Food Research International 190 (2024) 114548

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Food Research International

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Review

Activity and safety evaluation of natural preservatives

Huiying Dong a, 1, Yang Xu a, 1, Qingqing Zhang a, Hua Li a, b, *, Lixia Chen a, **
a
Wuya College of Innovation, Key Laboratory of Structure-Based Drug Design & Discovery, Ministry of Education, Shenyang Pharmaceutical University, Shenyang
110016, China
b
Institute of Structural Pharmacology & TCM Chemical Biology, Fujian Key Laboratory of Chinese Materia Medica, College of Pharmacy, Fujian University of Traditional
Chinese Medicine, Fuzhou 350122, China

A R T I C L E I N F O A B S T R A C T

Keywords: Synthetic preservatives are widely used in the food industry to control spoilage and growth of pathogenic mi­
Safety evaluation croorganisms, inhibit lipid oxidation processes and extend the shelf life of food. However, synthetic preservatives
Food additives have some side effects that can lead to poisoning, cancer and other degenerative diseases. With the improvement
Essential oils
of living standards, people are developing safer natural preservatives to replace synthetic preservatives,
Tea polyphenols
Nisin
including plant derived preservatives (polyphenols, essential oils, flavonoids), animal derived preservatives
Natamycin (lysozyme, antimicrobial peptide, chitosan) and microorganism derived preservatives (nisin, natamycin,
Phage ε-polylysine, phage). These natural preservatives exert antibacterial effects by disrupting microbial cell wall/
membrane structures, interfering with DNA/RNA replication and transcription, and affecting protein synthesis
and metabolism. This review summarizes the natural bioactive compounds (polyphenols, flavonoids and ter­
penoids, etc.) in these preservatives, their antioxidant and antibacterial activities, and safety evaluation in
various products.

1. Introduction essential nutrient (Gutiérrez-del-Río et al., 2018).

Food preservatives are compounds that occur naturally in foods or
are added directly to them. They inhibit the growth of spoilage micro­
organisms, pathogens, and lipid oxidation, extend the shelf life of food
and ensure product safety (Mittal et al., 2023). Various preservatives on
the market, including sodium benzoate and sodium propionate, potas­
sium sorbate, sorbic acid, nitrite, sulfite, nisin, natamycin, potassium
lactate, ascorbic acid, citric acid, tartaric acid, etc., have been approved
by regulatory authorities for use as food preservatives (Kuorwel et al.,
2011). However, the excessive use of some preservatives poses a threat
to the nutrition or health of consumers. For example, sorbic acid can
combine with sulfhydryl bonds in many microbial cell enzymes, inhibit
cell growth and microbial reproduction, slow down the production of
mold and yeast, but also inhibit probiotics, resulting in intestinal flora
disorder, low immunity (Ismail et al., 2020). Excessive intake of nitrite
and nitrate can increase the risk of cardiovascular disease, and sodium
nitrite can be carcinogenic when processed at high temperatures. Sulfite
can cause the degradation of vitamin B1 (thiamine) in food, which is an
Currently, natural preservatives are a research focus and people
generally believe that they have beneficial properties and few side ef­
fects (Pateiro et al., 2021). As a substitute for synthetic compounds,
essential oils and other natural substances have been widely used in the
meat industry and have been approved by the Food and Drug Admin­
istration (FDA) for using in food (Food and Drug Administration, 2023).
However, their natural source does not necessarily mean that they are
safe to use. It is necessary to evaluate important aspects such as dosage,
efficacy, mechanism of action and toxicity. Sometimes it is necessary to
use them in higher concentrations to achieve the same effect as com­
mercial additives, which can have adverse effects on the product and
even toxicity problems. Therefore, it is necessary to consider combina­
tion with other preservatives to achieve synergistic effects allowing
better activity at lower concentrations. The application of natural pre­
servatives in food has been reported, but the summary of their safety
evaluation in terms of dosage and toxicity is not comprehensive enough.
This article provides a more comprehensive review of the activity,
application, and safety of natural preservative, the combined use of

* Corresponding author at: Wuya College of Innovation, Key Laboratory of Structure-Based Drug Design & Discovery, Ministry of Education, Shenyang Phar­
maceutical University, Shenyang 110016, China.
** Corresponding author.
E-mail addresses: lihua@fjtcm.edu.cn (H. Li), syzyclx@163.com (L. Chen).
1
These authors have the equal contribution to the article.

https://doi.org/10.1016/j.foodres.2024.114548
Received 8 December 2023; Received in revised form 29 February 2024; Accepted 25 May 2024
Available online 29 May 2024
0963-9969/© 2024 Elsevier Ltd. All rights are reserved, including those for text and data mining, AI training, and similar technologies.
H. Dong et al.
multiple preservatives, and the combined production of preservatives
with other substances. The safety assessment of natural preservatives is
shown in Fig. 1.
2. Plant derived preservatives
2.1. Essential oils and their anti-corrosion activity
Essential oil (EO) is a lipid soluble secondary metabolite extracted
from plants. Many essential oils have a long history of use as natural
antimicrobials, health promoters, and food flavorings. EOs are
composed of hydrocarbons (monoterpenes and sesquiterpenes) and
oxygenated compounds (alcohols, esters, ethers, aldehydes, ketones,
lactones, phenols, and phenol ethers) (Da Silva et al., 2021). The anti­
bacterial activity of EOs is driven by complex interactions between these
components. EOs that contain certain aldehydes or phenols as their main
components have the highest antibacterial activity (Gurtler & Garner,
2022).
The antibacterial mechanism of EOs is shown in Fig. 2, essential oils
exert antibacterial effects by acting on the cell membrane of microor­
ganisms, disrupting the integrity of the cell membrane. The hydropho­
bicity of essential oils allows them to cross the cell cytoplasmic
membrane and mitochondria, increasing membrane permeability,
causing leakage of critical components in cells and eventual death of the
bacteria cells (Falleh et al., 2020). Several reports have suggested that
gram-positive bacteria are more sensitive to antibacterial compounds
compared with gram-negative bacteria. This reason may be due to
absence of lipopolysaccharide in cell wall of gram-positive bacteria
which prevented the entry of active compounds into the cytoplasmic
Fig. 1. The safety evaluation of natural preservatives.
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Food Research International 190 (2024) 114548
membrane (Rashidaie Abandansarie et al., 2019). Studies have shown
that beef patties containing copaiba essential oil (CEO) and oregano
essential oil (OEO) exhibit good antioxidant activity and reduce lipid
oxidation, and their antioxidant effect is stronger than synthetic additive
(Leite et al., 2022). The antibacterial effects of CEO and OEO are largely
attributed to their chemical components. As bioactive compounds with
antioxidant properties, polyphenols can provide a hydrogen atom to free
radicals and form a stable free radical intermediate, thereby delaying or
inhibiting lipid oxidation. The antibacterial and antioxidant effects
depend on the number and position of the hydroxyl groups (Martinengo
et al., 2021). OEO has the effect of inhibiting Enterococcus faecalis. OEO
induces an increase in reactive oxygen species levels, resulting in lipid
peroxidation of the cell membrane, damaging the permeability and
integrity of the membrane, thereby changing the shape of the bacteria
and inactivating E. faecalis. OEO can be used as a natural antibacterial
preservative to control E. faecalis in food production (Zhan et al., 2022).
OEO has been shown to cause potassium and phosphate leakage in
Staphylococcus aureus and Pseudomonas aeruginosa, leading to cell death
(Lambert et al., 2001). CEO had an obvious oxidative damaging effect on
Salmonella enteritidis, CEO treatment caused the disruption of the bal­
ance of the cellular antioxidant enzymes (SOD, CAT, POD) system,
resulting in an increase in the content of malondialdehyde, a membrane
lipid metabolite, and increased protein carbonylation, which ultimately
inhibited the growth of S. enteritidis (Zhang et al., 2022). The inhibitory
effect of cinnamon essential oil on the growth of S. enteritidis cells is also
related to cell membrane damage and the induction of oxidative stress,
and its antibacterial activity is related to phenolic substances. The
antioxidant effect of rosemary is also related to the content of phenolic
components (carnitine, rosemary phenol, and rosemary diphenol) as
H. Dong et al.
Fig. 2. The antibacterial mechanism of essential oils.
hydrogen donors (Kamel et al., 2022).
Garlic essential oil also has good antibacterial effects and inhibits the
growth of Trametes hirsuta and Laetiporus sulphureus. The most biologi­
cally active are organic sulfur compounds and their precursors, such as
allicin, ajoene, and diallyl disulfide (Gong et al., 2021). Among them,
sulfur compounds react with cysteine and inhibit mercaptan enzymes
(protease and alcohol dehydrogenase) or lipid metabolism. It also in­
hibits microbial respiration, alters RNA synthesis, and increases the
permeability of cell membranes, leading to ion leakage and cell death.
