Status, Antimicrobial Mechanism, and Regulation of Natural Preservatives in Livestock Food Systems
Status, Antimicrobial Mechanism, and Regulation of Natural Preservatives in Livestock Food Systems
Food Science of Animal Resources. 2016. Aug, 36(4): 547-557
Copyright © 2016, Korean Society for Food Science of Animal Resources
This is an open access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.
  • Received : June 30, 2016
  • Accepted : August 09, 2016
  • Published : August 31, 2016
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About the Authors
Na-Kyoung, Lee
Department of Food Science and Biotechnology of Animal Resources, Konkuk University, Seoul 05029, Korea
Hyun-Dong, Paik
Bio/Molecular Informatics Center, Konkuk University, Seoul 05029, Korea

This review discusses the status, antimicrobial mechanisms, application, and regulation of natural preservatives in livestock food systems. Conventional preservatives are synthetic chemical substances including nitrates/nitrites, sulfites, sodium benzoate, propyl gallate, and potassium sorbate. The use of artificial preservatives is being reconsidered because of concerns relating to headache, allergies, and cancer. As the demand for biopreservation in food systems has increased, new natural antimicrobial compounds of various origins are being developed, including plant-derived products (polyphenolics, essential oils, plant antimicrobial peptides (pAMPs)), animal-derived products (lysozymes, lactoperoxidase, lactoferrin, ovotransferrin, antimicrobial peptide (AMP), chitosan and others), and microbial metabolites (nisin, natamycin, pullulan, ε-polylysine, organic acid, and others). These natural preservatives act by inhibiting microbial cell walls/membranes, DNA/RNA replication and transcription, protein synthesis, and metabolism. Natural preservatives have been recognized for their safety; however, these substances can influence color, smell, and toxicity in large amounts while being effective as a food preservative. Therefore, to evaluate the safety and toxicity of natural preservatives, various trials including combinations of other substances or different food preservation systems, and capsulation have been performed. Natamycin and nisin are currently the only natural preservatives being regulated, and other natural preservatives will have to be legally regulated before their widespread use.
The food industry has developed along with globalization, resulting in an increased risk of foodstuffs being contaminated with pathogens, chemical residues, and toxins. The proliferation of pathogenic and spoilage bacteria should be controlled to guarantee food safety. Conventional preservatives are a group of synthetic chemical substances including nitrates/nitrites, sulfites, sodium benzoate, propyl gallate, and potassium sorbate. The use of these conventional preservatives in food has known side effects ( Sharma, 2015 ). Nitrites and nitrate have been linked to leukemia, colon, bladder, and stomach cancer. Sorbate and sorbic acid are rare; however, they are related to urticaria and contact dermatitis. Benozates have been suspected to relating to allergies, asthma, and skin rashes.
During recent decades, investigation on food preservation have focused on more natural and healthier food ( Caminiti ., 2011 ; Fangio and Fritz, 2014 ). Biopreservation has dealt with extending food shelf life and enhancing food safety using plants, animals, microorganisms, and their metabolites ( Settanni and Corsetti, 2008 ). Particularly, meat and meat products are perishable materials, and are controlled by the Hazard Analysis Critical Control Point (HACCP) approach. The risk of contracting foodborne illnesses is reduced by various food preservation methods; thermal processing, drying, freezing, refrigeration, irradiation, modified atmosphere packaging, and the addition of antimicrobial agents, salts, or other chemical preservatives. Unfortunately, these techniques cannot be applied to all food products because of undesired effects (texture, color, etc.) depending on food type, such as ready-to-eat foods and fresh foods. Especially, preserving meat products is more complex, with higher pH and mild pasteurization temperatures required.
Natural preservative are the chemical agents derived from plants, animals, and microorganisms, and are usually related to the host defense system ( Singh ., 2010 ; Tiwari ., 2009 ). As the demand for biopreservation in food systems has increased, new natural antimicrobial compounds of various origin are being developed, including animal-derived systems (lysozyme, lactoferrin, and magainins), plant-derived products (phytoalexins, herbs, and spices), and microbial metabolites (bacteriocins, hydrogen peroxide, and organic acids) ( Lavermicocca ., 2003 ). The requirements of natural preservatives are: safety, stability during food processing (pH, heat, pressure, etc.), and antimicrobial efficacy. The representative food pathogens are Escherichia coli, Salmonella spp., Listeria monocytogenes, Staphylococcus aureus, Bacillus cereus, Yersinia enterocolytica, Clostridium perfringens, Clostridium botulinum , and Campylobacter jejuni . The pathogenic fungi often related to food-borne diseases are toxin-producing Aspergillus flavus and Aspergillus paraciticus ( Prange ., 2005 ).
This review summarizes the current knowledge about natural preservatives regarding their antimicrobial effects, antimicrobial mechanism, application, and regulation in food systems.
Natural Preservatives of Plant Origin
Plant preservatives are composed to polyphenols and phenolics, essential oils, and plant antimicrobial peptides (pAMPs). These substances have evolved to possess antibacterial and antioxidant effect ( Dua ., 2013 ). Phenolics and polyphenols have various antimicrobial structures: simple phenols (caffeic acid, catechol, eugenol, and epicatechin) and phenolic acids (caffeic acid and cinnamic acid), quinones (hypericin), flavones, flavonols, flavonoids (epigallocatechin-3-gallate, catechin, and chrysin), tannins (pentagalloylglucose, procyanidine B-2), coumarins (coumarin, warfarin, and 7-hydroxycourmarin), terpenoids (menthol, artemisin, and capsaicin), and alkaloids (berberine and harmane) ( Table 1 ) ( Cowan, 1999 ; Hintz ., 2015 ). The pAMPs are represented by thionin, plant defensins, lipid transfer proteins (LTPs), myrosinase-binding proteins (MBPs), hevein- and knottin-like peptides, snakins, cyclotides, and peptides from hydrolysates ( Hintz ., 2015 ).
Major classes of natural preservatives of plant origin
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Major classes of natural preservatives of plant origin
- Status of plant preservatives
Plant polyphenol extracts have been used as natural meat preservatives, including extracts from oregano, cranberry, sage, rosemary, grape seed, and others. Polyphenols can act as reducing agents and metal ion chelators in the presence of various hydroxyl radicals.
