A protein spore coat and an exosporium, a membranous layer, form outside of the forespore membranes. At this time, the forespore undergoes internal changes.
Lastly, the forespore leaves the mother cell upon lysis of the mother cell Perez A mature endospore has no metabolic activity; it is inert. The interior of the endospore, the core, is very dry and resistant to moisture Schaechter Bacillus subtilis bacteria use their flagella for a swarming motility.
This motility occurs on surfaces, for example on agar plates, rather than in liquids. Bacillus subtilis are arranged in singles or chains. Cells arranged next to each other can only swarm together, not individually.
These arrangements of cells are called 'rafts'. In order for Bacillus subtilis bacteria to swarm, they need to secrete a slime layer which includes surfactin, a surface tension-reducing lipopeptide, as one of its components Schaechter Bacillus subtilis bacteria have been considered strictly aerobic, meaning that they require oxygen to grow and they cannot undergo fermentation.
However, recent studies show that they can indeed grow in anaerobic conditions making them facultative aerobes. The bacteria can make ATP in anaerobic conditions via butanediol fermentation as well as nitrate ammonification. Bacillus subtilis can use nitrite or nitrate as a terminal acceptor of electrons. Bacillus subtilis contains two unique nitrate reductases. One is used for nitrate nitrogen assimilation and the other is used for nitrate respiration.
However, there is only one nitrite reductase that serves both purposes. Nitrate reductase reduces nitrate to nitrite in nitrate respiration, which is then reduced to ammonia by nitrite reductase.
Bacillus subtilis is different from other facultative aerobes in that it undergoes fermentation without external acceptors of electrons Nakano Lactate dehydrogenase converts pyruvate to lactate Marino Bacillus subtilis contains catalase KatA and MrgA, an enzyme that is responsible in the catalysis of the decomposition of hydrogen peroxide to water and oxygen, and superoxide dismutase, an enzyme that catalyzes the breakdown of superoxide into oxygen and hydrogen peroxide Bandow Bacillus subtilis duplicates its single circular chromosome by initiating DNA replication at a single locus, the origin oriC.
Replication proceeds bidirectionally and two replication forks progress in the clockwise and counterclockwise directions along the chromosome halves. Chromosome replication is completed when the forks reach the terminus region, which is positioned opposite to the origin on the chromosome map, and contains several short DNA sequences Ter sites that promote replication arrest.
Specific proteins mediate all the steps in DNA replication. The comparison between the sets of proteins involved in chromosomal DNA replication in B. 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 genomes Graumann, The main habitat of endospore forming Bacillus organisms is the soil. Likewise Bacillus subtilis is most commonly found in soil environments and on plant undergrowth. These mesophilic microbes have historically been considered strict aerobes.
Thus they are likely to be found in O and A surface soil horizons where the concentration of oxygen is most abundant and temperatures are relatively mild. Consider how this organism functions in s competitive microbial community: when carbon-, nitrogen- and phosphorus-nutrient levels fall below the bacterium's optimal threshold, it produces spores.
Scientists have demonstrated that Bacillus subtilis concurrently produces antibiotics and spores. Antibiotic production increases B. Subtilis's cance at survival as the organism produces spores and a toxin that might kill surrounding gram positive microbes that compete for the same nutrients. These microbes form spores in times of nutrient exhaustion. When the nutrients required for the bacteria to grow are abundant, they exhibit metabolic activity.
These organisms can produce antibiotics during sporulation. Examples of the antibiotics that Bacillus subtilis can produce include are polymyxin, difficidin, subtilin, and mycobacillin. Many of the Bacillus microbes can degrade polymers such as protein, starch, and pectin, therefore, they are thought to be an important contributor to the carbon and nitrogen cycles.
When they cause contamination, they may result in decomposition. Quite a few of the Bacillus organisms are primarily responsible for the spoilage of food Todar. Bacillus subtilis supports plant browth. As a member of Bacillus , this bacterium often plays a role in replenishing soil nutrients by supplying the terrestrial carbon cycle and the nitrogen cycle.
Bacillus subtilis bacteria form rough biofilms, which are dense organism communities, at the air and water interface. Bacillus subtilis biofilms are beneficial. They allow for the control of plant pathogen infections. The plant benefits because B.