Diallyl trisulfide is the most effective component in inhibiting T. hirsuta
and L. sulphureus, and can be used for preventing food spoilage (Pranoto
et al., 2005). Some studies showed that EOs extracted from Rosmarinus
officinalis, Thymus capitatus, Origanum majorana, and Salvia officinalis,
which inhibited the survival, growth, and reproduction of Staphylo­
coccus aureus, Escherichia coli, and Candida albicans in fresh fruit juices,
indicating that these essential oils have antibacterial effects (El-Kased &
El-Kersh, 2022). Rosemary-magnolia composite essential oil was found
to have a significant inhibitory effect on Staphylococcus aureus, and its
antioxidant effect is attributed to the high content of β-pinene (Song
et al., 2023). Pectin and gelling biopolymer film rich in natamycin and
clove essential oil have an antibacterial effect against Staphylococcus
aureus and Candida parapsilosis in the packaging corn tortilla (Callejas-
Quijada et al., 2023).
The antibacterial activity of essential oils such as CEO, OEO, cinna­
mon essential oil, and rosemary essential oil may be related to the al­
dehydes and alcohols in their chemical components. The high antifungal
activity of garlic essential oil is due to the presence of sulfur ethers, of
which diallyl trisulfide (allicin) has the highest content. Essential oils
have a good antibacterial effect on Gram-positive bacteria, Staphylo­
coccus aureus is a common Gram-positive bacterium and also a food­
borne pathogen, most essential oils have a significant inhibitory effect
on Staphylococcus aureus, and their antibacterial activity is dose-
dependent. Some essential oils even have better antimicrobial effects
than some chemical preservatives. However, natural preservatives from
plants have many negative effects on food and have not yet been widely
used. There are several main issues that need further research: although
the biological characteristics of EOs have been extensively studied, few
studies have examined their toxicity and side effects. Some in vitro and in
vivo experiments have confirmed that EOs can be toxic even at low
concentrations, which can lead to health problems, such as skin, eye,
and mucosal irritation, and also cause allergic reactions when taken
orally (Lanzerstorfer, 2021). Research has shown that the essential oils
in Dysphania can affect mitochondrial function, related to the presence
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Food Research International 190 (2024) 114548
of carvacrol, caryophyllene oxide, and ascaridole, which induce sup­
pression of the respiratory function in the mitochondria, or in the
complex I of the mitochondrial electron transport chains, this toxic ef­
fect emerging on the kidneys, liver, and intestine (Dagni et al., 2022).
α-Pinene can induce cell death by regulating signaling pathways related
to oxidative stress (Jin et al., 2010). Rosemary essential oils (REOs) rich
in α-pinene are toxic to HaCaT and this toxicity is strongly related to the
α-pinene content (Huang et al., 2023). During the production process,
attention should be paid to the concentration and safety of REO with
high α-pinene content in food packaging. There are few experiments
evaluating the safety of EOs in humans, but some compounds in essen­
tial oils can be toxic and cause side effects. The lack of in vivo/in vitro
research on essential oils is one of the limiting factors for their appli­
cation. Therefore, well-designed studies in animals and humans are
demanded to assess efficacy and safety of EOs, and it is recommended to
conduct detailed toxicological assessments for each EOs and intended
application Due to the complex mixture of many volatile components in
essential oils, it is difficult to standardize the application amount of
essential oil preservatives for optimal in vitro effects in food preserva­
tion. Essential oils have a certain impact on the texture and sensory
properties of food themselves, and when researching or applying them,
it is necessary to comprehensively consider the preservative effect and
food quality. In addition, it has been found that essential oils can cause
skin irritation and allergies in some cases.
2.2. Flavonoids and their antimicrobial activity
Flavonoids are widely found in natural organic compounds and have
various pharmacological activities, including significant antibacterial
effects. Flavonoids belong to the category of polyphenols, and their
highly hydroxylated structures give them strong antioxidant properties,
which can eliminate free radicals and inhibit bacteria. Some studies
have shown that flavonoids with good hydrophilicity have better anti­
bacterial effects than flavonoids with poor hydrophilicity. The position
of free hydroxyl groups in the molecule is also important for activity,
and all flavonoid aglycones have better antibacterial and antifungal
activity than flavonoid glycosides (Rashed et al., 2014).
The flavone extract in licorice can destroy the cell membrane and
cause the contents to leak out. After the extract enters the cell, it can
disrupt the normal metabolism and physiological activities of the cell by
destroying the structure of macromolecular substances or inhibiting
protein synthesis, thereby achieving its bacteriostatic effect (Ahn et al.,
2014). Naringin is a natural flavonoid extracted from grapes and citrus
H. Dong et al.
fruits (Peng et al., 2024), which has an inhibitory effect on Pseudomonas
spp., inhibiting bacterial biofilm and clearing preformed biofilms. It has
a strong binding affinity for biofilm related proteins, thereby inhibiting
or reducing bacterial colonization. Bacteria associated with food
spoilage form biofilms on nonbiological surfaces, and cells with biofilms
exhibited greater resistance to antimicrobial agents. Extracellular
polysaccharides help maintain the structure of biofilms and protect them
from harmful environmental influences. Alginate is also crucial for the
biofilm of Pseudomonas spp., as it has a dual effect of protecting cells
from adverse external conditions and promoting bacterial adhesion to
the substrate. Naringin can effectively inhibit the production of extra­
cellular polysaccharides and alginate in Pseudomonas spp. (Husain et al.,
2021). The study found that luteolin has antibacterial activity against
Escherichia coli and Enterobacter cloacae. Luteolin can destroy the
integrity of the cell membrane and has a strong inhibitory effect on the
biofilm formation. The number and degree of cell damage are dose-
dependent (Qian et al., 2021).
Prescribed doses of flavonoids, such as those found in our daily diet,
may be safe for human health. Research has shown that dietary intake of
flavonoids is approximately 50–800 mg/day with no associated side
effects (Wu et al., 2023). Chrysin is a flavone that can be obtained from
different natural sources, such as Alpinia oxyphylla, mushrooms (Pleu­
rotus ostreatus), passionflower, and Indian trumpet flower (Tekeli et al.,
2023). Acute dosages of 400 mg chrysin do not show any observable
toxic effects in humans (Naz, 2019). Normal volunteers have received an
oral dose of 625 mg/day of chrysin for 28 days without any reported
toxicity. Dihydromyricetin is a natural flavonoid compound extracted
from the stems and leaves of Ampelopsis grossedentata (Brown et al.,
2001), which was administered to rats at doses of 1.5 g/kg and 0.3 g/kg
for 12 weeks without significant toxic effects (Wu et al., 2023). Senna
italica leaves contain various flavonoids such as kaempferol, quercetin
and apigenin, and it had no toxic effects or death in rats administered a
dose of 5000 mg/kg, indicating that they are non-toxic at the prescribed
dose (Towanou et al., 2023).
Some flavonoid compounds exhibit direct antibacterial effects by
destroying cell membranes, inhibiting nucleic acid synthesis, inhibiting
energy metabolism, and inhibiting cell wall and membrane synthesis.
They can destroy a variety of biofilm cells, inhibit the growth of various
bacteria, and play an important role in food preservation. Future
research should be conducted on a comprehensive assessment of the
safety and potential of flavonoids in conjunction with in vitro and in vivo
toxicity assessments. The food industry should pay attention to the
dosage of each type of flavonoids to avoid toxic situations. Although
flavonoids have high antimicrobial activity, they have problems such as
poor water solubility, poor stability and low bioavailability. In order to
fully exploit the antimicrobial effect of flavonoids, it is worth examining
their compatibility with other components (Han et al., 2022). At present,
protein–flavonoid complexes are widely used to improve the poor water
solubility and low bioavailability of flavonoids, and have stronger
oxidation capacity than flavonoids (Diao et al., 2021). With the in-depth
study of flavonoid structure and activity as well as their antibacterial
mechanisms, this will expand new space for the development of anti­
bacterial flavonoids.
2.3. Tea polyphenols and their antimicrobial activity
Tea polyphenol is the most important component of tea. Due to its
enzyme inhibition as well as antibacterial and antioxidant effects, it is
widely used in various foods such as meat, aquatic products, fruits and
vegetables (Dai et al., 2022). Tea polyphenol has potent antibacterial
and antioxidant properties, and studies have examined the antioxidant
activity of black tea. The results show that its antioxidant activity is
positively correlated with the content of phenolic hydroxyl groups
(Chen et al., 2023). Tea polyphenol contains potent antibacterial and
antioxidant compounds, particularly catechin compounds, including
(− )-epicatechin (EC), (− )-epigallocatechin (EGC), (− )-epicatechin
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Food Research International 190 (2024) 114548
gallate (ECG), and (− )-epigallocatechin gallate (EGCG) (Jia et al.,
2018). EGCG is the main component of green tea polyphenols, which is
widely used as food preservatives.
