Staphylococcus carnosus

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Staphylococcus carnosus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Bacillota
Class: Bacilli
Order: Bacillales
Family: Staphylococcaceae
Genus: Staphylococcus
Species:
S. carnosus
Binomial name
Staphylococcus carnosus
Schleifer and Fischer 1982

Staphylococcus carnosus is a bacterium from the genus Staphylococcus that is both Gram-positive and coagulase-negative. [1] It was originally identified in dry sausage and is an important starter culture for meat fermentation. [1] [2] Unlike other members of its genus, such as Staphylococcus aureus and Staphylococcus epidermidis , S. carnosus is nonpathogenic and safely used in the food industry. [3]

Contents

Taxonomy

Staphylococcus carnosus is classified under the domain Bacteria, phylum Bacillota, class Bacilli, order Bacillales, family Staphylococcaceae, genus Staphylococcus , and species S. carnosus. [4] The Staphylococcus genus currently comprises over 60 species and subspecies, including S. carnosus. [5]

Phylogeny

PCR and DNA sequencing along with 16s rRNA sequencing are commonly used to differentiate among the staphylococcal species. [5] Different methods of phylogenetic analysis have been used to examine the relationships between the different Staphylococcus species. One method divided the species into two distinct clades based on their status as oxidase positive or negative, and among the species lacking oxidase activity, the S. auricularis lineage emerged as the sister group to the other species. [5] The next lineage in this clade, which diverged basally, includes several species: S. simulans, S. condimenti , S. piscifermentans , and S. carnosus subspecies. The subspecies of S. carnosus exhibited a close clustering, forming the closest relative group to S. condimenti. [5] Within the Simulans-Carnosus group, comprising S. simulans, S. condimenti, S. carnosus, and S. piscifermentans , a total of four coagulase-negative species were identified, all particularly susceptible to novobiocin. [5] These species, including S. carnosus, formed part of one of the earliest diverging lineages in Staphylococcus. [5]

According to comparative phylogenetic analyses, S. pseudintermedius is the most basal lineage in the Staphylococcus genus, meaning that it evolved early from the root of the phylogenetic tree and is unbranched, while S. carnosus forms the next most basal lineage. [6] Both S. pseudintermedius and S. carnosus are not found on humans, thus differentiating them from other staphylococcal species like S. aureus and S. epidermidis , and phylogeny implies that it was only after the split from S. carnosus that adaptation to humans evolved among the staphylococci. [6] Other characteristics that distinguish S. carnosus from the other staphylococcal species include its ability to thrive in a high salt environment, reduce nitrate, and make acetoin. Its inability to make acid from sucrose and maltose also serves as an identifying characteristic. [1]

Discovery

S. carnosus was first isolated from dry sausage in 1982 by Schleifer and Fischer. [1] The discovery process included isolation of the research samples, cultivation, microscopic examination, and various biochemical tests. Initially, S. simulans strains were identified in dry sausage samples by plating them on a selective medium for Staphylococcus, as designed by Schleifer and Krämer, or alternatively on plate-count agar. [1] The medium specific to staphylococci ensured that only members of the genus Staphylococcus would grow while the growth of undesired groups would be inhibited. [1] The plate-count agar technique is used to estimate the total number of bacteria in a given sample. Most of the samples used in both methods were cultivated aerobically in peptone-yeast extract-glucose-NaCl broth. [1] Several tests were performed to determine carbohydrate and physiological reactions, peptidoglycan type, the chemical makeup of the teichoic acids in the cell wall, and cytochrome pattern. [1] All the strains were found to be facultative anaerobes and used glucose to produce equivalent amounts of D- and L-lactate. [1] Finally, DNA-DNA hybridization studies found that S. carnosus has the highest GC content among all Staphylococcus species. [2] In comparative studies, it was observed that DNA homology values between S. carnosus and S. simulans were notably high when contrasted with the homology values between S. carnosus and other species within the Staphylococcus genus. Although the strains of S. simulans and S. carnosus share a close genetic relationship based on DNA homology, this similarity was deemed insufficient to categorize them as a single species. [1]

Morphology

S. carnosus typically appears as single or paired spherical cells, known as cocci, with diameters ranging from 0.5 to 1.5 μm. [1] These cells form colonies with a round configuration, smooth margins, and a slightly raised elevation. Colonies of S. carnosus often exhibit a grayish-white coloration and a subtle shiny texture, making them easily distinguishable on agar plates. [1] The diameter of these colonies typically falls within the range of 1 to 3 mm. [1] Gram staining of S. carnosus cells reveals that they are Gram-positive due to the thickness of their peptidoglycan layer. Lastly, S. carnosus is non-motile and non-spore forming, indicating the absence of a flagellum for movement and its inability to form endospores for survival during adverse conditions. [1] Instead, its growth and survival depends on its metabolic capabilities and adaptations to the environment.

