Academic Editor: Lluís Ribas
de Pouplana
Received: 19 December 2024
Revised: 24 January 2025
Accepted: 27 January 2025
Published: 28 January 2025
Citation: Rodríguez-Marca, C.;
Domenech-Coca, C.; Nakamura, M.;
Ortega-Olivé, N.; Puigbò, P. Use of
Live Biopreservatives and
Bacteriophages to Enhance the Safety
of Meat Products. Life 2025, 15, 197.
https://doi.org/10.3390/
life15020197
Copyright: © 2025 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/).
Review
Use of Live Biopreservatives and Bacteriophages to Enhance the
Safety of Meat Products
Cristina Rodríguez-Marca 1 , Cristina Domenech-Coca 2 , Miho Nakamura 3,4,5, Nàdia Ortega-Olivé 6
and Pere Puigbò 1,7,8,*
1 Nutrition and Health Unit, Eurecat, Technology Centre of Catalonia, 43204 Reus, Catalonia, Spain;
cristina.rodriguez@eurecat.org
2 Department of Biochemistry and Biotechnology, University Rovira i Virgili, 43007 Tarragona, Catalonia, Spain;
cristina.domenech@urv.cat
3 Medicity Research Laboratory, Faculty of Medicine, University of Turku, 20520 Turku, Finland;
miho.nakamura@utu.fi
4 Institute of Biomaterials and Bioengineering, Tokyo Medical and Dental University, Tokyo 101-0062, Japan
5 Graduate School of Engineering, Tohoku University, Sendai 980-8577, Japan
6 Department of Food Technology, Engineering and Science, University of Lleida, 25198 Leida, Catalonia, Spain;
nadia.ortega@udl.cat
7 Department of Biology, University of Turku, 20014 Turku, Finland
8 Department of Animal and Food Science, Autonomous University of Barcelona,
08193 Bellaterra, Catalonia, Spain
* Correspondence: pepuav@utu.fi
Abstract: Critical health considerations for both raw and processed meats include address-
ing bacterial spoilage and ensuring safety. Nitrites and nitrates are widely used in the
meat industry to enhance color and flavor and extend shelf life. However, health concerns
linked to their use make reducing nitrites and nitrates in meat production a significant
challenge with potential benefits for both the food industry and consumer health. This
challenge has been addressed with the use of biopreservatives, i.e., substances extracted
from natural sources or produced by fermentation that can enhance food quality and safety.
In this article, we assess the use of live biopreservatives (LBs), defined here as microorgan-
isms that produce antimicrobial substances that can be used to preserve and extend the
shelf life of food. Moreover, the potential synergistic effects of LBs with bacteriophages
and biodegradable food packaging for meat is also explored. This innovative combina-
tion offers a comprehensive approach to meat preservation, enhancing both microbial
control and sustainability. Overall, the inclusion of LBs extends the shelf life of meat prod-
ucts through bacteriostatic mechanisms, whereas bacteriophages offer direct (lytic) action
against pathogens. Enhancing meat preservation and safety with mixed microbe-mediated
strategies requires deeper empirical and theoretical insights and further revision of laws
and ethical considerations.
Keywords: live biopreservatives; bacteriophages; lactic acid bacteria; meat safety; nitrites;
bacteriocins; natural antimicrobials; biodegradable food packaging; foodborne pathogens;
environmental microbes; microevolutionary models; interaction models
1. Introduction
The food industry faces a significant challenge in controlling foodborne pathogens
in meat and meat products [1]. Bacterial spoilage and safety are critical health concerns
for both raw and processed meats. [2,3]. The use of nitrites and nitrates as preservatives
in the meat industry has been associated with various functions, such as enhancing color
Life 2025, 15, 197 https://doi.org/10.3390/life15020197
Life 2025, 15, 197 2 of 14
and flavor and extending shelf life [1]. However, there are concerns regarding their safety
due to the formation of potentially carcinogenic compounds called N-nitrosamines [1]. The
European Food Safety Authority (EFSA)’s 2023 report warned about the potential DNA-
damaging and carcinogenic effects of 10 nitrosamine substances that are predominantly
present in processed and cured meat products [4]. However, technological changes are
necessary to eliminate nitrates/nitrites and ensure the production of high-quality salamis
while lowering the ripening temperature to ensure microbiological safety [5]. Thus, natural
sources and new technologies are being explored to develop alternative additives [6].
Reducing the use of nitrites and nitrates in meat production is a major challenge that
should substantially benefit the food industry and consumer health.
It has been suggested that these challenges can be successfully addressed by strate-
gically utilizing biopreservatives [1], i.e., natural substances derived from bacteria, fungi,