Tea polyphenol can be used to extend the shelf life of Pacific white
shrimp. They can inhibit the activity of peroxidase and protein enzyme
by attacking oxygen and free radicals and scavenging free radicals, such
as α-amylase, pepsin, trypsin, and lipase, and prevent lipid oxidation,
reduce the activity of enzymes in frozen Pacific white shrimp, and delay
lipid oxidation (Li et al., 2022). Tea polyphenol also plays an important
role in preserving fish, adding tea polyphenol can prolong the preser­
vation of fish by inhibiting lipid oxidation (Xu et al., 2023). Adding tea
polyphenol to the polysaccharide composite film can effectively
improve the physical properties and antibacterial/antioxidant functions
of the composite film. The films of sodium alginate and konjac gluco­
mannan mixed by Lycium ruthenicum anthocyanins and tea polyphenol
(TP-LRA-SA-KGM) exhibit a good preservative effect on milk and can
effectively inhibit the growth of bacteria in milk. LRA shows a signifi­
cant inhibitory effect on the proliferation of Staphylococcus aureus and
Escherichia coli. By adding tea polyphenol, the inhibitory effect on bac­
terial proliferation is significantly enhanced (Wang et al., 2023). The TP-
SA-KGM films have a good preservative effect on beef and apples by
effectively inhibiting microbial activity and food oxidation, thereby
significantly delaying the process of food spoilage. The TP-SA-KGM films
can be used as a natural bioactive film to replace plastic films for beef
preservation (Wang et al., 2023).
Tea polyphenols can inhibit most microorganisms, with a broad
antibacterial spectrum, and inhibit most food-borne pathogens (Li et al.,
2020), such as Costridium botulinum, Pseudomonas aeruginosa, Escherichia
coli, Salmonella typhimurium, Listeria monocytogenes, Staphylococcus
aureus and Candida albicans. Its main inhibitory effect is to destroy the
cell membrane, increase the permeability of the outer and inner mem­
branes of Pseudomonas aeruginosa, and destroy the cell membrane by
releasing of small cell molecules (Tang et al., 2022). Due to their anti­
bacterial properties, tea polyphenols have an inhibitory effect on pu­
trescine, cadaverine and histamine. Tea polyphenols retard spoilage
bacteria and inhibit bacterial growth (Xu et al., 2023).
More and more studies have shown that excessive consumption of tea
polyphenols can cause several harmful effects (Molinari et al., 2006.).
High doses of tea polyphenols not only cause cytotoxicity in vitro, but
also lead to hepatotoxicity, nephrotoxicity, and gastrointestinal disor­
ders (Keller & Wallace, 2021). When tea polyphenols reach high con­
centrations in the blood, they trigger cytotoxicity and oxidative stress. It
has been found that high levels of catechins can increase the carcino­
genic effects caused by carcinogens (Akagawa et al., 2003). Studies have
shown that EGCG has a protective effect on DNA at low concentrations,
but at higher concentrations, it exacerbates DNA oxidative damage
(Guan et al., 2012). Under the prescribed safe dose of tea polyphenols,
chronic and subchronic experiments were conducted on dogs for 9
months and 13 weeks, and no side effects were observed (Kapetanovic
et al., 2010). Taking tea polyphenols at the optimal dosage showed no
side effects on humans (Rashidinejad, 2021). The European Food Safety
Authority believes that intake of 800 mg/day EGCG may lead to an in­
crease in transaminases (Ouyang et al., 2020). Tea polyphenols can play
a preservative role in food, with a positive effect on the body and
maintaining redox homeostasis in the body. However, exceeding a
certain limit can cause damage to the body. In the food industry, when
using tea polyphenols, the dosage should be strictly controlled, and its
toxicity should be reduced as much as possible while exerting its anti­
microbial effect.
Many plant extracts, such as cinnamon extract, clove extract, rose­
mary extract, and garlic extract, have strong antioxidant and bacteri­
cidal effects and can be used as natural preservatives. Essential oils are
often used as seasonings and food additives, they are also used in the
cosmetic industry. Tea polyphenols also have excellent antimicrobial
and preservative effects and are widely used in food and medicine. An
overview of natural antimicrobial from plant sources is shown in
H. Dong et al. Food Research International 190 (2024) 114548

Table 1 Table 1 (continued )

The antibacterial effect of natural preservatives. Preservatives Microorganism References
Preservatives Microorganism References
Escherichia faecalis, Collins et al.
Essential oil Copaiba essential oil Enterococcus Leite et al. (2022) Staphylococcus (2019)
and oregano essential faecalis, aureus,
oil Staphylococcus Paenibacilluslarvae
aureus, spp.,
Pseudomonas Salmonella
aeruginosa, typhimurium,
Salmonella enteritidis Bacillus subtilis
Protamine Shewanella Li et al. (2016).
Cinnamon essential Salmonella enteritidis Kamel et al. putrefaciens,
oil (2022) Pseudomonas spp.
Garlic essential oil Trametes hirsuta, Gong et al. Bacteriocin Nisin Staphylococcus Wu et al. (2023)
Lactobacillus sulfate (2021) aureus,
Essential oils of Staphylococcus El-Kased and El- Escherichia coli
rosemary, thymus aureus, Kersh (2022) KCA018 bacteriocin Listeria Kim et al. (2022)
gland, oregano and Escherichia coli, monocytogenes
salvia miltiorrhiza Candida albicans Bacteriocin LFX01 Staphylococcus Xin et al. (2023)
Flavonoids Pseudomonas spp. Husain et al. aureus,
(2021) Escherichia coli
Escherichia coli, Qian et al. (2021) Lacticaseibacillus Staphylococcus Wu et al. (2022)
Enterobacteriaceae rhamnosus ZFM216 aureus
cloacae bacteriocin
Tea polyphenol Costridium Tang et al. (2022) Actinomycin Natamycin Candida tropicalis Agustín et al.
botulinum, Aspergillus (2019)
Pseudomonas ochraceopealiformis
aeruginosa, Polylysine Penicillium Bai et al. (2022)
Escherichia coli, expansum, Cai and Kuo
Salmonella Colletotrichum (2022)Zhu et al.
typhimurium, gloeosporioides, (2022)
Listeria Gloeophyllum
monocytogenes, trabeum,
Staphylococcus Rhodonia placenta,
aureus, Trametes versicolor,
Candida albicans Irpex lacteus,
Chitosan Escherichia coli Volmajer Valh Staphylococcus
O157:H7, et al. (2022)Díaz- aureus,
Listeria Cruz et al. (2022) Escherichia coli,
monocytogenes, Listeria
Staphylococcus monocytogenes,
aureus, Pseudomonas
Shewanella lundensis
putrefaciens, Phage Lytic bacteriophage Vibrio You et al. (2021)
Pseudomonas VPT02 parahaemolyticus
fluorescens Polyvalent phage Coagulase-negative Sofy et al. (2020)
Lysozyme Listeria Ajourloo et al. CoNShP-3 staphylococci
monocytogenes, (2021)Xu et al. Staphylococcus
Shewanella (2020) aureus,
putrefaciens, Staphylococcus
Pseudomonas aureus,
fluorescens, Bacillus cereus,
Staphylococcus Bacillus subtilis
aureus, Phage UAE_MI-01 Escherichia coli Sultan-Alolama
Escherichia coli, et al. (2021)
Pseudomonas Bacillus phage Klebsiella Sarjoughian et al.
aeruginosa endolysin pneumoniae, (2020)
Antimicrobial peptide Escherichia coli, Cournoyer et al. Salmonella
Listeria ivanovii, (2022)Abou-Diab typhimurium,
Mucor racemosus, et al. (2022)Shen Proteus,
Penicillium et al. (2022) Escherichia coli
chrysogenum,
Aspergillus versicolor,
Paecilomyces,
Table 1.
Rhodotorula
mucilaginosa,
Saccharomyces 3. Animal derived preservatives
boulardii,
Kluyveromyces
3.1. Chitosan and their antimicrobial activity
marxianus,
Candida
guilliermondii, Chitosan, a natural marine polysaccharide, consists of 1,4-linked
Salmonella glucosamine and N-acetylglucosamine, which is derived through
typhimurium deacetylation of chitin in alkaline medium (Mondéjar-López et al.,
Propolis Aspergillus spp., Vasilaki et al.
2022). Chitosan can be extracted from the cell walls of fungi, the shells
Macrospora spp., (2019)Postali
Listeria et al. (2022) of crustaceans, and the stratum corneum of insects (Yi et al., 2024).
monocytogenes, Chitosan is widely used in biomaterials, healthcare, environmental, and
other industrial fields due to its good biocompatibility, antibacterial

5
H. Dong et al.
activity, and safety (Ruan et al., 2022).
Hummus is a food paste commonly consumed in the Middle East. Its
major ingredients include boiled chickpeas and tahini pulp. Chitosan has
extensive antimicrobial properties and the addition of chitosan to
hummus can result in a significant reduction in the number of Escher­
ichia coli O157:H7 and Listeria monocytogenes (Osaili et al., 2022). Chi­
tosan coating can be used as an antibacterial agent and extend the shelf
life of packaged foods. Studies have shown that chitosan-coated pack­
aging materials can reduce the number of Staphylococcus aureus and
Escherichia coli, extending the shelf life of food (Volmajer Valh et al.,
2022). The inhibitory effect of chitosan in the corn starch-chitosan
composite film was confirmed on both Listeria monocytogenes and
Staphylococcus aureus (Díaz-Cruz et al., 2022). Chitosan can be used to
preserve fruits during storage, it can inhibit the production of ethylene,
delay the aging of fruits, and does not affect the taste or appearance of
food (Liu et al., 2021). Chitosan treatment was found to trigger anti­
oxidant systems, thereby scavenging reactive oxygen species (ROS) and
ensuring the integrity of the fruit membrane (Ackah et al., 2022).
