Trends in Food Science & Technology 131 (2023) 264–276

Contents lists available at ScienceDirect

Trends in Food Science & Technology

journal homepage: www.elsevier.com/locate/tifs

Current challenges in the application of the UV-LED technology for

food decontamination
Arturo B. Soro a, b, Sajad Shokri c, Iolanda Nicolau-Lapeña d, Daniel Ekhlas a, b,
Catherine M. Burgess a, Paul Whyte b, Declan J. Bolton a, Paula Bourke c, Brijesh K. Tiwari a, *
a
Teagasc Ashtown Food Research Centre, Ashtown, D15 DY05, Dublin, Ireland
b
UCD School of Veterinary Medicine, University College Dublin, Belfield, D04 V1W8, Dublin, Ireland
c
UCD School of Biosystems and Food Engineering, University College Dublin, Belfield, D04 V1W8, Dublin, Ireland
d
Universitat de Lleida, Food Technology Department, AGROTECNIO-CERCA Center, Rovira Roure 191, 25198, Lleida, Spain

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

Keywords: Background: Different strategies have been developed over the years to improve the safety of foods without
Ultraviolet light compromising on their quality. Ultraviolet (UV) light has shown potential to meet these expectations. However,
Stress response due to environmental and safety concerns regarding the use of UV mercury lamps, UV-light emitting diode (LED)
DNA repair
devices have emerged more recently, which may be more suited to applications in the food industry. Thus,
Microorganisms
Food quality
further evaluation of UV-LED technology is required.
Food safety Scope and approach: In this review, challenges involving the application of UV-LED technology as a disinfection
strategy in the food chain are described. Stress-related mechanisms induced by UV light exposure in microor­
ganisms and their potential consequences are reviewed. Moreover, other future challenges associated with the
inactivation of foodborne pathogens and spoilage microorganisms are considered, together with the impact of
the application of UV-LED technology on the quality of food.
Key findings and conclusions: UV-LEDs have been shown to inactivate a wide range of foodborne pathogens and
spoilage microorganisms. However, there are limited studies assessing the effectiveness of UV-LED on products to
enhance food safety. Additionally, further studies are required to explore the impact of sublethal exposure to UV-
LEDs and the potential consequences of UV-LED exposure in different microorganisms at various wavelengths.
Furthermore, UV-LEDs can negatively affect some physicochemical attributes of food. Thus, more attention
should be paid to the underlying mechanism of quality changes induced by UV-LEDs.

1. Introduction
In recent years, consumer expectations for high nutritional foods that
maintain their optimal organoleptic properties and are microbiologi­
cally safe have increased. Different approaches such as ultrasound, high
hydrostatic pressure, plasma and ultraviolet light, among others, have
been developed to reduce the microbial load in foodstuffs and prevent
undesirable physicochemical changes associated with traditional heat
treatments (Nicolau-Lapeña et al., 2019). The application of UV light has
been widely used for specific purposes, including food-contact surface
treatment and water and air disinfection. UV light occupies a wide band
of wavelengths in the non-ionizing region of the electro-magnetic
spectrum, between 200 nm (X-rays) and 400 nm (visible light). Irradi­
ation intensity is expressed in W/m2; and irradiation dose is dependent
on treatment times, distance to the source along with other parameters,
and is commonly expressed in J/m2. The application of UV light in food
production has gained interest due to the absence of toxic by-products or
impacts on organoleptic properties and the lower requirement of energy,
compared to high hydrostatic pressure processes and thermal pasteuri­
zation (Riganakos et al., 2017).
When used for disinfection purposes, several factors must be
considered, such as the type of light exposure (pulsed or continuous),
reactor type (continuous or flow through), distance from the light source
and incidence angle, and treatment conditions (eg. temperature) (Hinds

* Corresponding author. Teagasc Food Research Centre, Ashtown, Dublin, D15 KN3K, Ireland.
E-mail addresses: Arturo.blazquezsoro@ucdconnect.ie (A.B. Soro), sajad.shokri@ucd.ie (S. Shokri), iolanda.nicolau@udl.cat (I. Nicolau-Lapeña), daniel.ekhlas@
teagasc.ie (D. Ekhlas), kaye.burgess@teagasc.ie (C.M. Burgess), paul.whyte@ucd.ie (P. Whyte), declan.bolton@teagasc.ie (D.J. Bolton), paula.bourke@ucd.ie
(P. Bourke), brijesh.tiwari@teagasc.ie (B.K. Tiwari).