Metabolism

S. carnosus is a facultative anaerobic chemoorganotroph that utilizes respiratory metabolism. [1] In conditions where oxygen is absent, it can use nitrate as a terminal electron acceptor in anaerobic respiration. [7] When nitrate is reduced, the resulting nitrite first accumulates in the medium. [7] Once the nitrate has been depleted, the nitrite is absorbed by the cells and undergoes further reduction to ammonia, which is then incorporated into biomass. [7] The synthesis of both nitrite reductase and nitrate reductase is inhibited by oxygen, which is consistent with the status of S. carnosus as a facultative anaerobe that carries out aerobic respiration for energy in the presence of oxygen and switches to anaerobic nitrate respiration when oxygen is not available. [7]

When grown aerobically, strains of S. carnosus produced acid on some sugars such as glucose and fructose but did not produce acid on other sugars such as sucrose and lactose. [1] Compared to other staphylococci species, such as S. xylosus and S. equorum , S. carnosus has a greater tendency to degrade certain amino acids into methyl-branched aldehydes, their respective esters, and acids, and also produce more methyl ketone products through fatty acid β-oxidation. [8] All of these compounds contribute to aroma and affect the degree of maturity of fermented sausage. [8]

S. carnosus tested positive for catalase, an enzyme responsible for decomposing hydrogen peroxide into water and oxygen. [1] It also tested positive for benzidine, which confirms that it has a cytochrome system used during aerobic respiration. [1]

Physiology

S. carnosus is a mesophile because it displays a growth optimum at relatively high temperatures (32-37 °C). [9] S. carnosus has a high tolerance for salt and can persist in NaCl concentrations of up to 15%, [1] although growth begins to slow at 10% NaCl. [10] Due to its role in meat fermentation, S. carnosus can live in acidic conditions and adapt to a pH of 5.5. [10] However, its growth can be limited in the presence of more pH-tolerant microorganisms. [11] It has also been shown that S. carnosus, specifically strain TM300, is capable of altering the composition of its peptidoglycan in response to different incubation conditions. [12] When grown in the presence of high sugar levels, S. carnosus experienced overflow metabolism that led to the appearance of tetra stem peptides in its peptidoglycan. [12] S. carnosus also upregulates the production of catalase and superoxide dismutase, both of which provide important antioxidant functions, when incubated in a sausage fermentation environment (i.e., acidic conditions, available nitrite and nitrate, and minimal aeration). [13]

Genomics

The genome of S. carnosus TM300 has been sequenced and analyzed. [3] Out of all the sequenced staphylococcal genomes, S. carnosus is distinguished from other species because it contains the highest GC content at 34.6%. [3] Its genome size is one of the smallest among those in the Staphylococcus genus (2.56 Mbp) and has a high coding density (86.0%). The genome contains the genes required for a starter culture, including a nitrate- and nitrite-reduction pathway, appropriate metabolic pathways, and enzymes that mitigate oxidative stress. [3] Many of the open reading frames in the S. carnosus genome are truncated, reflecting the loss of gene functions as a result of living in a nutrient-rich environment. [3] The non-pathogenicity of S. carnosus is supported by the low number of mobile elements in its genome as well as a lack of toxins and superantigens found in pathogenic species like S. aureus and S. epidermidis. [3] [14] Although S. carnosus is avirulent, its genome encodes for several homologs of virulence factors found in other staphylococci. [15] These proteins, however, do not have a pathogenic effect in S. carnosus and may contribute to other important functions such as host colonization. [15]

Genomic sequencing and annotation of a different S. carnosus strain (LTH 7013) taken from South Tyrolean ham revealed that this strain could also catalyze the reduction of nitrate to nitrite and nitrite to ammonia, and no toxins nor superantigens were identified. [16] The genome of S. carnosus LTH 3730, which was obtained from a sample of fermented fish, has also been sequenced. In addition to a nitrate-nitrite reduction system, LTH 3730 also contained genes encoding catalase, peroxidases, and proteins involved in oxidative stress response. [17] However, unlike other strains of S. carnosus such as TM300, LTH 3730 demonstrated hemolytic activity. [17] This hemolytic activity, coupled with the presence of proteins identified in pathogenic strains of staphylococci, has prevented the use of LTH 3730 as a starter culture.