Chitosan shows a stronger inhibitory effect at lower pH and its
inhibitory effect decreases with the increase of pH. To improve anti­
bacterial activity, complexes of chitosan and certain materials were
prepared (Jana & Jana, 2019). Chitosan-nano-zinc oxide composite
films showed a good preservative effect on cherry tomatoes. The coating
formed a barrier on the fruit surface, which inhibited to a certain extent
the respiration intensity and growth of surface microorganisms, slowed
down the aging process and nutrient consumption, and had a preser­
vative effect (Li et al., 2021). Chitosan and gelatin form an edible
nanocomposite film that has antibacterial effects on Escherichia coli and
Staphylococcus aureus. It can be used to protect fruits and slow spoilage
(Fu et al., 2021). Due to the low antioxidant activity and weak water
resistance of chitosan, grafting phenol derivatives onto chitosan mac­
romolecules can effectively improve its antioxidant and hydrophobic
properties. It was also studied that lauryl gallate was grafted onto chi­
tosan, which significantly increased the oxidation resistance, hydro­
phobicity and moisture resistance (Liu et al., 2021). Chitosan combined
with apple polyphenols coating can effectively inhibit the growth of
microorganisms, delay the biochemical changes of ice stored large yel­
low croaker and extend its shelf life (Lan et al., 2022). The antioxidant
and antibacterial activities of chitosan could be enhanced by grafting
caffeic acid onto chitosan, which could inhibit microbial growth, reduce
lipid oxidation and protein degradation, and was suitable for extending
Fig. 3. Reaction mechanism for the synthesis of caffeic acid-g-chitosan by EDC/NHS chemical coupling.
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Food Research International 190 (2024) 114548
the shelf life of fish and meat. The reaction mechanism for the synthesis
of CS-g-CA by EDC/NHS is shown in Fig. 3 (Sun et al., 2022). Chitosan
and flaxseed gum form flaxseed gum/chitosan/flaxseed gum composite
coating, which has high total antioxidant capacity and DPPH scavenging
capacity. They have good antibacterial properties against Shewanella
putrefaciens and Pseudomonas fluorescens. The composite coating slows
the reduction of antioxidant enzyme activity and inhibits protein
oxidation in rainbow trout fillets, which can be used for food preser­
vation (Yang et al., 2022).
Chitosan is biocompatible and has been approved by the FDA as
generally recognized as safe (Khubiev et al., 2023). A series of toxico­
logical tests have been conducted with chitosan, including acute toxicity
(Liang et al., 2011), subacute toxicity (Sonawnae et al., 2012), chronic
toxicity, chromosomal malformation test, embryonic toxicity and tera­
togenic test (Khan et al., 2023), indicating that chitosan is non-toxic to
humans. Due to the lower antibacterial activity of chitosan under
neutral and alkaline conditions compared to acidic environmental
conditions, as well as its low antioxidant activity and water resistance,
the application of chitosan in food preservation is limited. Therefore,
materials modified with chitosan are widely used, mainly for fruit
preservation. They form a barrier on the fruit surface, limit the spread of
oxygen and extend the shelf life of the fruit. The antibacterial chitosan
film has great prospects as a widely used and environmentally friendly
material. Their unique properties, including biodegradability, biocom­
patibility and antibacterial activity, have been extensively studied for
food packaging applications to extend shelf life, maintain quality and
reduce spoilage due to microbial contamination.
3.2. Lysozyme and their antimicrobial activity
Lysozyme, also known as 1,4-β-D-N-acetyl cell wall amidase is an
alkaline enzyme that can hydrolyze mucopolysaccharides in bacteria.
This enzyme is widely present in various tissues of the human body and
is the main component of animal products (egg white, egg white lyso­
zyme). Microorganisms also contain this enzyme, with egg white being
the most abundant. Lysozymes have good antibacterial activity, espe­
cially against Gram-positive bacteria (Alhadrami et al., 2022). Gram-
positive bacteria have a single lipid membrane surrounded by a thick
layer of peptidoglycan cell wall. In Gram-negative bacteria, the cell wall
consists of a thin layer of peptidoglycan between the cytoplasm and the
outer lipid membrane. Lysozymes can hydrolyze peptidoglycans, which
H. Dong et al.
are the main components of the cell wall of Gram-positive bacteria. This
hydrolysis destroys the integrity of the cell wall and leads to lysis of the
bacteria (Primo et al., 2018). The antibacterial effect of lysozyme is
mediated by its cell wall amidase activity. It catalyzes the reaction be­
tween the 1-carbon atom of N-acetylmuramic acid (NAM) and the 4-car­
bon atom of N-acetylglucosamine (NAG) in the peptidoglycan layer of
the cell wall of Gram-positive bacteria β-1,4 hydrolysis of glycosidic
bonds. Lysozyme molecules are spherical structures with hydrophilic
groups on the outside and hydrophobic groups on the inside. There is
also a groove structure on the surface that serves as an active site for
binding and cutting bacteria. Polysaccharides consisting of six amino
sugars are used as binding substrates for active localization through
hydrogen bonding and hydrophobic interaction (Leśnierowski & Yang,
2021). Acid catalyst residues such as glutamic acid (Glu) and general
alkaline catalyst residues such as aspartic acid (Asp) or cysteine (Cys)
destroy the carbon–oxygen bonds between the fourth and fifth sugar
units, resulting in an increase in their glycosidic bonds. Glutamic acid
acts as a proton donor via free carbon groups on its side chains, while
aspartic acid as a nucleophilic reagent to form glycosyl enzyme in­
termediates that react with water molecules to produce hydrolysis
products (Fig. 4) (Nawaz et al., 2022). Lysozyme is a heat-stable enzyme
and can therefore be used to extend the shelf life and improve the safety
of various foods.
Although lysozyme has strong antibacterial activity against Gram-
positive bacteria, this ability is weaker in Gram-negative bacteria due
to the presence of an outer membrane to protect the cell wall. This
limitation can be eliminated by combining it with other preservatives
that can increase the antibacterial spectrum (Moshtaghi et al., 2018).
The combination of epigallocatechin gallate and lysozyme has a good
inhibitory effect on Pseudomonas spp., can extend the shelf life of large
yellow croakers (Pei et al., 2022). Ziziphora clinopodioides essential oil
(ZCEO) and lysozyme showed good antibacterial and antioxidant effects
against Listeria monocytogenes and could be used as a preservative in
fresh sausage (Ajourloo et al., 2021). The combined treatment with
lysozyme and cinnamaldehyde can significantly inhibit the growth of
microorganisms in refrigerated olive flounder (Paralichthys olivaceus),
Fig. 4. Hydrolytic mechanism of action of lysozyme on β-1,4 linkages between NAM and NAG residues of the bacterial cell wall backbone.
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enhance the antibacterial activity against Shewanella putrefaciens and
Pseudomonas fluorescens, delay lipid oxidation, and could be used for
preserving fish fillets (Xu et al., 2020). The chitosan-lysozyme coating
has a good antibacterial effect against Gram-positive bacteria (Staphy­
lococcus aureus) and Gram-negative bacteria (Escherichia coli and Pseu­
domonas aeruginosa), which can inhibit the lipid oxidation of
Pseudomonas aeruginosa, thus extending the shelf life of yellow croaker.
(Wu et al., 2018).
Lysozyme, as a natural antibacterial substance, can target the cell
wall of the cell without affecting other parts of the cell, and can be
digested and absorbed in the stomach without affecting human health.
Lysozyme generally does not affect the nutritional components in cold
and fresh meat and can be used to preserve chilled meat. Lysozyme is
widely used as a safe and effective preservative in dairy products, meat,
aquatic products and other foods. Lysozyme from egg white has been
reported to have insignificant acute, subacute and chronic toxicity in
animals. However, lysozyme has the problems of anaphylaxis and an­
tigenicity in the digestive process. Studies have shown that individuals
who are sensitive to eggs have no side effects or immune responses after
consuming cheese containing lysozyme (Rossi et al., 2012). Egg white
lysozyme has been declared as Generally Recognized As Safe (GRAS) by
the FDA for use in cheese. Due to the allergenic potential of lysozyme,
the FDA has concluded that “Egg white lysozyme” labeling is mandatory
for bulk and packaged foods containing lysozyme(Food and Drug
Administration, 2023) . In response to some of the problems that exist
with lysozyme, further research and investigation into the application of
lysozyme in the food industry is needed. Although the scope of appli­
cation of lysozyme is still relatively limited, it is expected to be an
important functional food additive for the food industry.
3.3. Antimicrobial peptide and their antimicrobial activity
Antimicrobial peptide (AMP) is a low molecular weight polypeptide
or protein synthesized by ribosomes and composed of 10–50 amino acid
residues (León Madrazo et al., 2022). Most AMPs are cationic peptides
with cationic and amphiphilic structures. The cationic structure
H. Dong et al.
electrostatically interacts with the negatively charged cell membrane,
destroying the integrity of the membrane and making it impossible for
bacteria to survive (Zhao et al., 2022). The amphiphilicity of AMP
contributes to AMP entering the cell membrane, destroying the lipo­
polysaccharide layer of the cell membrane, and leading to the death of
bacteria (Dong et al., 2022).
