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 and humans. 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
4 Chromosomal replication
8.1 In animals
8.2 In humans
10 See also
12 External links
Bacillus subtilis is a
Gram-positive bacterium, rod-shaped and
catalase-positive. It was originally named Vibrio subtilis by
Christian Gottfried Ehrenberg, and renamed
Bacillus subtilis by
Ferdinand Cohn in 1872 (subtilis being the Latin for 'fine'). B.
subtilis cells are typically rod-shaped, and are about 4-10
micrometers (μm) long and 0.25–1.0 μm in diameter, with a cell
volume of about 4.6 fL at stationary phase. As with other members
of the genus Bacillus, it can form an endospore, to survive extreme
environmental conditions of temperature and desiccation. B.
subtilis is a facultative anaerobe and had been considered as an
obligate aerobe until 1998. B. subtilis is heavily flagellated, which
gives it the ability to move quickly in liquids. B. subtilis has
proven highly amenable to genetic manipulation, and has become widely
adopted as a model organism for laboratory studies, especially of
sporulation, which is a simplified example of cellular
differentiation. In terms of popularity as a laboratory model
organism, B. subtilis is often considered as the Gram-positive
equivalent of Escherichia coli, an extensively studied Gram-negative
This species is commonly found in the upper layers of the soil, and
evidence exists that B. subtilis is a normal gut commensal in humans.
A 2009 study compared the density of spores found in soil (about 106
spores per gram) to that found in human feces (about 104 spores per
gram). The number of spores found in the human gut was too high to be
attributed solely to consumption through food contamination. B.
subtilis has been linked to grow in higher elevations and act as an
identifier for both eco-adaptability and honey bee health.
Sporulating B. subtilis.
Another endospore stain of B. subtilis.
B. subtilis can divide symmetrically to make two daughter cells
(binary fission), or asymmetrically, producing a single endospore that
can remain viable for decades and is resistant to unfavourable
environmental conditions such as drought, salinity, extreme pH,
radiation, and solvents. The endospore is formed at times of
nutritional stress, allowing the organism to persist in the
environment until conditions become favourable. Prior to the process
of sporulation the cells might become motile by producing flagella,
take up DNA from the environment, or produce antibiotics. These
responses are viewed as attempts to seek out nutrients by seeking a
more favourable environment, enabling the cell to make use of new
beneficial genetic material or simply by killing off
Under stressful conditions, such as nutrient deprivation, B. subtilis
undergoes the process of sporulation. This process has been very well
studied and has served as a model organism for studying
B. subtilis is a model organism used to study bacterial chromosome
replication. Replication of the single circular chromosome initiates
at a single locus, the origin (oriC). Replication proceeds
bidirectionally and two replication forks progress in clockwise and
counterclockwise directions along the chromosome. Chromosome
replication is completed when the forks reach the terminus region,
which is positioned opposite to the origin on the chromosome map. The
terminus region contains several short DNA sequences (Ter sites) that
promote replication arrest. Specific proteins mediate all the steps in
DNA replication. Comparison between the proteins involved in
chromosomal DNA replication in B. subtilis and in Escherichia coli
reveals similarities and differences. Although the basic components
promoting initiation, elongation, and termination of replication are
well-conserved, some important differences can be found (such as one
bacterium missing proteins essential in the other). These differences
underline the diversity in the mechanisms and strategies that various
bacterial species have adopted to carry out the duplication of their
B. subtilis has about 4,100 genes. Of these, only 192 were shown to be
indispensable; another 79 were predicted to be essential, as well. A
vast majority of essential genes were categorized in relatively few
domains of cell metabolism, with about half involved in information
processing, one-fifth involved in the synthesis of cell envelope and
the determination of cell shape and division, and one-tenth related to
The complete genome sequence of B. subtilis sub-strain QB928 has
4,146,839 DNA base pairs and 4,292 genes. The QB928 strain is widely
used in genetic studies due to the presence of various markers
[aroI(aroK)906 purE1 dal(alrA)1 trpC2].
Several noncoding RNAs have been characterized in the B. subtilis
genome in 2009, including Bsr RNAs. Microarray-based comparative
genomic analyses have revealed that B. subtilis members show
considerable genomic diversity.