Oregano and cranberry extracts were evaluated for antimicrobial activity against L. monocytogenes in laboratory media, beef, and fish ( Lin ., 2004 ). These phenolicbased plant extracts are widely used in food preparation and are classified as Generally Regarded as Safe (GRAS). The effects of neem oil on the meat pathogens Carnobacterium maltaromaticum, Brochothrix thermosphacta, E. coli , and Pseudomonas fluorescens , were investigated as a preservative for fresh retail meat ( Del Serrone ., 2015a , Del Serrone ., 2015b ). Citrus species extracts were investigated as antifungal agents against spoilage fungi including Mucor sp. and Rhizophus sp. ( Mohanka and Priyanka, 2014 ). Ethanol extract of Citrus species showed a higher antifungal effect than water extract did, and the minimum inhibitory concentration of the extract ranged from 6.25 to 25 mg/mL. Inula britannica ethanol extract showed an antimicrobial effect against five B. cereus strains in low fat milk, and the antimicrobial effect depended on terpene and polyphenol compounds ( Lee ., 2012 ). Brassica juncea extract showed an antiviral effect against influenza virus A/H1N1 in nonfat milk ( Lee ., 2014 ). Chestnut inner shell extract containing gallic acid and quercetin was shown an antimicrobial effect against C. jejuni in chicken meat at 1 and 2 mg/mL ( Lee ., 2016 ). Eight different flavonoids [quercetin, kaempferol, apigenin, luteolin, 5,4-dihydroxy-7-methnozyflavone (genkwanin), narigenin, hesperetin and hesperidin] were tested for antimicrobial effects against B. cereus strains (P14 and KCCM 40935) ( Lee ., 2011 ). Among these flavonoids, only kaempferol and apigenin were inhibitory, and kaempferol showed the greatest antimicrobial effect at 100 μM.
Essential oil or terpenes are secondary metabolites that provide flavor and aroma. The addition of adding essential oils from marjoram and rosemary was investigated in beef patties ( Mohamed and Mansour, 2012 ). These essential oils were beneficial for antioxidant activity and sensory evaluation.
Plant antimicrobial peptides (pAMPs) were discovered in 1942 as natural defense compounds against pathogens ( Hintz ., 2015 ). The pAMPs were named as thionins, plant defensins, lipid transfer proteins (LTPs), myrosinase-binding proteins (MBPs), hevein- and knottin-like peptides, snakins, cyclotides, and peptides from hydrolysates. These substances have been isolated from Triticum aestivum (wheat), Impatients balsamina, Hordeum vulgare (barley), Arabidopsis thaliana, Hevea brasiliensis, Solanum tuverosum (potato), Oldenlandia offinis , etc.
- Antimicrobial mechanisms of plant preservatives
The antimicrobial mechanisms of phenol compounds depend on their concentration. Phenols affect enzyme activity related to energy production at low concentrations; however, they cause protein denaturation at high concentrations ( Fig. 1 ). These abilities affect microbial cell permeability, thereby interfering with membrane function (material transport, nucleic acid synthesis, and enzyme activity) ( Bajpai ., 2008 ; Fung ., 1977 ; Rico-Munoz ., 1987 ). The high antibacterial activity of phenolic compounds can be due to alkyl substitution into the phenol nucleus, forming phenoxy radicals, which does not occur in more stable molecules such as the ethers myristicin or anethole ( Dorman and Deans, 2000 ; Gutierrez ., 2008 ). Catechol and pyrogallol possess phenolic toxicity to microorganisms through enzyme inhibition by oxidized compounds, possibly by reacting with sulfhydryl groups or through more nonspecific interactions with proteins ( Mason and Wasserman, 1987 ). The antimicrobial targets of quinones may include surface-exposed adhesins, cell wall polypeptide, and membrane-bound enzymes ( Cowan, 1999 ). The antimicrobial activities of isothiocynates derived from Brassicaceae vegetables, such as cauliflower, broccoli, mustard, and cabbage are related to 1) loss of cell membranes integrity, 2) inhibiting enzyme or regulatory activity by quorum sensing (in Helicobacter pylori, Pseudomonas aeruginosa, Chromobacterium violaceum , etc.), 3) inhibition of respiratory enzymes, 4) induction of heat-shock and oxidative stress, and 5) induction of a stringent response ( Dufort ., 2015 ). Carvacrol, (þ)-carvone, thymol, and trans-cinnamaldehyde decrease the intracellular ATP (adenosine triphosphate) content of E. coli O157:H7 cells ( Helander ., 1998 ).
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Antimicrobial mechanisms of natural preservatives. AMPs, antimicrobial peptides; pAMPs, plant antimicrobial peptides.
Essential oils have multiple cellular targets. Their hydrophobicity results in reactions with lipids on bacterial and fungal cell membranes, increasing membrane permeability and disturbing the original cell structure ( Hintz ., 2015 ; Pinto ., 2009 ). In addition, antiviral effects are achieved by inhibiting viral protein synthesis at multiple stages of viral infection and replication ( Wu ., 2010 ).
The antimicrobial mechanism of most pAMPs involves cell membranes of targeted organisms and is driven by net positive charge, flexibility, and hydrophobicity to enable interaction with bacterial membranes ( Jessen ., 2006 ). Their antifungal mechanisms are involved in cell lysis, interference with fungal cell wall synthesis, permeabilization, binding to ergosterol/cholesterol in the membrane, depolymerization of the actin cytoskeleton, and targeting intracellular organelles such as mitochondria. Antiviral activity is also related to viral adsorption and entry processes.
Natural Preservatives of Animal Origin
There are numerous antimicrobial systems of animal origin related to host defense mechanisms. Preservatives of animal origin include lysozymes, lactoperoxidase, lactoferrin, ovotransferrin, antimicrobial peptide (AMP), chitosan, and others ( Table 2 ).
Natural preservatives of animal origin
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Natural preservatives of animal origin
- Status of animal preservatives
Lysozyme is obtained from chicken egg whites, and is known as a bacteriolytic enzyme. Lysozyme has been used commercially under the name Inovapure, and can be used against a wide range of food spoilage organisms for extending the shelf life of various food products including raw and processed meats, cheese, and other dairy products ( Tiwari ., 2009 ).
The lactoperxoidase is a naturally active enzyme in milk with strong antimicrobial effects against both Gramnegative and -positive bacteria ( de Wit and van Hooydonk, 1996 ; Russell, 1991 ) and fungi. Lactoperoxidase catalyzes the hydrogen peroxide (H 2 O 2 ) oxidation of several acceptors; it has been utilized in industries such as dairy, oral care, cosmetics, cancer, and viral infection.
