International Journal of
Molecular Sciences
Review
Antimicrobial Compounds in Food Packaging
Aleksandra Duda-Chodak 1, * , Tomasz Tarko 1
Citation: Duda-Chodak, A.; Tarko, T.;
Petka-Poniatowska, K. Antimicrobial
Compounds in Food Packaging. Int.
J. Mol. Sci. 2023, 24, 2457. https://
doi.org/10.3390/ijms24032457
Academic Editor: Helena Felgueiras
Received: 30 December 2022
Revised: 22 January 2023
Accepted: 25 January 2023
Published: 27 January 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Int. J. Mol. Sci. 2023, 24, 2457. https://doi.org/10.3390/ijms24032457
and Katarzyna Petka-Poniatowska 2
1 Department of Fermentation Technology and Microbiology, Faculty of Food Technology, University of
Agriculture in Krakow, ul. Balicka 122, 30-149 Kraków, Poland
2 Department of Plant Products Technology and Nutrition Hygiene, Faculty of Food Technology, University of
Agriculture in Krakow, ul. Balicka 122, 30-149 Kraków, Poland
* Correspondence: aleksandra.duda-chodak@urk.edu.pl; Tel.: +48-12-662-4792
Abstract: This review presents current knowledge on antimicrobial agents that are already used in
the food packaging industry. At the beginning, innovative ways of food packaging were discussed,
including how smart packaging differs from active packaging, and what functions they perform. Next,
the focus was on one of the groups of bioactive components that are used in these packaging, namely
antimicrobial agents. Among the antimicrobial agents, we selected those that have already been
used in packaging and that promise to be used elsewhere, e.g., in the production of antimicrobial
biomaterials. Main groups of antimicrobial agents (i.e., metals and metal oxides, organic acids,
antimicrobial peptides and bacteriocins, antimicrobial agents of plant origin, enzymes, lactoferrin,
chitosan, allyl isothiocyanate, the reuterin system and bacteriophages) that are incorporated or
combined with various types of packaging materials to extend the shelf life of food are described.
The further development of perspectives and setting of new research directions were also presented.
Keywords: foodborne pathogens; smart packaging; active packaging; food coatings; antibacte-
rial agents
1. Introduction
Food is the source which provides essential nutrients to the human body. Unfortunately,
it is also a very good environment for numerous microorganisms to develop in, including those
that cause food spoilage. Many of these microorganisms are also human pathogens and pose a
real challenge for modern science. Among the most common foodborne pathogens are bacteria
(Escherichia coli, Listeria monocytogenes, Staphylococcus aureus, Bacillus cereus, Campylobacter jejuni,
Clostridium botulinum, C. perfringens, Cronobacter sakazakii, Salmonella spp., Shigella spp.,
Vibrio spp., and Yersinia enterocolitica), viruses (Hepatitis A and noroviruses) and parasites
(Cyclospora cayetanensis, Toxoplasma gondii and Trichinella spiralis) [1].
Pathogens are ubiquitous and cause a variety of human diseases which are often
life-threatening. Along with the development of medical science and progress in health
care, pathogen entry the human body as contamination of biomedical products began to be
an increasing burden for the system. The biomaterial introduced into the patient’s body
is intended to save human life or improve its quality, but instead it becomes a source of
infection that potentially threatens health or might be hazardous for life of the patient [2].
Hospital-acquired infections (HAI), which can be caused by bacteria, fungi and viruses, are
a significant complication for implantation procedures of various biomaterials, both during
planned surgical operations (various prostheses, implants, dentures, tissue scaffolds, stents,
heart valves, hip joint endoprostheses), as well as during life-saving actions in the intensive
care unit (vascular catheters, urinary catheters, fracture-fixation devices) [3]. The most
important pathogens causing biomaterial-assisted infections (BAI) include Staphylococcus
aureus, S. epidermidis, S. saprophyticus, Escherichia coli, Klebsiella pneumoniae, Proteus mirabilis,
P. vulgaris, Enterococcus faecalis, Streptococcus viridans, Propionibacterium acnes, Pseudomonas
aeruginosa, and Candida sp. [3,4]. HAI are amongst the main complications associated
https://www.mdpi.com/journal/ijms
Int. J. Mol. Sci. 2023, 24, 2457 2 of 34

with the implantation of any biomaterial. Hence, modern science strives to develop new
agents/substances with antibacterial (or, more broadly, biocidal) potential which can be
integrated with the biomaterial to prevent bacterial colonization and biofilm formation.
These compounds, in addition to a broad spectrum of antimicrobial properties, should be
characterized by minimal toxicity to humans. Bacteria can get into the biomaterial in various
ways, but usually the source of infection is the direct contamination of the biomaterial with
bacteria present in the operating room or with bacteria normally inhabiting the skin (of the
patient, doctor or nurse) during the introduction of the device into the subject’s body [4].
Less often, the infection occurs during hospitalization or later (e.g., even several years after
surgery), when bacteria originating from a local infection of the patient, via blood, get to
the site of biomaterial implantation and cause the development of a new contamination.
In the food production process, the situation is similar, and products can be con-
taminated at various stages of the food chain. Microorganisms get into the food from
the environment and primary production (raw material, production lines, equipment for
processing and transporting raw materials, semi-finished and ready-made food), as well as
from the skin of personnel who do not properly care about the hygiene of the process. They
can be present both in the raw material used for production, as well as being introduced at
subsequent stages, during improper storage, transport, distribution of finished products or
even during consumption [5,6].
Of course, food spoilage causes economic loss in the food industry and can damage a
producer’s reputation, even resulting in a punishment according to the local law. Not all
microorganisms that lead to food spoilage are dangerous for human, but the development
and the contamination of food with pathogens or their toxins impair the safety of food
and pose a risk to the health of consumers [6]. It is estimated that each year 48 million
Americans get sick, 128,000 are hospitalized, and 3000 die due to foodborne diseases [7].
The top five pathogens that cause domestically acquired foodborne diseases in USA are
noroviruses (58%), followed by non-typhoidal Salmonella spp. (11%), Clostridium perfringens
(10%), Campylobacter spp. (9%) and Staphylococcus aureus (3%). However, among the
causes of hospitalizations and deaths, Escherichia coli O157, Toxoplasma gondii and Listeria
monocytogenes are also mentioned [7]. According to WHO, more than 23 million Europeans
became ill (4654 of deaths) from eating contaminated food in 2010. The most frequent
causes were noroviruses, Campylobacter spp., non-typhoidal Salmonella enterica, Toxoplasma
gondii, Giardia spp., Cryptosporidium spp., Shiga toxin producing E. coli, enteropathogenic
E. coli, as well as hepatitis A virus, Shigella spp., L. monocytogenes, Brucella sp., Mycobacterium
bovis, and Echinococcus spp. [8]. These diseases generate huge costs related to treatment,
absence from work or compensation.
Therefore, considering both safety and economic considerations, manufacturers are
trying to develop new food packaging that, in addition to the “traditional” function, will
have antimicrobial properties. This review paper aims to gather in one place information
on the antimicrobial compounds that are used in the production of food packaging but
which have or potentially have a predisposition to be used in the production of antimicro-
bial biomaterials.

2. Food Packaging
The main functions of the food packaging are usually passive, chemical, biological,
and physical protection of food against secondary contamination, environmental influences,
mechanical damage during storage and transport, as well as against access to light, oxygen,
UV radiation, or moisture. However, the packaging also has an informative (name of
the product, information on chemical composition and allergens, expiration date, method
of preparation and administration, etc.) and promotional (information about the manu-
facturer) function [9]. In recent years, packaging has also started to play an active role
in preventing food waste by actively extending its shelf life while ensuring quality and
safety [10].
Int. J. Mol. Sci. 2023, 24, 2457 3 of 34

Traditionally, the main kinds of materials used for food packaging are glass, metals,
paper, and plastic, and their main common characteristics were neutrality to the packaged
food [9]. What was initially the main advantage of plastic (its chemical resistance, low price
and weight) has become a disadvantage, causing these issues to dominate the packaging
market and have a huge negative impact on the environment. The main polymers used in
food packaging industry are polyethylene (HDPE, LDPE, and LLDPE), polystyrene (PS),
polypropylene (PP), polyvinylchloride (PVC), polyamide (PA), and polyethylene tereph-
thalate (PET) [11,12]. To minimize environmental impact, some kinds of food packaging
have started to be recycled or even reused (mainly glass and metals), but the recycling of
plastic is still insufficient. Therefore, fully biodegradable, safe biopolymers are constantly
under development and undergoing a process of improvement [13].
Biopolymers that are gaining popularity in the packaging industry are polysaccharides
(e.g., starch and modified starches, cellulose, pectin, carrageenan, alginate, chitin, chitosan),
lipids (resins, bee waxes, natural resins, lecithin, emulsifiers, etc.), proteins (casein, colla-
gen, gelatin, gluten, zein, soy proteins, etc.), and microbial polymers (pullulan, curdlan,
gellan, xanthan polyhydroxyalkanoates/PHA/, polyhydroxybutyrate/PHB/), polylactic
acid/PLA/, etc.) [10,12–14]. Although they are all biodegradable and renewable, their
“packaging values” are not perfect (e.g., due to weak mechanical properties, hydrophilic-
ity, crystallization, aggregation) and still require improvement. However, many of these
biopolymers can be variously modified by methods that improve their barrier function or
completely change their characteristics. Thanks to that, bio-based polymers easily found
their place in the industry and have different applications. Among the examples of this
are the derivatives of cellulose [13] or chitosan [15]. Moreover, new bio-based materials
were introduced recently and, in what is exactly the opposite of traditional food packaging,
they should interact with the packaged food, albeit in a desirable and planned way. Such
innovative packaging systems were created from various biopolymers used as a packag-
ing matrix, to which additional ingredients with an important function were introduced.
Among such innovative systems are ‘active’ and ‘smart’ packaging and their characteristics
are presented below.

2.1. Smart Packaging

“Smart materials and articles” means materials and articles that monitor the condition
of packaged food or its environment [Commission Regulation (EC) No 450/2009]. Smart
packaging is designed to inform about the product in a more advanced way than traditional
packaging. It not only gives knowledge about the composition or the manufacturer, it
allows the tracking of the fate of a food product throughout the chain, including monitoring
changes that have taken place in the environment and/or in the product (e.g., changes
in temperature, pH, acidity). Smart packaging communicates to the consumer whether
the product still meets all the requirements, whether it is safe, for example, whether the
cold chain has been disturbed during the manufacturing process, storage and transport of
the food product, or whether anything else has happened that could adversely affect the
safety and quality of that food [16,17]. The new generation of smart packaging is equipped
with biosensors (e.g., ToxinGuard® ; Food Sectinel System® ). They may communicate if
any harmful or undesirable metabolites (such as toxins or biogenic amines) have appeared
in food product or if dangerous food pathogens have developed [18,19]. Thus it is an
intelligent way to monitor food quality. Furthermore, smart packaging indirectly allows
us to extend shelf life. First, it is a useful tool that enhances product traceability, allows to
control the shelf life of the product throughout the supply chain, and thus allows to make
better decisions regarding the order of transport, as well as in terms of sales policy and
product promotion in the store. Secondly, especially in the case of products with a ‘best
before’ date, it can prevent a healthy and wholesome product from being thrown away.
Consumers, seeing that a product has ‘exceeded’ the date given on the packaging, often
throw food away without thinking because they cannot distinguish between the term ‘best
before date’ from the term ‘use-by-date’. This leads to a lot of food loss. The stated ‘best
Int. J. Mol. Sci. 2023, 24, 2457 4 of 34

before date’ only defines the minimum period for which all product characteristics are
guaranteed, while the actual shelf life is longer, although it largely depends on how the
product has been stored. Therefore, smart packaging, by informing the consumer about
the condition of the product (e.g., whether the pH has dropped too much, whether the
product has been refrigerated all the time, whether harmful components are present in the
packaging) can prevent food from being thrown away and the waste of something that still
could be safely used [19].