Ecology

Although it was originally isolated from sausage, the natural habitat of S. carnosus remains contested. [2] While many species of Staphylococcus have been found in humans, S. carnosus has never been collected from human sources. [3] It has been speculated that S. carnosus may live on the skin of animals because of its common presence in meat. [3] Others have proposed that S. carnosus could be derived from fish based on its phylogenetic proximity to S. piscifermentans and the similarities in their 16S rRNA and CydA and CydB proteins. [3]

A study investigated the surface characteristics of different S. carnosus strains and found that they could adhere to surfaces commonly found in food processing factories such as stainless steel but could not accumulate, likely due to their inability to synthesize the polysaccharides that are important for adhesion. [18] This finding provides an explanation for why S. carnosus is not usually isolated from the environment, such as in the food industry and the clinical setting, and why its true ecological niche is still uncertain.

Applications

S. carnosus is one of the main species of Staphylococcus used in food fermentation. [19] The practice of using S. carnosus as a meat starter culture originated in the 1950s because of its nitrate- and nitrite-reducing ability, contributing to the desired coloring and flavoring of the meat while reducing odors. [3] Additionally, S. carnosus has an important role in preventing the growth of undesirable bacteria and hence guards against food spoilage. [2] The beneficial properties of S. carnosus make it particularly useful to the food industry, such as preserving fresh meat products. [20]

Other potential applications of S. carnosus outside of food fermentation are also being explored. Gram-positive bacteria like S. carnosus can be engineered to express metal-binding peptides that allow it to absorb metal ions, which can be applied to bioremediation of wastewater contaminated with toxic metals. [21] Additionally, its extensive use in meat preparation and status as a GRAS (generally regarded as safe) organism makes S. carnosus a possible candidate for delivering live vaccines as it poses very little danger to the host. [2] Thus, the unique characteristics of S. carnosus reveal avenues for its further development in medical advancements and environmental impact.

Related Research Articles

<i>Staphylococcus aureus</i> Species of Gram-positive bacterium

Staphylococcus aureus is a Gram-positive spherically shaped bacterium, a member of the Bacillota, and is a usual member of the microbiota of the body, frequently found in the upper respiratory tract and on the skin. It is often positive for catalase and nitrate reduction and is a facultative anaerobe that can grow without the need for oxygen. Although S. aureus usually acts as a commensal of the human microbiota, it can also become an opportunistic pathogen, being a common cause of skin infections including abscesses, respiratory infections such as sinusitis, and food poisoning. Pathogenic strains often promote infections by producing virulence factors such as potent protein toxins, and the expression of a cell-surface protein that binds and inactivates antibodies. S. aureus is one of the leading pathogens for deaths associated with antimicrobial resistance and the emergence of antibiotic-resistant strains, such as methicillin-resistant S. aureus (MRSA), is a worldwide problem in clinical medicine. Despite much research and development, no vaccine for S. aureus has been approved.

The nitrite ion has the chemical formula NO−
2. Nitrite is widely used throughout chemical and pharmaceutical industries. The nitrite anion is a pervasive intermediate in the nitrogen cycle in nature. The name nitrite also refers to organic compounds having the –ONO group, which are esters of nitrous acid.

<i>Staphylococcus haemolyticus</i> Species of bacterium

Staphylococcus haemolyticus is a member of the coagulase-negative staphylococci (CoNS). It is part of the skin flora of humans, and its largest populations are usually found at the axillae, perineum, and inguinal areas. S. haemolyticus also colonizes primates and domestic animals. It is a well-known opportunistic pathogen, and is the second-most frequently isolated CoNS. Infections can be localized or systemic, and are often associated with the insertion of medical devices. The highly antibiotic-resistant phenotype and ability to form biofilms make S. haemolyticus a difficult pathogen to treat. Its most closely related species is Staphylococcus borealis.

<i>Staphylococcus saprophyticus</i> Species of bacterium

Staphylococcus saprophyticus is a Gram-positive coccus belonging to the genus Staphylococcus. S. saprophyticus is a common cause of community-acquired urinary tract infections.

Staphylococcus hominis is a coagulase-negative member of the bacterial genus Staphylococcus, consisting of Gram-positive, spherical cells in clusters. It occurs very commonly as a harmless commensal on human and animal skin and is known for producing thioalcohol compounds that contribute to body odour. Like many other coagulase-negative staphylococci, S. hominis may occasionally cause infection in patients whose immune systems are compromised, for example by chemotherapy or predisposing illness.