Natural bacterial transformation involves the transfer of DNA from one
bacterium to another through the surrounding medium. In B. subtilis
the length of transferred DNA is greater than 1271kb (more than 1
million bases). The transferred DNA is likely double-stranded DNA
and is often more than a third of the total chromosome length of 4215
kb. It appears that about 7-9% of the recipient cells take up an
In order for a recipient bacterium to bind, take up exogenous DNA from
another bacterium of the same species and recombine it into its
chromosome, it must enter a special physiological state called
competence. Competence in B. subtilis is induced toward the end of
logarithmic growth, especially under conditions of amino-acid
limitation. Under these stressful conditions of semistarvation,
cells typically have just one copy of their chromosome and likely have
increased DNA damage. To test whether transformation is an adaptive
function for B. subtilis to repair its DNA damage, experiments were
conducted using UV light as the damaging agent. These
experiments led to the conclusion that competence, with uptake of DNA,
is specifically induced by DNA-damaging conditions, and that
transformation functions as a process for recombinational repair of
Gram-stained B. subtilis
Cultures of B. subtilis were popular worldwide before the
introduction of antibiotics as an immunostimulatory agent to aid
treatment of gastrointestinal and urinary tract diseases. It was used
throughout the 1950s as an alternative medicine, which upon digestion
has been found to significantly stimulate broad-spectrum immune
activity including activation of secretion of specific antibodies IgM,
IgG and IgA and release of CpG dinucleotides inducing INF A/Y
producing activity of leukocytes and cytokines important in the
development of cytotoxicity towards tumor cells. It was marketed
throughout America and Europe from 1946 as an immunostimulatory aid in
the treatment of gut and urinary tract diseases such as
Since the 1960s B. subtilis has had a history as a test species in
spaceflight experimentation. Its endospores can survive up to 6 years
in space if coated by dust particles protecting it from solar UV
rays. It has been used as an extremophile survival indicator in
outer space such as Exobiology
EXPOSE orbital missions.
Wild-type natural isolates of B. subtilis are difficult to work with
compared to laboratory strains that have undergone domestication
processes of mutagenesis and selection. These strains often have
improved capabilities of transformation (uptake and integration of
environmental DNA), growth, and loss of abilities needed "in the
wild". And, while dozens of different strains fitting this description
exist, the strain designated '168' is the most widely used.[citation
Colonies of B. subtilis grown on a culture dish in a molecular biology
B. globigii, a closely related but phylogenetically distinct species
now known as
Bacillus atrophaeus was used as a biowarfare
Project SHAD (aka Project 112). Subsequent genomic
analysis showed that the strains used in those studies were products
of deliberate enrichment for strains that exhibited abnormally high
rates of sporulation.
A strain of B. subtilis formerly known as
Bacillus natto is used in
the commercial production of the Japanese food nattō, as well as the
similar Korean food cheonggukjang.
As a model organism, B. subtilis is commonly used in laboratory
studies directed at discovering the fundamental properties and
Gram-positive spore-forming bacteria. In
particular, the basic principles and mechanisms underlying formation
of the durable endospore have been deduced from studies of spore
formation in B. subtilis.
Its surface-binding properties play a role in safe radionuclide waste
[e.g. thorium (IV) and plutonium (IV)] disposal.
Recombinant strains pBE2C1 and pBE2C1AB were used in production of
polyhydroxyalkanoates (PHA), and malt waste can be used as their
carbon source for lower-cost PHA production.
Due to its excellent fermentation properties, with high product yields
(20 to 25 gram per litre) it is used to produce various enzymes, such
as amylase and proteases.
Other enzymes produced by B. subtilis and B. licheniformis are widely
used as additives in laundry detergents.
It is used to produce hyaluronic acid, which is used in the joint-care
sector in healthcare[full citation needed] and cosmetics.
B. subtilis is used as a soil inoculant in horticulture and
agriculture.[full citation needed][full citation
needed][full citation needed]
It may provide some benefit to saffron growers by speeding corm growth
and increasing stigma biomass yield.