Lactoferrin and ovotranferrin are glycoproteins derived from bovine milk and hen egg respectively, that can bind iron, thereby restricting or preventing bacterial growth. Lactoferrin shows strong antimicrobial effects against various Gram-negative and -positive bacteria, fungi, and parasites in neutral pH and refrigeration temperature ( Al-Nabulsi and Holley, 2005 ). Ovotranferrin peptide fragment OTAP-92 has strong bactericidal activity against both S. aureus and E. coli strains through membrane damage ( Ibrahim ., 2000 ). However, these transferrin peptides cannot be utilized in food systems because of their high cost.
AMPs are widely distributed in nature and are essential components of nonspecific host defense systems ( Park ., 1997 ; Tossi ., 2000 ). The AMPs produced by animal cells include magainin ( Zasloff ., 1988 ), MSI-78 ( Ge and Yan, 2002 ), PR-39 ( Shi ., 1996 ), pleurocidin ( Cole ., 1997 ), and dermaseptin S4 ( Mor and Nicolas, 1994 ). AMPs are considered a promising solution for antibiotic resistance because of their non-specific molecular targets and fast membrane destruction. Pleurocidin is isolated from the winter flounder ( Pleuronectes americanus ) is active against Gram-negative and -positive bacteria ( Cole ., 2000 ). It is stable in heat and salt; however, it is unstable in supraphysiological concentrations to magnesium and calcium. An antimicrobial effect of pleurocidin was reported in foodborne organisms including Vibrio parahemolyticus, L. monocytogenes, E. coli O157: H7, Saccharomyces cerevisiae , and Penicillium expansum ( Burrowes ., 2004 ). Defensins are widely found in mammalian epithelial cells from chicken, turkey, and others ( Brockus ., 1998 ). Their antimicrobial spectrum included Gram-negative and -positive bacteria, fungi, and enveloped viruses ( Ganz, 2003 ; Murdock ., 200 7).
Chitosan is a natural biopolymer obtained from the exoskeletons of crustaceans and arthropods, and has been used as an antifungal and antimicrobial agent ( Ben-Shalom ., 2003 ; Je and Kim, 2006 ; Liu ., 2006 ). Chitooligosaccharides have a bacteriostatic effect on Gramnegative bacteria, E. coli, Vibrio cholera, Shigella dysenteriae , and Bacteriodes fragilis ( Benhabiles ., 2012 ). Chitosan (0.25, 0.5, and 1%) was studied as an antimicrobial ingredient in fresh pork sausage ( Bostan and ’Isin Mahan, 2011 ).
Lipids of animal origin have antimicrobial activity against a wide range of microorganisms. Free fatty acids at mucosal surfaces have been shown to inactivate S. aureus ( Bibel ., 1989 ). Milk lipids are active against Gram-positive bacteria including S. aureus, C. botulinum, B. subtilis, B. cereus , and L. monocytogenes , and Gramnegative bacteria such as P. aeruginosa, E. coli , and Salmonella enteritidis ( Isaacs ., 1990 ; Lampe ., 1998 ; Wang and Johnson, 1997 ).
- Antimicrobial mechanisms of animal preservatives
AMPs, transferrins, and lipids can influence cell membranes and peptide synthesis ( Fig. 1 ) ( Brogden, 2005 ; Zasloff, 2002 ). AMPs can interact directly with the microbial cell membrane and result in the leaching out of vital cell components ( Cole ., 2000 ; Hancock, 1997 ). Lipids mainly inhibit bacterial cell walls or membranes, intracellular replication, or intracellular targets. Lysozymes inhibit bacterial cell membranes by hydrolyzing β-1,4-glycosidic linkages between N-acetylmuramic acid and N-acetylglucosamine in bacterial peptidoglycan.
Natural Preservatives from Microorganisms
The preservative of microbial origin include nisin, natamycin, pullulan, ε-polylysine, organic acid, and others ( Singh ., 2010 ) ( Table 3 ).
Natural preservatives from microorganisms
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Natural preservatives from microorganisms
- Status of microbial preservatives
Lactic acid bacteria produce antimicrobial compounds like organic acids, diacetyl, hydrogen peroxide, and proteinaceous bacteriocins ( Lee ., 2013 ). Bacteriocins are antimicrobial peptides or proteins produced by mainly lactic acid bacteria; these compounds are small and ribosomally synthesized. Most bacteriocins have potential as food preservatives because of their antimicrobial effect against food pathogens. The representative bacteriocins are nisin, diplococcin, acidophilin, bulgaricin, helveticin, lactacin, pediocin, and plantarin. Of these bacteriocins, nisin and pediocin have been used as commercial natural preservatives.
Nisin is a representative bacteriocin produced by various Lactococcus lactis strains, and has activity against food pathogens including Alicyclobacillus spp., L. monocytogenes, Bacillus spp., Micrococcus spp., Clostridium spp., Pediococcus spp., Desulfotomaculus spp., S. aureus, Enterococcus spp., Streptococcus haemolyticus, Lactobacillus spp., Sporolactobacillus spp., and Leuconostoc spp. Nisin is proteinaceous polypeptide that is most stable in acidic conditions. Nisin is soluble in aqueous conditions and is unstable in alkali solutions and heat. It has been used in various food products alone or in combination with other compounds. Nisin is the most widely used bacteriocin approved by the FDA as a food preservative. Dairy and meat products are applied with doses of 50-200 mg/kg. In the USA, nisin is used to inhibit outgrowth of C. botulinum spores and toxin formation in pasteurized processed cheese spread with fruits, vegetables or meats with a limited dose of about 250 ppm in finished products.
Pediocin is GRAS bacteriocin produced by strains of Pediococcus acidilactici (AcH, PA-1, JD, and 5) and P. pentosaceus (A, N5p, ST18, and PD1) ( Anastasiadou ., 2008 ). Most pediocins are stable in heat and a wide range of pH vlues. Pediocin AcH is effective against both spoilage and pathogenic organisms, including L. monocytogenes, Enterococcus faecalis, S. aureus , and Clostridium perfringens ( Bhunia ., 1988 ).
Natamycin is an antifungal substance produced by Streptomyces natalensis that is effective against almost all molds and yeasts; however, it has little or no effect on bacteria ( EFSA, 2009 ). Natamycin has been used in dairy, meats, and other foods for antifungal effects, and its use as a surface preservative is regulated (E 235).
Reuterin (β-hydroxypropionaldehyde), an antimicrobial compound produced by Lacotobacillus reuteri , is a watersoluble nonproteinaceous metabolite of glycerol ( Axelsson ., 1989 ). Its broad antimicrobial spectrum includes Gram-negative and -positive bacteria, yeasts, and filamentous fungi ( Nom and Rombouts, 1992 ).