2.2. Active Packaging

The term “active” means that it has additional functions consisting of actively caring
for the quality and safety of food and extending its shelf life, mainly by providing additional
protection against the growth of microorganisms or the oxidation of a product. According to
Commission Regulation (EC) No 450/2009, “active materials and articles” means materials
and articles intended to extend the shelf life or to maintain or improve the condition of
packaged food, and they are intentionally designed in such a way that the ingredients
they contain release or absorb substances into the packaged food or its surroundings.
Consequently, active packaging can act by: (i) managing the atmosphere inside the package
(for example, by binding oxygen or emitting CO2 ); (ii) preventing the development of
foodborne pathogens and/or by inhibiting or killing food spoilage microorganisms, (iii);
inhibiting the formation of biofilm; (iv) exerting anti-radical and antioxidant effects; or
(v) by absorbing substances which can cause food spoilage or have an unpleasant odor
(ethanol, acids, oxygen, carbon oxide, ammonia) [14,16].
Since microorganisms are one of the main reasons why food loses its quality or ceases
to be safe, the greatest interest is aroused by active packaging, whose task is to inhibit the
growth of microorganisms or limit the release of harmful metabolites by them (including
toxins, gases, acids).

3. Antimicrobial Agents and Their Application in Food Industry

Striving to produce healthy, high-quality food while limiting chemical preservatives,
combined with the desire to reduce the impact of packaging on the environment, causes
scientists and food producers to focus on producing the most natural packaging containing
natural antimicrobial agents. It is not possible to find a single antimicrobial agent able to
inhibit all microorganisms that cause food spoilage or food poisoning. However, scien-
tists have checked hundreds of potential candidates to find the most potent. Moreover,
also some packaging materials can exert antimicrobial activity (e.g., chitosan). Usually,
antimicrobial packaging is developed by loading or coating an antimicrobial agent onto
polymeric packaging films, and an antimicrobial component can migrate to the food’s
surface (migrating film) or can act against surface microbial growth without migration
(non-migrating film) [16].
Active agents used in food packaging can be classified according to various criteria,
such as origin (plant, bacterial, animal), mode of action (scavengers, emitters, antioxidants,
antimicrobial agents, antibiofilm agents, etc.), structure (organic, inorganic, metals, an-
timicrobial peptides, enzymes, antibiotics, etc.) and others. According to Vilela et al. [20],
active agents include antioxidants (polyphenols, α-tocopherols, lignin, essential oils, plant
extracts), CO2 emitters (sodium bicarbonate, citric acid, ferrous carbonate), O2 scavengers
(iron, palladium, ascorbic acid, pyrogallol, gallic acid, glucose oxidase, laccase), ethylene
scavengers (KMnO4 , activated carbon, metal oxides, titanium dioxide) and antimicrobial
agents (metals, nisin, essential oils, lysozyme, chitosan, nisin, lactoferrin).
Below, the main group of antimicrobial agents are described together with their
application in the food packaging industry. First, inorganic agents, i.e., metals and metal
oxides, are described. Then, various groups of organic antimicrobial agents, such as organic
acids, antimicrobial peptides, enzymes, and natural antimicrobial agents of plant origin
(like polyphenols, essential oils components and complex plant extracts) are described.
Int. J. Mol. Sci. 2023, 24, 2457 5 of 34

Additionally, at the final part of the manuscript, other agents that are used in the packaging
industry, but cannot be classified into other groups, are described.

3.1. Metal-Based Nanoparticles as Antimicrobial Agents

A new generation of packaging materials are nanocomposite films which combine
the bio-based polymer (acts as a matrix) with fillers (e.g., antimicrobial agents) that are
dispersed therein to improve the functional properties of material. Bio-based polymers
can be, as mentioned earlier, both natural (various polysaccharides, proteins, lignins,
lipids, etc.) and synthetic (natural-based or bio-based synthetic polymers like polylactic
acid/PLA/and partially bio-based polymers such as polyethylene/PE/, poly(ethylene
terephtalate)/PET/and polyamide/PA/)) [21], while as nanofillers metal or metal oxides
are often utilized.
Metals and metal oxides are used as nanoparticles (NPs) due to their small size,
favoring penetration into the interior of the bacterial cells. Although the exact mechanism
of the toxicity of metal NPs and metal oxides NPs against bacteria is not fully understood,
it has been postulated that it includes the generation of free radicals and reactive oxygen
species (ROS), as well as the direct contact with bacteria surface that results in cell wall
damage and the interruption of transmembrane electron transport [22–24]. ROS then
induce oxidative stress, modulate the gene expression and cause the damage of important
cell components, such as DNA and proteins, affects the mitochondria function and causes
the cell membrane and cell wall disruption, leading finally to bacteria cell death [25]. The
antibacterial activity of metal NPs is also dependent on the release of metal ions, especially
when metal ions play the physiological role (such as Cu2+ ) [26]. The differences between
cell wall structure and composition of Gram-positive and Gram-negative bacteria are
important for the final antibacterial impact of metals NPs. It has been proved that Gram-
positive bacteria are more resistant to the NPs than Gram-negative bacteria because they
contain more peptidoglycan layers and teichoic acids. Metal NPs can therefore interact
with numerous units of amino acids and carbohydrates of the peptidoglycan and can also
be more effectively trapped by negatively charged peptidoglycan [24,27]. The importance
of the charge of metal nanoparticles for its antimicrobial activity was demonstrated by
Abbaszadegan [28]. Among different silver NPs tested, the positively charged NPs had the
highest antimicrobial activity against all tested bacterial species, both Gram-positive and
Gram-negative.

3.1.1. Metal Nanoparticles

Among metal nanoparticles (NPs) which exert an antimicrobial effect, the most com-
mon used are NPs of copper, silver, selenium, and gold [16,21,29–32]. Metal NPs can
be added to the polymer matrix. However, not all metal NPs are incorporated in food
coating, as sometimes they are used alone or with combinations of other antimicrobials.
For example, copper nanoparticles exert anti-biofilm activity against E. coli, P. aeruginosa,
Salmonella Typhi, and Shigella flexneri at the concentration of 12.5 µM, while the minimum
inhibitory concentration (MIC) is 25 µM [33].
In vitro studies showed that silver nanoparticles (Ag NPs), synthesized from Corynebac-
terium glutamicum, had the highest antimicrobial activity against Klebsiella pneumoniae (in-
hibition zone 18 mm in diameter) and were effective against E. coli, S. aureus, Salmonella
enterica, S. flexneri, P. aueroginosa, Bacillus subtilis and B. flexus (clear zones of inhibition with
16 mm to 4 mm in diameter) [34]. Gold nanoparticles exerted strong inhibitory activity
(with MIC value lower than that of antibiotics) against E. coli ATCC25922, S. aureus ATCC
29213, Bacillus subtilis ATCC 11774, P. aeruginosa ATCC27853 and Candida albicans [35].
Chitosan nanoparticles and chitosan nanoparticles loaded with Fe2+ or Fe3+ , showed signif-
icantly higher antimicrobial activity (against E. coli, S. aureus and C. albicans) than chitosan
and related metal ions, and Fe2+ ions were more effective than Fe3+ [36]. Table 1 presents ex-
amples of food packaging with the addition of metal nanoparticles which gave or increased
the antimicrobial properties of the packaging.
Int. J. Mol. Sci. 2023, 24, 2457 6 of 34

Table 1. Examples of antimicrobial metals application for food preservation.

Metal Nanoparticles Polymer/Matrix Characteristics References

antimicrobial activity against mold and total bacteria in
LDPE [37]
barberries packaged in Ag-LDPE film with 1 wt% AgNP
the Ag-based packaging system was effective in
agar hydrogel inhibiting the growth of Pseudomonas spp. in Fior di [38]
Latte cheese
bio-based coating with MAP the elongation of the shelf life of Fior di Latte cheese [39]
antimicrobial effect against
Ag NPs polystyrene (PS) matrix [40]
Gram-positive and Gram-negative bacteria and yeast
prepared modified antibacterial membranes could kill
almost 100% of bacteria under certain conditions and
carboxymethyl CS [41]
inhibition zone still existed after more than 7 cycles
of tests
CS with different molar masses and deacetylation
degrees, and their modifications were used; all
CS/montmorillonite nanocomposite films [42]
nanocomposite-AgNPs films inhibited the growth of
E. coli and Bacillus subtilis
good antibacterial activity against bacteria (S. aureus,
Ag NPs and Au NPs CS film P. aeruginosa), fungi (Aspergillus niger) and yeast [43]
(Candida albicans)
PVA crosslink composite films (as a
the improvement of banana shelf life with
Au NPs crosslinking agent in film production glyoxal [44]
PVA-glyoxal-AuNPs composite film
and/or glutaraldehyde (GA) were employed)
active packaging based on selenium nanoparticles
Se NPs multilayer laminates made of PET and LDPE prevented the oxidation of tested real food products and [45]
extended their shelf life
inhibited growth of Salmonella Typhimurium and E. coli,
Se NPs potato starch film [46]
slightly inhibited B. cereus, did no inhibit Listeria innocua
CS—chitosan; LDPE—low-density polyethylene; PVA—polyvinyl alcohol; PHBV—poly(3-hydroxybutyrate-co-3-
hydroxyvalerate); PLA—polylactic acid.

3.1.2. Metal Oxide Nanoparticles

Additionally, metal oxide nanoparticles can be used as nanofillers. The most common
to be used in this way include zinc oxide (ZnO), titanium oxide (TiO2 ), silica (SiO2 ),
aluminum oxide (Al2 O3 ), iron oxide (Fe2 O3 ), and copper oxide (CuO) (Table 2).
Cerium (Ce)-doped SnO2 nanoparticles were obtained by mixing SnCl2 ·2H2 O and
CeCl3 solutions with hydrogen peroxide [47]; they were proven to show antimicrobial
activity against E. coli. SnO2 nanoparticles showed inhibitory activity against E. coli ATCC
25922 and—to a lesser extent—against S. aureus ATCC 29213 in MilliQ water [48]. In other
studies, it was proven that the addition of the ZnO nanoparticles resulted in the emergence
of antibacterial activities that were not found in the pure PLA film [49]. The most potent
was biocomposite PLA with 5% ZnO which reduced the E. coli concentration by 99.99%
already after 24 h, while the 1% ZnO and 3% ZnO reached the same antimicrobial activity
after 5 days. According to Sirelkhatim et al. [25], mechanisms of toxicity of NPs and their
ions (e.g., silver and zinc) against bacteria include the ROS generation, with the subsequent
induction of oxidative stress and irreversible damage of cellular structures resulting in
bacterial death.
Although NiO nanoparticles were proven to exert antimicrobial activity against E. coli
and S. aureus growth [50], and the use of chitosan-based film with the incorporation of
NiO NPs had improved thermal stability and crystallinity and exhibited good antibacterial
activity against S. aureus and S. typhimurium [51], NiO NPs are not often used in food
packaging. This is caused by the possible negative impact of nickel on the health of
consumers, especially due to the growing frequency of nickel allergies [52].
Int. J. Mol. Sci. 2023, 24, 2457 7 of 34

Table 2. Examples of antimicrobial metal oxides used in food packaging industry.