Monsanto has isolated a gene from B. subtilis that expresses cold
shock protein B and spliced it into their drought-tolerant corn hybrid
MON 87460, which was approved for sale in the US in November
It is used as an "indicator organism" during gas sterilization
procedures, to ensure a sterilization cycle has completed
successfully.[full citation needed][full citation needed] This
is due to the difficulty in sterilizing endospores.
Novel strains of B. subtilis that could use 4-fluorotryptophan (4FTrp)
but not canonical tryptophan (Trp) for propagation were isolated. As
Trp is only coded by a single codon, there is evidence that Trp can be
displaced by 4FTrp in the genetic code. The experiments showed that
the canonical genetic code can be mutable.
B. subtilis has been found to act as a useful bioproduct fungicide
that prevents the growth of Monilinia vaccinii-corymbosi, a.k.a. the
mummy berry fungus, without interfering with pollination or fruit
B. subtilis was reviewed by the US FDA Center for Veterinary Medicine
and found to present no safety concerns when used in direct-fed
microbial products, so the Association of American Feed Control
Officials has listed it approved for use as an animal feed ingredient
under Section 36.14 "Direct-fed Microorganisms". The
Canadian Food Inspection Agency
Canadian Food Inspection Agency Animal Health and Production Feed
Section has classified
Bacillus culture dehydrated approved feed
ingredients as a silage additive under Schedule IV-Part 2-Class 8.6
and assigned the International Feed Ingredient number IFN
B. subtilis spores can survive the extreme heat during cooking. Some
B. subtilis strains are responsible for causing ropiness — a
sticky, stringy consistency caused by bacterial production of
long-chain polysaccharides — in spoiled bread dough. For a long
time, bread ropiness was associated uniquely with B. subtilis species
by biochemical tests. Molecular assays (randomly amplified polymorphic
DNA PCR assay, denaturing gradient gel electrophoresis analysis, and
sequencing of the V3 region of 16S ribosomal DNA) revealed greater
Bacillus species variety in ropy breads, which all seems to have a
positive amylase activity and high heat resistance. B. subtilis
CU1 (2 × 109 spores per day) was evaluated in a 16-week study (10
days administration of probiotic, followed by 18 days wash-out period
per each month; repeated same procedure for total 4 months) to healthy
subjects. B. subtilis CU1 was found to be safe and well-tolerated in
the subjects without any side effects.
B. subtilis and substances derived from it has been evaluated by
different authoritative bodies for their safe and beneficial use in
food. In the United States, an opinion letter issued in the early
1960s by the
Food and Drug Administration
Food and Drug Administration (FDA) recognized some
substances derived from microorganisms as generally recognized as safe
(GRAS), including carbohydrase and protease enzymes from B. subtilis.
The opinions were predicated on the use of nonpathogenic and
nontoxicogenic strains of the respective organisms and on the use of
current good manufacturing practices. The FDA stated the enzymes
derived from the B. subtilis strain were in common use in food prior
to January 1, 1958, and that nontoxigenic and nonpathogenic strains of
B. subtilis are widely available and have been safely used in a
variety of food applications. This includes consumption of Japanese
fermented soy bean, in the form of Natto, which is commonly consumed
in Japan, and contains as many as 108 viable cells per gram. The
fermented beans are recognized for their contribution to a healthy gut
flora and vitamin K2 intake; during this long history of widespread
use, natto has not been implicated in adverse events potentially
attributable to the presence of B. subtilis. The
natto product and the B. subtilis natto as its principal component are
FOSHU (Foods for Specified Health Use) approved by the Japanese
Ministry of Health, Labour, and Welfare
Ministry of Health, Labour, and Welfare as effective for preservation
B. subtilis has been granted "Qualified Presumption of Safety" status
by the European Food Safety Authority.
Below is a table that helps identify B. subtilis through certain
Acid from Glucose
Acid from Lactose
Acid from Sucrose
Acid from Mannitol
Gas from Glucose
Hydrogen Sulfide Production
Adenylosuccinate lyase deficiency
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Media related to
Bacillus subtilis at Wikimedia Commons
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