- Antimicrobial mechanisms of microbial preservatives
The antimicrobial mechanism of bacteriocin involves pore formation in the cytoplasmic membrane of target microorganisms ( Fig. 1 ). This leads to cell death by loss of intracellular molecules and a collapse of the proton motive force ( Driessen ., 1995 ). Bacteriocin originating from Gram-positive bacteria is only effective for Grampositive bacteria, and is less effective on Gram-negative bacteria due to their selective membrane permeability ( Lee ., 2003 ). These disadvantages could be compensated for by using other preservatives and preservative methods.
Natamycin has an antimicrobial effect by binding to ergosterol, a cell membrane sterol, in fungal membranes ( EFSA, 2009 ). The structure of natamycin contains a large lactone ring with a rigid lipophilc chain containing conjugated double bonds, and a flexible hydrophilic portion bearing several hydroxyl groups. The hydrophobic groups form a polar pore with ergosterol in the membrane, and this complex affects material passage (K + , H + , amino acids, and other metabolites) ( Deacon, 1997 ).
Application of Natural Preservatives toward Livestock Food Systems
Raw meat, meat products, milk, and milk products are major sources of foodborne pathogens, and a variety of methods have been considered to reduce bacterial contaminants. These methods include (a) chemical decontamination with organic acids ( Gill and Badoni, 2004 ; Goncalves ., 2005 ; Nissen ., 2001 ) and trisodium phosphate ( Bashor ., 2004 ; Okolocha and Ellerbroek, 2005 ); (b) physical processes such as irradiation ( Badr, 2005 ; Kim ., 2004 ), high pressure processing ( Oliveira ., 2015 ), steam ( Kang ., 2001a ; Kang ., 2001b ; Logue ., 2005 ; Stivarius ., 2002 ), and UV; (c) natural antimicrobials such as bacteriocins ( de Martinez ., 2002 ; Gogus ., 2004 ) and iron chelating compounds; and (d) combination treatment ( Bashor ., 2004 ; Koohmaraie., 2005 ).
Challenge studies using meat samples mainly reported efficacy against L. monocytogenes, B. cereus, C. jejuni , and S. aureus ( Barman ., 2014 ). The efficacy of natural preservatives was tested against commercial formulation (Microgard 100, Microgard 300, nisin, Altak 2002, Perlack 1902) ( Lemay ., 2002 ). These natural preservatives were investigated in an acidified chicken meat model (pH 5.0). E. coli ATCC 25922 and Brochothrix thermosphacta CRDAV452 were inhibited, however Lactobacillus alimentarius BJ33 (FloraCarn L-2) was not inhibited.
The use of fruit byproducts, including rinds of grapefruit, orange, and mandarin with or without γ-irradiation, was applied in raw ground beef ( Abd El-khalek and Zahran, 2013 ). These substances demonstrated antioxidant and antimicrobial properties on microbial growth, lipid oxidation, and color change of raw ground beef meat. The antimicrobial effects on the survival of Salmonella typhimurium, E. coli and B. cereus were demonstrated.
A combination of plant extracts and MAPs was applied in meat products. Thymol and thymol-MAP were applied in sausage to inhibit Pseudomonas spp.; however, the performance is unacceptable respect to sensory acceptability ( Mastromatteo ., 2011 ). Bay essential oil with MAP (20% CO 2 and 80% N 2 ) was applied in ground chicken meat, and extend the shelf life without L. monocytogenes and E. coli ( Irkin and Esmer, 2010 ). Oregano oil was added to fresh chicken breast meat under MAP ( Chouliara ., 2007 ). At 1%, oregano oil had a very strong taste in the sensory evaluation; however 0.1% oregano oil and MAP extended the shelf life by 5-6 d without strong taste.
Plant preservatives like clove oil showed a synergistic effect with lactic acid and vitamin C for antioxidant and antimicrobial effects ( Naveena ., 2006 ). Ntzimani . (2010) used a mixture of EDTA, lysozymes, rosemary, and oregano oil, and the shelf life of semi-cooked coated chicken fillets was extended under vacuum packaging at 4℃ to more than 2 wk.
Nisin was applied with lactoferrin in Turkish-style meatballs. Counts of total aerobic bacteria, coliform, E. coli , and other species were decreased by lactoferrin alone and by the mixture of lactoferrin and nisin ( Colak ., 2008 ). A mixture of lysozyme, nisin, and EDTA effectively inhibited L. monocytogenes and meat-borne spoilage bacteria in ostrich patties packaged in air and vacuum ( Kim ., 2002 ; Mastromatteo ., 2010 ).
Regulation of Natural Preservatives in Livestock Foods
Preservatives permitted in livestock foods are sodium acetate, natamycin, pimamycin, nisin, nitrites (potassium nitrite and sodium nitrite), nitrates (potassium nitrate and sodium nitrate), sorbates (sorbic acid, sodium sorbate, potassium sorbate, and calcium sorbate), and sulphites (sulfur dioxide, sodium sulfite, sodium bisulfite, sodium metabisulfite, potassium metabisulfite, potassium sulfite, and potassium bisulfite) ( Food and Drug Administration, 2016 ).
Natural food preservatives are regulated by maximum permitted levels for food safety and health ( Table 4 ). The only natural preservatives regulated by legislation are natamycin and nisin. Natamycin (E235) is permitted for use in over 150 countries in the surface treatment of hard, semi-hard and semi soft cheeses and dried, cured sausages with a maximum permitted level of 6-40 mg/kg. Nisin (E234) is permitted for use in over 80 countries worldwide, including the United States and European Union, and has been in use as a food preservative for over 50 years ( Adams, 2003 ; EFSA, 2006 ). The maximum permitted levels in meat, poultry, game products are 5.5-7 mg/kg.
Representative natural preservatives and their maximum permitted level from codex
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ESFA (2006, 2009); GSFA (1995); KFDA (2016).
Natural preservatives are considered safer than synthetic preservatives because of their existence in nature and long history of use. However, the use of natural preservatives in food is not powerful enough when considering added amounts in food system. Therefore, effective use levels of conventional and plant extracts/oils against microorganisms are less than 0.1% and 10-20%, respectively ( Browne ., 2012 ). Therefore, the regulation of these natural preservatives as food additives is necessary regarding their safety, toxicity, and effectiveness.