plants or animals that can be used to extend the shelf life of food products while guar-
anteeing their safety [7]. Natural preservatives offer a promising alternative to combat
microbial resistance [8] and mitigate the adverse effects associated with certain synthetic
compounds [7]. Moreover, studies on the mechanism of action, safety evaluations, and the
establishment of appropriate regulations for natural preservatives are needed before they
are used [1].
2. Antimicrobial Innovations
Antimicrobial innovations encompass the development and implementation of novel
strategies, compounds, and technologies designed to prevent and control the growth of
harmful microorganisms [9]. With the rise of antimicrobial resistance, these innovations
are essential to protect people’s health, extend products’ shelf lives, and reduce reliance
on traditional antibiotics [10]. Research on antimicrobial innovations calls for actions
that offer a comprehensive assessment of novel biopreservation strategies, such as plant-
based extracts, peptides, nanomaterials, and bacteriophages [11]. Each of these approaches
offers unique advantages in addressing microbial threats and combating the spread of
antimicrobial resistance. This review aims to provide guidance regarding all aspects
that should be thoroughly addressed in the special issue. In this article, we provide an
overview of the critical intersection between live biopreservatives (LBs), bacteriophages,
and biodegradable food packaging for meat biopreservation, addressing key considerations
from scientific, technological, and socioeconomic perspectives. Furthermore, we provide a
theoretical and a mathematical model that can be used as guidance to quantify the effect of
LBs and bacteriophages in real-world scenarios.
3. Use of Biopreservatives to Enhance Food Quality and Safety
Biopreservatives are antimicrobial substances extracted from natural sources or pro-
duced by fermentation that enhance food quality and safety [12]. Bacteriocins are antimi-
crobial peptides produced by microorganisms like LAB to gain a competitive advantage in
their environment, while being harmless to the producing strains [13]. These biologically
active compounds include peptide structures that exhibit antimicrobial activity [14]. Bacte-
riocins are naturally produced by certain bacterial strains; however, their use as additives
may be limited by their high cost and limited effectiveness against specific pathogenic
microorganisms in the final product [1]. Among all the purified bacteriocins tested thus
far, nisin (E-234) stands out as the only one permitted as a food additive in the European
Union, as stated by the European Commission in 2011 [1,7]. Furthermore, Pediocin PA-1, a
crude extract obtained from bacteriocin-producing strains, is also available commercially.
Although nisin has relatively broad-spectrum antimicrobial properties, including the in-
hibition of the overgrowth of sporulated bacteria, these agents are not effective in meat
Life 2025, 15, 197 3 of 14
production [2]. Bacteriocins and bacteriocin-producing bacteria have been proposed for
use as potential natural preservatives [15]. However, the major effect of bacteriocins on
food is obtained when they are combined with other preservation techniques [16] such
as the use of essential oils (EOs). A wide range of EO applications, such as incorporation
in active packaging or edible films and direct encapsulation, have been described [16].
The combined use of EOs and bacteriocins or bacteriocin-producing bacteria can not only
improve food safety and the shelf lives of meat and meat products, but also stabilize sensory
and nutritional quality [16,17].
4. Live Biopreservatives (LBs)
LBs are defined here as microorganisms capable of secreting antimicrobial substances
that can be utilized as a way of preserving food and extending its shelf life (Figure 1). LBs
have a bacteriostatic effect; i.e., they prevent the growth of bacteria. LBs possess diverse
characteristics beyond their antimicrobial properties [18]. These include minimal potential
for pathogenic activity, in compliance with regulatory protocols; limited impact on product
characteristics, such as sensory attributes and color [18]; biopreservation action spanning a
wide range of temperatures and pH levels [14,19]; and versatile activity against a spectrum
of microorganisms [20]. LBs also exhibit resistance through mechanisms of action that
impede microbial adaptation [21]. Furthermore, LBs may have a lower impact on the
gut microbiota than synthetic preservatives [22]. The potential for a decreased impact on
beneficial bacteria in the gut makes LBs an attractive option for those concerned about the
effects of food preservatives on overall gut health.


