PMID 16959967.
  14. ^ Cohn F (1872). "Untersuchungen über Bakterien" [Studies on Bacteria.]. Beiträge zur Biologie der Pflanzen [Contributions to the Biology of Plants] (in German). 2 (1): 127–224.
  15. ^ Bacillus in LPSN; Parte AC, Sardà Carbasse J, Meier-Kolthoff JP, Reimer LC, Göker M (1 November 2020). "List of Prokaryotic names with Standing in Nomenclature (LPSN) moves to the DSMZ". International Journal of Systematic and Evolutionary Microbiology. 70 (11): 5607–5612. doi:10.1099/ijsem.0.004332.
  16. ^ Pei AY, Oberdorf WE, Nossa CW, Agarwal A, Chokshi P, Gerz EA, et al. (June 2010). "Diversity of 16S rRNA genes within individual prokaryotic genomes". Applied and Environmental Microbiology. 76 (12): 3886–3897. Bibcode:2010ApEnM..76.3886P. doi:10.1128/AEM.02953-09. PMC 2893482. PMID 20418441.
  17. ^ Travers RS, Martin PA, Reichelderfer CF (June 1987). "Selective Process for Efficient Isolation of Soil Bacillus spp". Applied and Environmental Microbiology. 53 (6): 1263–1266. Bibcode:1987ApEnM..53.1263T. doi:10.1128/aem.53.6.1263-1266.1987. PMC 203852. PMID 16347359.
  18. ^ Foysal MJ, Lisa AK (December 2018). "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. doi:10.1016/j.jgeb.2018.01.005. PMC 6353715. PMID 30733751.
  19. ^ Caldwell FE (2011-04-06). The Students Reference Guide to Bacteria. Lulu.com. ISBN 978-1-257-42416-0.
  20. ^ "Bacillus subtilis strain WS1A". GenBank. U.S. National Library of Medicine. 19 May 2019.
  21. ^ "Bacillus subtilis strain YBS29". GenBank. U.S. National Library of Medicine. 19 June 2020.
  22. ^ "Bacillus subtilis strain KSP163A". GenBank. U.S. National Library of Medicine. 19 May 2019.
  23. ^ "Bacillus subtilis strain OA122". GenBank. U.S. National Library of Medicine. 20 June 2020.
  24. ^ "Bacillus subtilis strain ISP161A". GenBank. U.S. National Library of Medicine. 20 June 2020.
  25. ^ "Bacillus subtilis strain OI6". GenBank. U.S. National Library of Medicine. 19 June 2020.
  26. ^ "Bacillus subtilis strain WS11". GenBank. U.S. National Library of Medicine. 19 May 2019.
  27. ^ "Bacillus subtilis strain KSP151E". GenBank. U.S. National Library of Medicine. 19 June 2020.
  28. ^ "Bacillus subtilis strain S8". GenBank. U.S. National Library of Medicine. 20 June 2020.
  29. ^ a b Ash C, Farrow J, Wallbanks S, Collins M (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. S2CID 82988953.
  30. ^ Xu D, Côté JC (May 2003). "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. doi:10.1099/Ijs.0.02346-0. PMID 12807189.
  31. ^ Yarza P, Richter M, Peplies J, Euzeby J, Amann R, Schleifer KH, et al. (September 2008). "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. doi:10.1016/j.syapm.2008.07.001. hdl:10261/103580. PMID 18692976.
  32. ^ Yarza P, Ludwig W, Euzéby J, Amann R, Schleifer KH, Glöckner FO, et al. (October 2010). "Update of the All-Species Living Tree Project based on 16S and 23S rRNA sequence analyses". Systematic and Applied Microbiology. 33 (6): 291–299. doi:10.1016/j.syapm.2010.08.001. hdl:10261/54801. PMID 20817437.
  33. ^ Munoz R, Yarza P, Ludwig W, Euzéby J, Amann R, Schleifer KH, et al. (May 2011). "Release LTPs104 of the all-species living tree" (PDF). Systematic and Applied Microbiology. 34 (3): 169–70. doi:10.1016/j.syapm.2011.03.001. PMID 21497273. Archived from the original (PDF) on 23 September 2015.
  34. ^ Salvetti E, Harris HM, Felis GE, O'Toole PW (September 2018). Björkroth J (ed.). "Comparative Genomics of the Genus Lactobacillus Reveals Robust Phylogroups That Provide the Basis for Reclassification". Applied and Environmental Microbiology. 84 (17). Bibcode:2018ApEnM..84E.993S. doi:10.1128/AEM.00993-18. PMC 6102987. PMID 29915113.
  35. ^ a b Alcaraz LD, Moreno-Hagelsieb G, Eguiarte LE, Souza V, Herrera-Estrella L, Olmedo G (May 2010). "Understanding the evolutionary relationships and major traits of Bacillus through comparative genomics". BMC Genomics. 11: 332. doi:10.1186/1471-2164-11-332. PMC 2890564. PMID 20504335. 1471216411332.
  36. ^ a b Gupta RS, Patel S, Saini N, Chen S (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". International Journal of Systematic and Evolutionary Microbiology. 70 (11): 5753–5798. doi:10.1099/ijsem.0.004475. ISSN 1466-5026. PMID 33112222.
  37. ^ a b Patel S, Gupta RS (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". International Journal of Systematic and Evolutionary Microbiology. 70 (1): 406–438. doi:10.1099/ijsem.0.003775. ISSN 1466-5026. PMID 31617837.