The small molecule peptide porcine myeloid antimicrobial peptide
37 (PMAP-37) isolated from pig bone marrow has broad-spectrum
antibacterial activity. The peptide has been shown to inhibit the activ­
ity of Gram-positive and Gram-negative bacteria. The interaction be­
tween positively charged PMAP-37 and negatively charged bacterial cell
membranes disrupted the integrity of the cell membrane and resulted in
leakage of bacterial contents (Dong et al., 2023). Antimicrobial peptides
can be produced from porcine hemoglobin after pepsin hydrolysis. Pork
hydrolysates have been shown to exhibit antibacterial activity,
including bacteria (Gram-positive and Gram-negative bacteria), fila­
mentous molds and yeasts (Cournoyer et al., 2022). Similar to porcine
hemoglobin, bovine hemoglobin hydrolysate also has significant anti­
bacterial activity against many bacterial strains (Gram-positive and
Gram-negative bacteria) and fungal strains (mold and yeast) (Abou-Diab
et al., 2022). Shen et al. designed and synthesized four antimicrobial
peptides that have an inhibitory effect against Salmonella typhimurium,
preventing the formation of biofilms, interacting with the bacterial cell
membrane and penetrating, resulting in loss of membrane potential,
damage to membrane integrity and bacterial death (Shen et al., 2022). It
can also bind to the DNA of Salmonella typhimurium, causing DNA ag­
gregation or precipitation. A peptide with a molecular weight of 9.77
kDa was isolated from the supernatant of Saccharomyces cerevisiae strain
ATCC 36858. It inhibited the growth of Escherichia coli and Staphylo­
coccus aureus (Thyab Gddoa Al-sahlany et al., 2020). Chitin has shown
antibacterial activity against Listeria monocytogenes, Staphylococcus
aureus, Enterococcus faecalis, Escherichia coli, and Salmonella enteritidis,
with the best antibacterial effect against Salmonella enteritidis in chilled
pork samples. Toxic effects of this antimicrobial peptide on human fi­
broblasts and red blood cells have not been reported, indicating its
safety for consumption (León Madrazo & Segura Campos, 2023). The
results of cytotoxicity tests of antimicrobial peptides show that they are
toxic only at higher concentrations (20–25 µM), while their toxicity is
very low at lower concentrations (<15 µM) (Arasu & Al-Dhabi, 2023).
Although antimicrobial peptides have good antibacterial potential, their
toxicity as food preservatives is rarely researched. It is hoped that future
research can focus more on the application of antimicrobial peptides in
food and maximize their antimicrobial effects within the allowable
range of toxicity.
Antibacterial peptides are widely distributed in organisms and have
broad-spectrum antibacterial properties, green and anti-pollution
properties, and even nutritional and health benefits. As a natural pre­
servative, it has achieved good results and is widely used in the field of
food preservation. However, due to the disadvantages of low yield,
complex manufacturing process, high cost, and the fact that antimicro­
bial peptides cannot be produced on a large scale, this limits their
commercialization process. Therefore, developing the manufacturing
process of antimicrobial peptides, increasing production, reducing costs,
and studying the details of their dynamic mechanism of action can
effectively promote the entry of antimicrobial peptides into the field of
food preservation and preservation.
3.4. Propolis and their antimicrobial activity
Propolis is a kind of bee product that contains a variety of organic
compounds and is a resinous material. It consists of tree and plant se­
cretions, pollen and enzymes collected by bees. The antibacterial
properties of propolis are attributed to its flavonoids and phenolic acids
(Sánchez-Martín et al., 2022).
Propolis has an antibacterial effect by increasing the permeability
between bacterial cell walls and cell membranes, causing the outflow of
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internal substances and disrupting the normal metabolism of bacteria,
thereby achieving an antibacterial effect. It can be used as a preservative
in food processing and aquaculture. Research has shown that propolis
extract has a stronger inhibitory effect against mold, particularly
Aspergillus spp. and Macrospora spp. (Vasilaki et al., 2019). Red propolis
hydroalcoholic extract can effectively delay the decay and lipid oxida­
tion of tilapia salami. It has obvious antibacterial activity against Gram-
positive bacteria and strong inhibitory activity against Listeria mono­
cytogenes and Escherichia faecalis (Mafra et al., 2022). Studies have
shown that propolis can inhibit the growth of Staphylococcus aureus and
Listeria monocytogenes (Postali et al., 2022). Caffeic acid esters are active
ingredients extracted from propolis. The results indicate that caffeic acid
esters can eliminate bacterial growth through oxidative stress mecha­
nisms and have inhibitory effects on Paenibacilluslarvae spp. (Collins
et al., 2019).
The combination of propolis and edible coatings can also be used for
meat preservation. Carboxymethyl cellulose containing 3 % ethanol
propolis extract can be coated on chicken breast meat and stored in cold
storage, which can extend the storage time of chicken breast meat (El
Sheikha et al., 2022). Propolis combined with other preservatives can
improve its antimicrobial effect as a composite material. The composite
material formed by chitosan nanoparticles, propolis and nano selenium,
as a potential antibacterial nano compound, was found to have a strong
antibacterial effect on Escherichia coli, Staphylococcus aureus and Sal­
monella typhimurium (Youssef et al., 2022). Propolis ethanol extract and
glycerin can significantly inhibit the growth of Listeria monocytogenes
when added to milk stored at 4 ◦ C (Michailidis et al., 2021). A sodium
alginate bilayer coating incorporated with green propolis extract can
reduce microbial deterioration and extend the shelf life of Colossoma
macropomum fillet (Cruz et al., 2021).
Propolis has an efficient and broad-spectrum bacteriostatic effect.
Propolis has significantly greater bacteriostatic ability to Gram-positive
bacteria than Gram-negative bacteria, and has a good antibacterial ef­
fect on Salmonella spp., Staphylococcus aureus, Enterococcus spp., Strep­
tococcus spp., Bacillus subtilis, and almost no bacteriostatic activity to
Escherichia coli. As the concentration of propolis alcohol extract in­
creases, the antibacterial effect becomes more apparent. A series of
toxicological tests have been conducted on Brazilian red propolis,
including cytotoxicity, genetic toxicity and in vivo genotoxicity, indi­
cating that it has no toxicity (Aldana-Mejía et al., 2021). With long-term
oral administration of 1000 mg/kg Brazilian red propolis to males,
biochemical changes in liver and kidney parameters were detected, but
no histopathological changes in any tissue or organ (Aldana-Mejía et al.,
2023). Clinical and in vivo animal studies have shown that propolis is
safe and non-toxic (Castaldo & Capasso, 2002). However, at higher
doses, some side effects and allergies may occur (Choudhary et al.,
2023). The main sensitizers in propolis are 3-methyl-2-butenyl caffeic
acid ester, phenylethyl caffeic acid ester, benzyl salicylate ester, benzyl
cinnamate ester, and 1,1-dimethylallyl caffeic acid (Burdock, 1998).
Propolis also has various biological activities and functional health
benefits, such as clearing reactive oxygen species and playing an anti­
oxidant role. Therefore, propolis has broad application prospects in food
preservation and preservation. At present, research has shown that
propolis has good effects on the preservation of poultry eggs, aquatic
products, meat products, dairy products, and fruits.
3.5. Protamine and their antimicrobial activity
Protamine is a cationic peptide rich in proline and arginine. It is an
alkaline protein that is tightly bound to DNA in the form of nuclear
protamine. At present, protamine has been extracted from mature testis
tissues of various fish species (such as salmon, red trout, herring, etc.)
and mammals (such as mice, humans, wild boars, etc.). In neutral and
alkaline media, protamine has a strong antibacterial ability and thermal
stability, a broad spectrum of antibacterial activity and a strong inhib­
itory effect on yeasts, molds and Gram-positive bacteria such as bacillus
H. Dong et al.
and lactic acid bacteria (Aspedon & Groisman, 1996).
Protamine can inhibit the metabolism process dependent on the cell
plasma membrane, target the cell plasma membrane by destroying en­
ergy and inhibiting the uptake of nutrients, and inhibit certain compo­
nents of the mitochondrial electron transfer system without causing cell
lysis or changing the permeability of the cell plasma membrane. The
mode of action of protamine is the electrostatic attraction between
positively charged molecules and negatively charged cell envelopes,
which leads to inhibition or cell death due to the leakage of K+, ATP and
intracellular enzymes (Truelstrup Hansen et al., 2001). The activity of
protamine is affected by the size of the electric membrane potential, and
its antibacterial effect can be reduced by low pH culture or treatment
with respiratory toxins that reduce membrane potential.
Protamine is widely used for fish preservation. Shewanella putrefa­
ciens is the main spoilage bacteria of grass carp slices during the whole
storage period, which may lead to the spoilage of fish stored in aerobic
condition under cold storage. Protamine can effectively inhibit the
growth of Shewanella putrefaciens, improve the quality of grass carp fillet
and extend its shelf life (Li et al., 2016). Protamine also has a strong
inhibitory effect on Pseudomonas spp., but has a weak inhibitory effect
on Psychrobacter, Acinetobacter and Brevibacterium (Duan et al., 2018).