Chemical preservative have side effects related to the emergence of drug-resistant strains and chronic toxicity. Traditional methods of preservation including refrigeration, pasteurization, and low pH are not completely effective in controlling food pathogens. Therefore, the efficacy of combining natural preservatives with traditional methods has been tested. Combination with other substances or different food preservation systems, coatings, or microand nano-capsulation should be tested to assure safety and nontoxicity of natural preservatives. In addition, the use of natural preservatives must be regulated by law for safety, toxicity, and effectiveness.
This work was supported by Priority Research Centers Program through the National Research Foundation of Korea (NRF) funded by the Ministry of Education, Science and Technology (2012-0006686).
Abd El-khalek H. H. , Zahran D. A. (2013) Utilization of fruit by-product in ground meat preservation Food Sci. Qual. Man. 11 49 - 60
Adams M. , Roller S. (2003) Natural antimicrobials for the minimal processing of foods CRC Press LLC Boca Raton, FL Nisin in multifactorial food preservation 11 - 33
Al-Nabulsi A. A. , Holley R. A. (2005) Effect of bovine lactoferrin against Carnobacterium viridans Bioresource Technol. 22 179 - 187
Anastasiadou S. , Papagianni M. , Filiousis G. , Ambrosiadis I. , Koidis P. (2008) Pediocin SA-1, an antimicrobial peptide from Pediococcus acidilactici NRRL B5627: Production conditions, purification and characterization Bioresources Technol. 99 5384 - 5390    DOI : 10.1016/j.biortech.2007.11.015
Axelsson L. T. , Chung T. C. , Dobrogosz W. J. , Lindgren S. E. (1989) Production of a broad spectrum antimicrobial substance by Lactobacillus reuteri Microb. Ecol. Health D. 2 131 - 136    DOI : 10.3109/08910608909140210
Badr H. M. (2005) Elimination of Escherichia coli O157:H7 and Listeria monocytogenes from raw beef sausage by γ-irradiation Mol. Nutr. Food Res. 49 343 - 349    DOI : 10.1002/mnfr.200400095
Bajpai V. K. , Rahman A. , Dung N. T. , Huh M. K. , Kang S. C. (2008) In vitro inhibition of food spoilage and foodbourne pathogenic bacteria by essential oil and leaf extracts of Magnolia liliflora Desr. J. Food Sci. 73 M314 - M320    DOI : 10.1111/j.1750-3841.2008.00841.x
Barman S. , Ghosh R. , Mandal N. C. (2014) Use of bacteriocin producing Lactococcus lactis subsp. lactis LABW4 to prevent Listeria monocytogenes induced spoilage of meat Food Nutr. Sci. 5 2115 - 2123    DOI : 10.4236/fns.2014.522224
Bashor M. P. , Curtis P. A. , Keener K. M. , Sheldon B. W. , Kathariou S. , Osborne J. A. (2004) Effects of carcass washers on Campylobacter contamination in large broiler processing plants Poultry Sci. 83 1232 - 1239    DOI : 10.1093/ps/83.7.1232
Benhabiles M. S. , Salah R. , Lounici H. , Drouiche N. , Goosen M. F. A. , Mameri N. (2012) Antibacterial activity of chitin, chitosan and its oligomers prepared from shrimp shell waste Food Hydrocolloid. 29 48 - 56    DOI : 10.1016/j.foodhyd.2012.02.013
Ben-Shalom N. , Ardi R. , Pinto R. , Aki C. , Fallik E. (2003) Controlling gray mould caused by Botytis cinerea in cucumber plants by means of chitosan Crop Prot. 22 285 - 290    DOI : 10.1016/S0261-2194(02)00149-7
Bhunia A. K. , Johnson M. C. , Ray B. (1988) Purification, characterization and antimicrobial spectrum of a bacteriocin produced by Pediococcus acidilactici J. Appl. Microbiol. 65 261 - 268
Bibel D. J. , Miller S. J. , Brown B. E. , Pandey B. B. , Elias P. M. , Shinefield H. M. , Aly R. (1989) Antimicrobial activity of stratum corneum lipids from normal and essential fatty acid-deficient mice J. Invest. Dermatol. 92 632 - 638    DOI : 10.1111/1523-1747.ep12712202
Bostan K. , ’Isin Mahan F. (2011) Microbiological quality and shelf-life of sausage treated with chitosan J. Fac. Vet. Med. Istanbul Univ. 37 117 - 126
Brogden K. A. (2005) Antimicrobial peptides: pore formers or metabolic inhibitors in bacteria? Nat. Rev. Microbiol. 3 238 - 250    DOI : 10.1038/nrmicro1098
Browne B. A. , Geis P. , Rook T. (2012) Conventional vs. natural preservatives HPPI. 2012 69 - 73
Brockus C. W. , Jackwood M. W. , Hamon B. G. (1998) Characterization of β-defensin prepropeptide mRNA from chicken and turkey bone marrow Anim. Genet. 29 283 - 289    DOI : 10.1046/j.1365-2052.1998.00338.x
Burrowes O. J. , Hadjicharalambous C. , Diamond G. , Lee T. C. (2004) Evaluation of antimicrobial spectrum and cytotoxic activity of pleurocidin for food application J. Food Sci. 69 66 - 71
Caminiti I. M. , Noci F. , Muñoz A. , Whyte P. , Morgan D. J. , Cronin D. A. , Lyng J. G. (2011) Impact of selected combinations of non-thermal processing technologies on the quality of an apple and cranberry juice blend Food Chem. 124 1387 - 1892    DOI : 10.1016/j.foodchem.2010.07.096
Chouliara E. , Karatapanis A. , Savvaidis I. N. , Kontominas M. G. (2007) Combined effect of oregano essential oil and modified atmosphere packaging on shelf-life extension of fresh chicken breast meat, stored at 4℃ Food Microbiol. 24 607 - 617    DOI : 10.1016/
Colak H. , Hampikyan H. , Bingol E. B. , Aksu H. (2008) The effect of nisin and bovine lactoferrin on the microbiological quality of Turkish-style meatball (Tekirdag k öfte) J. Food Safety 28 355 - 375    DOI : 10.1111/j.1745-4565.2008.00105.x