Metal Oxide Nanoparticles Polymer/Matrix Characteristics References

in vitro inhibitory effect on the growth of E. coli and
CS-based coatings [53]
S. aureus
reduction in the E. coli
LDPE colony on fresh lettuce packed with TiO2 [54]
TiO2 NPs
nanoparticle-coated films
various organic and inorganic
bactericidal property of TiO2 coatings against E. coli
binders (e.g., PVA, polyethylene [55]
O157:H7
glycol, polyurethane)
treatments with CCEO and TiO2 extended the shelf life
TiO2 NPs and Cymbopogon citratus
CS film of minced meat as total bacterial count was in [56]
essential oil (CCEO)
acceptable range after 10 days of storage
the microbial load of fresh orange juice kept in
ZnO NPs and Ag NPs LDPE films packages with Ag and ZnO was below the limit of [57]
microbial shelf life (6 log CFU/mL) up to 28 days
ZnO NPs were active against S. aureus, L.
monocytogenes, E. coli O157:H7, and S. Typhimurium,
while Ag NPs were more active against S. aureus than
ZnO NPs and Ag NPs and L. monocytogenes;
pullulan films [58]
essential oils pullulan films containing the compounds effectively
inhibited the pathogens associated with
vacuum-packaged meat and poultry products stored at
4 ◦ C for up to 3 weeks
nanocomposites show great antimicrobial activity in
ZnO-Ag nanocomposite PHBV-CS [59]
the food packaging of poultry items
the bionanocomposite PHBV/Ag–ZnO (10%) was the
most potent against S. aureus and E. coli when
Ag-ZnO NPs PHBV [60]
compared with bionanocomposites Ag-ZnO 5%, 3%
and 1%
reduction in E. coli growth by 3.14 log for 0.5% ZnO
ZnO NPs PLA [61]
loading in the PLA coating layer
antibacterial activity against S. aureus S33R and E. coli
ZnO-SiO2 CS, PVA [62]
IRAQ 3; increased the shelf life of bread
the maximum antibacterial activity (about 94.7%
SiO2 PHBV growth inhibition for E. coli and 92% for S. aureus) was [63]
obtained for PHBV/SiO2 (2.0%)
CM-CS/MgO composites exhibited antimicrobial
carboxymethyl-CS [64]
activity against L. monocytogenes and Shewanella baltica
MgO NPs biofilms with 2% MgO NPs caused progressive
PLA damage and death of nearly 46% of E. coli bacterial [65]
culture after 12 h treatment
Aluminum-doped zinc
PLA strong antibacterial activity against E. coli [66]
oxide (AZO)
Al2 O3 –Ag and TiO2 –Ag disc diffusion assays proved antimicrobial potential
epoxy polymer [67]
composite NPs against E. coli DH5α and S. epidermidis NCIMB 12,721
CS—chitosan; LDPE—low-density polyethylene; PVA—polyvinyl alcohol; PHBV—poly(3-hydroxybutyrate-co-3-
hydroxyvalerate); PLA—polylactic acid.

3.2. Organic Acids

Many organic acids have a long history as natural or traditional food additives because
they decrease pH in solutions, inhibiting the growth of undesired microorganisms. One of
the best known examples of organic acids used for food preservation is lactic acid, which
is produced by the lactic acid bacteria during fermentation of plant raw materials (silage,
pickled vegetables and fruits) or milk.
Organic acids exist in two forms—dissociated and undissociated. It has been proved
that the efficiency of an organic acid in inhibiting the microbial growth depends on its pKa
value, which describes the pH value at which the acid has available 50% in its dissociated
and 50% in its undissociated form. Some studies revealed that only undissociated form of
Int. J. Mol. Sci. 2023, 24, 2457 8 of 34

organic acid can pass through the walls of microorganism and alter their metabolism or
exert an inhibitory effect [68,69]. It is believed that weak organic acids, such as lactate or
acetate, can permeate the lipid bilayer and release their protons inside the cell, causing a
drop in the intracellular pH. This causes that bacteria try to eliminate excessive protons
from their cells. As protons can be effluxed from the bacteria using special proton pumps
that require energy (ATP), bacteria consume energy for pH stabilization instead of cell
growth [70,71]. Furthermore, the impaired transport across membranes and increased
membrane permeability can even result in cell death. This means that the organic acids are
good candidates to be antimicrobial agents.
Many organic acids, e.g., butyric acid, valeric acid, monopropionin, monobutyrin,
monovalerin, monolaurin, sodium formate, were proved to exert antimicrobial activity [72];
however, only several of them are used in food packaging. Salts of some organic acids, such
as sodium lactate, potassium lactate as well as sodium citrate, have been proved to reduce
the growth of L. monocytogens in beef frankfurters [73]. The effectiveness of antimicrobial
activity against L. monocytogenes can be enhanced by using a UVC treatment of the surface
of frankfurters that contained potassium lactate and sodium diacetate [74].
The microbiological stability of fresh salmon slices, preserved by dipping in 2.5%
(w/v) aqueous solution of sodium lactate (NaL), sodium citrate (NaC) and sodium acetate
(NaA) and then stored at 1 ◦ C was examined by Sallam [75]. All tested salts inhibited
the proliferation of the food spoilage microorganisms (Pseudomonas spp., H2 S-producing
bacteria, lactic acid bacteria, and Enterobacteriaceae), with the order of their efficacy being
NaA > NaL > NaC. The results were similar to those obtained for the rainbow trout
fillets dipped for 10 min in the same solutions (2.5% NaA, NaL, and NaC) before storage
at 4 ◦ C [76]. The shelf life of the treated fillets was extended by 3 days, as all tested
salts prevented the proliferation of total mesophilic bacteria, psychrotrophs, and coliform
bacteria. Additionally, in this experiment, the order of antibacterial activity of organic salts
used was NaA > NaL > NaC.
Salts of benzoic acid, sorbic acid and propionic acid have been used as food preser-
vatives for many years. Although they have GRAS status, recently some of them began
to raise doubts about its safety for consumer health. For example, propionic acid acts
mainly against molds and can be added to food products (calcium propionate and sodium
propionate are used as preservatives in bread and bakery products, grains, meat prepara-
tions, processed meat and processed fish, and animal feed) or is produced in situ during
fermentation (e.g., in cheese) [77]. Recently, it was demonstrated that oral consumption
of propionic acid can lead to inappropriate activation of the insulin counterregulatory
hormonal network [78]. Therefore, propionate has rarely been investigated as an additive
to active packaging.
Sodium benzoate and potassium sorbate are well known antimicrobial agents, and
in 1995 they were reported to suppress the growth of Zygosaccharomyces bailli in mayon-
naise [79]. Active ethylene vinyl alcohol copolymer (EVOH) film, containing the addition
of 3% (w/w) sorbic acid–chitosan microcapsules (S-MPs), was applied to fish fillets and had
stronger inhibitory capacity against Salmonella Enteritidis and E. coli than against L. monocy-
togenes [80]. The combination of sodium nitrite and sodium benzoate exerted inhibitory
activity on E. coli, S. aureus, Bacillus mucoides and Candida albicans, while the combination of
potassium sorbate with sodium nitrite had antimicrobial effects against Bacillus mucoides,
P. aeruginosa and E. coli [81]. The fermentation of black olives in brine with the addition of
500 ppm potassium sorbate (PS) and 1000 ppm sodium benzoate (SB) caused the reduction
of the yeast count, and its effect was stronger than when brine with the half lower level
of PS and SB or the control brine with no preservative were used. The growth of molds
was completely inhibited in all treatments during fermentation [82]. Active starch films
with glycerol and potassium sorbate inhibited the growth of Candida spp., Penicillium spp.,
S. aureus and Salmonella spp. and extended the shelf life of refrigerated cheese by 21% [83].
Antibacterial packaging films, prepared by combining polyethylene (PE) with antibacterial
Int. J. Mol. Sci. 2023, 24, 2457 9 of 34

agents sodium dehydroacetate and potassium sorbate, had excellent physical properties
and inhibited the growth of E. coli more strongly than that it did the growth of S. aureus [84].
No Listeria monocytogenes were detected (8-log reduction) after 24 h of exposure to a
corn zein film with lauric acid (LA) incorporated [85]. In another study, soy-based biofilms,
enhanced with lauric acid (8%, wt/wt) and/or 2.5% pure nisin (4%, wt/wt), were used to
protect the turkey bologna surface [86]. Films containing both LA and nisin completely
eliminated L. monocytogenes from a 106 culture to an undetectable level after 8 h of exposure
at 22 ◦ C, while films with LA alone reduced L. monocytogenes culture from 106 to <102 after
48 h.

3.3. Antimicrobial Peptides and Bacteriocins

Antimicrobial peptides (AMPs) are the host defence oligopeptides composed of five
to over a hundred amino acids produced by various organisms, both prokaryotic and
eukaryotic [87,88]. There are thousands of AMPs. The first AMPs isolated and characterized
were those produced by bacteria, i.e., bacteriocins. Bacteriocins include nisin, leucocin,
lactocin, enterocin, lactococcin A, bifidin, pediocin, while AMPs of other origin include,
among others, cathelicidins, defensins, pleurocidin, LL-37, plectasin, protegrins, cecropins
and magainins [89–92].
Most of the known and studied AMPs are cationic (positively charged) and am-
phiphilic (hydrophilic and hydrophobic) α-helical peptide molecules, but β-sheet structures
can also be present [4]. AMPs are characterized by a broad spectrum of targeted organisms,
protecting against bacteria, fungi, parasites as well as viruses. They act with high efficacy at
very low concentrations and perform synergistic action with antibiotics. Due to their highly
cationic character, AMPs can bind with the negatively charged bacterial cell membranes,
leading to the change of their electrochemical potential. This results in cell membrane dam-
age and the leakage of larger molecules such as proteins, destroying cell morphology and
membranes, and eventually in cell death [88]. For example, cathelicidins are active against
both Gram-negative (E. coli, Salmonella Typhimurium, P. aeruginosa), and Gram-positive
bacteria (S. aureus, L. monocytogenes, B. subtilis). They bind to lipopolysaccharide in the
bacterial cell wall [93]. The antibacterial activity of magainin I molecules which were an-
chored to the gold surfaces was tested against three Gram-positive bacteria (Listeria ivanovii,
Enterococcus faecalis and S. aureus), and the results revealed that the adsorbed magainin I
reduced the adhesion of bacteria to the surface by more than 50%, together with killing the
bacteria that nonetheless adhered to the surface [94].
Despite their name, antimicrobial peptides not only have a bactericidal effect, but they
also inhibit the formation of biofilm, modulate the immune system, and have anti-cancer
or anti-viral properties [89,95,96].
AMPs are quite commonly used in the food packaging industry (Table 3). Depending
on the type of food and the structure of the AMPs, various strategies can be used for this
purpose. For example, AMPs can be directly incorporated into the packaging polymer, can
be coated onto the polymeric surface, or can be immobilized in the polymer [96]. AMPs can
be stabilized by nanoencapsulation using various biopolymers and then can be delivered
to food via head space or via the direct contact of active packaging system with food [92].
Int. J. Mol. Sci. 2023, 24, 2457 10 of 34

Table 3. Examples of AMPs and bacteriocins application in active food packaging systems.

Peptide Polymer/Packaging Material Characteristics References

Nisin
protein isolates, soy protein
isolates, egg albumen and
Nisin and ε-polylysine
Nisin with calcium propionate
Nisin and bacteriocin-like
substance (BLS) P34
Nisin combined with Grape
Seed Extract (GSE) or Green
Tea Extract (GTE)
Bacteriocin KU24
Enterocin 416K
Enterocins A and B
Lactocin 705 and AL705
inserts and anti-microbial
Lacticin 3147 and nisin
significantly inhibited growth of
L. monocytogenes and Salmonella Enteritidis in
PLA [97]
liquid egg white and of E. coli O157:H7 in
orange juice
completely inactivation of L. monocytogenes
PLA [98]
growth in skim milk and liquid egg white
reduction of E. coli O157:H7 count in
PLA [99]
strawberry puree at 22 ◦ C
films with 6 g/100 g nisin completely
Starch–halloysite
inhibited L. monocytogenes on Minas frescal [100]
nanocomposites
cheese surface
inhibited growth of L. monocytogenes and
cellulose [101]
S. aureus in minimally processed mangoes
antimicrobial effect in vitro against S. aureus
cellulose [102]
and L. monocytogenes
reduction of L. monocytogenes level in
plastic film [103]
cold-smoked salmon samples
edible films from whey
inhibitory activity against L. monocytogenes
[104]
strain V7
wheat gluten
decreased growth of microorganism (yeast
and mold, total viable counts, total coliforms
chitosan coating [105]
counts, S. aureus and Pseudomonas spp.) in
fresh-cut carrots
strong inhibitory impact on the growth of
edible zein film coatings L. monocytogenes in film-coated [106]
ready-to-eat chicken
reduced population of L. monocytogenes
liposome-encapsulated during 21 days of storage of Minas frescal [107]
cheese at 7 ◦ C
the greatest inhibitory effect against
L. monocytogenes was observed in turkey
soy protein film [108]
frankfurters coated with film with 1% GSE
and 10,000 IU/mL nisin
in vitro inhibitory effect against
[109]
methicillin-resistant S. aureus
significant decrease in L. monocytogenes
LDPE film [110]
viable counts in frankfurters
in combination with high-pressure
alginate films processing (HPP) reduction of [111]
L. monocytogenes level in sliced cooked ham
in vitro antimicrobial activity against
LDPE Lactobacillus plantarum CRL691 and [112]
Listeria innocua 7
reduced the population of lactic acid bacteria
cellulose-based bioactive in sliced cheese and ham stored in modified
atmosphere packaging (MAP) at
[113]
polyethylene/ refrigeration temperatures; nisin-adsorbed
polyamide pouches bioactive inserts reduced levels of L. innocua
and S. aureus
Int. J. Mol. Sci. 2023, 24, 2457 11 of 34

Table 3. Cont.