https://doi.org/10.1016/j.tifs.2022.12.003
Received 28 September 2022; Received in revised form 3 December 2022; Accepted 5 December 2022
Available online 12 December 2022
0924-2244/© 2022 Elsevier Ltd. All rights reserved.
A.B. Soro et al.
et al., 2019). The target microorganisms and their physiological status
must also be considered. It has been shown that UV light doses necessary
for inactivation of enteric bacteria (20–80 J/m2) are lower than those of
cocci and micrococci, followed by bacterial spores, enteric viruses,
yeasts, fungi, protozoans, and algae which require up to 6000 J/m2 for
inactivation (Koutchma & Popović, 2019). Growth phase, conditions
and exposure to stressors are also parameters that may influence the
effectiveness of UV light to inactivate microorganisms (Gayán et al.,
2013). However, as the mode of action of UV light is primarily attributed
to DNA damage, its efficacy against microorganisms may vary according
to the spectrum range that is being used. The UV spectrum is typically
divided in short-wave UV light (UVC, 200–280 nm), medium-wave UV
light (UVB, 280–320 nm), and long-wave UV light (UVA, 320–400 nm)
(Song et al., 2016). The most germicidal part is considered to be UVC, as
this wavelength coincides with the highest absorbance by DNA (254
nm), compromising cell function in microorganisms (Singh et al., 2021).
DNA damage is primarily caused by reactions between adjacent thymine
bases to form thymine dimers, formation of cyclobutene-pyrimidine
dimers (CPDs) and (6-4) pyrimidine–pyrimidone photoproducts (6-4
PPs), that are cytotoxic and mutagenic (Banas et al., 2020). Although
UVB also produces CPDs and 6-4 PPs, these photoproducts are induced
at lower levels compared to UVC (Gayán et al., 2013). DNA can also be
indirectly altered by UVA, as it can trigger photoreactions via the gen­
eration of singlet oxygen (Koutchma & Popović, 2019). Additionally,
UVA has higher penetrability, so it can achieve greater efficacy when
used in opaque liquids and biomaterials with a high absorptivity coef­
ficient (Kebbi et al., 2020). In addition to DNA damage, photooxidation
can act against other targets, which include proteins, lipids, and sterols.
In particular, UVA and UVB can damage a small number of amino acid
residues such as tryptophan, tyrosine, histidine and cystine (Gayán
et al., 2013). Moreover, the production of reactive oxygen species (ROS)
generated by UV light treatment affects some protein chain residues, and
once this reaction has started, multiple secondary processes occur,
resulting in damage to other intra- and inter-molecular sites, leading to
protein fragmentation or aggregation and loss of their functionality (Gao
& Garcia-Pichel, 2011). ROS can also cause the oxidation of lipid bi­
layers in fatty acids causing poration, rearrangements in composition,
and alteration of transmembrane ion gradients, which eventually result
in the disruption of the cell membrane (Smith et al., 2009).
To date, the most common UV light sources have been mercury
lamps, consisting of an UV-transmitting envelope (tube of silica glass)
sealed at both ends, where an electrode is located and connected to the
outside through a seal. Mercury and inert gas (e.g. argon) fill the lamp
and are typically powered with alternating current. Mercury is used as it
is the most volatile metal element, for which activation in the gas phase
can be obtained at temperatures compatible with the lamps (Song et al.,
2016; Koutchma, 2009. However, the use of these lamps in food pro­
cessing settings may cause some concern, and alternatives such as
excimer lamps, broadband pulsed lamps, microwave UV lamps and
UV-light-emitting diodes (UV-LEDs) have emerged. Despite their lower
UV intensity and higher cost, UV-LEDs are of particular interest
compared to UV mercury lamps due to their: (1) higher energy effi­
ciency, which minimizes energy consumption, (2) longer lifetime, (3)
minimized environmental impact, (4) lower heat generation, and (5)
consistent light intensity (Muramoto et al., 2014). UV-LEDs convert
electrical current directly into irradiation using a structure based on the
junction of two-terminal semiconductors (p-n junction). Depending on
the nature of the semiconductors, the emitted UV wavelength can be
adjusted for specific purposes (Hinds et al., 2019). Moreover, different
wavelengths can be generated simultaneously, with the possibility to
exert additive or synergistic effects (Kheyrandish et al., 2018).