  38. ^ a b Nikolaidis M, Hesketh A, Mossialos D, Iliopoulos I, Oliver SG, Amoutzias GD (August 2022). "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. doi:10.3390/microorganisms10091720. PMC 9505155. PMID 36144322.
  39. ^ Økstad OA, Kolstø AB (December 2010). "Chapter 2: Genomics of Bacillus species". In Wiedmann M, Zhang W (eds.). Genomics of Foodborne Bacterial Pathogens. Food Microbiology and Food Safety. Vol. 29. New York, NY: Springer. pp. 29–53 (34–35). doi:10.1007/978-1-4419-7686-4_2. ISBN 978-1-4419-7686-4.
  40. ^ Brenner DJ (1984). "Family I. Enterobacteriaceae Rahn 1937, Nom. fam. cons. Opin. 15, Jud. Com. 1958, 73; Ewing, Farmer, and Brenner 1980, 674; Judicial Commission 1981, 104.". In Krieg NR, Holt JG (eds.). Bergey's Manual of Systematic Bacteriology. Vol. 1 (first ed.). Baltimore: The Williams & Wilkins Co. pp. 408–420.
  41. ^ Loshon CA, Beary KE, Gouveia K, Grey EZ, Santiago-Lara LM, Setlow P (March 1998). "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. doi:10.1016/S0167-4781(97)00204-2. PMID 9540829.
  42. ^ Ding Y, Wang J, Liu Y, Chen S (2005). "Isolation and identification of nitrogen-fixing bacilli from plant rhizospheres in Beijing region". Journal of Applied Microbiology. 99 (5): 1271–1281. doi:10.1111/j.1365-2672.2005.02738.x. PMID 16238759. S2CID 19917931.
  43. ^ Xie G, Su B, Cui Z (Dec 1998). "Isolation and identification of N2-fixing strains of Bacillus in rice rhizosphere of the Yangtze River Valley". Wei Sheng Wu Xue Bao = Acta Microbiologica Sinica (in Chinese). 38 (6). Chinese Academy of Sciences: 480–483. PMID 12548929.
  44. ^ War Nongkhla F, Joshi S (2014). "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. doi:10.15517/rbt.v62i4.12138. PMID 25720168.
  45. ^ Jooste M, Roets F, Midgley GF, et al. (2019). "Nitrogen-fixing bacteria and Oxalis – evidence for a vertically inherited bacterial symbiosis". BMC Plant Biology. 19 (1): 441. doi:10.1186/s12870-019-2049-7. PMC 6806586. PMID 31646970.
  46. ^ Ramesh A, Sharma SK, Sharma MP, Yadav N, Joshi OP (2014). "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. Bibcode:2014AppSE..73...87R. doi:10.1016/j.apsoil.2013.08.009. ISSN 0929-1393.
  47. ^ Slonczewski JL, Foster JW (2011). Microbiology: An Evolving Science (2nd ed.). Norton.
  48. ^ Jeong H, Jeong DE, Kim SH, Song GC, Park SY, Ryu CM, et al. (August 2012). "Draft genome sequence of the plant growth-promoting bacterium Bacillus siamensis KCTC 13613T". Journal of Bacteriology. 194 (15): 4148–4149. doi:10.1128/JB.00805-12. PMC 3416560. PMID 22815459.
  49. ^ Keen EC, Bliskovsky VV, Adhya SL, Dantas G (November 2017). "Draft Genome Sequence of the Naturally Competent Bacillus simplex Strain WY10". Genome Announcements. 5 (46): e01295–17. doi:10.1128/genomeA.01295-17. PMC 5690344. PMID 29146837.
  50. ^ Ryan KJ, Ray CG, eds. (2004). Sherris Medical Microbiology (4th ed.). McGraw Hill. ISBN 978-0-8385-8529-0.
  51. ^ "History of Bt". University of California San Diego. Retrieved 28 June 2024.
  52. ^ James C (1996). "Global Review of the Field Testing and Commercialization of Transgenic Plants: 1986 to 1995" (PDF). The International Service for the Acquisition of Agri-biotech Applications. Retrieved 17 July 2010.
  53. ^ Peairs FB (2013). "Bt Corn: Health and the Environment – 0.707" (PDF). Colorado State University Extension Office. Archived (PDF) from the original on 2022-10-09.
  54. ^ Program HF (2024-09-09). "Microorganisms & Microbial-Derived Ingredients Used in Food (Partial List)". FDA. Retrieved 2024-11-05.
  55. ^ Schallmey M, Singh A, Ward OP (January 2004). "Developments in the use of Bacillus species for industrial production". Canadian Journal of Microbiology. 50 (1): 1–17. doi:10.1139/w03-076. PMID 15052317.
  56. ^ Graumann P, ed. (2012). Bacillus: Cellular and Molecular Biology (2nd ed.). Caister Academic Press. ISBN 978-1-904455-97-4. [1].
  57. ^ Ash C, Priest FG, Collins MD (1994). "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. doi:10.1007/BF00873085. PMID 8085788. S2CID 7391845.
  58. ^ Heyndrickx M, Lebbe L, Kersters K, De Vos P, Forsyth G, Logan NA (January 1998). "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. doi:10.1099/00207713-48-1-99.
  • Bacillus genomes and related information at PATRIC, a Bioinformatics Resource Center funded by NIAID