Peptide Polymer/Packaging Material Characteristics References

PLA/sawdust particle (SP)
Pediocin
biocomposite film
Pediocins cellulose acetate
PET/polyvinylidene
chloride/retort casting
Plantaricin BM-1
polypropylene (PPR) plastic
multilayer film
multilayer biofunctionalized
Gramicidin A thin films cationic
poly(L-lysine)
Antimicrobial peptide
mitochondrial-targeted PET
peptide 1 (MTP1)
PLA—polylactic acid; CS—chitosan; LDPE—low-density polyethylene; ALG—alginate; PET—polyethylene
terephthalate.
potential inhibition ofL. monocytogenes (99%
of total listerial population) on raw sliced [114]
pork during the chilled storage
inhibiting growth of L. innocua and
[115]
Salmonella sp. on slices of ham
decreased the viable counts of
[116]
L. monocytogenes in meat at 4 ◦ C
inhibitory activity against Enterococcus faecalis [117]
reduction in total aerobic mesophilic bacteria
(APC) and yeasts on ricotta cheese and meat [118]
samples

3.4. Natural Antimicrobial Agents of Plant Origin

It is estimated that there are more than 1300 plants that contain known components
with antibacterial activity against spoilage microorganism and foodborne pathogens. Plant
raw material is very rich in antimicrobial agents because plants naturally synthetize a
wide range of compounds that protect their tissue against microorganisms and other
predators [119]. Therefore, they are used as plant extracts, but also as individual fractions
(purified by various extraction processes) containing mixtures of biologically active ingre-
dients (e.g., essential oils) or as a single compound (e.g., specific polyphenolic compound).
All these compounds are usually also strong antioxidants, therefore there is a great demand
for them in food industry.
Antioxidants in food products play a dual role. First of all, they preserve food,
protecting its ingredients from oxidation and, consequently, from spoilage and changes in
flavor and aroma. However, they might also have a microbiological effect because they
inactivate or inhibit the growth of microorganisms that cause spoilage of the product, as
well as pathogenic microorganisms.
Antioxidants can be enzymatic and non-enzymatic. One of their most important
activities includes the scavenging of free radicals (such as, among others, hydroxyl radical
• OH and the superoxide anion radical O −• ) by donating a single electron (SET) or by
2
hydrogen atom transfer (HAT) [120–123]. Among strong free radical scavengers are mainly
polyphenols and some antioxidant vitamins. Carotenoids are antioxidants known for their
ability to quench the reactive oxygen species (ROS), mainly 1 O2 [124,125]. Antioxidants
include also the enzymes that catalyze the reactions of radical and/or ROS degradation.
Superoxide dismutase (SOD) catalyzes the dismutation of O2 −• , catalase is responsible for
conversion of H2 O2 into H2 O and O2 , while peroxidases catalyze the reduction of H2 O2 ,
organic hydroperoxides and peroxynitrite [126,127]. The antioxidant functions of enzymes
also include the regeneration of glutathione (GSH) from the oxidized form (GSSG) by GSSG
reductase [128]. Vitamin C, apart from its ability to donate a hydrogen atom or electron, can
regenerate other antioxidants, such as α-tocopherol [129–131]. Besides, the activity of some
antioxidants consists of preventing the ROS production, mainly by the chelation of metal
ions (like Fe and Cu), inhibiting in this way the Fenton and Haber-Weiss reactions [130].
The earliest documented and most important examples of plant materials whose dual
effects have been used in industrial practice are essential oils. Essential oils (EOs) are the
secondary metabolites of plants. EOs are a mixture of various volatile organic compounds,
such as ketones (e.g., camphor, carvone), aldehydes (e.g., citral, citronellal, cinnamic alde-
Int. J. Mol. Sci. 2023, 24, 2457 12 of 34

hyde), phenols (e.g., thymol, eugenol, carvacrol, myristicin), esters (e.g., geraniol acetate,
cedryl acetate), ethers or oxides (e.g., eucalyptol, linalool oxide), terpenes (e.g., cymene,
limonene, myrcene), monoterpene and sesquiterpene alcohols (e.g., linalool, citronellol,
geraniol, menthol, nerol, bisabolol), and many other substances [132]. For industrial ap-
plications, EOs are extracted mainly from herbs and spices (rosemary, cloves, horseradish,
mustard, cinnamon, cardamom, anis, sage, onion, garlic, parsley, oregano, basil, marjoram,
savory, thyme, lavender, turmeric, mint, ginger), but also from flowers (jasmine, rose,
violet), and citrus fruit [133–139]. Due to their antibacterial, antifungal, anti-inflammatory,
anesthetic, insecticidal, and antiviral properties, EOs have many applications in medicine,
but also in the cosmetic, perfumery, pharmaceutic, beverage, feed, and food industries.
The use of herbs and spices, as well as EOs extracted from them, for food preserva-
tion is aimed at extending food storage time and shelf life, reducing the number of or
causing the complete elimination of pathogens and improving the overall quality of food
products [133,138,140–143]. Spices and herbs are commonly added to various types of
food products. They can be found in meat and meat products, fish, fruit and vegetable
processing products, but also in cheese, milk, yoghurts and butter, where they inhibit
pathogens (e.g., S. aureus, E. coli O157:H7, C. perfringens, L. monocytogenes, Salmonella sp.,
Shigella sp., Pseudomonas sp., Vibrio parahaemolyticus) and other microorganisms, causing
food spoilage and the significant financial losses associated with it (e.g., Botrytis cinerea,
Alicyclobacillus acidoterrestris, molds) [134,135,137,138,143–145].
In addition to essential oils, other ingredients of plant origin with antimicrobial
potential may also be used in food technology, including polyphenols, terpenoids, alkaloids,
saponins, polyamines, isothiocyanates, thiosulfinates and glucosinolates [119]. It should be
highlighted that some polyphenols and terpenoids are also components of essential oils.
Moreover, the biosynthesis pathways of some of EOs components have common parts with
polyphenols biosynthesis; for example, some stages of the eugenol biosynthesis are the
same as those for p-coumaric acid, caffeic acid and ferulic acid [146].
In the food industry, particular attention is paid to polyphenols, which are usually
divided into 4 or 5 classes: phenolic acids, flavonoids (these include flavonols, flavanones,
flavanols, flavones, anthocyanins and isoflavones), stilbenes, lignans, and sometimes
coumarins [147,148]. Thanks to their structure, polyphenols are ideally suited to act as
antioxidants so they can inhibit the formation of free radicals, quench already formed free
radicals, inhibit initiation and interrupt initiated radical reactions [149–151]. In this way,
polyphenols effectively inhibit lipid peroxidation, the oxidation of proteins and sugars, and
oxidative damage to nucleic acids. Polyphenolic compounds contain numerous hydroxyl
groups, which have an inhibitory effect because they react with the bacterial cell wall, dam-
age the structure of cell membranes and cause the leakage of cell components. Furthermore,
as antioxidants, they inhibit the formation and action of reactive oxygen species (ROS) and
scavenge free radicals, and consequently reduce the oxidation–reduction potential of the
environment or growth medium [139]. Thanks to this, in addition to antioxidant activity,
polyphenols have a bactericidal or bacteriostatic effect against undesirable microorganisms.
Among the most important mechanism of the antibacterial action of polyphenols are: reac-
tions with proteins; inhibition of nucleic acid synthesis by bacterial cells; DNA damage;
interaction with the bacterial cell wall or inhibition of cell wall formation; alteration of cyto-
plasmic membrane function; inhibition of energy metabolism; changes in cell attachment
and inhibition of biofilm formation; and substrate and metal deprivation [148].
Many ingredients of essential oils have already been recognized as safe in the European
Union (e.g., carvacrol, carvone, cinnamaldehyde, citral, eugenol, p-cymene, limonene,
menthol, thymol) and new sources of compounds are still being sought with antioxidant
and/or antimicrobial activity. Still, not all discovered substances, the addition of which to
food could bring measurable benefits, have been checked for their impact on the human
body, for example in terms of toxicity and allergenicity. The values of MIC have not been
assessed for these characteristics, and possible synergistic or antagonistic reactions have
 deprivation [148].
Many ingredients of essential oils have already been recognized as safe in the Euro-
pean Union (e.g., carvacrol, carvone, cinnamaldehyde, citral, eugenol, p-cymene, limo-
nene, menthol, thymol) and new sources of compounds are still being sought with anti-
oxidant and/or antimicrobial activity. Still, not all discovered substances, the addition of
Int. J. Mol. Sci. 2023, 24, 2457 which to food could bring measurable benefits, have been checked for their impact13 onofthe
34
human body, for example in terms of toxicity and allergenicity. The values of MIC have
not been assessed for these characteristics, and possible synergistic or antagonistic reac-
tions
not have
been not been
assessed assessed
with withingredients.
other food other food ingredients.
The necessity Theofnecessity
such studiesof such studies is
is confirmed
byconfirmed
the resultsbyobtained
the results byobtained by various
various authors authors [152–154].
[152–154].
As in
As in the case
caseof ofantioxidant
antioxidantpotential,
potential,thethe
location
location of hydroxyl
of hydroxyl groups is also
groups is of great
also of
importance
great importancefor antimicrobial
for antimicrobial properties. Thymol
properties. (2-isopropyl-5-methylphenol)
Thymol (2-isopropyl-5-methylphenol) andandcar-
carvacrol (5-isopropyl-2-methylphenol)are
vacrol (5-isopropyl-2-methylphenol) areisomers;
isomers;theytheyhave
haveaasimilar
similar molecular structure,
structure,
but the
but the –OH groups in thymol are located in the meta position, and
groups in thymol are located in the meta position, and in carvacrol in the in carvacrol inortho
the
ortho position.
position. DormanDorman and Deans
and Deans [155] [155]
showed showed thathas
that this thisahas a significant
significant impact impact
on theoneffec-
the
effectiveness of action
tiveness of action againstagainst Gram-positive
Gram-positive and Gram-negative
and Gram-negative bacteria,
bacteria, with thymol
with thymol having
having a stronger
a stronger inhibitory
inhibitory effect.
effect. The The presence
presence of thegroup
of the –OH –OH group at the
at the C5 C5 carbon
carbon atom isatom
also
isimportant
also important
for thefor the activity
activity of flavanones
of flavanones and flavones
and flavones against against MRSA (methicillin-
MRSA (methicillin-resistant
resistant
strains ofstrains of S. [139].
S. aureus) aureus) [139]. Esterification
Esterification of the hydroxyl
of the hydroxyl group
group (e.g., in(e.g., in carvacrol
carvacrol methyl
methyl ester) causes changes in the delocalized electron cloud.
ester) causes changes in the delocalized electron cloud. Such a compound loses itsSuch a compound loses its
ability
ability to inhibit the growth of Bacillus cereus [124], and its activity against
to inhibit the growth of Bacillus cereus [124], and its activity against other bacteria is neg- other bacteria is
negligible compared
ligible compared to that
to that of carvacrol
of carvacrol [155].
[155].
The
Theco-occurrence
co-occurrenceof of–OH
–OHgroups
groupsandandthe
theelectron
electrondelocalization
delocalizationsystem
system(in(inthe
theform
form
ofof double bonds in the ring) is important here. The lack of this second “element” ofthe
double bonds in the ring) is important here. The lack of this second “element” of the
molecular
molecularstructure,
structure,as asininmenthol
mentholmolecule,
molecule,prevents
preventsthe thedetachment
detachmentof ofthe
theproton
protonfrom
from
the
the–OH
–OHgroup
groupand andmakes
makesthe theantimicrobial
antimicrobialactivity
activity much
much lower
lower (Figure
(Figure 1).
1). Additionally,
Additionally,
ring
ring rupture weakens the antibacterial potential, which is why geraniolnerol
rupture weakens the antibacterial potential, which is why geraniol and and are much
nerol are
weaker inhibitors of bacterial growth than carvacrol or thymol
much weaker inhibitors of bacterial growth than carvacrol or thymol [139,155]. [139,155].