	

	

	













	










	





	
	

	

		
	

	
	
Figure 1. Main characteristics of live biopreservatives (LBs). LBs are microbial strains that can be
utilized as food preservatives because they have the capacity to secrete antimicrobial substances
(such as lactic and acetic acids and bacteriocins). Furthermore, they need to present the following
desirable characteristics: antimicrobial properties; absence of pathogenicity elements; limited impact
on product characteristics; biopreservation action spanning a wide range of temperatures, water
activity and pH levels; and lower impact on the gut microbiota.
5. Lactic Acid Bacteria to Enhance Meat Product Safety
Research on meat protection has focused on the selection of lactic acid bacteria (LAB)
that do not cause meat spoilage prevents meat spoilage and that enhance product safety [2].
However, the use of LAB is controversial because they can either contribute to spoilage
Life 2025, 15, 197 4 of 14
(through the generation of harmful metabolites and the subsequent organoleptic down-
grading of meat) or act as stabilizing agents that prevent the growth of other harmful
bacteria [23]. In addition to LAB, Pseudomonas species have been suggested to be key
organisms for consideration when evaluating the impact of specific chilling durations and
temperature conditions on the growth of spoilage bacteria [24]. Certain probiotics have
also been utilized as LBs in the preservation of fermented meat products [25–27].
LAB protect meats by outcompeting other microorganisms and producing inhibitory
substances such as lactic and acetic acids and bacteriocins [2,13,28,29]. LAB-preservation
methods, such as spray drying, are effective methods that allow products to be obtained
at a low cost compared with lyophilization [30]. Ongoing scientific efforts have led to the
emergence of a growing market for food products that incorporate protective cultures to
enhance food safety and extend shelf life [31]. Notably, the Lactobacillales order, which
includes various genera, plays a significant role in this context; for example, a starter
culture (Latilactobacillus sakei CTC494) has been investigated for its ability to protect against
Listeria monocytogenes in two varieties of chicken-based dry-fermented sausages during
fermentation and ripening [32]. Moreover, mathematical models are also utilized in anti-
Listeria biopreservation strategies [33].
Several studies have tested the effectiveness of combining different LAB or their
metabolites with novel preservative techniques, such as gas packing, natural extracts, es-
sential oils, and bacteriocins, to create more selective and defensive systems to overcome
pathogenic and spoilage microorganisms [34] while also enhancing the physicochemical
and sensory properties of the bacteria. Specifically, mixed methods, such as the adoption of
higher concentrations of bioprotective starter cultures along with a low sugar concentration,
have been shown to improve textural features and reduce oxidation levels in salamis sub-
jected to challenges with Listeria innocua, Salmonella enterica, and Clostridium botulinum [5].
Furthermore, LAB can promote the decomposition of proteins and lipids to produce flavor-
precursor substances such as free amino acids or free fatty acids, give food a unique flavor,
and have a certain positive impact on the overall flavor of the finished product (they thus
yield flavor improvement along with product preservation and, ultimately, the extension of
shelf life [35]). LAB play a key role in the production of fermented meat products, resulting
in texture and flavor improvement, product preservation and, ultimately, the extension of
shelf life [34].
6. Bacteriophages to Enhance Food Safety
Bacteriophages, also known as phages, are the most abundant organisms in the
biosphere and replicate in bacterial cells [36]. Bacteriophages are naturally occurring
predators of bacteria and are presumably harmless to humans and animals; thus, they
have been utilized as antimicrobial agents in food products [37–41]. Lytic bacteriophages,
with their strict host specificity, are promising biocontrol agents for specifically targeting
and controlling pathogenic bacteria in the food industry [42]. Moreover, natural and
genetically modified bacteriophages may also play a future role in meat preservation [15],
preventing pathogen growth and increasing shelf life. The role of bacteriophages includes
the control of the following bacterial pathogens: (i) L. monocytogenes: bacteriophages have
recently been used commercially for pathogen biocontrol in the food industry (in meat and
poultry products) [37]; (ii) Salmonella sp.: bacteriophages have also been used in ground
beef [43], ground pork [43] and fresh-cut fruits under different storage temperatures; and
(iii) Shiga toxin-producing Escherichia coli (STEC) [44]. Furthermore, bacteriophages have
been employed as effective biomarkers for fecal contamination at various processing stages
in poultry and beef [38].
Life 2025, 15, 197 5 of 14
While recent advancements in biological interventions suggest bacteriophages have
potential for use as effective microbial regulators, further research is needed to optimize