Some studies have shown that salmon protamine can kill the growing
Gram-positive bacteria, and significantly inhibit the growth of Gram-
negative bacteria in Tryptone Soy Broth (Johansen et al., 1995). EDTA
and protamine have a synergistic effect. The combination of protamine
and EDTA enhances the inhibition of Gram-negative bacteria including
Aeromonas hydrophila, Morganella morganii and Yersinia enterocolitica).
The most sensitive organisms are Brochothrix thermosphacta and Lacto­
bacillus sakei (Truelstrup Hansen et al., 2001).
Protamine exhibits strong antibacterial activity in neutral and alka­
line media and has high thermal stability. It has a broad-spectrum
antimicrobial effect on bacteria, yeasts and molds, especially against
Gram-positive bacteria. As a food preservative, protamine has the ad­
vantages of broad-spectrum antibacterial activity, high safety, good
thermal stability, and no smell or taste, making it a promising natural
preservative. Protamine is very effective in extending the shelf life of fish
cake products and flour and rice foods. Combining fish sperm with other
natural preservatives can enhance the antibacterial effect and expand
the scope of food preservation applications. Compared to other natural
preservatives, protamine is relatively expensive and less commonly used
in meat products. Currently, the main research focuses on the prepara­
tion of protamine and its preparation into nanocapsules. So far, no toxic
incidents have occurred to the human body when using protamine.
Studies on the safety of protamine have shown that the protamine LD50
is above 10 g/kg. According to the toxicity classification of chemical
substances, protamine can be considered a non-toxic substance (Soko­
lowska et al., 2016). No teratogenic or mutagenic effects on the chro­
mosomes of primary spermatocytes in the testis of mice were detected.
Through various toxicological experiments, it has been proven that
protamine is actually non-toxic, has no chromosomal aberration effect,
and has no genetic mutation effect. Therefore, protamine can be
considered as a new type of food preservative, which is harmless, highly
safe, and highly nutritious to the human body (Lee et al., 2023). An
overview of natural antimicrobial from animal sources is shown in
Table 1.
4. Microorganism derived preservatives
4.1. Bacteriocin and their antimicrobial activity
Bacteriocin is a safe and effective antibacterial peptide synthesized
by ribosomes during bacterial metabolism (Jiang et al., 2022). The study
found that Lactococcus lactis can inhibit pathogenic and putrefactive
bacteria, including 10 Gram-positive bacteria and 16 Gram-negative
bacteria. The hemolytic activity test of bacteriocin showed that it has
no toxicity to eukaryotic cells (Fotso Techeu et al., 2022).
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Nisin is a bacteriocin produced by lactic acid bacteria. Nisin can kill
Gram-positive bacteria and bacterial spores, but not Gram-negative
bacteria, yeasts, molds, and other fungi. Nisin has no toxic effect on
the human body and is very sensitive to proteolytic enzymes (pancre­
atin, salivary enzyme, digestive enzyme, etc.)(Patil & Kunda, 2022).
After human consumption, it is quickly hydrolyzed into amino acids in
the digestive tract, and does not change the normal bacterial colony in
the intestinal tract (Zimet et al., 2018).
The mode of action of nisin on microorganisms is to inhibit cell wall
formation, causing cytoplasmic leakage, which may be related to the
ability of nisin to adhere to lipid II. Lipid II is a precursor material used
for bacterial wall biosynthesis. Nisin infiltrates into lipid II, hindering
the growth of peptidoglycan network. When the N-terminal part of nisin
is connected with the carbohydrate pyrophosphate part of lipid II, pores
appear on the cell membrane, thus inhibiting microorganisms (Popa
et al., 2022). The antimicrobial mechanism of nisin is shown in Fig. 5. It
was found that the nanogel containing nisin also has an antibacterial
effect and can inhibit the growth of Staphylococcus aureus and Escherichia
coli (Wu et al., 2023). Although a single natural preservative can inhibit
microorganisms, its range of action on microorganisms is limited. The
composite preservative has a broader antibacterial spectrum, stronger
antibacterial activity. Studies have demonstrated that the tender coco­
nut water (TCW) quality can be extended up to 10 days after treatment
with nisin, and the TCW quality can be maintained for up to 15 days
after treatment with polylysine. The combination of nisin and polylysine
showed good antimicrobial performance, which could extend the shelf
life of TCW to 20 days (Pandiselvam et al., 2022). Chitosan, tea poly­
phenols, and nisin have better antimicrobial effects than single pre­
servatives when used to treat plant-based meat products, demonstrated
that compared with a single preservative, the compound preservatives
were more effective in preventing lipid oxidation in plant-based meat
products, and its inhibitory effect on microorganisms was also better
than that of a single preservative (Dai et al., 2022). The combination of
bacteriophages and nisin is more effective than single bacteriophages or
nisin, and the antibacterial activity of nisin increases with the increase of
concentration. The results show the potential of combining phages with
nisin and polylysine for the biocontrol of Salmonella enteritidis, Salmo­
nella typhimurium, and Escherichia coli O157:H7 in food industry (Wu
et al., 2023).
Nisin is a natural preservative that is non-toxic, efficient, and has
strong bactericidal effects, and has been recognized internationally.
Nisin, as a bacteriocin preservative used in food, is widely used in dairy
products such as fresh milk, yogurt, milk powder, and cheese, as well as
in beverages, meat products, canned foods, and other fields. Nisin does
not affect the flavor, texture, and product structure of food. When nisin
is used to preserve chilled meat, the total number of bacteria in the
chilled meat significantly decreases. At the same time, the higher the
concentration of nisin the more obvious its preservation effect. The
antibacterial effectiveness of nisin in food preservation is compromised
by the emergence of resistance to nisin in various bacteria. Antibacterial
agents that work synergistically with nisin were proved to minimize its
resistance. The combination of nisin and cinnamaldehyde can not only
serve as a promising natural food preservative but also reduce bacterial
resistance problems (Shi et al., 2017).
The bacteriocin produced by Carnobacterium maltaromatium KCA018
isolated from raw milk has higher antibacterial activity against Gram-
positive bacteria than against Gram-negative bacteria and can effec­
tively inhibit the activity of Listeria monocytogenes (Kim et al., 2022).
Studies have shown that the bacteriocin LFX01, produced by Lactoba­
cillus plantarum LF-8 isolated from the intestine of tilapia (Oreochromis
mossambicus), has an antibacterial effect against the growth and biofilm
formation of Staphylococcus aureus and Escherichia coli. And bacteriocin
LFX01 could reduce the metabolic activity of cells, disrupts the perme­
ability and integrity of the cell membrane, and lead to the leakage of
intracellular material (Xin et al., 2023). The bacteriocin from Lactica­
seibacillus rhamnosus ZFM216 has broad-spectrum antibacterial activity
H. Dong et al.
Fig. 5. Antimicrobial mechanism of nisin killing bacteria.
and can inhibit Gram-positive bacteria and Gram-negative bacteria.
Bacteriocin has an antibacterial effect on Staphylococcus aureus,
destroying the membrane and wall of Staphylococcus aureus cells,
resulting in cytoplasm leakage (Wu et al., 2022). A new antibiotic
(Plantaricin Bio-LP1) produced by Lactiplantibacillus plantarum NWAFU-
BIO-BS29. It shows strong inhibitory activity against both Gram-positive
bacteria and Gram-negative bacteria, and has a broad antibacterial
spectrum, with no biological human risk, it can use as a potential bio-
preservative in food and feed (Ismael et al., 2023). Bacteriocin XJS01
derived from Lactobacillus salivarius can inhibit the growth of foodborne
Staphylococcus aureus, proving the inhibitory effect of XJS01 on the
spoilage of raw pork (Ying et al., 2023). Bacillocin 490 is a low molec­
ular weight (2 kDa) bacteriocin produced by a thermophilic strain iso­
lated from dairy products. But Bacitracin 490 only has inhibitory effects
on some Gram-positive bacteria (Shleeva et al., 2023).
Bacteriocins have properties that can be used for a wide range of food
and medical applications. Therefore, the safety of bacteriocins requires
high attention. Since 2011, nisin has been widely used as a food pre­
servative (Soltani et al., 2021) and has no chronic, subchronic toxicity,
reproductive toxicity, genetic toxicity, carcinogenic and teratogenic ef­
fects (Hagiwara et al., 2010). However, there have been reports of high
concentrations of nisin causing some toxicity or side effects (De Almeida
Vaucher et al., 2011). Researches have shown that bacteriocins possess
low toxicity. The Spizizenii strain Ba49 isolated from Allium cepa showed
low toxicity to 293 mammalian cell lines when administered at con­
centrations several times higher than the minimum inhibitory concen­
tration (Taggar et al., 2021). Most bacteriocins are non-toxic to
eukaryotic cells, but enterococcal cytolysin has been found to be toxic
(Cox et al., 2005). More comprehensive studies on the safety of bacte­
riocins are needed, and various measurements are required to evaluate
their cytotoxicity, ability to induce apoptosis, growth inhibition, he­
molytic activity, and acute and subchronic toxicity in eukaryotic cells.