Figure1.1.Antimicrobial
Figure Antimicrobial activity
activity of aromatic
of aromatic essential
essential oils compounds
oils compounds in dependence
in dependence of their [156].
of their structure struc-
ture [156].
Polyphenols can be used individually or in combination (mixture or as an extract
Polyphenols
ingredient), they actcanonbe used
both individually and
Gram-positive or inGram-negative
combination (mixture
bacteria, or
andastheir
an extract
effec-
ingredient),
tiveness they among
depends, act on both
otherGram-positive
factors, on the and
dose,Gram-negative bacteria,
pH, temperature and their
and oxygen effec-
availabil-
tiveness
ity depends, among
[133,136,137,140,143]. For other factors,
example, on the dose,
xanthohumol, pH, flavonoid
the main temperature and exerted
in hops, oxygen
high antimicrobial activity against Bacteroides fragilis, Clostridium perfringens and C. diffi-
cile [157]. Xanthohumol, naringenin, chalconaringenin and 4-hydroxy-40 -methoxychalcone
inhibited the growth of S. aureus [158], and the presence of at least one –OH group, es-
pecially at the C-4 position, was crucial for that activity. Inactivation of the compounds
might be achieved by the replacement of hydroxyl group by a halogen atom, nitro group,
ethoxy group, or aliphatic groups. Hydroxycinnamic acids induced greater potassium and
phosphate leakage than hydroxybenzoic acids across the membranes of Oenococcus oeni and
Lactobacillus hilgardii [159]. Caffeic acid had a higher antimicrobial activity than p-coumaric
acid due to the additional –OH group attached to the phenolic ring in the molecule [160].
Flavonoids can occur in two forms: free as “aglycons” or in the form of “glycosides”,
and it has been postulated that glycosylation enhances antimicrobial activity, in contrast
to their antioxidant, anti-inflammatory, anticancer and cardioprotective properties [161].
However, it has been demonstrated that flavonoid aglycones, but not their glycosides, may
inhibit the growth of some intestinal bacteria [162]. The study in question revealed that
quercetin 3-O-rutinoside had no inhibitory influence on the intestinal bacteria analyzed, and
even slightly stimulated Lactobacillus spp. growth, whereas its aglycone quercetin exerted a
dose-depended inhibitory effect on intestinal bacteria Ruminococcus gauvreauii, Bacteroides
galacturonicus and Lactobacillus spp. The same was true for flavanones; glycosides naringin
Int. J. Mol. Sci. 2023, 24, 2457
Antimicrobial Agent
Caffeic acid (CA)
Salicylic acid (SA)
Lauric acid (LA)
Ellagic acid
Ferulic acid (FA)
Gallic acid (GA)
Quercetin and Ag NPs
Hydroxyapatite-Quercetin (H-Q)
complexes
Curcumin
Curcumin
Curcumin + ZnO
Fenugreek essential oil (FEO) +
Curcumin
Cinnamaldehyde (CIN)
Resveratrol (RS) and its inclusion
complex (IC) with
hydroxypropyl-γ-cyclodextrin
Resveratrol (RS) and eugenol
(EUG)
Lignin nanoparticles
14 of 34
and hesperidin (flavanone 7-O-glycosides) had no impact, but their aglycones (naringenin
and hesperetin, respectively) inhibited the growth of almost all the bacteria analyzed. The
fact that polyphenols do not only act selectively against undesirable microorganisms should
be considered when food packaging is designed.
Some properties of EOs, such as their volatility, low solubility in water, and sensitivity
to oxidation limit their applications in producing food packaging films. Therefore, various
methods were developed, such as encapsulation or incorporation into polymer matrix, to
eliminate these disadvantages.
Table 4 presents the examples of active packaging in which antimicrobial agents of
plant origin (polyphenols, essential oils, plant extracts) were used for food applications,
together with targeted microorganisms. Although the antibacterial activity was reported in
hundreds of plant species [163], extracts of only a few of them have been used in the design
of active packaging. Plant extracts are complex mixtures that contain a varied range of
components, such as terpenoids, phenolic compounds, alkaloids, glucosinolates, and many
others, so synergistic and antagonistic interactions between the ingredients can be present.
Therefore, plant extracts together with polyphenols and essential oils were presented below
as a large group of antimicrobial agents of plant origin used for food preservation.
Table 4. Examples of antimicrobial agents of plant origin and their application for food preservation.
Polymer/Matrix Characteristics References
Polyphenols
CS-CA postharvest treatment of mulberry fruit improved the
CS [164]
quality and extend the shelf life during cold storage
CS-SA coating inhibited chilling injury better than SA or CS alone
CS [165]
and increased the antioxidant enzyme activities
incorporation LA increased the inhibitory effects against the
spoilage bacteria growth, and LA almost completely protected the
CS edible coating [166]
fresh beef samples against the discoloration after 21 days
of storage
significant reduction of the damage caused by Colletotrichum
candelilla wax [167]
gloesporioides and extended the shelf life of avocado
FA-CS coating effectively extended the shelf life of refrigerated
CS films pork to 7 days; FA-CS had higher antibacterial activity than CS [168]
coatings (reduction of total viable counts by 1–2 log)
coating with CS-GA film improved quality of white button
CS and PE films [169]
mushroom (Agaricus bisporus) in comparison to CS and PE films
PVC-based films with quercetin and AgNPs proved to be highly
PVC-based film effective in inhibiting bacterial growth of food pathogens (E. coli, [170]
S. Typhimurium and L. monocytogenes)
H-Q alginate coatings inhibited the spoilage bacteria (Pseudomonas
ALG edible coatings spp. and Enterobacteriaceae) growth and preserved the quality of [171]
chicken fillets for 11 days at 6 ◦ C
number of bacteria decreased by 1 to 2 logarithmic
PLA-based composite films [172]
cycles
only curcumin-PBAT film showed a slight antibacterial activity
PBAT film [173]
against E. coli and L. monocytogenes (reduction of 1–2 log CFU)
CMC film only ZnO and ZnO/curcumin films had antibacterial properties [174]
good antibacterial and antioxidant properties of
PLA composite film [175]
PLA-FEO-curcumin composite film
monolayer and bilayer films with CIN had bacteriostatic effects to
PLA and starch mono- and bilayer films [176]
E. coli and L. inocua
film based on cellulose derivatives
antimicrobial activity of films against Campylobacter jejuni and
(hydroxyethylcellulose and [177]
Campylobacter coli
cellulose acetate)
increasing the concentration of RS or EUG in CMC films caused an
increase in antimicrobial effects against L. monocytogenes, S. aureus,
CMC films [178]
Salmonella Enteritidis, and E. coli—the antagonistic effects of the
combined use of RES and EUG compared with their use alone
the inhibition of the bacterial growth of Erwinia carotovora subsp.
PVA, CS [179]
carotovora and Xanthomonas arboricola pv. pruni
Int. J. Mol. Sci. 2023, 24, 2457
Antimicrobial Agent
Thymol and montmorillonite
D43B
Thymol (T) and eugenol (E)
Thyme essential oils (TO)
Thymol and
carvacrol
Thyme oil (TEO) and clove oil
(CEO)
Thyme volatile oil (TVO),
chitosan (and chitosan
nanoparticles (CS_NP)
Mediterranean propolis (EEP)
and Thymus vulgaris essential oil
(TV-EOs)
D-limonene nanoemulsion (DLN)
and
D-limonene essential oil (DLEO)
D-limonene
Limonene–liposome
Monolaurin, eugenol, oregano,
and thyme essential oil
Oregano oil (OO)
Origanum vulgare essential oils
(OO) and 1 or 2% of grape seed
extract (GSE)
Carvacrol (CRV)
Clove essential oil
(CEO)—encapsulated in
PLA and polycaprolactone composite films
mesoporous silica nanoparticles
(MSN)
Lemongrass oil, citronella oil and
cajeput oil
Lemongrass oil
15 of 34
Table 4. Cont.
Polymer/Matrix Characteristics References
Essential oils and their ingredients
addition of 8 wt.% thymol and 2.5 wt.% D43B significantly
PLA increased the antibacterial activity against E. coli and S. aureus [180]
8325-4 at all tested temperatures
antibacterial activity, determined using disk diffusion method,
PLA,
proved that PBS/E was the most efficient against E. coli, S. aureus,
poly (adipic acid succinate), and poly [181]
Bacillus tequilensis, B, subtilis and B. pumilis, while against
(succinic acid succinate) (PBS).
Stenotrophomonas maltophilia PLA/E was better
the growth of Alternaria alternata was inhibited significantly by the
β-Cyclodextrin capsule addition and exposure to TO:β-CD as measured by both the agar [182]
dilution and the headspace method
carvacrol and thymol presented a fungistatic effect on
edible starch films [183]
C. gloeosporioides growth on coated mango and papaya
complete killing of S. aureus, that is a reduction from
6.5 log CFU/mL to 0 log CFU/mL, was observed on the 10 wt%
PLA-PBAT film [184]
CEO incorporated composite film; CEO and TEO composite films
inhibited E. coli biofilm by 93.43% and 82.30%, respectively
edible coating of CS, CS_NP and thyme edible CS coating considerably extended the shelf life of basil
volatile oil encapsulated CS_NP leaves, especially TVO-CS_NPs coating (2.4-fold higher shelf life [185]
(TVO-CS_NP) than the control)
EEP showed the best inhibitory effect on S. aureus and
Penicillium sp. (the diameters of the inhibition zones were 12.1 mm
and 11.58 mm, respectively); antimicrobial activity of the films
PLA film [186]
showed that films containing 10 wt% EEP inhibited the growth of
Candida albicans and the combination of EEP and TV-EOs in the
PLA matrix showed a synergistic effect against E. coli
antibacterial activity was proved against S. enterica, E. coli,
- S. aureus and L. monocytogenes, MBC value of DLN was lower than [187]
MBC of DLEO
edible fish gelatin- CS film strong antibacterial activity against E. coli [188]
the limonene–liposome-treated blackberries had significantly
lower visible mold incidence compared to non-coated berries on
ALG coating days 16 and 20 days of storage, whereas the ALG-coated berries [189]
had significantly lower visible mold incidence compared to the
control on the 16th day only
zein films with thyme were not active against the tested
microorganisms; significant inhibitory effect was observed against
zein-based films S. aureus, when oregano, monolaurin, or eugenol were added; [190]
films with eugenol were active against E. coli, Aspergillus niger and
Candida albicans
growth rate of total flora (total viable count) and pseudomonads
sorbitol-plasticized whey protein isolate were significantly reduced by a factor of two with the use of
[191]
(WPI) films OO-films, while the growth of lactic acid bacteria was
completely inhibited
turkey breast meat coated with CS-based coating with GSE and
OO showed reduced count of total viable count,
CS coatings [192]
Enterobacteriaceae, Pseudomonas spp., lactic acid bacteria (and
yeast-mold on day 20 of cold storage
microencapsulated CRV and the non-encapsulated CRV
pectin/sodium
treatments significantly reduced the populations of yeast, mold, [193]
alginate matrix
E. coli and mesophilic aerobic bacteria
coatings with CRV reduced the counts of E. coli and Salmonella
Typhimurium by ~ 5 log CFU/g; Aeromonas hydrophila was
edible coating of cassava starch [194]
reduced by ~ 8 log CFU/g, and S. aureus was reduced by ~2 log
CFU/g on the 7 day of storage of minimally processed pumpkin
CRV, as well as CRV-CS or CRV-GA coatings on poultry reduced
C. jejuni from day 0 through 7 by up to 3.0 log10 CFU/sample;
gum arabic (GA) and cCS [195]
CRV-CS coatings reduced total aerobic counts when compared
with non-coated samples for a majority of the storage times
the higher the content of MSN/CEO/PLA, the better the
[196]
antibacterial effect against E. coli and S. aureus
lemongrass oil was the most effective inhibitor of A. niger (MIC of
10 µL/mL), citronella oil and cajeput oil MIC were of 20 µL/mL,
edible CS or ALG film [197]
with elongation of shelf life of mango coated by ALG film with
lemon grass oil from 10 to 14 days
antimicrobial activity of ALG-based films against E. coli ATCC
ALG edible films 25922 was not dependent on the type of encapsulating agent; [198]
lemongrass oil-ALG film has MIC 0.5%, while ALG film 0.6%
Int. J. Mol. Sci. 2023, 24, 2457
Antimicrobial Agent
Encapsulated citronella oil
(complex coacervation of gelatine
with
carboxymethylcellulose or with
gum Arabic)
Lemongrass oil (LMO)
microcapsules
Citronella essential oil (CEO)
Peppermint (menthol) and
Origanum vulgare (carvacrol)
essential oil
Kaffir Lime Leaf Essential Oil
(Citrus hystrix DC)
Cinnamon-perilla essential oil
(C-PEO)
Cinnamon leaf (CLO) and garlic
oils (GO)
Green tea extract (GTE)
Green tea water extracts (GTWE)
Extract from red grape seeds
(Vitis vinifera) (GSE)
Grape seed extract (GSE)
Flavonoids extracted from
Moringa oleifera seed coat
Paeonia rockii extracts (PPR)
dispersed in chitosan (CS)
Tomato Plant Extract (TPE)
Blueberry fruit and leaf extracts
(BLE)
Pomegranate Peel Extract (PPE)
16 of 34
Table 4. Cont.
Polymer/Matrix Characteristics References
the antimicrobial activity of released citronella vapors on E. coli
paper coatings [199]
and S. cerevisiae (for paper coating 30 g/m2 )
films containing LMO at concentrations of 1250, 2500 and
edible coating based on sodium caseinate
5000 ppm were able to inhibit growth of E. coli ATCC 25922 and [200]
as wall material
L. monocytogenes ISP 65–08 in liquid cultures
the strongest antimicrobial activity was exhibited by the
CS film with ZnO and Ag NPs CS-ZnO-AgNPs membrane, with the inhibition diameter being [201]
~30 mm for S. aureus and E. coli and over 20 mm for C. albicans
increment of menthol and carvacrol concentration in PE film
PE-coated films coating leads to an increase in the antimicrobial activity of films [202]
against E. coli, S. aureus, L. innocua, and S. cerevisiae
edible coating, with the addition of kaffir lime leaves
edible coating with tapioca flour essential oils, decreased the microbial growth (total plate count) of [203]
beef sausage
the shelf life of fresh red sea bream fillets wrapped in tested
edible composite films based on CS_NP [204]
composite film was extended to 6–8 days
good antifungal activity of CLO:β-CD and GO:β-CD
β-cyclodextrin (β-CD) microcapsules [205]
microcapsules against Alternaria alternata
Plant extracts
GTE-CS film inhibited growth of L. inocua and E. coli K12 to
CS film [206]
undetectable levels in tryptic soy broth after 24 h exposure
GTWE inclusion in pork samples reduced growth of mesophilic
CS coating [207]
and psychrotrophic bacteria
GSE-CS film caused strong inhibition of S. aureus, E. coli,
CS film L. monocytogenes, Brochothrix thermosphacta, Acinetobacter guillouiae, [208]
Enterobacter amnigenus, and P. aeruginosa.
edible coatings and films based on CS GSE-chitosan film inhibited growth of L. inocua and E. coli K12 [209]
exhibited antibiofilm potential against S. aureus, P. aeruginosa and
- [210]
Candida albicans.
coatings has antifungal activity and was able to prolong the shelf
polysaccharide gels [211]
life of strawberries to about 16 days
both CS and TPE alone reduced the mesophyll count of sierra
edible CS coating fillets stored on ice during 15 days compared to control fillets, but [212]
a synergistic activity TPE-CS was the best antimicrobial treatment
BLE showed good antimicrobial activity against S. aureus,
CS coatings L. monocytogenes, Salmonella Typhimurium and E. coli, with a [213]
minimum inhibition concentration from 25 to 50 g L−1 .
pure PPE and PPE-loaded CS_NPs effectively retarded the growth
CS_NPs of S. aureus with MIC of 0.27 and 1.1 mg/mL, respectively; E. coli [214]
was not sensitive
CS—chitosan; PE—polyethylene; PVC—poly(vinyl chloride); ALG—alginate; PLA—polylactic acid; PBAT—
poly(butylene adipate-co-terephthalate); CMC—carboxymethyl cellulose; PVA—polyvinyl alcohol; NPs—
nanoparticles.
3.5. Enzymes
Enzymes are increasingly used as antimicrobial agents in food due to their ability to
inhibit bacterial biofilm formation, either direct attacking the microorganisms and/or un-
dergoing catalyzing reactions that result in formation of antibacterial compounds. There are
three main groups of enzymes with antimicrobial and antibiofilm potential (Figure 2) [215].
As was presented by Alves [4], enzymes can act against biofilm using various mecha-
nisms, like the degradation of structures involved in the cell adhesion and attachment to the
surfaces, by the inhibition of biofilm formation, by the detachment of established biofilm or
by increasing its susceptibility to antimicrobials. Among the enzymes with well-known
anti-biofilm potential are lysozyme (responsible for peptidoglycan hydrolysis), amylase
dispersin B (hydrolysis of poly-N-acetylglucosamine), and alginate lyase, being the most
common used, followed by lysostaphin, proteinase K, and DNAse I.
 3.5. Enzymes
Enzymes are increasingly used as antimicrobial agents in food due to their ability to
inhibit bacterial biofilm formation, either direct attacking the microorganisms and/or un-
dergoing catalyzing reactions that result in formation of antibacterial compounds. There
Int. J. Mol. Sci. 2023, 24, 2457 17 of 34
are three main groups of enzymes with antimicrobial and antibiofilm potential (Figure 2)
[215].

