Bacillus

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Bacillus
Bacillus subtilis Gram.jpg
Bacillus subtilis , stained
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Bacillota
Class: Bacilli
Order: Bacillales
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]

Contents

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) Bacillus subtilis 2.jpg
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 typeTestCharacteristics
Colony charactersSizeMedium
TypeRound
ColorWhitish
ShapeConvex
Morphological charactersShapeRod
Physiological charactersMotility+
Growth at 6.5% NaCl+
Biochemical charactersGram's staining+
Oxidase-
Catalase+
Oxidative-FermentativeO/F
Motility+
Methyl Red+
Voges-Proskauer-
Indole-
H2S Production+/–
Urease-
Nitrate reductase+
β-Galactosidase+
Hydrolysis ofGelatin+
Aesculin+
Casein+
Tween 40+
Tween 60+
Tween 80+
Acid production fromGlycerol+
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

Three proposals have been presented as representing the phylogeny of the genus Bacillus. The first proposal, presented in 2003, is a Bacillus-specific study, with the most diversity covered using 16S and the ITS regions. It divides the genus into 10 groups. This includes the nested genera Paenibacillus, Brevibacillus, Geobacillus, Marinibacillus and Virgibacillus . [29]

The second proposal, presented in 2008, [30] constructed a 16S (and 23S if available) tree of all validated species. [31] [32] The genus Bacillus contains a very large number of nested taxa and majorly in both 16S and 23S. It is paraphyletic to the Lactobacillales (Lactobacillus, Streptococcus, Staphylococcus, Listeria, etc.), due to Bacillus coahuilensis and others. [33]

A third proposal, presented in 2010, was a gene concatenation study, and found results similar to the 2008 proposal, but with a much more limited number of species in terms of groups. [34] (This scheme used Listeria as an outgroup, so in light of the ARB tree, it may be "inside-out").

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 due to medical reasons, they are considered separate species [35] (an issue also present for four species of Shigella and Escherichia coli ). [36]

A phylogenomic study of 1104 Bacillus proteomes was based on 114 core proteins and delineated the relationships among the various species, defined as Bacillus from the NCBI taxonomy. [37] The various strains were clustered into species, based on Average Nucleotide identity (ANI) values, with a species cutoff of 95%. [37]

Bacillus phylogenetics
Root
Phylogeny of the genus Bacillus according to [34]

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. [39] [40] [41] [42] [43] B. thuringiensis produces a toxin that can kill insects and thus has been used as insecticide. [44] 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. [45] Some species of Bacillus are naturally competent for DNA uptake by transformation. [46]

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 (and cotton) crops. The resulting GMOs are resistant to some insect pests.[ citation needed ]

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.[ citation needed ]

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. [48]

Use as model organism

Colonies of the model species Bacillus subtilis on an agar plate Bacillus subtilis colonies.jpg
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. [49]

See also

Related Research Articles

<i>Bacillus cereus</i> Species of bacterium

Bacillus cereus is a Gram-positive rod-shaped bacterium commonly found in soil, food, and marine sponges. The specific name, cereus, meaning "waxy" in Latin, refers to the appearance of colonies grown on blood agar. Some strains are harmful to humans and cause foodborne illness due to their spore-forming nature, while other strains can be beneficial as probiotics for animals, and even exhibit mutualism with certain plants. B. cereus bacteria may be anaerobes or facultative anaerobes, and like other members of the genus Bacillus, can produce protective endospores. They have a wide range of virulence factors, including phospholipase C, cereulide, sphingomyelinase, metalloproteases, and cytotoxin K, many of which are regulated via quorum sensing. B. cereus strains exhibit flagellar motility.

<span class="mw-page-title-main">Endospore</span> Protective structure formed by bacteria

An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form, but it is not a true spore. It is a stripped-down, dormant form to which the bacterium can reduce itself. Endospore formation is usually triggered by a lack of nutrients, and usually occurs in gram-positive bacteria. In endospore formation, the bacterium divides within its cell wall, and one side then engulfs the other. Endospores enable bacteria to lie dormant for extended periods, even centuries. There are many reports of spores remaining viable over 10,000 years, and revival of spores millions of years old has been claimed. There is one report of viable spores of Bacillus marismortui in salt crystals approximately 25 million years old. When the environment becomes more favorable, the endospore can reactivate itself into a vegetative state. Most types of bacteria cannot change to the endospore form. Examples of bacterial species that can form endospores include Bacillus cereus, Bacillus anthracis, Bacillus thuringiensis, Clostridium botulinum, and Clostridium tetani. Endospore formation is not found among Archaea.