Cole A. M. , Darouiche R. O. , Legarda D. , Connell N. , Diamond G. (2000) Characterization of a fish antimicrobial peptide: gene expression, subcellular localization and spectrum of activity Antimicrob. Agents Chemother. 44 2039 - 2045    DOI : 10.1128/AAC.44.8.2039-2045.2000
Cole A. M. , Weis P. , Diamond G. (1997) Isolation and characterization of plurocidin, an antimicrobial peptide in the skin secretions of winter flounder J. Biol. Chem. 272 12008 - 12013    DOI : 10.1074/jbc.272.18.12008
Cowan M. M. (1999) Plant products as antimicrobial agents Clin. Microbiol. Rev. 12 564 - 582
de Martinez Y. B. , Ferrer K. , Salas E. M. (2002) Combined effects of lactic acid and nisin solution in reducing levels of microbiological contamination in red meat carcasses J. Food Prot. 65 1780 - 1783
de Wit J. N. , van Hooydonk A. C. M. (1996) Structure, functions and applications of lactoperoxidase in natural antimicrobial systems Neth. Milk Dairy J. 50 227 - 244
Deacon J. W. (1997) Modern Mycology 3rd Ed. Blackwell Science Oxford Prevention and control of fungal growth 289 - 290
Del Serrone P. , Toniolo C. , Nicoletti M. (2015a) Neem (Azadirachta indica A. Juss) oil to tackle enteropathogenic Escherichia coli BioMed Res. Int. Article ID 343610
Del Serrone P. , Toniolo C. , Nicoletti M. (2015b) Neem (Azadirachta indica A. Juss) oil: A natural preservative to control meat spoilage Foods 4 3 - 14    DOI : 10.3390/foods4010003
Dorman H. J. D. , Deans S. G. (2000) Antimicrobial agents from plants: Antibacterial activity of plant volatile oils J. Appl. Microbiol. 88 308 - 316    DOI : 10.1046/j.1365-2672.2000.00969.x
Driessen A. J. M. , van den Hoov H. W. , Kuiper W. , van der Kamp M. , Sahl H. G. , Konings R. N. H. , Konings W. N. (1995) Mechanistic studies of lantibiotic-induced permeabilization of phospholipids vesicles Biochem. 34 1606 - 1614    DOI : 10.1021/bi00005a017
Dua A. , Garg G. , Mahajan R. (2013) Polyphenols, flavonoids and antimicrobial properties of methanolic extract of fennel (Foeniculum vulgare Miller) Euro. J. Exp. Bio. 3 203 - 208
Dufort V. , Stahl M. , Baysse C. (2015) The antibacterial properties of isothiocyantes Microbiol. 161 229 - 243    DOI : 10.1099/mic.0.082362-0
EFSA (2006) Opinion of the scientific panel on food additives, flavorings, processing aids and material in contact with food on the safety in use of nisin as a food additive in an additional category of liquid eggs and on the safety of nisin produced using a modified production process as a food additive EFSA J. 314 1 - 8
EFSA (2009) Scientific opinion on the use of natamycin (E 235) as a food additive EFSA panel on food additives and nutrient sources added to food (ANS) EFSA J. 1412 1 - 25
Fangio M. F. , Fritz R. (2014) Potential use of a bacteriocin-like substance in meat and vegetable food biopreservation Int. Food Res. J. 21 677 - 683
Food and Drug Administration (2016) Food additive status list Available from: Accessed April 8, 2016
Fung D. Y. C. , Taylor S. , Kahan J. (1977) Effects of butylated hydroxyanisole (BHA) and butylated hydroxitoluene (BHT) on growth and aflatoxin production of Aspergillus flavus J. Food Safety 1 39 - 51    DOI : 10.1111/j.1745-4565.1977.tb00258.x
Ganz T. (2003) Defensins: antimicrobial peptides of innate immunity Nat. Rev. Immunol. 3 710 - 720    DOI : 10.1038/nri1180
Garba S. , Okeniyi S. O. (2012) Antimicrobial activities of total alkaloids extracted from some Nigerian medicinal plants J. Microbiol. Antimicrob. 4 60 - 63
Ge Y. , Yan H. (2002) Extraction of natural vitamin E from wheat germ by supercritical carbon dioxide J. Agric. Food Chem. 50 685 - 689    DOI : 10.1021/jf010615v
Gill C. O. , Badoni M. (2004) Effects of peroxyacetic acid, acidified sodium chlorite or lactic acid solutions on the microflora of chilled beef carcasses Int. J. Food Microbiol. 91 43 - 50    DOI : 10.1016/S0168-1605(03)00329-5
Gogus U. , Bozoglu F. , Yurdugul S. (2004) The effects of nisin, oil-wax coating and yogurt on the quality of refrigerated chicken meat Food Control 15 537 - 542    DOI : 10.1016/j.foodcont.2003.08.007
Goncalves A. C. , Almeida R. C. C. , Alves M. A. O. , Almeida P. F. (2005) Quantitative investigation on the effects of chemical treatments in reducing Listeria monocytogenes populations on chicken breast meat Food Control 16 617 - 622    DOI : 10.1016/j.foodcont.2004.06.026
GSFA (1995) General Standard for Food Additives. Codex STAN 192
Gutierrez J. , Rodriguez G. , Barry-Ryan C. , Bourke P. (2008) Efficacy of plant essential oils against foodborne pathogens and spoilage bacteria associated with ready-to-eat vegetables: Antimicrobial and sensory screenin J. Food Prot. 71 1846 - 1854
Hancock R. E. W. (1997) Peptide antibiotics Lancet 349 418 - 422    DOI : 10.1016/S0140-6736(97)80051-7
Helander I. M. , Alakomi H. L. , Latva-Kala K. , Mattila-Sandholm T. , Pol I. , Smid E. J. , Gorris L. G. M. , von Wright A. (1998) Characterisation of the action of selected essential oil components on gram-negative bacteria J. Agric. Food Chem. 46 3590 - 3595    DOI : 10.1021/jf980154m
Hintz T. , Matthews K. K. , Di R. (2015) The use of plant antimicrobial compounds for food preservation Biomed Res. Int. Article ID 246264
Ibrahim H. R. , Sugimoto Y. , Aoki T. (2000) Ovotranferrin antimicrobial peptide (OTAP-92) kills bacteria through a membrane damage mechanism Biochim. Biophys. Acta. 1523 196 - 205    DOI : 10.1016/S0304-4165(00)00122-7