Bacteriocins have broad application prospects in food processing,
preservation, and food safety. With the progress of molecular biotech­
nology and the maturity of genetic engineering technology, new types of
bacteriocins can be produced through genetic and protein engineering,
thereby enhancing the application potential of bacteriocins. Although
some bacteriocins such as nisin have been commercially applied in
specific food systems, there may be limitations to use bacteriocins alone,
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including high separation and purification costs, narrow inhibition
spectra, susceptibility to enzymatic degradation, and susceptibility to
complex food environments. Different types of bacteriocins may not
develop cross-resistance against bacteria because they have different
structures and targets of action. Numerous studies have shown that the
combining treatment of bacteriocins and various chemicals has a syn­
ergistic antibacterial effect on spoilage microorganisms and foodborne
pathogens-causing, effectively extending the shelf life of food and
ensuring food safety (Yu et al., 2023).
4.2. Actinomycin
4.2.1. Natamycin and their antimicrobial activity
Natamycin is a natural polyene macrolide antifungal agent fer­
mented by Streptomyces natalensis (Mohamadi et al., 2021). The anti­
bacterial mechanism of natamycin is that it can bind with ergosterol on
the cell membranes of fungi and yeast, change the permeability of the
cell membrane, and is ineffective against bacteria and viruses without
ergosterol in the cell membrane (Chen et al., 2021). The eukaryotic cell
membrane composed of lipids, phospholipids, proteins, and sterols is
called ergosterol. Sterols provide selective binding sites for Natamycin.
Due to its high affinity for ergosterol, Natamycin irreversibly combines
with ergosterol in fungal cell membrane to form a polyene sterol com­
plex. This complex alters the permeability of the membrane, causing the
rapid leakage of essential ions and small peptides, thus preventing the
growth of yeast and fungi, however, natamycin is ineffective against
bacteria and viruses without ergosterol in the cell membrane. The
schematic diagram representing the mode of action of Natamycin on a
fungal cell is illustrated in Fig. 6 (Meena et al., 2021). Natamycin has
been approved as a food additive in over 40 countries and has been
considered as a GRAS product by FDA (He et al., 2019).
Natamycin is commonly used as an additive in the food industry and
has been registered as a biological fungicide for postharvest use of citrus
and other fruits. Citrus sour rot caused by Geotrichum citri-aurantii is one
of the most important postharvest diseases in citrus fruit, after using
natamycin, the incidence rate is significantly reduced, indicating that
natamycin can be used to sour rot control (Fernández et al., 2022). The
study showed that that natamycin significantly reduced the incidence
and lesion size of fruit Botrytis cinerea, and reduced the conidial
H. Dong et al.
Fig. 6. Mode of action of Natamycin on a fungal cell. (A) The schematic diagram of fungal cells and location of sterol and membrane channels within cells. (B)
Affinity of Natamycin towards sterol resulting in formation of polyene sterol complex. (C) Leakage of membrane channel with loss of ions and peptides from
fungal cells.
germination and mycelial growth of Botrytis cinerea (Saito et al., 2020).
Natamycin can reduce the decay rate of mulberry fruit. Mulberries with
increased concentration of natamycin could significantly inhibit the
growth of fungal pathogens in mulberry fruit and improve the quality of
mulberry (Wen et al., 2019). Natamycin is a safe preservative that can
effectively inhibit the growth of mold and yeast in mulberry fruits and
improve their quality throughout the storage period after harvest.
Natamycin can also effectively inhibit fungal growth in soft Galotyri
cheese. At high concentrations, natamycin leads to a significant reduc­
tion in the number of Candida tropicalis biofilm cells and damages the
biofilm structure. The combination of farnesol and natamycin, effec­
tively reduced cell counts and disrupted the structure of C. tropicalis
biofilms. The combination of natamycin and other substances to prepare
composite materials also has an antifungal effect (Agustín et al., 2019).
The gum acacia-gold nanocomposite fabricated with antifungal preser­
vative agent natamycin, has good antifungal activity against the fungus
strain Aspergillus ochraceopealiformis (Karthick Raja Namasivayam et al.,
2022). Natamycin blocks the growth of fungi through specific combi­
nation with ergosterol, without membrane permeability. Natamycin can
also timely inhibit the lateral transport of amino acids and glucose to the
plasma membrane, leading to complete damage to fungal hyphae.
Natamycin has a wide range of applications, with a small dosage,
significant effect, and long antibacterial time. Natamycin can exert
antibacterial activity at lower concentrations, and its antibacterial ac­
tivity is almost unaffected by pH. When natamycin acts together with
nisin, its antibacterial spectrum is complementary. Several studies
conducted toxicity tests with natamycin and found that it had no sub­
chronic toxicity, chronic toxicity (Levinskas, 1966), cytotoxicity, or
genetic toxicity (Rasgele & Kaymak, 2013). However, long-term
administration of high doses may result in some side effects such as
nausea, vomiting and diarrhea (Meena et al., 2021). Natamycin is
difficult to be digested and absorbed in the human body and consuming
natamycin at the prescribed dose has no toxic side effects on the human
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body. When natamycin is directly added to yogurt, it only inhibits the
growth of mold and yeast, without affecting the bacteria (bifidobacteria)
in the yogurt. In addition, it is odorless and colorless, and does not affect
the taste when consumed. It is now the only internationally recognized
antifungal food preservative and is used in the production and storage of
baked goods, fruit and vegetable drinks, dairy products and meat
products.
4.2.2. Polylysine and their antimicrobial activity
Polylysine is formed by the combination of ε-amino and α-carboxyl
groups of lysine in alkaline solution through peptide bonds. Polylysine is
a natural polycationic peptide produced by Streptomyces albicans. When
the degree of polymerization is lower than decapeptide, polylysine will
lose its antibacterial activity. Polylysine has broad-spectrum antibacte­
rial activity and strong inhibitory effects on microorganisms such as
fungi, bacteria, and yeast (Wang et al., 2021). The antibacterial activity
of ε-polylysine is due to its positive charge can be adsorbed on the
surface of microorganisms with negative charge, and its combination
can change the permeability of bacterial cell membrane (Wang et al.,
2023). The cytoplasm entered through membrane pores by polylysine
and two parallel reactions occurred afterward, that is, an increase in
reactive oxygen species (ROS) levels and the DNA genome interactions,
resulting in cell death (Zarrintaj et al., 2022). In 2013, polylysine was
approved by the FDA as a safe food preservative, and in 2014, it was
approved by China for use in food preservation.
Polylysine can be used for storage and preserve of fruits after harvest,
it was demonstrated that ε-polylysine has a strong inhibitory effect on
the fruit pathogenic bacteria Penicillium expansum and Colletotrichum
gloeosporioides and to delay the decline in fruit quality during storage of
juniper apples and guava fruits, inducing resistance to fruit diseases and
being used as a preservative for post-harvest storage (Bai et al., 2022).
ε-Polysine treatment can significantly inhibit the black spot rot of apple,
jujube, and tomato, and has antifungal activity against Alternaria rot. By
H. Dong et al.
interfering with the integrity of pathogen membrane and inducing the
formation of endogenous reactive oxygen species in pathogens, it can
inhibit the growth of colonies and spore germination and induce the
ultrastructural changes of pathogens. ε-Polysine not only shows anti­
bacterial activity, but also enhances the expression of genes involved in
the pathogenesis resistance pathway in the host (Shu et al., 2021). When
malt agar was used as a substrate, it was found that Epsilon poly-L-lysine
can inhibit the growth of Gloeophyllum trabeum, Rhodonia placenta,
Trametes versicolor and Irpex lacteus and other fungi (Cai & Kuo, 2022).
Compared to using each natural preservative alone, mixed natural pre­
servatives have better antibacterial effects. ε-Polylysine, 70 % ethanol
extract of chestnut and hydrothermal extract of cinnamon showed sig­
nificant synergistic antibacterial effect on the growth of Staphylococcus
aureus (Lee et al., 2022). Polylysine and chitosan both have antibacterial
activity themselves, but ε-polylysine combined with chitosan-based
coatings has better antibacterial and antioxidant activity, and can
effectively inhibit the growth of spoilage microorganisms (Zhang et al.,
2022). Cobalt-chromium-molybdenum and ε-polylysine composite
coatings have antibacterial and antibiofilm effects, and has obvious
inhibitory effects on Staphylococcus aureus and Escherichia coli, leading to
the reduction of a bacterial cell aggregation and activity (Guo et al.,
2021). Poly (L-diaminopropionic acid) and poly (ε-L-lysine) showed
strong antibacterial activity in fresh grape juice and has great potential
as a biological preservative(Wang & Rong, 2022). ε-Polylysine hydrox­
ide (ε-PLH) is a homopolymer of ε-polylysine and also has antibacterial
activity. Research shows that ε-PLH can inhibit the formation of Listeria
monocytogenes biofilms, significantly enhances the inhibition of biofilms
when combined with cinnamon essential oil, and shows a synergistic
anti-biofilm effect on Listeria monocytogenes and Pseudomonas lundensis,
which can be used in food preservatives (Zhu et al., 2022).
Polylysine is a broad-spectrum antibacterial, it can be decomposed
into L-lysine in the human body and completely digested and absorbed
by the human body. Moreover, polylysine has good thermal stability and
does not decompose at high temperatures (120 ◦ C, 120 min). It can be
sterilized together with the raw materials, so it has broad application
prospects in meat products. Although polylysine has a slightly bitter
taste, an addition of micrograms can achieve ideal antimicrobial effects.