their use as antimicrobial agents [38]. This includes determining the threshold level of hosts
necessary for efficient bacteriophage replication and identifying the optimum temperature
conditions that enhance bacteriophage performance [38]. A common practice in phage
therapy, i.e., the use of phages for the treatment of pathogenic bacterial infection, is to
use phage therapy in combination with antibiotics that can interact synergistically [45].
A parallel approach would involve incorporating bacteriophages into mixed strategies
for food preservation, combining the use of LBs to prolong shelf life through bacterio-
static mechanisms with the use of bacteriophages to provide direct (lytic) action against
pathogens. However, a more comprehensive empirical and theoretical understanding of
microbe-mediated strategies, specifically of those incorporating LBs and bacteriophages, is
necessary to effectively address challenges in meat preservation.
7. Biodegradable Food Packaging for Meat Products
Spoilage control in meat and meat products presents a critical challenge in the food
industry due to the high water content, water activity, pH, and nutrient availability in
meat [34]. For this reason, specific packaging materials are needed to ensure food safety and
product stability in terms of water content and oxidation. Maintaining color stability in fresh
meat requires gas-permeable films, whereas gas-barrier films are preferred to prevent the
oxidation of fats and proteins in meat products [46]. However, petroleum-derived materials
used for meat packaging are becoming obsolete because they are not environmentally
sustainable since the European Union (EU) adopted the strategy of making all plastic
packing recyclable or reusable in the EU market by 2030. This strategy aims to protect
our environment and reduce marine litter and greenhouse gas emissions, and dependence
on fossil fuels is considered a key element of Europe’s transition to a carbon-neutral and
circular economy [47].
Furthermore, environmental awareness has increased in recent years. Consumers
demand new ecologically friendly and sustainable ways of preserving food [1]. In this
context, biodegradable materials obtained from natural sources represent an alternative
in meat production and packaging, although they do not have sufficient mechanical and
barrier properties. These biopolymers can be categorized based upon the source of the
material. The most well-known natural biopolymers are polysaccharides and proteins
such as starch, cellulose, chitosan, and gelatin [46]. On the other hand, some synthetic
biopolymers, e.g., polyhydroxyalkanoate (PHA) and polyhydroxybutyrate (PHB), can be
generated by fermentation, while others, such as polylactic acid (PLA), can be produced by
chemical synthesis from biomass [46].
Current research on natural biopolymers is focused on developing biodegradable
materials from agrifood byproducts as a strategy to reduce environmental impact and
contribute to a circular economy. This provides an opportunity for agricultural and food
industries to revalorize byproducts and create added value in the market. Although
biodegradable food packaging for meat [48–50] has been used for a long time, the use of
microbial biopreservatives [23] represents an unexplored challenge in the field. In this
context, LAB are highly interesting due to their ability to be encapsulated by extrusion to
produce antimicrobial films [20]. Therefore, as soon as biodegradable materials overcome
current technical limitations, additional studies will be needed to validate the integration of
microbial bioprotectors in emergent materials. Furthermore, to the best of our knowledge
there are no studies that integrate the use of LBs and bacteriophages with that of upcycled
agrifood for packaging.
Life 2025, 15, 197 6 of 14
8. Future Perspectives
To address future challenges in the use of LBs and bacteriophages, both short-term
and long-term goals are essential. In the short term, the application of LAB and bacteriocins
is ready for real-world scaling (Section 8.2). In the mid-term, computational models are
required to evaluate the implementation of bacteriophages in the meat industry (Section 8.2)
and assess the potential impact of LBs and bacteriophages on the global antimicrobial-
resistance crisis (Section 8.3). In the long term, the integration of LBs and bacteriophages
into various types of biodegradable packaging (Section 8.4) and the evaluation of their
economic and environmental impacts are key objectives.
8.1. Scaling Up: Use of LAB and Bacteriocins in Real Meat Products
The future integration of LAB and bacteriocins in real meat products holds substantial
promise for revolutionizing the food industry (Figure 2C). Despite their proven efficacy
in in vitro settings, the full potential of these natural antimicrobial agents in real meat
applications remains largely untapped due to complexities inherent in meat matrices [1].
The approval of nisin as a purified bacteriocin additive in European and US food regu-
lations [1] and the commercial availability of crude extracts, such as Pediocin PA-1 from
bacteriocin-producing strains, highlights its potential as an effective biopreservative [51,52].
Life 2025, 15, 197 6 of 15 
 