Preemptive colonization prevents other pathogens from infecting the plant because B. The biofilm communities form a mutualistic interaction with plant rhizome systems. Bacillus subtilis biofilms found in the rhizosphere of plants promote growth and serve as a biocontroller. In this sense, B.
Bacillus subtilis strains can act as biofungicides for benefiting agricultural crops and antibacterial agents. Bacillus subtilis also reduces mild steel corrosion Morikawa Bacillus subtilis bacteria are non-pathogenic. They can contaminate food, however, they seldom result in food poisoning. They are used on plants as a fungicide. They are also used on agricultural seeds, such as vegetable and soybean seeds, as a fungicide. In agriculture, studies have shown that adding an appropriate amount of B.
In this paper, we reviewed recent progress in the metabolic engineering and protein expression systems, as well as industrial, agricultural, and biomaterial applications of B. Finally, we analyzed the factors that hinder the further application of this strain and discussed the reasons. This review provides a reference for researchers who want to gain a general understanding of B.
Application of B. It can also be used to produce various chemicals, such as riboflavin, menaquinone-7, inositol, or N-acetylglucosamine. In agriculture, it can be used as a feed additive. Biofilms of B. In medicine, B. Classical genome modification relies on the insertion of a selectable marker, usually an antibiotic resistance gene, into the chromosome of the target strain [ 11 ].
The most commonly used scarless genetic manipulations systems for B. CSM are often used for the markerless construction of engineered strains and have been used to construct Bacillus cell factories for various industrial applications. Selectable markers can generally be divided into positive and negative selection markers, whereby the former are most commonly antibiotic-resistance markers.
In this classical approach, antibiotic-resistant strains are selected on appropriate agar plates Fig. In addition to the genomically integrated markers, Jeong et al.
In the first recombination, P xyl - lacI and neo are integrated into the genome as a selectable marker. When xylose is added to the medium, the lacI gene is expressed then the chloramphenicol resistant gene is repressed. Consequently, the cell will survive only when the P xyl - lacI and neo are deleted through a second round of recombination.
Finally, the plasmid can be removed after several rounds of culture without chloramphenicol [ 15 ]. This is a highly efficient method for genome engineering in B. Other counter-selectable markers commonly used in B. Fabret et al. Brans et al. However, CSM-based strategies require host pre-modification and have a low success rate due to the leaky expression of the CSM. Schematic overview of genome editing methods based on counter-selectable markers.
Left: genome editing gene knockout as an example with two integration steps. Step 1, an exogenous artificial DNA plasmid or fragment with up- and downstream homologous sequences is integrated into the genome, replacing the target gene. The recombinant clone can be selected under condition A. Examples of selectable markers A include: cat chloramphenicol , phleo phleomycin , or spe spectinomycin. Examples of toxic genes include: upp , pyrF , or mazF. Examples of repressor genes include: xylR , blaI , araR , or lacI.
The repressor can inhibit the expression of the selectable marker B, which can be integrated into the genome or a plasmid. The resulting duplexes bind Cas9 protein to form a targeted cutting complex, specifically cutting foreign sequences to achieve the goal of identifying and eliminating invading foreign genes such as plasmids and viruses [ 21 , 22 , 23 ]. In this system, Cas9, gRNA, and donor DNA are assembled on two different plasmids, which are respectively used to produce Cas9 protein and deliver the gRNA transcription module and donor DNA template; 3 The chromosomally integrated system is more stable and effective than the first two systems, but it requires the use of engineered strains.
The CRISPRi system is composed of a deactivated Cas9 dCas9 protein and gRNA, enabling the targeting of dCsa9 to any target gene on the genome under the guidance of gRNA to inhibit its transcription without inducing a double-strand break, which can be applied to gene repression in metabolic engineering [ 24 ]. So et al. This method has wide applicability for various types of site-directed mutagenesis in B.
Liu et al. To enhance and appropriately adjust gene expression levels in B. Inducible and constitutive promoters are usually applied for the expression of heterologous genes in B. In addition to the promoters, the protein expression level is also influenced by the strength of the ribosome binding site, while plasmids with different copy numbers also provide choices for adjusting the level of gene expression [ 31 ].