UV-LED treatments have shown potential in the food safety area and
hygiene for sanitation purposes, and might effectively replace mercury
UV lamps. Their mode of action, configurations and efficacy on target
microorganisms and treatment parameters, as well as the impact on food
quality have already been reviewed (Hinds et al., 2019; Kebbi et al.,
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Trends in Food Science & Technology 131 (2023) 264–276
2020). However, there is evidence that microorganisms can respond to
the stress posed by UV treatments in several ways, including biofilm
formation (Kaushik et al., 2022), expression of stress related genes
(Baharoglu & Mazel, 2014; van der Veen & Abee, 2011), increase
tolerance to pH, temperature or salt (Alvarez-Ordonez et al., 2015; Voth
& Jakob, 2017) and DNA repairing mechanisms (Prado, 2018; Rastogi
et al., 2010). The study of such responses is vital to have a better un­
derstanding of the outcomes UV-LEDs may have on the fate of micro­
organisms. Moreover, the use of UV-LED technology for the inactivation
of foodborne pathogens and spoilage microorganisms in food matrices
requires further evaluation, with a special focus on the impact of this
technology on the quality of foodstuffs. For this reason, our review fo­
cuses on current challenges involving the application of the UV-LED
technology in the food chain with perspectives to future
implementation.
2. Microbial stress responses to UV light exposure
Many food products are considered stressful environments for mi­
croorganisms due to intrinsic factors (e.g. composition, pH, water ac­
tivity) and extrinsic chemical and physical processes (e.g. refrigeration,
dehydration, acidification). However, some foodborne microorganisms
are able to tolerate these stresses and may be able to grow and/or sur­
vive in food matrices, potentially influencing their safety or shelf life.
(Wesche et al., 2009). Currently, food producers consider the use of
minimal processing technologies, such as UV light to ensure the safety of
food products without compromising quality in order to meet consumer
demands for healthier and more sustainable food. The use of minimal
process technologies, could theoretically increase the tolerance of mi­
croorganisms to severe stress, as a result of exposure to sub-lethal
stresses. This phenomenon is also known as stress hardening or stress
adaptive response and is based on resistance to homologous or heter­
ologous stresses (Ravishankar & Juneja, 2002; Wesche et al., 2009).
Although any type of stress can induce differentially-regulated genes to
express specific proteins and pathways, some responses to stress are
condition-specific in order to protect the cells from damage and enable
them to adapt to different conditions (Nezhad et al., 2015). Hence,
microbial resistance to UV light, or the induction of responses to other
stresses, requires further assessment and investigation to determine the
specific microbial mechanisms involved in the adaptive response to
stress.
2.1. Gene and pathway regulation following UV light-induced stress: DNA
repair mechanisms
Microbial cells are capable of detecting environmental changes and
adapting to them through the upregulation and downregulation of
certain genes. This genetic regulation can derive from random transient
or permanent mutations that could trigger stress-counteracting modifi­
cations in the cell physiology (e.g. proteins, DNA, membrane), repair
activities, reduction or elimination of the stress source, and/or induced
actions to increase the resistance of the cell to future stress agents
(Alvarez-Ordonez et al., 2015). However, these stress response mecha­
nisms are dependent on the type of microorganisms and the stressor
(Marmion et al., 2022). Stress tolerance responses (e.g. to cold, acid, and
heat) are commonly considered as transient responses. Regulation of
these fast-adaptive mechanisms is based on the activation of specific
response regulators to induce expression cascades of particular genes,
together with complex systems (e.g. enzyme production, synthesis of
stress-shock protein, and modification of the properties of the cellular
envelopes) (Alvarez-Ordonez et al., 2015; Marmion et al., 2022). These
fast response mechanisms can also occur in microbial cells when novel
preservation technologies such as UV light are applied to food products
(Cebrian et al., 2016; Gayán et al., 2013) Factors influencing the effec­
tiveness of UV light technology, are for instance intra- and inter-specific
microbial differences, the growth rate of microbial cells, and optical
A.B. Soro et al.
properties of the medium, which may contribute to facilitate some
survival in the cell population and therefore, allow microbial cells to
establish mechanisms against UV light-induced stress (Cebrian et al.,
2016).