 
knowledge there are no studies that integrate the use of LBs and bacteriophages with that 
of upcycled agrifood for packaging. 
8. Future Perspectives 
To address future challenges in the use of LBs and bacteriophages, both short-term 
and long-term goals are essential. In the short term, the application of LAB and bacterioc-
ins is ready for real-world scaling (Section 8.2). In the mid-term, computational models 
are required to evaluate the i plementation of bacteriophages in the meat industry (Sec-
tion 8.2) and assess the potential impact of LBs and bacteriophages on the global antimi-
crobial-resistance crisis (Section 8.3). In the long term, the integration of LBs and bacteri-
ophages into various types of biodegradable packaging (Section 8.4) and the evaluation 
of their economic and environ ental impacts are key objectives. 
            
  i tegration of LAB and bacteriocins in real meat products holds substan-
tial promise for revolutio izing the foo  industry (Figure 2C). Despit  their proven effi-
cacy in in vitro settings, the full po ential of th se natural antimicrobial agents in real  
       t i  t i s [ ]. 
 roval of nisin as a purified bacterioc n additive in European and US food regulations 
[1] a d the commercial avail bility of crude extracts, such as Pediocin PA-1 from bacteri-
ocin-producing strains, highlights its potential as an effective biopreservative [51,52]. 
 