<i>Staphylococcus xylosus</i> Species of bacterium

Staphylococcus xylosus is a species of bacteria belonging to the genus Staphylococcus. It is a Gram-positive bacterium that forms clusters of cells. Like most staphylococcal species, it is coagulase-negative and exists as a commensal on the skin of humans and animals and in the environment.

<i>Staphylococcus epidermidis</i> Species of bacterium

Staphylococcus epidermidis is a Gram-positive bacterium, and one of over 40 species belonging to the genus Staphylococcus. It is part of the normal human microbiota, typically the skin microbiota, and less commonly the mucosal microbiota and also found in marine sponges. It is a facultative anaerobic bacteria. Although S. epidermidis is not usually pathogenic, patients with compromised immune systems are at risk of developing infection. These infections are generally hospital-acquired. S. epidermidis is a particular concern for people with catheters or other surgical implants because it is known to form biofilms that grow on these devices. Being part of the normal skin microbiota, S. epidermidis is a frequent contaminant of specimens sent to the diagnostic laboratory.

Lysostaphin is a Staphylococcus simulans metalloendopeptidase. It can function as a bacteriocin (antimicrobial) against Staphylococcus aureus.

<span class="mw-page-title-main">Staphylococcal infection</span> Medical condition

A staphylococcal infection or staph infection is an infection caused by members of the Staphylococcus genus of bacteria.

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<i>Staphylococcus capitis</i> Species of bacterium

Staphylococcus capitis is a coagulase-negative species (CoNS) of Staphylococcus. It is part of the normal flora of the skin of the human scalp, face, neck, scrotum, and ears and has been associated with prosthetic valve endocarditis, but is rarely associated with native valve infection.

<i>Staphylococcus</i> Genus of Gram-positive bacteria

Staphylococcus is a genus of Gram-positive bacteria in the family Staphylococcaceae from the order Bacillales. Under the microscope, they appear spherical (cocci), and form in grape-like clusters. Staphylococcus species are facultative anaerobic organisms.

<i>Staphylococcus condimenti</i> Species of bacterium

Staphylococcus condimenti is a Gram-positive, coagulase-negative member of the bacterial genus Staphylococcus consisting of single, paired, and clustered cocci. Strains of this species were originally isolated from fermenting soy sauce mash and are positive for catalase, urease, arginine dihydrolase, nitrate reductase, beta-galactosidase, and phosphatase activity.

<i>Latilactobacillus sakei</i> Species of bacterium

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<i>Staphylococcus hyicus</i> Species of bacterium

Staphylococcus hyicus is a Gram-positive, facultatively anaerobic bacterium in the genus Staphylococcus. It consists of clustered cocci and forms white circular colonies when grown on blood agar. S. hyicus is a known animal pathogen. It causes disease in poultry, cattle, horses, and pigs. Most notably, it is the agent that causes porcine exudative epidermitis, also known as greasy pig disease, in piglets. S. hyicus is generally considered to not be zoonotic, however it has been shown to be able to cause bacteremia and sepsis in humans.

Staphylococcus massiliensis is a Gram-positive, coagulase-negative member of the bacterial genus Staphylococcus consisting of clustered cocci. Strains of this species were first isolated from a human brain abscess and were found to be most closely related to Staphylococcus piscifermentans, Staphylococcus condimenti, Staphylococcus carnosus subsp. carnosus, Staphylococcus carnosus subsp. utilis, and Staphylococcus simulans. A subsequent study found that S. massiliensis may actually be part of the human skin microbiome and may have been a contaminant of brain abscess-derived samples.

Staphylococcus piscifermentans is a Gram-positive, coagulase-negative member of the bacterial genus Staphylococcus consisting of clustered cocci. This species was originally isolated from fermented fish in Thailand. A later study found a strain of S. piscifermentans in dog feces. The species is used in the preparation of fermented foods along with Staphylococcus carnosus; both species reduce nitrate and produce ammonia.

Staphylococcus schleiferi is a Gram-positive, cocci-shaped bacterium of the family Staphylococcaceae. It is facultatively anaerobic, coagulase-variable, and can be readily cultured on blood agar where the bacterium tends to form opaque, non-pigmented colonies and beta (β) hemolysis. There exists two subspecies under the species S. schleiferi: Staphylococcus schleiferi subsp. schleiferi and Staphylococcus schleiferi subsp. coagulans.