<i>Bacillus subtilis</i> Catalase-positive bacterium

Bacillus subtilis, known also as the hay bacillus or grass bacillus, is a gram-positive, catalase-positive bacterium, found in soil and the gastrointestinal tract of ruminants, humans and marine sponges. As a member of the genus Bacillus, B. subtilis is rod-shaped, and can form a tough, protective endospore, allowing it to tolerate extreme environmental conditions. B. subtilis has historically been classified as an obligate aerobe, though evidence exists that it is a facultative anaerobe. B. subtilis is considered the best studied Gram-positive bacterium and a model organism to study bacterial chromosome replication and cell differentiation. It is one of the bacterial champions in secreted enzyme production and used on an industrial scale by biotechnology companies.

A Bacillus phage is a member of a group of bacteriophages known to have bacteria in the genus Bacillus as host species. These bacteriophages have been found to belong to the families Myoviridae, Siphoviridae, Podoviridae, or Tectiviridae. The genus Bacillus includes the model organism, B. subtilis, and two widely known human pathogens, B. anthracis and B. cereus. Other strains of Bacillus bacteria that phage are known to infect include B. megaterium, B. mycoides, B. pseudomycoides, B. thuringiensis, and B. weihenstephanensis. More than 1,455 bacillus phage have been discovered from many different environments and areas around the world. Only 164 of these phages have been completely sequenced as of December 16, 2021.

<i>Bacillus anthracis</i> Species of bacterium

Bacillus anthracis is a gram-positive and rod-shaped bacterium that causes anthrax, a deadly disease to livestock and, occasionally, to humans. It is the only permanent (obligate) pathogen within the genus Bacillus. Its infection is a type of zoonosis, as it is transmitted from animals to humans. It was discovered by a German physician Robert Koch in 1876, and became the first bacterium to be experimentally shown as a pathogen. The discovery was also the first scientific evidence for the germ theory of diseases.

The exosporium is the outer surface layer of mature spores. In plant spores it is also referred to as the exine. Some bacteria also produce endospores with an exosporium, of which the most commonly studied are Bacillus species, particularly Bacillus cereus and the anthrax-causing bacterium Bacillus anthracis. The exosporium is the portion of the spore that interacts with the environment or host organism, and may contain spore antigens. Exosporium proteins, such as Cot protein, are also discovered related to strains of B. anthracis and B.cereus. This Cot protein share similar sequences with other spore coat proteins, and their putative determinants are believed to include bxpC, lunA, exsA, etc.

Metabacillus is a genus of rod-shaped bacteria exhibiting Gram-positive or Gram-variable staining in the family Bacillaceae within the order Bacillales. The type species for this genus is Metabacillus fastidiosus.

Neobacillus is a genus of rod-shaped bacteria that show Gram-positive or Gram-variable staining. This genus belongs under the family Bacillaceae within the order Bacillales. The type species of Neobacillus is Neobacillus niacini.

<span class="mw-page-title-main">Cytobacillus</span> Genus of rod-shaped bacteria

Cytobacillus is a genus of rod-shaped bacteria that stain either Gram-positive or Gram-variable in the family Bacillaceae within the order Bacillales. The type species for this genus is Cytobacillus firmus.

Alteribacter is a genus of Gram-positive or Gram-variable, rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Alteribacter auranticus.

Ectobacillus is a genus of Gram-positive, rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Ectobacillus panaciterrae.

Evansella is a genus of Gram-positive rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Evansella cellulosilytica.

Ferdinandcohnia is a genus of rod-shaped bacteria that generally display Gram-positive staining in the family Bacillaceae within the order Bacillales. The type species for this genus is Ferdinandcohnia humi.

Gottfriedia is a genus of gram-positive or Gram-variable rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Gottfriedia luciferensis.

Heyndrickxia is a genus of gram-positive rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Heyndrickxia oleronia.

<i>Lederbergia</i> Genus of bacteria

Lederbergia is a genus of gram-positive or Gram-variable, rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Lederbergia galactosidilyticus.

Litchfieldia is a genus of Gram-positive rod-shaped bacteria in the family Bacillaceae within the order Bacillales. The type species for this genus is Litchfieldia alkalitelluris.

Margalitia is a genus of gram-positive or gram-variable rod-shaped bacteria in the family Bacillaceae from the order Bacillales. The type species of this genus is Margalitia shackletonii.

Robertmurraya is a genus of mostly Gram-Positive rod-shaped bacteria in the family Bacillaceae from the order Bacillales. The type species of this genus is Robertmurraya siralis.

Rossellomorea is a genus of Gram-Positive or Gram-variable staining rod-shaped bacteria in the family Bacillaceae from the order Bacillales. The type species of this genus is Rossellomorea aquimaris.

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