Figure 2. Future perspectives. The utilization of live biopreservatives (LBs) as bacteriostatic agents, 
combined with the use of lytic bacteriophages, poses future research and innovation challenges for 
the meat industry. (A) Biodegradable food packaging has been in the market for a few years already. 
However, its potential interactions with LBs are not yet fully understood. Moreover, the combina-
tion of LBs (bacteriostatic effect), bacteriophages (bacteriolytic effect), and biodegradable food pack-
aging has never been tested. (B) A reduction in the use of nitrites and nitrates will bring potential 
benefits to the food industry, the environment, and consumer health. However, before they can reach 
the market, LBs require further investigations into their economic challenges. Furthermore, bacterio-
phages are not yet regulated as biopreservatives by the major regulatory agencies, including those in 
the European Union and the United States. (C) Lactic acid bacteria and bacteriocins have performed 
well in controlled settings. However, antimicrobial agents need to be scaled up to demonstrate their 
usability in highly complex meat matrices. (D) Antimicrobial resistance is a major global challenge. 
Interspecific horizontal gene transfer (HGT), including the exchange of antimicrobial resistance genes, 
Figure 2. Future perspectives. The utilization of live biopreservatives (LBs) as bacteriostatic agents,
combined with the use of lytic bacteriophages, poses future research and innovation challenges for
the meat industry. (A) Biodegradable food packaging has been in the market for a few years already.
However, its potential interactions with LBs are not yet fully understood. Moreover, the combination
of LBs (bacteriostatic effect), bacteriophages (bacteriolytic effect), and biodegradable food packaging
has never been tested. (B) A reduction in the use of nitrites and nitrates will bring potential benefits to
the food industry, the environment, and consumer health. However, before they can reach the market,
LBs require further investigations into their economic challenges. Furthermore, bacteriophages
are not yet regulated as biopreservatives by the major regulatory agencies, including those in the
Europe n Union and the United States. (C) Lactic acid bacteria and bact riocins have performed
w ll in controlled setti gs. However, antimicrobi l agents need to be scaled up to demonstrate th ir
usability in highly complex meat matrices. (D) Antimicrobial resistance is a major global challenge.
Interspecific horizontal gene transfer (HGT), including the exchange of antimicrobial resistance genes,
is pervasive in bacteria. Future experiments on the use of LBs and bacteriophages need to explore the
potential highways for HGT that can be created between LBs and environmental microbes.
Advancements in understanding the behavior of LAB and bacteriocins in different
meat environments are shaping tailored approaches. For instance, Pseudomonas species
Life 2025, 15, 197 7 of 14
thrive in aerobic environments, while LAB flourish in anaerobic settings, such as in vacuum-
packed meat [24]. This knowledge aids in optimizing the effectiveness of these products
against specific spoilage or pathogenic microorganisms present in different meat types.
Moreover, the use of LAB cocktails or synergistic combinations of bacteriocins has the
potential to create robust antimicrobial systems that can combat a broader spectrum of
microbes, ensuring product safety and quality of meat products, as well as prolonging
shelf life and preserving sensory attributes [7,53]. Innovations in processing technologies,
such as controlled fermentation temperatures, targeted bioprotective starter cultures, and
emerging techniques such as high-pressure processing (HPP), are also proving instrumental
in harnessing the antimicrobial capabilities of LAB and bacteriocins [20].
Looking ahead, the future landscape of meat preservation and safety is likely to shift
toward greater reliance on natural antimicrobial agents such as LAB and bacteriocins. As
research and technological advancements continue to unfold, the effective integration of
these natural compounds in meat processing is advised to address food-safety concerns,
extend shelf life, and satisfy consumer demands for cleaner labels and natural preservatives
in meat products. This evolution represents a paradigm shift toward sustainable and
innovative food-preservation methodologies in the meat industry.
8.2. Computational Models for Assessing the Use of LBs and Bacteriophages
Computational models may play a crucial role in evaluating the potential synergistic
effect of LBs and bacteriophages on meat safety and preservation. Predictive microbiology
relies on the use of deterministic and stochastic models and is based on the idea that the
responses of microbial populations to environmental elements can be replicated [54,55].
Thus, the application of predictive microbiology techniques allows for determination of
the potential behavior of pathogens under diverse conditions within a certain degree
of probability. The use of predictive biological models through databases with trained
models offers a robust mechanism for enhancing quality support and ensuring food-safety
measures in the food industry [56]. These models, grounded in evidence, signify progress
in the effective evaluation and management of food safety [56].
Counting models of pathogen growth can be written as general birth-and-death pro-
cesses (Figure 3) [57]. Birth-and-death models focus on the internal dynamics of the size of
a single population over time. These models offer versatility in their representations, allow-
ing for diverse forms that encompass a wide spectrum of microevolutionary scenarios [58].
Nevertheless, birth-and-death models require a minimum of two parameters, λ (birth
rate) and µ (death rate), to determine the effect of LBs and bacteriophages on pathogen
growth. Thus, the population of a bacterial pathogen of size n increases at a rate of nλ and
decreases at a rate of nµ, with the parameters (λ, µ) being different for each microbe and
environmental condition.
Modelling interactions among a bacterial pathogen, an LB, a bacteriophage, and the
physicochemical properties of meat production and storage can be achieved by combining
aspects of competitive growth models and predator−prey dynamics based on the classic
Lotka−Volterra equations [59] and the Monod and logistic growth models [60,61]. The
concentration of the bacterial pathogen at time t, S(t), can be measured using Equation (1);
the concentration of the LB at time t, B(t), can be measured using Equation (2); and the
concentration of a bacteriophage targeting the pathogen at time t, P(t), can be measured
using Equation (3). Furthermore, stochastic versions of these equations can account for
real-world randomness, which is important for understanding population dynamics in
small or variable populations.
Life 2025, 15, 197 8 of 14
  




	