Irkin R. , Esmer O. K. (2010) Control of Listeria monocytogenes in ground chicken breast meat under aerobic, vacuum and modified atmosphere packaging conditions with or without the presence of bay essential oil at 4℃ Food Sci. Technol. Res. 16 285 - 290    DOI : 10.3136/fstr.16.285
Isaacs C. E. , Kashyap S. , Heird W. C. , Thormar H. (1990) Antiviral and antibacterial lipids in milk and infant formula feeds Arch. Dis. Child 65 861 - 864    DOI : 10.1136/adc.65.8.861
Je J. Y. , Kim S. K. (2006) Chitosan derivatives killed bacteria by disrupting the outer and inner membrane J. Agric. Food Chem. 54 6629 - 6633    DOI : 10.1021/jf061310p
Jessen H. , Hamill P. , Hancock R. E. W. (2006) Peptide antimicrobial agents Clin. Microbiol. Rev. 19 479 - 491
Kang D. H. , Koohmaraie M. , Dorsa W. J. , Siragusa G. R. (2001a) Development of a multiple-step process for the microbial decontamination of beef trim J. Food Prot. 64 63 - 71
Kang D. H. , Koohmaraie M. , Siragusa G. R. (2001b) Application of multiple antimicrobial interventions for microbial decontamination of commercial beef trim J. Food Prot. 64 168 - 171
Kim B. H. , Jang A. R. , Lee S. O. , Min J. S. , Lee M. H. (2004) Combined effect of electron-beam (beta) irradiation and organic acids on shelf life of pork loins during cold storage J. Food Prot. 67 168 - 171
Kim Y. M. , Paik H. D. , Lee D. S. (2002) Shelf-life characteristics of fresh oysters and ground beef as affected by bacteriocin-coated plastic package film J. Sci. Food Agric. 82 998 - 1002    DOI : 10.1002/jsfa.1125
Koohmaraie M. , Arthur T. M. , Bosilevac J. M. , Guerini M. , Shackelford S. D. , Wheeler T. L. (2005) Post-harvest interventions to reduce/eliminate pathogens in beef Meat Sci. 71 79 - 91    DOI : 10.1016/j.meatsci.2005.03.012
KFDA (2015) Korea Food Additives Code Available from: . Accessed Feb. 20, 2016
Lampe M. F. , Ballweber L. M. , Isaacs C. E. , Patton D. L. , Stamm W. E. (1998) Killing of Chlamydia trachomatis by novel antimicrobial lipids adapted from compounds in human breast milk Antimicrob. Agents Chemother. 42 1239 - 1244
Lavermicocca P. , Valerio F. , Visconti A. (2003) Antifungal activity of phenyllactic acid against molds isolated from bakery products Appl. Environ. Microbiol. 69 634 - 640    DOI : 10.1128/AEM.69.1.634-640.2003
Lee D. U. , Heinz V. , Knorr D. (2003) Effects of combination treatments of nisin and high-intensity ultrasound with high pressure on the microbial inactivation in liquid whole egg Innov. Food Sci. Emerg. Technol. 4 387 - 393    DOI : 10.1016/S1466-8564(03)00039-0
Lee J. H. , Lee Y. J. , Ahn S. H. , Lee N. K. , Paik H. D. (2012) Antimicrobial properties of whole milk with Inula britannica extract against Bacillus cereus strains during storage Milchwissenschaft 67 315 - 317
Lee K. A. , Moon S. H. , Kim K. T. , Nah S. Y. , Paik H. D. (2011) Antimicrobial effect of kaempferol on psychrotrophic Bacillus cereus strains outbreakable in dairy products Korean J. Food Sci. An. 31 311 - 315    DOI : 10.5851/kosfa.2011.31.2.311
Lee N. K. , Han E. J. , Han K. J. , Paik H. D. (2013) Antimicrobial effect of bacteriocin KU24 produced by Lactococcus lactis KU24 against methicillin-resistant Staphylococcus aureus J. Food Sci. 78 M465 - M469    DOI : 10.1111/1750-3841.12053
Lee N. K. , Jung B. S. , Na D. S. , Yu H. H. , Kim J. S. , Paik H. D. (2016) The impact of antimicrobial effect of chestnut inner shell extracts against Campylobacter jejuni in chicken meat LWT-Food Sci. Technol. 65 746 - 750    DOI : 10.1016/j.lwt.2015.09.004
Lee N. K. , Lee J. H. , Lim S. M. , Lee K. A. , Kim Y. B. , Chang P. S. , Paik H. D. (2014) Antiviral activity of subcritical water extract of Brassica juncea against influenza virus A/H1N1 in nonfat milk J. Dairy Sci. 97 5383 - 5386    DOI : 10.3168/jds.2014-8016
Lemay M. J. , Choquette J. , Delaquis P. J. , Gariépy C. , Rodrigue N. , Saucier L. (2002) Antimicrobial effect of natural preservatives in a cooked and acidified chicken meat model Int. J. Food Microbiol. 78 217 - 226    DOI : 10.1016/S0168-1605(02)00014-4
Lin Y. T. , Labbe R. G. , Shetty K. (2004) Inhibition of Listeria monocytogenes in fish and meat systems by use of oregano and cranberry phytochemical synergies Appl. Environ. Microbiol. 70 5672 - 5678    DOI : 10.1128/AEM.70.9.5672-5678.2004
Liu N. , Chen X. G. , Park H. J. , Liu C. G. , Liu C. S. , Meng X. H. , Yu L. J. (2006) Effect of MW and concentration of chitosan on antibacterial activity of Escherichia coli Carbohydr. Polym. 64 60 - 65    DOI : 10.1016/j.carbpol.2005.10.028
Logue C. M. , Sheridan J. J. , Harrington D. (2005) Studies of steam decontamination of beef inoculated with Escherichia coli O157:H7 and its effect on subsequent storage J. Appl. Microbiol. 98 741 - 751    DOI : 10.1111/j.1365-2672.2004.02511.x
Mason T. L. , Wasserman B. P. (1987) Inactivation of red beet beta-glucan synthase by native and oxidized phenolic compounds Phytochem. 26 2197 - 2202    DOI : 10.1016/S0031-9422(00)84683-X
Mastromatteo M. , Incoronato A. L. , Conte A. , Del Nobile M. A. (2011) Shelf life of reduced pork back-fat content sausages as affected by antimicrobial compounds and modified atmosphere packaging Int. J. Food Microbiol. 150 1 - 7.    DOI : 10.1016/j.ijfoodmicro.2011.07.009
Mastromatteo M. , Lucera A. , Sinigaglia M. , Corbo M. R. (2010) Synergic antimicrobial activity of lysozyme, nisin, and EDTA against Listeria monocytogenes in ostrich meat patties J. Food Sci. 75 M422 - M429    DOI : 10.1111/j.1750-3841.2010.01732.x