Figure 2. Main categories

Figure 2. of antimicrobial
Main categories ofenzymes based
antimicrobial on [215].
enzymes based on [215].

As was presented Lysozyme

by Alves is a[4],
natural antimicrobial
enzymes compound
can act against that can
biofilm hydrolyze
using various β-(1,4)
mech- glycosidic
linkage between N-acetylmuramic acid and N-acetylglucosamine in the peptidoglycan,
anisms, like the degradation of structures involved in the cell adhesion and attachment to
which is the main component of the bacterial cell walls. Therefore, lysozyme can be used
the surfaces, by the inhibition of biofilm formation, by the detachment of established bio-
to inhibit bacterial growth in various food products, especially Gram-positive spoilage
film or by increasing
bacteria itswhich
susceptibility
have more to peptidoglycan
antimicrobials.[216]. Among the enzymes
Lysozyme is used with well-bacterial
to inhibit
known anti-biofilmgrowth potential are lysozyme
in unpasteurized (responsible
beer [217], meat [218],for peptidoglycan
sausage [219], as well hydrolysis),
as in dairy products.
amylase dispersin Hen egg white lysozyme (300 mg/L) was tested for antibacterial activitybeing
B (hydrolysis of poly-N-acetylglucosamine), and alginate lyase, against lactic
the most common acid used,
beer followed by lysostaphin,
spoilage bacteria: Pediococcus proteinase
inopinatus, K,Lactobacillus
and DNAsebrevis, I. L. brevisimilis and
Lysozyme is L. alindneri
natural [220]. It was revealed
antimicrobial that P. inopinatus
compound was the most
that can hydrolyze sensitive,
β-(1,4) while the most
glycosidic
resistant was L. lindneri.
linkage between N-acetylmuramic acid and N-acetylglucosamine in the peptidoglycan,
Lysozyme is the enzyme most commonly utilized in food packaging. Coatings with
which is the main component of the bacterial cell walls. Therefore, lysozyme can be used
lysozyme extended the shelf life of mozzarella cheese [221], ‘coalho’ cheese [222], halloumi
to inhibit bacterial growth in various food products, especially Gram-positive spoilage
cheese [223] or gouda cheese [224], reducing the microorganism level. Edible films, based
bacteria which on have moreand
chitosan peptidoglycan
sodium alginate[216]. Lysozyme
with lysozyme is used tohad
incorporated, inhibit
higherbacterial
inhibitory activity
growth in unpasteurized beer [217], meat [218], sausage [219], as well as in dairy
against the fish spoilage bacteria Pseudomonas fluorescens and Shewanella putrefaciens than the products.
Hen egg white lysozyme
monolayer(300 filmmg/L)
[225]. A was tested for of
combination antibacterial activity against
chito-oligosaccharides (COS)lactic acid
and lysozyme was
beer spoilage bacteria:
effectivePediococcus
against bacteriainopinatus, Lactobacillus
in the meat brevis,resulting
model system, L. brevisimilis and L. lind-
in the complete elimination
neri [220]. It wasof E. coli, P.
revealed fluorescens
that and B.
P. inopinatus wascereus
the and
mosta sensitive,
reduced load while S. aureus.
of the The shelf life of
most resistant
was L. lindneri. minced meat containing COS–lysozyme mixture was extended up to 15 days at chilled
temperatures [226]. The chitosan composite films with 60% lysozyme incorporation had
Lysozyme is the enzyme most commonly utilized in food packaging. Coatings with
enhanced inhibition efficacy against Streptococcus faecalis and Escherichia coli when compared
lysozyme extended the shelf life of mozzarella cheese [221], ‘coalho’ cheese [222], halloumi
to chitosan films [227].
cheese [223] or gouda cheese [224],
Enzymes that can reducing
dissolvethe themicroorganism
established biofilm level.areEdible
DNAse films, based K and
I, proteinase
on chitosan anddispersin
sodium B, alginate with lysozyme incorporated, had higher
but this ability depends on the type of biofilm [228,229]. Dispersin inhibitory ac-
B is an en-
tivity against thezyme
fish that
spoilage bacteria
has ability Pseudomonas
to cleave fluorescens
the polysaccharide and Shewanella putrefaciens
poly-N-acetylglucosamine, a component
than the monolayerof thefilm [225].
biofilm A combination
matrix produced byof chito-oligosaccharides
several Gram-positive bacteria (COS) such and lyso-
as Staphylococcus
epidermidis S. aureus.
zyme was effective against bacteria in the meat model system, resulting in the completedispersin
and Pavlukhina et al. [230] produced the coatings by binding
elimination of E.Bcoli,
to surface-attached
P. fluorescens and polymer matrices
B. cereus and(poly(allylamine
a reduced loadhydrochloride),
of S. aureus. The hydrogel
shelf matrices
and poly(methacrylic
life of minced meat containing COS–lysozyme acid)), proving that they
mixture was inhibited
extended biofilm
up formation
to 15 days(atatleast 98%
reduction) by two strains of S. epidermidis.
chilled temperatures [226]. The chitosan composite films with 60% lysozyme incorpora-
The extracellular DNA plays an important role in the formation and aggregation of bac-
tion had enhanced inhibition
terial efficacyand
biofilm [231,232], against Streptococcus
so DNAse faecalis
I can be used as a and
biofilmEscherichia
inhibitor. coli when
Swartjes et al. [233]
compared to chitosan films [227].
demonstrated that the inclusion of DNAse I in surface coating was effective in enzymatic
Enzymes that can dissolve
cleavage and disruptingthe established biofilm
the extracellular are DNAse
polymeric substancesI, proteinase
produced by K bacteria,
and and
dispersin B, butyielded
this ability
a >90% depends
reduction oninthe type ofPseudomonas
adhering biofilm [228,229].
aeruginosa Dispersin B is an aureus.
and Staphylococcus
Kim et al. [234] demonstrated that DNase I significantly inhibited the biofilm formation by
Campylobacter jejuni and C. coli when isolated from commercially bought raw chickens.
Int. J. Mol. Sci. 2023, 24, 2457 18 of 34

Extracts of Raphanus sativus var. longipinnatus (Brassicaceae plant) combined with

proteinase K have been shown to exert anti-biofilm activity and reduce biofilm formation
by E. coli O157:H7 on stainless steel surfaces [235]. Cellulase and proteinase K were able
to degrade Salmonella Enteritidis biofilms, while proteinase K mixed with chlorine had
the potential to effectively control the Salmonella biofilms formed on the food contact
surface [236]. The combined enzymes of lipase, cellulase and proteinase K inhibited the
biofilm formation by Vibrio parahaemolyticus on different carriers, with the highest inhibition
rate of 59% on nonrust steel plate [237].
Lysostaphin (Lys) catalyzes the hydrolysis of cross-linking pentaglycine bridges of the
cell wall of staphylococci and so it seems to be a good candidate for a antimicrobial agent.
Unfortunately, only several studies have addressed food preservation by this enzyme
application. A lysostaphin-producing strain of Lactobacillus curvatus (Lys+) was used as
starter culture in fermenting sausages contaminated with S. aureus and S. carnosus [238]. The
residual activity of Lys after fermentation was sufficiently high to reduce staphylococci level
by 104 to 105 CFU/g within 2 to 3 days to below the level of detection. Milk from transgenic
cows secreting over 3 µg Lys/mL of milk was pasteurized and then processed into cheese.
Although the quantity and activity of the lysostaphin decreased during cheesemaking, its
level was sufficient to protect against staphylococci in resulting dairy foods [239].