Different genes have specific expression features, and their expression must be adjusted to a level appropriate for a specific metabolic pathway, which necessitates the use of promoters with different strengths for engineering purposes [ 32 ].
Researchers have broadened the scope of target gene transcription levels by constructing promoter libraries. In a recent study, Liu et al. To enable the efficient and accurate co-expression of multiple genes in metabolic networks, a recent review discussed the construction promoter libraries by site-directed mutagenesis [ 34 ], such as error-prone PCR, saturation mutagenesis and directional design. RBS sequences can be used to fine-tune gene expression at the translational level.
When inducible promoters are used, different ribosomal binding sites can be used to fine-tune the dynamic range of gene expression. In addition, a proteolysis tag can be used to control the degradation rate of a protein at the post-translational level [ 35 ]. A synthetic gene expression toolbox consisting of promoter libraries, RBS libraries, and different proteolytic tags can realize gene regulation with a dynamic range of 5 orders of magnitude [ 36 ].
Other elements other than the promoter and RBS are also used to adjust gene expression in B. Tian et al. They found that NCS substitution is more efficient and convenient than promoter substitution for gene expression improvement [ 37 ]. Naseri et al. However, the examples shown in their review indicate that E.
Therefore, future studies should focus on developing new tools for this important Gram-positive model bacterium. Bacillus subtilis has a strong capacity for protein expression and secretion, which has led to its wide use in the production of industrial enzyme preparations.
In addition to the abundant promoters and plasmid expression systems described above, B. There are three classical protein secretion pathways in B. The Sec pathway is the main transport channel, which can transport a large number of exported proteins. The essential elements of the Sec pathway are the signal recognition particle SRP , Sec translocase, type I signal peptidase, and chaperones.
After the synthesis of the precursor protein, there are two routes through the Sec pathway. In the first one, the signal peptide is recognized by the signal recognition particle SRP with the help of a cytoplasmic chaperone, and then transferred to the membrane to bind with FstY, the receptor protein of the SRP, and transported to the channel of the Sec translocase complex translocation channel.
Next, the N-terminal signal peptide sequence is cut off by signal peptidases. Finally, the translocating protein is folded in the extracellular space with the aid of extracellular chaperones [ 30 , 40 ]. In contrast to the Sec pathway, which relies on unfolded substrates, the Tat pathway transports tightly folded proteins that contain a conserved twin-arginine motif in the signal peptide sequences.
Before translocation, the precursors fold in the cytoplasm with the help of cofactors, and are then excreted through the Tat protein complex Tat translocase utilizing energy from the pH gradient across the cytoplasmic membrane. After translocation, the type I signal peptidase processes the signal peptide, and finally, the folded mature proteins are secreted out of the cell. They are relatively specific for their substrates and can export or import various molecules ions, amino acids, peptides, antibiotics, polysaccharides, proteins, etc.
Schematic diagram of protein secretion pathways in B. The mechanism of the non-classical secretion pathway is not clear. In addition to the three classical protein secretion pathways mentioned above, researchers also found that many non-classical secretion pathways are used to secrete non-classical proteins lacking any known signal peptides or secretion motifs [ 41 ]. Wang et al. Chen et al.
The fusion proteins not only retained the corresponding enzymatic or biological activities, but also had the activity of RDPE [ 43 ] Fig. The industrial application of B.
Chemicals produced by B. Here, we will focus on vitamins and other chemicals produced by B. Table 1 also lists some other representative chemicals produced by B. In a recent review, microbial cell factories for the production of B vitamins were described in detail [ 48 ]. Vitamin B 2 is one of the most successful microbial fermentation products on an industrial scale [ 49 ].
Also known as riboflavin RF , Vitamin B 2 is the precursor of flavin mononucleotide and flavin adenine dinucleotide [ 50 ], and it is widely used for its antioxidant, immunity enhancing, anticancer, as well as food and feed enhancing effects [ 51 ].
Metabolic engineering strategies based on the riboflavin metabolic pathway of B. The precursor supply in the RF biosynthesis pathway can be enhanced by redirecting the carbon flux through the PPP pentose phosphate pathway from the EMP Embden-Meyerhof-Parnas pathway, and increasing the expression of purine biosynthesis genes pur operon , thus increasing GTP production [ 53 , 54 ].