Mohamed M. H. , Mansour H. A. (2012) Incorporating essential oils of marjoram and rosemary in the formulation of beef patties manufactured with mechanically deboned poultry meat to improve the lipid stability and sensory attributes LWT-Food Sci. Technol. 45 79 - 87    DOI : 10.1016/j.lwt.2011.07.031
Mohanka R. , Priyanka. (2014) Plant extract as natural food preservative against spoilage fungi from processed food Int. J. Curr. Microbiol. App. Sci. 3 91 - 98
Mor A. , Nicolas P. (1994) Isolation and structure of novel defensive peptides from frog skin Eur. J. Biochem. 219 145 - 154    DOI : 10.1111/j.1432-1033.1994.tb19924.x
Murdock C. A. , Cleveland J. , Matthews K. R. , Chikindas M. L. (2007) The synergistic effect of nisin and lactoferrin on the inhibition of Listeria monocytogenes and Escherichia coli O157:H7 Lett. Appl. Microbiol. 44 255 - 261    DOI : 10.1111/j.1472-765X.2006.02076.x
Naveena B. M. , Muthukumar M. , Sen A. R. , Babji Y. , Murthy T. R. K. (2006) Improvement of shelf life of buffalo meat using lactic acid, clove oil and vitamin C during retail display Meat Sci. 74 409 - 415    DOI : 10.1016/j.meatsci.2006.04.020
Nissen H. , Maugesten T. , Lea P. (2001) Survival and growth of Escherichia coli O157:H7, Yersinia enterocolitica and Salmonella enteritidis on decontaminated and untreated meat Meat Sci. 57 291 - 298    DOI : 10.1016/S0309-1740(00)00104-2
Nom M. J. R. , Rombouts F. M. (1992) Fermentative preservation of plant foods Appl. Bacterial Symp. Suppl. 73 1365 - 1478
Ntzimani A. G. , Giatrakou V. I. , Savvaidis I. N. (2010) Combined natural antimicrobials treatments (EDTA, lysozyme, rosemary and oregano oil) on semi cooked chicken meat stored in vacuum packages at 4℃: Microbiological and sensory evaluation Innov. Food Sci. Emerg. Technol. 11 187 - 196    DOI : 10.1016/j.ifset.2009.09.004
Okolocha E. C. , Ellerbroek L. (2005) The influence of acid and alkaline treatments on pathogens and the shelf life of poultry meat Food Control 16 217 - 225    DOI : 10.1016/j.foodcont.2004.01.015
Oliveira T. L. C. , Junior B. R. C. , Ramos A. L. S. , Ramos E. M. , Piccoli R. H. , Cristianini M. (2015) Phenolic carvacrol as a natural additive to improve the preservative effects of high pressure processing of low-sodium sliced vacuumpacked turkey breast ham LWT-Food Sci. Technol. 64 1297 - 1308    DOI : 10.1016/j.lwt.2015.06.011
Park C. B. , Lee J. H. , Park I. Y. , Kim M. S. , Kim S. C. (1997) Novel antimicrobial peptide from loach, Misgurnus anguillicaudatus FEMS Lett. 411 173 - 178
Pinto E. , Vale-Silva L. , Cavaleiro C. , Salgueiro L. (2009) Antifungal activity of the clove essential oil from Syzygium aromaticum on Candida, Aspergillus and dermatophyte species J. Med. Microbiol. 58 1454 - 1462    DOI : 10.1099/jmm.0.010538-0
Prange A. , Birzele B. , Hormes J. , Modrow H. (2005) Investigation of different human pathogenic and food contaminating bacteria and mould grown on selenite/selenate and tellurite/tellurate by X-ray absorption spectroscopy Food Control 16 713 - 728
Rico-Munoz E. , Eriotou E. , Davidson P. M. (1987) Effect of selected phenolic compounds on the membrane-bound adenosine triphosphate of Staphylococcus aureus Food Microbiol. 4 239 - 249    DOI : 10.1016/0740-0020(87)90006-2
Russell A. D. (1991) Mechanisms of bacterial resistance to nonantibiotics: Food additives and food and pharmaceutical preservatives J. Appl. Bacteriol. 71 191 - 201    DOI : 10.1111/j.1365-2672.1991.tb04447.x
Settanni L. , Corsetti A. (2008) Application of bacteriocins in vegetable food biopreservation Int. J. Food Microbiol. 121 123 - 138    DOI : 10.1016/j.ijfoodmicro.2007.09.001
Sharma S. (2015) Food preservatives and their harmful effects Int. J. Sci. Res. Pub. 5 1 - 2
Shi J. , Ross C. R. , Chengappa M. M. , Style M. J. , McVey D. S. , Blecha F. (1996) Antibacterial activity of a synthetic peptide (PR-26) derived from PR-39, a proline-argininerich neutrophil antimicrobial peptide Antimicrob. Agents Chemother. 40 115 - 121
Singh A. , Sharma P. K. , Garg G. (2010) Natural products as preservatives Int. J. Pharm. Bio Sci. 1 101 - 612
Stivarius M. R. , Pohlman F. W. , McElyea K. S. , Waldroup A. L. (2002) Effects of hot water and lactic acid treatment of beef trimmings prior to grinding on microbial, instrumental color and sensory properties of ground beef during display Meat Sci. 60 327 - 334    DOI : 10.1016/S0309-1740(01)00127-9
Tiwari B. K. , Valdramidis V. P. , O’Donnell C. P. , Muthukumarappan K. , Bourke P. , Cullen P. J. (2009) Application of natural antimicrobials for food preservation J. Agric. Food Chem. 57 5987 - 6000    DOI : 10.1021/jf900668n
Tossi A. , Sandri L. , Giangaspero A. (2000) Amphipathic, - helical antimicrobial peptides Biopolymers 55 4 - 30    DOI : 10.1002/1097-0282(2000)55:1<4::AID-BIP30>3.0.CO;2-M
Wang L. L. , Johnson E. A. (1997) Control of Listeria monocytogenes by monoglycerides in foods J. Food Prot. 60 131 - 138
Wu S. , Patel K. B. , Booth L. J. , Metcalf J. P. , Lin H. K. , Wu W. (2010) Protective essential oil attenuates influenza virus infection: an in vitro study in MDCK cells BMC Complement. Altern. Med. article 69 10
Zasloff M. , Martin B. , Chen H. C. (1988) Antimicrobial activity of synthetic magainin peptides and several analogues Proc. Natl. Acad. Sci. USA 85 910 - 913    DOI : 10.1073/pnas.85.3.910
Zasloff M. (2002) Antimicrobial peptides of multicellular organisms Nature 415 389 - 395    DOI : 10.1038/415389a