Microbial DNA lesions derived from UV light exposure (CPDs and 6-
4 PPs) may trigger several DNA repair mechanisms, such as photore­
activation, recombinational and excision repair, and the SOS response
which are dependent on the genomic location and type of the lesion
(Rastogi et al., 2010). All of these DNA repair mechanisms in response to
UV light-induced stress are graphically presented in Fig. 1. Together
with these DNA repair processes, the so-called cell-cycle checkpoint
arrest is induced to avoid any potential malfunction in the cell division
and/or loss of genetic information after UV light exposure. Briefly, the
cell triggers a series of molecular sensors, signal transducers, mediators
and effectors in order to stop the progress of the cell cycle and prevent its
advance to a different cycle phase. Thus, repair mechanisms can restore
DNA damaged by UV light without comprising vital cell functions
(Anand et al., 2020; Rastogi et al., 2010). The photoreactivation repair
phenomenon is mainly caused by the action of photolyases in light or
dark conditions. These enzymes are capable of recognizing the presence
of pyrimidine dimers (CPDs and 6-4 PPs) on DNA strands. Photolyases
first bind to the pyrimidine dimer through their binding pocket. Sub­
sequently, they split the pyrimidine dimer into an individual pyrimidine
by transferring sufficient energy to break the dimer bond thereby
creating two separate rings. Finally, the undamaged monomeric bases
are restored and incorporated back in the DNA helix (Banas et al., 2020).
Another microbial repair mechanism is excision or dark repair, which
is a much more complex system and involves two different pathway
categories: base (BER) or nucleotide (NER) excision repair. Both path­
ways are based on three steps: recognition, excision, and resynthesis.
This mechanism is also capable of repairing CPDs and 6-4 PPs. In
contrast to the photoreactivation repair, NER and BER can remove helix
distorting (bulky) and non-bulky DNA lesions, respectively. In partic­
ular, BER is involved in fixing single-base lesions to avoid any potential
interference in the function of the microbial replication and transcrip­
tion processes. In the recognition and excision steps, DNA glycosylases
are specifically used to detect and eliminate DNA lesions caused by UV
light exposure in two steps: flipping the damaged base out of the DNA
Fig. 1. Schematic illustration of the different DNA repair mechanisms triggered after UV light exposure in microbial cells. PR: photoreactivation, NER: nucleotide
excision repair, BER: base excision repair, HR: homologous repair and NHEJ: non-homologous end joining. If severe DNA lesions occur in the cell, the induced SOS
response promotes NER, HR and translesion synthesis (TLS) pathways.
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Trends in Food Science & Technology 131 (2023) 264–276
helix followed by breaking the bond that is attached to the base to
release it and leave an empty site (abasic). Thereafter, the resynthesis
process takes place with different enzymes and chemical reactions to
generate a new DNA base. In the case of NER, the unpaired
single-stranded DNA opposite to the UV light-induced lesion is detected.
The NER process can follow two different pathways: global genome
(GG-NER) and transcription coupled repair (TC-NER). GG-NER detects
and eliminates DNA lesions within the whole genome and TC-NER is
responsible for detecting and removing UV light-induced lesions in the
transcribing strands of active genes. A final resynthesis step common to
both sub-pathways is followed where the processes of gap-filling and
ligation take place using various enzymes (Lee & Kang, 2019; Sinha &
Hader, 2002). Recombinational repair is an efficient mechanism that
repairs double and single-strand breaks of damaged DNA caused by UV
light. The double-strand breaks (DSBs) can be repaired using two
pathways: homologous recombination (HR) and non-homologous end
joining (NHEJ). In summary, the HR process initiates with the DNA
resection of the 5′ -ends of the DSBs by an exonuclease to generate a
3′ -ended single-stranded DNA. Subsequently, this single-strand is
aligned and exchanged through the action of a cascade of enzymes. In
this way, a D-loop is created in the DNA structure, which is stabilised by
DNA synthesis. In some cases, the D-loop intermediate can be formed
and produce a crossover between the broken and the template chain
(also known as double-Holliday junction, HJ). As a result, this crossover
needs to be dissolved or resolved to continue the DNA synthesis process
and repair the damaged DNA. If HR is inactivated, the NHEJ mechanism
proceeds to repair the damaged DSBs. This process however is more
prone to generate mutations compared to the HR mechanism. NHEJ
involves the binding of specific enzymes to the broken ends, bridging of
repair proteins to the DNA breakage site, and processing of the repair
proteins through DNA polymerization and ligation mechanisms. Thus,
broken chromosomal ends are joined together independently of their
homology (Prado, 2018; Rastogi et al., 2010).