Therefore, the addition of trace amounts of polylysine has no effect on
the taste of foods. Two-week oral administration of polylysine to rats
showed no signs of toxicity and no changes in the organs, suggesting that
polylysine is not toxic when administered orally (Titlow et al., 2013).
Subchronic oral toxicity experiments also showed it is no toxicity (Hiraki
et al., 2003). In the chronic feeding study in rats, oral administration of
high doses of polylysine had no significant toxicological effects on the
health of the experimental animals. The oral dose of polylysine in the fed
mice corresponds to 100 times the maximum allowable content of
polylysine in human food (Song et al., 2017). Therefore, at the allowable
level of polylysine in the human body, polylysine is safe, and oral
administration of polylysine does not cause systemic organ or systemic
toxicity. Polylysine has excellent biological activity and high safety. In
addition to being approved as a food preservative by countries such as
the United States, Japan, and South Korea, China has also added poly­
lysine to the list of approved food preservatives (Yang et al., 2023).
4.3. Phage and their antimicrobial activity
Phage is a virus that specifically infects bacteria and can be used as a
natural food preservative. The antibacterial mechanism is due to the
high specificity, self-replication, and rapid killing characteristics of
phages on host cells. Phages attach specifically to their host bacterial
strains, inject DNA or RNA genome, and replicate and express their
genes through the mechanism of hijacking host cells. Then the new virus
particles are assembled in the host cell, the cell is split, and the virus of
offspring is released (Ibarra-Sánchez et al., 2018).
Lytic phages are viruses that cause specific bacterial hosts to lyse, not
disrupting the normal microbial community of humans. They will not
12
Food Research International 190 (2024) 114548
infect or colonize eukaryotic cells, nor do they have harmful effects on
human health. A bacteriophage extracted from sewage, and adding DNA
extracted from Shiga toxin-producing Escherichia coli (STEC) to these
bacteriophages can effectively combat STEC (El-Sayed et al., 2022).
VPT02, a new cleaving bacteriophage isolated from oysters, was
confirmed to kill the seafood pathogen Vibrio parahaemolyticus and
enhance the survival rate of brine shrimp infected with
V. parahaemolyticus. The VPT02 genome does not contain toxin and
antibiotic resistance genes that can be used as food preservatives (You
et al., 2021). Phages have lytic activity against pathogenic bacteria, not
altering the quality of food. It was demonstrated that DNA extracted
from enteropathogenic Escherichia coli in beef lung was added to bac­
teriophages, which could achieve antibacterial effects on lettuce and
milk (Waturangi et al., 2021). Bacteriophage VVP001 can inhibit the
growth of Vibrio vulnificus and protect the host from the invasion of
V. vulnificus, and can be used as a seafood preservative (Kim et al.,
2021). Studies show that bacteriophage P100 can inhibit the growth of
Listeria monocytogenes in ready-to-eat sliced pork ham and fresh cheeses
(Figueiredo & Almeida, 2017). The bacteriophage Spp001 reduces the
level of Streptococcus putrescens in marine fish Paralichthys olivaceus
under cold storage to prevent spoilage and achieve antiseptic effect. It
was demonstrated that the multivalent bacteriophage CoNShP-3 can not
only lyse Coagulase-negative staphylococci (CoNS), but also lyse other
genera including Staphylococcus aureus, methicillin-resistant Staphylo­
coccus aureus (MRSA), vancomycin-resistant Staphylococcus aureus
(VRSA), Bacillus cereus and B. subtilis. Moreover, the CoNShP-3 phage
also exhibited anti biofilm effects on Staphylococcus epidermidis CFS79
and Staphylococcus hemolyticus CFS43, respectively, while the biofilm of
Staphylococcus saprophyticus CFS28 was completely removed (Sofy et al.,
2020).
Enteropathic Escherichia coli (EPEC) is a leading cause of foodborne
illness and poses a threat to human health. Phages have the potential to
be used as a natural food preservative that can be used to combat
foodborne pathogens in food preservatives. Phage UAE_MI-01 is a
bacteriophage of Escherichia coli O157: H7, which is isolated from the
feces of wild pigeons (Columba livia home) and inhibits the growth of
Escherichia coli by destroying the host cell membrane (Sultan-Alolama
et al., 2021). The Bacillus phage endolysin has antibacterial activity
against Klebsiella pneumoniae, Salmonella typhimurium, Proteus, and
Escherichia coli (Sarjoughian et al., 2020). Phage can also be used with
other preservatives to exert synergistic antibacterial effects. Phage-
antibiotic synergism can inhibit food borne disease bacteria (Salmo­
nella, Staphylococcus aureus and Escherichia coli). Bacteriophages and
chemical disinfection have a synergistic effect on the opportunistic
pathogen Pseudomonas aeruginosa. Research has shown that the combi­
nation of bacteriophage and chemical disinfectant can improve the
removal of surface wet biofilm and bacteria and prevent the regenera­
tion of dry biofilm (Stachler et al., 2021).
Phages have excellent food preservation properties and are crucial
for preclinical safety and toxicity testing of mammalian cells and tissues.
The FDA has approved the use of phages as food preservatives to combat
foodborne bacteria and treat bacterial infections (Jault et al., 2019).
Phage preparations against Listeria monocytogenes (Listshield TM), Sal­
monella enterica (SalmFresh TM), and Escherichia coli (Ecoshield TM)
have received Generally Recognized As Safe (GRAS) designation from
the FDA for direct use on food (Nazir et al., 2023). Studies have evalu­
ated the safety of the pneumococcal endolysins Cpl-1 and Pal, including
an in vivo safety assessment of mouse models and gene expression
profiling, all of which confirmed no adverse effects in mice (Harhala
et al., 2018). Phages with high specificity and a narrow lysis spectrum,
can specifically cleave harmful microorganisms and have no infective
ability against humans, animals and plants outside the host range (Gyles,
2018). Phages have rapid self-reproduction and short screening cycles
and are widespread in nature. In recent years, phages have been
increasingly recognized for their efficient, rapid and safe properties. In
addition, while phages kill bacteria, they have no harmful effects on
H. Dong et al.
food sensory organs (Khan et al., 2023). Although safety research results
so far support the claim that phages are a potentially safe antibacterial
agent. Further research is needed to confirm that different types of
bacteriophages do not have significant effects on humans. Phages have
the potential to become natural preservatives in food, and their use in
the food industry is a viable, efficient and promising method for corro­
sion protection and preservation. An overview of natural antimicrobial
from microorganism sources is shown in Table 1.
5. Conclusions and prospect
Chemical preservatives have side effects and may cause adverse re­
actions such as cancer, asthma, and allergic reactions (Li et al., 2022).
Natural preservatives are extracted from plants, animals, and microor­
ganisms, with a wide range of sources and strong antibacterial activity,
providing a new way to extend the shelf life of food (Zhang et al., 2023).
Natural preservatives have been widely studied and reported to have
inhibitory effects on these food pathogenic bacteria (such as Salmonella,
Escherichia coli, Listeria monocytogenes, and Staphylococcus aureus).
Combining natural antibacterial agents with other technologies can
improve their performance. Combining with other substances or
different food preservation systems, coatings, and nano encapsulation to
ensure the safety of natural preservatives. The single use of some natural
preservatives, its amount is large, easy to have problems such as color
and smell. In addition to screening natural preservatives that are more
sensitive to microorganisms and use less, the compounding of pre­
servatives has also become a research topic.
In recent decades, the use of natural preservatives in food processing
has received great attention, and their safety assessment has also
received increasing attention. However, insufficient data on the safety
and toxicity of natural preservatives hinder their wider use in the food
industry. In future research, in vitro experiments on the potential toxicity
of natural preservatives to the human body should be carried out to
ensure the safety of natural preservatives after absorption by the human
body. The article focuses on the antibacterial mechanism and safety
evaluation of natural antibacterial substances from plants, animals and
microorganisms. It briefly introduces the influence of different anti­
bacterial ingredients on different strains and their inhibitory effects, and
provides a certain idea and reference value for future research devel­
opment and wide application of natural antibacterial substances in food
field.
CRediT authorship contribution statement
Huiying Dong: Writing – original draft. Yang Xu: Writing – original
draft. Qingqing Zhang: Writing – original draft. Hua Li: Writing – re­
view & editing. Lixia Chen: Writing – review & editing.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
No data was used for the research described in the article.
Acknowledgments
The authors thank Chunhui Program-Cooperative Research Project
of the Ministry of Education, Liaoning Province Natural Science Foun­
dation (No. 2022-MS-241), Liaoning Revitalization Talents Program
(No. XLYC1807182), and Shenyang Young and Middle-aged Innovative
Talents Support Program (No. RC210446) for financial supports. And we
acknowledged the support from National-Local Joint Engineering
13
Food Research International 190 (2024) 114548
Research Center for Molecular Biotechnology of Fujian & Taiwan TCM,
Fujian Key Laboratory of Chinese Materia Medica at Fujian University of
Traditional Chinese Medicine.
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