The yield of riboflavin in fed-batch fermentation using B. The excellent RF productivity of B. In addition, the yield of RF can be increased by improving the host characteristics.
Recent studies found that the introduction of heat shock proteins from thermophilic bacteria can improve the heat resistance and osmotic tolerance of B. In addition to B vitamins, B. In the fermentation process of MK-7, glycerol, glucose, sucrose, or starch are used as carbon sources, and yeast extract, peptone, sodium nitrate, or soybean peptone as nitrogen sources [ 56 ].
Using ethanol to extract MK-7 directly from the cells after fermentation produced a MK-7 yield of 1. Wu and Ahn optimized the medium components via a three-step response surface methodology RSM approach, and the yield of vitamin K increased to Yang et al. Knocking out dhbB can promote the production of MK-7, and it was further improved by adopting high-density fermentation technology [ 59 ]. In addition, MK-7 is an important component of the microbial membrane, where it plays an important role in the process of electron transport and oxidative phosphorylation.
Cui et al. However, this compound is far less abundant than its analog myo -inositol MI , which is the most abundant inositol stereoisomer in nature and can be obtained from rice bran. Consequently, many studies have investigated strategies to convert MI into SI using microorganisms. Tanaka et al. Through the genetic modification of inositol metabolism and phytase secretion pathway of B. At the same time, the pnt AB gene of E.
Hyaluronic acid HA is a high-value glycosaminoglycan, which is widely used in the biomedical, pharmaceutical, cosmetic, and food industries. Notably, HA preparations with different molecular weight show different effects [ 63 ].
Peng Jin et al. They used the ribosome binding site engineering strategy to regulate the translational level of hyaluronidase and optimized the HA synthesis pathway, which led to the specific production of low-molecular-weight HA [ 64 ]. Li et al. They found that when the biomass increased, the molecular weight of the produced HA decreased while the titer increased [ 63 ]. In addition to naturally occurring products, scientists are also introducing new pathways into B.
N-acetylglucosamine is an acetylated amine derivative of glucose, which plays an important role in the maintenance and repair of cartilage and joint tissue function. By applying modular pathway engineering, they increased the production of GlcNAc in B. Then, by knocking out acetolactate synthase AlsS and acetolactate decarboxylase AlsD , the formation of the neutral byproduct acetoin was reduced, and the carbon flux from fructosephosphate toward the GlcNAc synthesis pathway was increased.
Consequently, the titer and yield of GlcNAc increased to The catabolism of GlcNAc in B. Recently, Liu et al. Using this system, a genetic feedback circuit was constructed to fine-tune the metabolic flow toward GlcNAc synthesis and competing modules, which increased the titer of GlcNAc in a L fed-batch bioreactor to Due to its rapid growth on inexpensive substrates, strong protein secretion ability, non-pathogenicity, and favorable downstream processing, B.
According to incomplete statistics, enzymes produced using B. Many enzymes have been successfully expressed in B. These enzymes play important roles in the food, feed, detergent, textile, leather, paper, and pharmaceutical industries [ 46 ].
Here, we will introduce illustrative examples of engineered B. Alpha-amylase EC 3. It is widely used in the textile and paper industries, and B. Ma et al. Other studies focused on regulating protein transport and transcription levels by integrating signal peptides and promoter engineering [ 73 ]. PrsA is an effective folding catalyst for proteins expressed in B. Overexpression of native PrsA from B. QuesadaGanuza et al.
Xylanases EC. They have significant application value in biotechnology and can be used to modify lignocellulosic materials. Xylanases are used in animal feed manufacturing, the paper and textile industries, and biofuel production.
Commercial xylanases are mainly produced in B. Sanchez-Alponti et al. When the five mutants were combined in a random combinatorial library, a double mutant with improved specific activity and thermal stability was obtained [ 77 ].
It is possible that proteins such as CotH and CotO act synergistically to enforce a specific width on the outer coat, analogous to the 'molecular rulers' of flagella and injectisomes The coat morphogenetic proteins may be important targets for evolutionary adaptation across species, providing a pathway for rapid modification of coat architecture. Thus, an especially interesting area of future research will be to compare the coat protein assembly network of B.