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Figure 3. Hypothetical model of the action of live biopreservatives (LBs) and bacteriophages. Birth-
and-death models for assessing the effect of LBs and bacteriophages on meat production. λ: birth
rate and µ: death rate. The parameters (λ, µ) vary based on the microbes and experimental con-
ditions (physicochemical properties, bacteriophages, and LBs). The synergistic interaction of LBs,
bacteriophages, and the physicochemical properties of the food matrix, processing, and packaging
on pathogen growth can be quantified using variations of the classic dynamic interaction equations
(Equations (1)–(3)).
Equation (1). Pathogen growth.
dS
dt = XS −YS − ZS when XS > YS + ZS
XS = (v · i) · S ·
(
1− S+BK
)
YS = γ · S · B ZS = φ · S · P
i = it · ipH · iw · im · ix 0 ≥ i ≤ 1
(1)
where ν is the growth rate of the pathogen; K is the carrying capacity of the environment
for bacteria; γ is the competition rate between B (LB) and S (pathogen); φ is the phage ad-
sorption rate; and i is the interaction with environmental properties, including temperature
(it), water activity (iw), pH (ipH), material (im), and other potential factors (ix).
Equation (2). Live biopreservative (LB) growth
dB
dt = XB −YB when XB > YB
XB = (ω · j) · B ·
(
1− S+BK
)
YB = α · S · B
j = jt · jpH · jw · jm · jx 0 ≥ j ≤ 1
(2)
where ω is the growth rate of the LB; α is the competition coefficient between (S) pathogen
and B (LB); and j is the interaction with environmental properties, including temperature
(jt), water activity (jw), pH (jpH), material (jm), and other potential factors (jx).
Equation (3). Bacteriophage growth
dP
dt = XP −YP
XP = β · φ · S · P ·
(
1− PKp
)
YP = δ · P (3)
Life 2025, 15, 197 9 of 14
where β is the burst size (number of new phages produced per lysed host cell), δ is the
natural decay rate of the phage, P is the bacteriophage concentration, and Kp is the carrying
capacity of the environment for the bacteriophage.
8.3. Assessment of the Potential Impact of LBs on Antimicrobial Resistance
Antimicrobial resistance is a complex and emerging global health challenge charac-
terized by the ability of microorganisms to withstand the effects of antimicrobial agents.
The extensive utilization of antibiotics and other chemicals has the potential to induce
multiresistance in microbes [62]. Horizontal gene transfer (HGT, i.e., the exchange of
genetic material between different species) of antimicrobial resistance genes is a major
source of antimicrobial resistance [63]. Thus, suitable LBs must not only exhibit the gen-
eral characteristics outlined earlier in the text but must also be free from antimicrobial
resistance genes. One may speculate that the use of LBs and bacteriophages could serve
as an alternative to help mitigate the arms race. However, previous studies have shown
that the use of antimicrobials may contribute to the formation of multiresistant bacterial
strains [62,64–66]. Furthermore, the use of preservatives can accelerate plasmid-mediated
HGTs of antimicrobial resistant genes (AMR) in bacteria [64]. Therefore, we hypothesize
that interactions and competition within microbial communities, along with a potential
increase in the frequency of HGTs, may accelerate the spread of AMR within a microbial
community. In addition, considering that AMR mechanisms of bacterial resistance are
interlinked [62], further investigation is needed into the potential impact of employing
LBs and bacteriophages in meat production on the ongoing arms race of antimicrobial
resistance [8] (Figure 2D).
8.4. Uses of LBs and Bacteriophages with Biodegradable Packaging for Meat
Active packaging or microencapsulation techniques are considered valuable alter-
natives for ensuring microbiological food safety and enhancing the quality of food prod-
ucts [34,67]. The application of antimicrobial and active compounds allows persistent
migration to the food matrix during storage. In this regard, the application of these ma-
terials has various advantages, including controlling spoilage; restricting the growth of
pathogenic microorganisms; reducing lipid, protein and pigment oxidation; and prevent-
ing off-odors [68]. Moreover, the incorporation of LAB and their metabolites in active
packaging for meat and meat products is more efficient than their direct application [34].
However, to date, there are a limited number of studies that combine LBs [23] in sus-
tainable/biodegradable food packaging for meat [48,49]. Some studies have shown the
effectiveness of plantaricin BM-1 and pediocin BA-1/AcH for pork meat as coatings in new
biocomposite films [1]. All the treatments reduced the population of L. monocytogenes by
approximately 1.5–2 log units after 14 days of storage at 4 ◦C [1].
Even though the combination of different microencapsulation techniques, natural
extracts, and LAB cocktails with novel techniques such as MAP, HPP, or active packaging
increases the preservative effect [34], the synergistic potential of these combinations with
bacteriophages in biodegradable materials has not yet been tested (Figure 2A). Moreover,
biodegradable plastic packing is exposed to the action of environmental plastic-degrading
microorganisms [69], leading to potential interactions with LBs and bacteriophages.
8.5. Socioeconomic Implications of LBs and Bacteriophages
The integration of LBs and bacteriophages in food-preservation processes has sig-
nificant economic and environmental implications, reshaping the landscape of food
safety and sustainability. Beyond technical considerations, addressing safety issues is