3.6. Lactoferrin
Milk is the first nutritious product consumed by mammals, and therefore it plays
a very important role in the development of the young organism. It owes its valuable
properties to the right proportions of nutrients, but also to the wealth of compounds that
modulate the immune system. Among the numerous protein components of milk, it is
certainly worth to mention alpha-lactalbumin, beta-lactoglobulin, bovine serum albumin,
immunoglobulins, glycomacropeptides, and minor proteins, i.e., lysozyme, lactoperox-
idase, and lactoferrin [240]. The antimicrobial properties of milk are related mainly to
lactoperoxidase, lysozyme, lactoferrin, and immunoglobulins.
Lactoferrin is an iron-binding protein in milk. It therefore has antibacterial properties,
especially against bacteria requiring high iron in the environment (like coliforms) [241].
In food packaging industry lactoferrin was used in combination with various polymers.
For example, lactoferrin-gellan nanoparticles (L:G) were mixed in various proportions
to produce efficient antimicrobial coating [242]. The 9L:1G particles showed a MIC of
0.3 mg/mL against S. aureus, which was six times lower than that of pure lactoferrin
(2 mg/mL). Moreover, fresh strawberries coated with lactoferrin nanoparticles showed the
reduction in the weight loss and growth of mesophilic bacteria during storage, and lactofer-
rin coating in the presence of carboxymethylcellulose (improves nanoparticle adherence
to the fruit surface) extended fruit shelf life up to 6 days. A polyethylene terephthalate
film containing lysozyme and lactoferrin was used to coat salmon fillet samples before
storage up to 4 days at 0 and 5 ◦ C [243]. It was demonstrated that such a coating was
able to decrease H2 S-producing bacteria at longer storage times and higher temperatures
(2.7 instead of 4.7 log CFU/g in the control sample). Edible chitosan film with lactoferrin
with and without lysozyme significantly reduced the growth of E. coli O157:H7 and L. mono-
cytogenes [244]. The functionalized films made from bacterial cellulose with lactoferrin were
used as edible antimicrobial packaging for fresh sausage as a model of meat products [245].
Film had bactericidal activity against E. coli (mean reduction 69% in the films per se versus
94% in the sausages) and S. aureus (mean reduction 97% in the films per se versus 36% in
the sausages).

3.7. Chitosan
Chitosan plays a double role in food packaging, being able to be used both as a matrix
material (polymer) and as a antimicrobial agent. Chitosan is a nontoxic natural antimicro-
bial polymer, produced from chitin, that was approved by GRAS (Generally Recognized as
Safe) by the United States Food and Drug Administration. Chitin is a polymer composed of
Int. J. Mol. Sci. 2023, 24, 2457 19 of 34

β-(1,4)-linked N-acetyl-β-D-glucosamine, and is one of the most abundant natural polysac-

charides (after cellulose). It can be found in the exoskeletons of crustaceans and molluscs
and in insect cuticles as well as in fungal cells [15]. Chitosan is a deacylated derivative of
chitin. Due to its biodegradability, biocompatibility, and low toxicity, good film-forming
ability and high versatility, chitosan has been extensively investigated and is commonly
used in various industries [246–249]. Chitosan-based polymeric materials can be formed
into fibers, films, flakes, gels, powders, sponges, beads or even nanoparticles [15].
It is believed that the polycationic structure of chitosan is a prerequisite for its antibac-
terial activity. The most accepted explanation for chitosan’s antimicrobial activity is that
its positively charged molecule binds to the negatively charged bacterial cell wall, leading
to cell wall damage, altered membrane permeability, osmotic imbalance, and ultimately
cell death [248–251]. It was also proved that chitosan can affect protein biosynthesis and
membrane fluidity, causes an increase in intracellular ROS, damages the integrity of the cell
surface architecture and might be involved in energy metabolism [246]. The antibacterial
potential of chitosan depends on various factors, such as pH, temperature, molecular
weight of chitosan, degree of acetylation, its physical state, concentration, ionic strength,
positive charge density, chelating capacity, as well as environmental and microorganism
characteristics [15,249]. The chitosan-sensitive microorganism include bacteria: E. coli,
S. aureus, Bacillus subtilis, B. cereus, Pseudomonas aeruginosa, P. fluorescens, Salmonella ty-
phimurium, S. choleraesuis, Streptococcus mutans, S. sobrinus, S. sanguis, S. salivarius, Klebsiella
pneumoniae, Enterobacter aerogenes, Xanthomonas campestris, Erwinia carotovora, as well as
yeast and mold (Candida albicans, Aspergillus sp. and many more [15,251]. In order to
enhance the antimicrobial activity of chitosan, various modifications of its molecule were
conducted to obtain chitosan derivatives with higher antimicrobial properties and better
water solubility [248,252]. Chitosan has been proved to also exert antifungal activity; it is
able to inhibit spore germination [253], can change the expression of genes, and can alter
the activity of enzymes responsible for the hyphae growth as well as intracellular ROS
level [254]. The minimal inhibitory concentration of chitosan differs depending on the
microorganism tested, the chitosan form or its derivatization. Values obtained in various
experiments range from 10 ppm (Botrytis cinerea) or 20 ppm (E. coli, S. aureus) to >2000 ppm
for Aspergillus fumigatus and lactobacilli; however, the MIC values can significantly differ,
even for the same tested species if experimental parameters vary [250]. For example, chi-
tosan in a solution form and chitosan nanoparticles were tested against Candida albicans,
Fusarium solani and Aspergillus niger [255]. It has been revealed that nanoparticles of chi-
tosan had stronger inhibitory activity against C. albicans and F. solani than the solution form
(MIC values were from 3 to up to 12 times lower, depending on the microorganism and
the molecular weight of the nanoparticles). Fungal (from champignon) chitosan edible
coatings caused a total growth inhibition of Saccharomyces cerevisiae and Escherichia coli at
concentrations of 1% and 2%, respectively. When edible coatings were applied on fresh-cut
melons, the microbial counts were reduced (up to 4 log CFU/g) [256].
It is worth mentioning that chitosan nanoparticles seem to be efficient antibacterial
agent, even against pathogens with antibiotic resistance. The nanoparticles of chitosan
exerted antibacterial activity against 13 WHO Neisseria gonorrhoeae reference strains, in-
cluding those resistant to multiple antibiotics, with a minimum inhibitory concentration
(MIC90 ) of 0.16 to 0.31 mg/mL and a minimum bactericidal concentration (MBC) of 0.31 to
0.61 mg/mL [257]. Chitosan was also highly effective in inhibiting methicillin-sensitive
S. aureus (MSSA) and methicillin-resistant S. aureus (MRSA) strains, both in planktonic form
and sessile settings, reaching biofilm inhibition percentages as high as 90% for MRSA [258].
Chitosan, extracted from Portunus pelagicus, had antibiofilm activity against MRSA [259].
When edible chitosan membranes were applied to whole cuts of beef and mutton, a de-
crease in both methicillin-resistant S. aureus (x = −1.95 log10 CFU/g) and L. monocytogenes
(x = −1.07 log10 CFU/g) counts were observed [260]. Fatty acids, coated by chitosan
nanoparticles or glycol-chitosan nanoparticles, were proven to be as effective as antibiotics
against both Gram-positive (e.g., S. aureus, S. pneumoniae, Lactobacillus, C. difficile) and
Int. J. Mol. Sci. 2023, 24, 2457 20 of 34

Gram-negative (E. coli, Shigella spp., Salmonella spp.) bacteria [261]. Moreover, Staphy-
locooccus was unable to evolve resistance to ethyl dodecanoate in chitosan, a fact which
was proven in a 6-month experiment. Wasp chitosan nanoparticles showed inhibitory
activity against the growth of beta-lactamase- and carbapenemase-producing Klebsiella
pneumoniae, E. coli, and P. aeruginosa [262]. With all this in mind, chitosan as well as chitosan
nanoparticles can be used as antibacterial agents, with strong effectiveness even against
antibiotic-resistant strains.
Examples of chitosan usage as a coating material in combination with various compo-
nents are also described in other chapters of this review.

3.8. Allyl Isothiocyanate

Allyl isothiocyanate (AIT) is a compound present in the seeds, stem, leaves, and roots
of cruciferous plants, such as mustard, wasabi, and horseradish mustard. It is characterized
by a strong antimicrobial activity in its vapor form against a wide range of spoilage
and pathogenic microorganisms at low concentrations. Hence, it can be used to design
antimicrobial packaging with a slow release of AIT to prolong the shelf life of foods [263].
An antimicrobial sachet, containing AIT encapsulated in calcium alginate beads, was
developed and the effect of released AIT vapor was assessed against E. coli O157:H7 on
spinach leaves [264]. The number of E. coli O157:H7 on spinach leaves (5.6 log CFU/leaf)
decreased by 1.6–2.6 log CFU/leaf at 4 ◦ C and 2.1–5.7 log CFU/leaf at 25 ◦ C within
5 days, and the reduction level was significantly dependent on the relative humidity.
Stickers containing various concentration of AIT were used against the growth of Penicillium
nordicum in pizza at 4 ◦ C for 30 days [265]. All tested levels significantly delay visual
fungal growth in pizza in a dose-dependent manner. The AIT label, in combination with
atmospheric air present in the packaging, extended the shelf life of Danish Danbo cheese
from 4 12 to 13 weeks, while two AIT labels extended the shelf life from 4 12 to 28 weeks [266].
When AIT labels were combined with A modified atmosphere in the packaging, the shelf
life of the cheese was extended from 18 to 28 weeks.
Biodegradable composite films, prepared from polylactic acid (PLA) and sugar beet
pulp, were coated with coating solutions (PLA+AIT or chitosan+AIT). The films signifi-
cantly inhibited the growth of Salmonella Stanley during 24 h of incubation at 22 ◦ C, while
the populations of Salmonella in controls increased [267].

3.9. The Reuterin System (3-Hydroxypropionaldehyde/3-HPA/, 3-HPA Dimer, Acrolein, HPA

Hydrate, and 3-Hydroxypropionic Acid)
Reuterin (3-hydroxypropionaldehyde, 3-HPA) is an intermediate formed by Limosi-
lactobacillus reuteri during the coversion of glycerol into 1,3-propanediol, with interesting
antimicrobial activity against bacteria, fungi or even viruses [268–270]. In fact, it is not
one compound, but a whole set, the so-called the reuterin system, which includes 3-HPA,
3-HPA dimer, acrolein, HPA hydrate, and 3-hydroxypropionic acid (Figure 3).
Antimicrobial activity of the reuterin system is not clear, but its proposed mechanisms
involve the induction of oxidative damage in cells by acrolein and 3-HPA (due to depletion
of free –SH groups in proteins like glutathione) or DNA synthesis inhibition due to the
competitive inhibition of ribonucleotide reductases by 3-HPA dimers [269].
 3.9. The Reuterin System (3-Hydroxypropionaldehyde/3-HPA/, 3-HPA Dimer, Acrolein, HPA
Hydrate, and 3-Hydroxypropionic Acid)
Reuterin (3-hydroxypropionaldehyde, 3-HPA) is an intermediate formed by
Limosilactobacillus reuteri during the coversion of glycerol into 1,3-propanediol, with
interesting antimicrobial activity against bacteria, fungi or even viruses [268–270]. In fact,
Int. J. Mol. Sci. 2023, 24, 2457 21 of 34
it is not one compound, but a whole set, the so-called the reuterin system, which includes
3-HPA, 3-HPA dimer, acrolein, HPA hydrate, and 3-hydroxypropionic acid (Figure 3).

acrolein 3-HPA
hydrate

3-HPA 3-hydroxypropionic
dimer acid
Figure3.3.The
Figure Thereuterin
reuterinsystem
systembased
basedon
on[269,271].
[269,271].