For example, SpoIVA is necessary to anchor the coat to the spore surface in both species. By contrast, deletion of cotE in B. The dynamics of spore encasement may also be largely conserved among spore-forming bacteria. In early electron micrographs, assembly of the exosporium in B. These electron microscopy observations appear to agree with the localization dynamics of individual exosporium proteins, which form an initial cap on the MCP pole before transitioning to a complete shell of protein that surrounds the spore In cells deleted of exsY , a homologue of B.
Through a combination of genetic, biochemical and microscopy studies, much progress has been made in understanding how the spore protective layers are assembled in B. By examining how spores interact with various surfaces, biofilms and microbial consortia, it should be possible to connect what has been learned in the laboratory to what occurs in nature.
Future studies that assess how well-conserved the proteins and assembly mechanisms are should help us to determine whether the phenomena observed in B. These future goals also bring to mind a key gap in our knowledge: we currently know very little about the diversity in spore coat structure and composition among species.
Progress in addressing some of these outstanding questions could be substantially facilitated by analysis of species beyond the familiar models. Importantly, knowledge of the coat assembly mechanisms could also be exploited in various applications with public health implications, such as vaccine development Box 2.
A variety of metabolically dormant bacterial cells, called resting cells, have been found in nature 1. Endospores form inside a mother cell after the separation of the sporangium into distinct forespore and mother cell compartments; however, endospore formation is not limited to the production of a single spore per mother cell. Metabacterium polyspora , which is found in the gastrointestinal microbiota of guinea pigs, forms endospores at both poles of the mother cell.
Genetic manipulation of Bacillus subtilis can also result in the formation of two viable endospores in a single mother cell Various other resting cell types that form 'outside' the original vegetative cell have also been described.
Streptomyces species sporulate by forming a multinucleate sporogenic cell at the leading tip of an aerial hypha filament. Divisomes areassembled along the length of the sporogenic cell to partition each chromosome into an individual compartment that will eventually become an exospore During myxospore formation in Myxococcus species and akinete formation in heterocyst-forming species of Cyanobacteria, an entire individual cell transforms its morphology to form a resting cell in the absence of division 1.
To our knowledge, there has been no comprehensive parallel study of the relative resistance properties of the known types of resting cells; however, meta-analysis suggests that all resting cell types appear to be resistant to desiccation, whereas resistance to other types of stress such as extreme heat and predation by protozoa is variable The spore coat of myxospores has been investigated to some extent , but it appears to be unrelated to the spore coat of endospores, both in terms of composition it is made essentially of exopolysaccharides and mechanism of assembly, which requires a dedicated protein machinery.
An understanding of spore coat organization may benefit the development of microbial cell surface display technologies, which have been used to produce biocatalysts, biosorbents and vaccines Bacillus subtilis is edible and has been used in Japan for centuries to ferment soy beans for the production of natto. Spores are also naturally heat-stable, so spore-based therapies may not require refrigeration, eliminating a major cost of vaccine distribution to developing nations Several groups have attempted to generate vaccines against Clostridium tetani by engineering the surface of B.
However, the antigens were fused to the outer coat proteins CotB and CotC, which are covered by the crust layer in B. In order to avoid using protein fusions to coat proteins, attempts have been made to use heterologous proteins that can be adsorbed to the spore surface. Nevertheless, this approach would also require a detailed characterization of the spore surface to determine how the different coat layers influence adsorption properties. Another avenue to explore in surface-display technologies is the generation of synthetic vesicles encased by selected coat proteins that could be tailor-made to display specific properties.
Eukaryotic vesicles are readily produced with purified components in vitro , so it should also be possible to produce vesicles coated with spore coat proteins. A first step in that direction was accomplished with the coat morphogenetic protein SpoVM fused to green fluorescent protein SpoVM—GFP , which appears to bind preferentially to sites of positive membrane curvature and displays affinity for vesicles close to the size of B.
Our understanding of the mechanisms of coat assembly in bacteria can be integrated with what is known about coated vesicle formation in eukaryotes. The first step in both events is the localization of proteins to a site of membrane curvature , Second, a scaffold of coat material is assembled at the initial site of localization, and protein polymerization proceeds around the circumference of the developing vesicle.
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