Additionally, multiple single-stranded DNA lesions caused by an
external factor like UV light may disrupt cell replication and trigger the
SOS response. This repair process initiates when presynaptic complexes
(RecBCD or RecFOR) detect the DNA lesion and recruit the RecA acti­
vator to promote its binding to the location of the damaged DNA. The
A.B. Soro et al.
binding of RecA to the single-stranded DNA causes proteolysis of a LexA
repressor. which is responsible for repressing genes associated with LexA
by binding their promoter sequences. Induction of the SOS response
promotes activation of three main DNA repair mechanisms: HR, NER,
and translesion synthesis (TLS). The two former repair pathways are
usually repair mechanisms free of mutations. If the SOS signal persists,
the RecA catalyses proteolysis of the UmuD repressor that triggers the
TLS process. Thus, different DNA polymerases replicate DNA on the
generated lesions. Although this process is very effective in incorpo­
rating DNA bases in lesions, it is also highly mutagenic due to the low
specificity of these enzymes to integrate the correct nucleotides in the
lesion. If the stress dissipates, the catalysis of the LexA repressor will
desist together with the SOS response (Baharoglu & Mazel, 2014; van
der Veen & Abee, 2011). Microbial cells use these different DNA repair
mechanisms to cope with lesions produced by stressors such as UV light.
However, exposure to such a stressor can be difficult for microbial cells
and the repair process may be not sufficient. Thus, the cell can induce a
permanent and irreversible cycle arrest (cellular senescence) or an
instant programmed cell death (apoptosis).
The majority of studies investigating the effect of UV light on mi­
crobial DNA repair mechanisms used UV lamp-based devices. UV-LED
devices on the other hand have not been extensively explored yet
(Song et al., 2016). Li et al. (2017) studied the levels of photoreactiva­
tion and excision or dark repair in E. coli cells when treated with low
pressure (LP) UV lamps and an UV-LED device. These authors concluded
that the studied repair mechanisms were induced at a much lower level
in E. coli cells treated with UV-LED than LP-UV lamps and speculated
that damage of repair-inducing enzymes may be the main cause (Li et al.,
2017). UV-LED devices may trigger lower levels of DNA repair responses
thereby reducing the tolerance of microbial cells towards UV light.
However, further investigation is needed to understand these DNA
repair mechanisms in response to UV light stress in depth, when
different UV light wavelengths and combinations, types of UV systems
and microorganisms are explored, simulating realistic conditions.
Furthermore, emerging techniques, such as next-generation sequencing
(NGS) methods have been applied to microbial food safety and could
contribute to our understanding of microbial responses to UV light
exposure (Bergholz et al., 2014). Some of the molecular techniques that
could be interesting to explore for this purpose are whole genome
sequencing (WGS) and transcriptomics and non-sequencing methods
like proteomics, and metabolomics. Using NGS to explore the generation
of UV-induced mutations in the microbial genome could help to un­
derstand the microbial stress response to this stressor. In particular,
single nucleotide polymorphism (SNP) analysis may be useful in WGS to
identify when changes in the microbial genome occur. This approach is
based on the mapping of sequencing reads to a known sequenced
reference genome together with the identification of nucleotide differ­
ences in coding and non-coding regions of the genome. Thus, mutated
DNA regions derived from the action of these DNA repair mechanisms
may be detected using the SNP approach (de Boer, 2002; Jagadeesan
et al., 2019). However, such analysis for this purpose remains in its
infancy.
2.2. Potential consequences of inducing DNA repair mechanisms in
microbial cells
The DNA repair process can lead to modifications in cellular struc­
tures and functions, as well as inducing cell recovery. Most of these
changes are attributed to the SOS response. The activated SOS regulon
may also promote expression of other genes related to adaptation and
resistance factors and therefore alter the genomic plasticity of the mi­
crobial cell (Baharoglu & Mazel, 2014). Despite the generation of
non-beneficial mutations, DNA repair mechanisms prone to induce
mutations may facilitate production of phenotypes associated with
increased survival to external stress (i.e. food processing), also known as
adaptive mutagenesis (van der Veen & Abee, 2011). Consequently,
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several protective strategies may be potentially expressed by the mi­
crobial cell to survive UV light exposure.