Staphylococcus pseudintermedius is a gram positive coccus bacteria of the genus Staphylococcus found worldwide. It is primarily a pathogen for domestic animals, but has been known to affect humans as well. S. pseudintermedius is an opportunistic pathogen that secretes immune modulating virulence factors, has many adhesion factors, and the potential to create biofilms, all of which help to determine the pathogenicity of the bacterium. Diagnoses of Staphylococcus pseudintermedius have traditionally been made using cytology, plating, and biochemical tests. More recently, molecular technologies like MALDI-TOF, DNA hybridization and PCR have become preferred over biochemical tests for their more rapid and accurate identifications. This includes the identification and diagnosis of antibiotic resistant strains.

Staphylococcus borealis is a bacterial species, member of the genus Staphylococcus, closely related to Staphylococcus haemolyticus and described in 2020. Its cells are Gram positive, coccoid in shape, with a diameter of 0.65 to 1.23 µm and form clusters. Additionally, they are facultative anaerobic, coagulase negative and catalase positive. The type strain was isolated from human blood culture at the University Hospital of North Norway, in 1997. Four additional strains included in the description were isolated from skin swabs, from healthy volunteers. The genome sequence of the type strain is deposited in DNA Data Bank of Japan, European Nucleotide Archive and GenBank under the accession number JABVEJ000000000.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 SCHLEIFER, K. H.; FISCHER, U. (1982). "Description of a New Species of the Genus Staphylococcus: Staphylococcus carnosus". International Journal of Systematic and Evolutionary Microbiology. 32 (2): 153–156. doi:10.1099/00207713-32-2-153. ISSN   1466-5034.
  2. 1 2 3 4 5 Löfblom, John; Rosenstein, Ralf; Nguyen, Minh-Thu; Ståhl, Stefan; Götz, Friedrich (2017). "Staphylococcus carnosus: from starter culture to protein engineering platform". Applied Microbiology and Biotechnology. 101 (23): 8293–8307. doi:10.1007/s00253-017-8528-6. ISSN   0175-7598. PMC   5694512 . PMID   28971248.
  3. 1 2 3 4 5 6 7 8 9 10 Rosenstein, Ralf; Nerz, Christiane; Biswas, Lalitha; Resch, Alexandra; Raddatz, Guenter; Schuster, Stephan C.; Götz, Friedrich (2009-02-01). "Genome Analysis of the Meat Starter Culture Bacterium Staphylococcus carnosus TM300". Applied and Environmental Microbiology. 75 (3): 811–822. doi:10.1128/AEM.01982-08. ISSN   0099-2240. PMC   2632126 . PMID   19060169.
  4. ↑ taxonomy. "Taxonomy browser (Staphylococcus carnosus)". www.ncbi.nlm.nih.gov. Retrieved 2024-04-09.
  5. 1 2 3 4 5 6 Lamers, Ryan P.; Muthukrishnan, Gowrishankar; Castoe, Todd A.; Tafur, Sergio; Cole, Alexander M.; Parkinson, Christopher L. (2012-09-06). "Phylogenetic relationships among Staphylococcus species and refinement of cluster groups based on multilocus data". BMC Evolutionary Biology. 12 (1): 171. doi: 10.1186/1471-2148-12-171 . ISSN   1471-2148. PMC   3464590 . PMID   22950675.
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  7. 1 2 3 4 Neubauer, H; Götz, F (1996-04-01). "Physiology and interaction of nitrate and nitrite reduction in Staphylococcus carnosus". Journal of Bacteriology. 178 (7): 2005–2009. doi:10.1128/jb.178.7.2005-2009.1996. ISSN   0021-9193. PMC   177897 . PMID   8606176.
  8. 1 2 Søndergaard, Anne K.; Stahnke, Louise H. (2002-05-05). "Growth and aroma production by Staphylococcus xylosus, S. carnosus and S. equorum—a comparative study in model systems". International Journal of Food Microbiology. 75 (1–2): 99–109. doi:10.1016/S0168-1605(01)00729-2. PMID   11999121.