Antimicrobial
Therefore, activity
reuterin has of the large
quite reuterin systemasisan
potential notantimicrobial
clear, but itsagent
proposedin themecha-
food
nisms involve
industry. Purifiedthereuterin
inductionandof oxidative
the reuterin damage in cells by acrolein
system compounds have been and 3-HPA
already (dueto
added to
depletion
various of free
kinds –SHproducts
of dairy groups in proteins
and they werelike proved
glutathione) or DNA
to inhibit synthesis
the growth inhibition
of pathogens
or
duespoilage microorganisms.
to the competitive For example,
inhibition reuterin produced
of ribonucleotide reductases bybyLactobacillus
3-HPA dimers reuteri[269].
ATCC
53,608Therefore,
was an effective
reuterinprotectant
has quite against Escherichia
large potential as ancoliantimicrobial
DH5α, Salmonellaagententerica
in the food subsp.
in-
enterica, Listeria monocytogenes,
dustry. Purified reuterin and theStaphylococcus
reuterin system aureus and Penicillium
compounds have been expansum without
already added to
changing the quality
various kinds parameters
of dairy products(pH, and acidity,
they were soluble
provedsolids, colour,the
to inhibit andgrowth
rheological aspects)
of pathogens
of
orthe fermented
spoilage milk product
microorganisms. For[272]. The antimicrobial
example, reuterin produced effectbyofLactobacillus
reuterin combined
reuteri ATCCwith
diacetyl against E. coli O157:H7, Salmonella Enteritidis, L. monocytogenes
53,608 was an effective protectant against Escherichia coli DH5α, Salmonella enterica subsp. was demonstrated
in fermented
enterica, milkmonocytogenes,
Listeria [273]. Langa etStaphylococcus
al. [274] studied cheeses
aureus and artificially
Penicilliumcontaminated
expansum without with
L.changing
monocytogenes and E.parameters
the quality coli O157:H7 (pH,and then ripened
acidity, soluble for 30 days.
solids, Both
colour, andpathogens
rheological wereas-
not detected
pects) of theinfermented
25 g cheese made
milk with reuterin-producing
product [272]. The antimicrobial L. reuteri INIA
effect P572 andcombined
of reuterin 100 mM
glycerol from day
with diacetyl 7 onwards,
against which means
E. coli O157:H7, that in
Salmonella situ production
Enteritidis, of reuterinwas
L. monocytogenes maydemon-
be an
interesting alternative to improve cheese safety instead of reuterin addition
strated in fermented milk [273]. Langa et al. [274] studied cheeses artificially contaminated to food.
withAs L. mentioned
monocytogenes above,
andreuterin can be added
E. coli O157:H7 directly
and then to liquid
ripened food
for 30 or produced
days. Both pathogensin situ
by properly selected strains. However, attempts are also being made
were not detected in 25 g cheese made with reuterin-producing L. reuteri INIA P572 and to add it to coatings
or
100food
mMpackaging.
glycerol fromPectin-based edible which
day 7 onwards, coatings with that
means lemon essential
in situ oils and
production ofreuterin
reuterin
decreased viable Penicillium spp. conidia counts in strawberries
may be an interesting alternative to improve cheese safety instead of reuterin addition[275]. In meat and fish
to
products,
food. the reuterin system decreased the level of foodborne pathogens, e.g., Pseudomonas
spp., L.Asmonocytogenes,
mentioned above, Salmonella Enteritidis,
reuterin E. coli O157:H7
can be added directly [269]. Preservation
to liquid of sea bass
food or produced in
fillets active sodium alginate film with Limosilactobacillus
situ by properly selected strains. However, attempts are also being made to add(source
by coating with reuteri of
it to coat-
reuterin) and glycerol showed good antibacterial activity and caused delayed proliferation
ings or food packaging. Pectin-based edible coatings with lemon essential oils and reu-
of the main spoilage microorganisms [276]. The impact of post-chill spray of the mixtures of
terin decreased viable Penicillium spp. conidia counts in strawberries [275]. In meat and
reuterin, microcin J25, and lactic acid on the viability of Salmonella enterica and total aerobes
fish products, the reuterin system decreased the level of foodborne pathogens, e.g., Pseu-
on broiler chicken carcasses were also studied [277]. The results proved that a mixture of
domonas spp., L. monocytogenes, Salmonella Enteritidis, E. coli O157:H7 [269]. Preservation
reuterin and lactic acid, sprayed onto chilled chicken carcasses, was more efficient in the
of sea bass fillets by coating with active sodium alginate film with Limosilactobacillus reuteri
reduction of Salmonella spp. counts than the mixture of reuterin and microcin J25. Taking
(source of reuterin) and glycerol showed good antibacterial activity and caused delayed
into account recent studies, reuterin in combinations with various bacteriocins (microcin
proliferation of the main spoilage microorganisms [276]. The impact of post-chill spray of
J25, nisin, pediocin) seems to be a promising strategy for extending the shelf life of various
food products [278–281].

3.10. Bacteriophages
One of the most interesting but still somewhat controversial ideas is the use of bac-
teriophage for the development of future packaging materials. Bacteriophage is a virus,
so it must find a host through which could replicate in the environment, meaning that
coatings with bacteriophage must have a physical contact with food. The whole cycle of
Int. J. Mol. Sci. 2023, 24, 2457 22 of 34

bacteriophages includes finding the host, infecting it, replication of bacteriophage, bacteria
lysis and the release of new phage particles into the medium. There are many commercially
available bacteriophage products that are ‘generally recognized as safe’ and approved
by European Food Safety Authority (EFSA) or the US Food and Drug Administration
(FDA) [282,283].
Cellulose acetate films, incorporated with a solution of bacteriophages (BFSE16,
BFSE18, PaDTA1, PaDTA9, PaDTA10 and PaDTA11), showed antimicrobial activity against
Salmonella enterica subsp. enterica serovar Typhimurium ATCC 14028 [284]. The antimicro-
bial activity of the film was shown in both the diffusion method in a liquid medium and in
the method of solid medium diffusion, respectively, when incubated at 35 ◦ C, evidenced by
the growth curve and the inhibition formed.
The commercially available anti-Listeria phage preparation LISTEX™P100 was used
for limiting the growth of L. monocytogenes on ready-to-eat roast beef and cooked turkey
in the presence or absence of the chemical antimicrobials potassium lactate and sodium
diacetate [285]. LISTEX™P100, applied without chemical antimicrobials, was an effec-
tive inhibitor and caused the reduction of L. monocytogenes of 2.1 log10 CFU/cm2 and
1.7 log10 CFU/cm2 (respectively, for cooked turkey and roast beef) during incubation at
4 ◦ C. In turkey samples cooked and stored at 10 ◦ C, both with and without chemical antimi-
crobials, the phage-treated samples had significantly lower numbers of L. monocytogenes
when compared to the untreated controls throughout the 28-day storage period. In other
studies, bacteriophages and their enzymes endolysins were used for the growth control
of L. monocytogenes, E. coli O157:H7, Shigella sonei, P. aeruginosa, S. aureus, S. epidermidis,
Campylobacter jejuni, Bacillus cereus, and Salmonella spp. [282,283,286].

4. Challenges and Possible Directions of Development in Food Packaging Industry

Food packaging is becoming more and more important. By protecting the product, it
supports the producer’s interests, while by maintaining the quality and safety of food it
also protects the health of the consumer. On the other hand, the impact of packaging on the
environment has also become important. Apart from the aspect of reuse or biodegradability,
food packaging allows producers to extend the shelf life, thus preventing food waste or
the throwing away of still-good products. Taking into account that food is not always
kept in the right conditions (e.g., cold chain, exposure to light) or that packaging can be
damaged or unsealed, it may happen that, although the shelf life has not been reached, the
product is unsuitable for consumption (it will spoil or harmful metabolites and pathogens
develop). In the authors’ opinion, food producers should strive to create such innovative
food packaging that will actively support the extension of the shelf life, but at the same
time will provide more and more accurate information about the condition of the product.
Perhaps in the future, instead of a specific date on the food packaging, there will be an
inscription “fit for consumption as long as the indicator is blue” or “do not consume if the
indicator turns red”, etc. Additionally, ideally, monitoring and signalling would not be
conducted in a zero–one way, but the colour or other scale used for the marking will be
more complex (multi-level). In such a way, the kinetics of changes could also be observed,
including, for example, that the expiration day is approaching, after which the consumption
of a given product will be risky or even dangerous.
The direction of further development will certainly be the search for new antimicrobial
agents that can be effectively used to design active packaging, and more precisely, to find
out which of the currently known ingredients are suitable for this purpose. The sheer
number of plant species showing antimicrobial activity [163] prompts the search in this
area. Examples of interesting plants include: Haplophyllum tuberculatum [287], Caryophyllus
aromaticus and Syzygyum joabolanum [288], Phytolacca dodecandra fruits, Rumex nepalensis
leaves, Grewia ferruginea [289], as well as Sambucus nigra, Murraya koenigii, Murraya koenigii
or Mikania glomera, which are characterized by low MIC values against bacteria [163].
It is also worth paying attention to one more issue. Some foodborne pathogens (E. coli,
S. aureus, C. perfringens) can enter the bloodstream, causing bacteraemia (the presence
Int. J. Mol. Sci. 2023, 24, 2457 23 of 34

of microorganisms in bloodstream) or sepsis (defined as “a life-threatening organ dys-

function caused by dysregulated host response to infection” [290]). Depending on the
methodology used, between 4.6% (explicit sepsis codes) and 33.2% (Angus sepsis codes)
of hospitalizations were sepsis related to a foodborne pathogen [291]. Among the main
causes of hospitalizations, the authors listed: Clostridium perfringens, Vibrio vulnificus, Vibrio
parahaemolyticus, V. cholerae, Listeria monocytogenes, Cyclospora cayetanensis and noroviruses,
but important were also E. coli (STEC, diarrheagenic, enterotoxigenic), Streptococcus spp.
group A, Salmonella typhi and non-typhoidal, Campylobacter sp., Shigella spp., and Yersinia en-
terocolitica. It is known that T. gondii can be acquired by humans in several ways, such as the
ingestion of contaminated food, drinking water containing the oocyst, but also by contami-
nated blood transfusion or organ transplantation, or by transplacental transmission [1]. This
means that some foodborne pathogens can also cause BAI or HAI. Considering that these
infections are largely caused by the same or related species, the presence of which is also a
challenge in food, substances suspected of having antimicrobial activity can be—before
being used in the human body—tested in another research model, which is food packaging.
In this case, it is also important that the antimicrobial agent, after being combined with
the packaging matrix, maintains its ability to inhibit the growth of microorganisms and to
limit the formation of biofilm, while not affecting the quality characteristics of food and its
safety for the consumer. Therefore, all antimicrobial agents mentioned in this review are
good candidates to be “medical” antimicrobial agents and to be applied in the future for
biomaterial coating.

Author Contributions: Conceptualization, A.D.-C. and T.T.; writing—original draft preparation,

A.D.-C., T.T. and K.P.-P.; writing—review and editing, A.D.-C., T.T. and K.P.-P.; visualization, A.D.-C.,
T.T. and K.P.-P.; supervision, A.D.-C. All authors have read and agreed to the published version of
the manuscript.
Funding: This research received no external funding.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.

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