Biofilms are structured microbial communities in the form of
microcolonies embedded in a self-produced extracellular polymeric
matrix. This substance provides microbial cells with a survival mecha­
nism to adhere to solid surfaces, protect them from external stresses, and
facilitate the recruitment of nutrients. Biofilm formation is induced by a
cell-to-cell communication process known as ‘quorum sensing’. This
phenomenon of multicellular responses is triggered when several signal
molecules (autoinducers) are produced and secreted by the microbial
cells which accumulate extracellularly. Thus, the expression of specific
cellular functions is induced and regulated in order to initiate the for­
mation of a biofilm and its subsequent growth and maturation (Alvar­
ez-Ordonez et al., 2019; Lianou et al., 2020). For instance, SOS responses
could be associated with biofilm production due to the role of RecA and
LexA in favouring this mechanism. The mutations that originate as a
consequence of inducing a SOS response following exposure to a stress
agent may contribute to biofilm formation phenotypes and generate
microbial cell variants associated with biofilms and their formation
(Kaushik et al., 2022; van der Veen & Abee, 2011; Yin et al., 2019).
Future efforts should be made to examine the impact of UV-LED tech­
nology on biofilm formation phenotypes in microbial species of interest.
Another example of a survival strategy is the viable but-
nonculturable (VBNC) state. Microbial cells subjected to external
stress can alter their morphology by reduction of cell size and/or change
in cell shape that can be a consequence of changes in the cell wall (e.g.
fatty acid composition and peptidoglycan cross linking). In the VBNC
state, cell growth on conventional culture medium declines, although
metabolic and virulence activity is maintained, enabling resuscitation
when conditions improve. Quorum sensing is also connected with the
regulation of the VBNC formation and resuscitation, and allows adaptive
phenotypes to promote tolerance to a particular stress (Li & Zhao, 2020).
Zhang et al. (2015) investigated the impact of UV light on E. coli and
Pseudomonas aeruginosa cells and examined how UV light induced the
VBNC state in cells of both species. In another study, the survival of
antibiotic resistant E. coli cells to UV light was investigated. VBNC cells
of E. coli induced by UV light were reactivated by photoreactivation and
dark repair mechanisms (Guo & Kong, 2019). Most of the studies eval­
uating the effect of UV light on VBNC have found that UVC induces
VBNC cells the most (Cai et al., 2021). The generation of VBNC cells and
cell recovery after UV light exposure are still underexplored across
UV-LED applications.
Development of antimicrobial resistance (AMR) is another
possible consequence of inducing mutations via the DNA repair mech­
anisms. In particular, the SOS response is associated with AMR and
thereby, a cross-protection for bacteria towards other external stresses
may induce the expression of antimicrobial resistance genes (ARGs).
AMR is an increasing public health concern and ARGs have been
demonstrated to transfer from food containing AMR bacteria to the
human gastrointestinal tract (Liao et al., 2020). Zhang et al. (2017)
observed that the survival of antibiotic-resistant E. coli cells was greater
at lower doses of UV light (below 20 mJ/cm2) than higher doses and
hypothesised that UV light could be used to select for AMR bacteria. In
another study, Staphylococcus aureus cells treated with a pre-treatment
of UV light (8.6 J/m2) were found to induce a SOS response (Cirz
et al., 2007). The SOS response can cause the inhibition of protein
synthesis and DNA transcription in microbial cells and this might pro­
mote resistance to antibiotics like streptomycin and rifampin which act
by disrupting the same metabolic pathways (Liao et al., 2020). Other
studies by Templeton et al. (2009) and Zhang et al. (2019) found no
increased tolerance of E. coli and Pseudomonas cells when UV light
treated cells were subsequently exposed to antibiotics. Moreover, the
combination of UV light and chlorination in drinking water has been
shown to cause a reduction in AMRs and ARGs (Zhang et al., 2019).
These inconsistencies in results suggest further studies are required to
evaluate the potential effects of UV light on ARGs and AMR.
A.B. Soro et al.
The biological process of transcriptional regulation may also be
affected by the action of UV light on the cell. Many molecules and
pathways are regulated in the transcription mechanism and are involved
in several steps, which range from the recruitment and assembly of the
transcription machinery to transcription termination and RNA matura­
tion (Casamassimi & Ciccodicola, 2019). When DNA is damaged by
exposure to UV light and cannot be restored by DNA repair mechanisms,
complex transcriptional cascades can be triggered in the cell to repair
the DNA lesions and/or promote survival mechanisms (Sampath &
Lloyd, 2019). Although the transcription step is usually affected by UV
light-induced lesions in the DNA, a process known as transcription
coupled NER (TC-NER) will try to counteract the blocked transcription
and restart it. However, little information is available in the scientific
literature about these transcriptional mechanisms (Lans et al., 2019).