  9. ↑ Sindelar, J. J. (2014-01-01), "CURING | Natural and Organic Cured Meat Products in the United States", in Dikeman, Michael; Devine, Carrick (eds.), Encyclopedia of Meat Sciences (Second Edition), Oxford: Academic Press, pp. 430–435, doi:10.1016/b978-0-12-384731-7.00116-1, ISBN   978-0-12-384734-8 , retrieved 2024-04-09
  10. 1 2 Guo, H. L.; Chen, M. T.; Liu, D. C. (2000-03-01). "Biochemical Characteristics of Micrococcus varians, Staphylococcus carnosus and Staphylococcus xylosus and Their Growth on Chinese-Style Beaker Sausage". Asian-Australasian Journal of Animal Sciences. 13 (3): 376–380. doi: 10.5713/ajas.2000.376 . ISSN   2765-0189.
  11. ↑ Coventry, J.; Hickey, M. W. (1991-01-01). "Growth characteristics of meat starter cultures". Meat Science. 30 (1): 41–48. doi:10.1016/0309-1740(91)90033-M. ISSN   0309-1740.
  12. 1 2 Deibert, Julia; Kühner, Daniel; Stahl, Mark; Koeksoy, Elif; Bertsche, Ute (2016-09-23). "The Peptidoglycan Pattern of Staphylococcus carnosus TM300—Detailed Analysis and Variations Due to Genetic and Metabolic Influences". Antibiotics. 5 (4): 33. doi: 10.3390/antibiotics5040033 . ISSN   2079-6382. PMID   27669322.
  13. ↑ Barrière, C.; Leroy-Sétrin, S.; Talon, R. (2001-09-12). "Characterization of catalase and superoxide dismutase in Staphylococcus carnosus 833 strain". Journal of Applied Microbiology. 91 (3): 514–519. doi:10.1046/j.1365-2672.2001.01411.x. ISSN   1364-5072. PMID   11556918.
  14. ↑ Wagner, Elke; Doskar, Jirí; Götz, Friedrich (1998-02-01). "Physical and genetic map of the genome of Staphylococcus carnosus TM300". Microbiology. 144 (2): 509–517. doi: 10.1099/00221287-144-2-509 . ISSN   1350-0872. PMID   9493387.
  15. 1 2 Rosenstein, Ralf; Götz, Friedrich (2010-02-01). "Genomic differences between the food-grade Staphylococcus carnosus and pathogenic staphylococcal species". International Journal of Medical Microbiology. 300 (2–3): 104–108. doi:10.1016/j.ijmm.2009.08.014.
  16. ↑ Müller, Anne; Huptas, Christopher; Wenning, Mareike; Schmidt, Herbert; Weiss, Agnes (2015-06-25). "Draft Genome Sequence of Staphylococcus carnosus subsp. utilis LTH 7013, Isolated from South Tyrolean Ham". Genome Announcements. 3 (3). doi:10.1128/genomeA.00456-15. ISSN   2169-8287. PMC   4432338 .
  17. 1 2 Müller, Anne; Klumpp, Jochen; Schmidt, Herbert; Weiss, Agnes (2016-10-27). "Complete Genome Sequence of Staphylococcus carnosus LTH 3730". Genome Announcements. 4 (5). doi:10.1128/genomeA.01038-16. ISSN   2169-8287. PMC   5043556 . PMID   27688338.
  18. ↑ Planchon, S.; Gaillard-Martinie, B.; Leroy, S.; Bellon-Fontaine, M. N.; Fadda, S.; Talon, R. (2007-02-01). "Surface properties and behaviour on abiotic surfaces of Staphylococcus carnosus, a genetically homogeneous species". Food Microbiology. 24 (1): 44–51. doi:10.1016/j.fm.2006.03.010. hdl: 11336/54154 . ISSN   0740-0020.
  19. ↑ Heo, Sojeong; Lee, Jong-Hoon; Jeong, Do-Won (2020-08-01). "Food-derived coagulase-negative Staphylococcus as starter cultures for fermented foods". Food Science and Biotechnology. 29 (8): 1023–1035. doi:10.1007/s10068-020-00789-5. ISSN   2092-6456. PMC   7347722 . PMID   32670656.
  20. ↑ Xu, Michelle M.; Kaur, Mandeep; Pillidge, Christopher J.; Torley, Peter J. (2021-11-01). "Evaluation of the potential of protective cultures to extend the microbial shelf-life of chilled lamb meat". Meat Science. 181: 108613. doi:10.1016/j.meatsci.2021.108613. ISSN   0309-1740. PMID   34218124.
  21. ↑ Samuelson, Patrik; Wernérus, Henrik; Svedberg, Malin; Ståhl, Stefan (2000-03-01). "Staphylococcal Surface Display of Metal-Binding Polyhistidyl Peptides". Applied and Environmental Microbiology. 66 (3): 1243–1248. doi:10.1128/AEM.66.3.1243-1248.2000. ISSN   0099-2240. PMC   91973 . PMID   10698802.
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