crucial for ensuring the optimal and appropriate use of bacteriophages for maximum
efficacy [15]; there is also the need for further legal revisions and weighting of ethical
Life 2025, 15, 197 10 of 14
considerations. Bacteriophages are at the forefront of major innovations in biotech-
nology (e.g., as biopreservatives within the food industry [38]) and biomedicine (e.g.,
as phage therapy, a promising field of research with the aim of overcoming antibiotic
resistance [70]). However, significant regulatory bodies in the EU and the United States
have not overseen the utilization of bacteriophages in the fields of biomedicine and
biotechnology [71]. Furthermore, consumers are not yet fully aware of the potential
impact of these technologies. Although a limited recent survey in the United Kingdom
showed that the lay public has a moderate level of acceptance for phage therapy [72],
future, wider studies on public acceptance are still needed.
Economically, these natural preservatives offer a potential solution for industries where
substantial financial losses are incurred due to product contamination [1]. By implementing
biopreservation strategies, companies can potentially reduce waste, minimize recalls, and
extend shelf life, leading to cost savings and improved profitability [31]. Research in this
area aims not only at combating foodborne pathogens such as L. monocytogenes [33] but
also at addressing critical environmental concerns by potentially curbing greenhouse gas
emissions associated with food waste [31]. Moreover, the adoption of bioprotective starter
cultures and other natural preservation methods can streamline production processes,
reducing the reliance on synthetic additives and simplifying labeling, meeting consumer
preferences for clean and natural ingredients (Figure 2B). This shift aligns with market
demands for healthier, minimally processed foods, potentially enhancing market competi-
tiveness for food producers [73]. For example, the implementation of higher concentrations
of bioprotective starter cultures combined with low sugar concentrations has shown tangi-
ble benefits, such as improved textural qualities and reduced oxidation levels in products
such as salamis, even when faced with challenges from various harmful bacteria, such as L.
innocua, S. enterica, and C. botulinum [5]. Finally, the emergence of metagenomic techniques
in food safety, particularly in identifying and characterizing environmental microorganisms
in food-processing chains, represents a promising avenue for further investigation [1]. Such
approaches not only enhance our understanding of food-preservation processes but also
contribute to refining biopreservation strategies for better food quality and safety.
9. Conclusions
Biopreservation with LBs and bacteriophages is a promising sector of food processing
and food research, with a specific spectrum of action against targeted bacterial species
or even strains. Furthermore, biopreservation will be used in combination with physical
factors such as heat treatment, chilling/cooling, or food drying, which results in a decrease
in water activity. The emerging field of combining LBs and bacteriophages promises
enhanced meat preservation and longevity. Adding LBs will yield meat products with
an extended shelf life due to bacteriostatic mechanisms, whereas bacteriophages provide
more direct (lytic) action against pathogens. However, achieving meat preservation with
LBs and bacteriophages requires further empirical and theoretical understanding of these
microbe-mediated strategies. A compiled list of outstanding questions regarding the use of
LBs, bacteriophages, and biodegradable food packaging, to be addressed in future studies,
is provided in Box 1. Moreover, the utilization of microbe-mediated approaches will create
a need for additional revisions to the law and weighting of ethical considerations.
Life 2025, 15, 197 11 of 14
Box 1. Outstanding questions
• Is the combination of LBs (bacteriostatic) and bacteriophages (bacteriolytic) the most suitable
strategy to improve quality and meat safety?
• What is the impact of combining LBs and bacteriophages on the physicochemical, technological,
and sensory properties of both fresh and fermented meat?
• How do LBs and bacteriophages perform in real meat products, considering food complexities?
• How does the combination of LBs with biodegradable food packaging improve meat safety
and sustainability?
• What potential interactions can occur between LBs (and bacteriophages) with environmental
plastic-degrading microbes on biodegradable plastic packaging materials?
• Does the use of LBs (and bacteriophages) pose a potential risk for the spread of antimicrobial
resistance?
• Can mixed strategies involving LBs and bacteriophages effectively address the issue of reduc-
ing nitrites and nitrates in meat preservation?
Author Contributions: Conceptualization, P.P. and C.R.-M.; methodology, C.R.-M. and P.P.; com-
putational model, P.P.; investigation, P.P., C.R.-M. and C.D.-C.; resources, N.O.-O., C.R.-M. and P.P.;
writing—original draft preparation, P.P. and C.R.-M.; writing—review and editing, C.R.-M., C.D.-C.,
M.N., N.O.-O. and P.P.; writing the final version of the manuscript, P.P. 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.
Acknowledgments: We thank members of the Eurecat’s Nutrition and Health Unit for their help-
ful discussions.
Conflicts of Interest: The authors declare no conflict of interest.
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