Transcriptional recovery is a stressful process that can lead to cell death.
If successful, the transcriptional initiation step may resume as normal.
However, transcript elongation and other successive steps involved in
co-transcriptional RNA-processing may be affected and therefore,
changes in RNA processing may be expected in surviving cells (Gre­
gersen & Svejstrup, 2018). Uesugi et al. (2016) studied gene expression
derived from UV light exposure in Listeria monocytogenes using whole
genome microarray analysis. These authors observed that levels of
stress-related proteins, transcriptional regulators, cell membrane, and
motility genes were increased in cells subjected to UV lamp treatment
(Uesugi et al., 2016). Despite this, there is a lack of evidence regarding
the transcriptional effect of UV light on microbial cells. RNA sequencing
techniques (RNA-seq) may help to understand the transcriptional re­
sponses enabling the survival of microbial cells exposed to UV light in
future studies. The application of RNA-seq in food microbiology,
together with other molecular techniques such as proteomics and
metabolomics are needed to understand the breadth of UV stress-related
mechanisms in microbial cells and their consequences in the food chain
(Lans et al., 2019).
UV light, as well as other external stressors, can produce cellular
damage in other structures, such as proteins. When proteins are affected,
a group of proteins called chaperones detect and locate the damage,
and in response activate a specific chaperone function. Briefly, the
mechanism of action of chaperones is to regulate the folding and
unfolding of cellular proteins (Voth & Jakob, 2017). According to Awan
et al. (2018), UV light-induced lesions in proteins can trigger the syn­
thesis of chaperones and therefore, regulate the protein homeostasis of
the cell. Moreover, some bacteria (Clostridium sp., Bacillus sp., Micro­
coccus sp., etc) can activate the production of polyhydroxyalkanoate
(HAP) after UV light exposure, which is the result of metabolising fatty
acids to endure stress (Muhammadi et al., 2015). Other metabolites (e.
g. scytonemin, mycosporines, melanins, and carotenoids) have been
observed to be generated in response to UV light exposure in microor­
ganisms, which act as physical protection. In other words, these mole­
cules can absorb UV light and reduce its impact on the microbial cell
(Gao & Garcia-Pichel, 2011). The production of compounds as a result of
UV light-induced stress remains however an underexplored field. More
information on this topic could shed light on the phenotypic changes
induced by UV-LED light exposure in microorganisms.
2.3. Cross-protection
To conclude the list of potential consequences associated with UV
light exposure, it is necessary to highlight the action of UV light in
inducing cross-protection to other stressors such as heat, acid, ethanol,
and hydrogen peroxide (Alvarez-Ordonez et al., 2015). Several mole­
cules and pathways (e.g. sigma factors, glutamate decarboxylase system,
chaperones, shock proteins, etc.) have been associated with
cross-protection to multiple stresses, but the exact mechanisms of
cross-protection remain unclear (Wu et al., 2022). Studies that associ­
ated UV light exposure of microbial cells with cross-protection towards
other stresses were reviewed by Gayán et al. (2013). These authors
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Trends in Food Science & Technology 131 (2023) 264–276
concluded that more studies are required for a better understanding of
cross-protection to evaluate the safety of UV light in food production
(Gayán et al., 2013). NGS and other molecular techniques could play an
important role in unravelling the cross-protection machinery and facil­
itate their study (Wu et al., 2022).
3. Challenges in UV-LED application to inactivate foodborne
microorganisms
3.1. UV-LED effect on foodborne pathogens
One of the main challenges in the food industry is to control and
minimise foodborne pathogens, whilst maintaining satisfactory physi­
cochemical, nutritional, and sensory qualities. The following section
reviews the most recent studies on UV-LEDs applications for inactivation
of foodborne pathogenic bacteria as it is shown in Table 1 and highlights
the main factors affecting the efficacy of UV-LED.
The nature of the target pathogens can affect UV-LED germicidal
efficiency. Substantial variations in resistance among pathogens to UV
light have been reported (Gayán et al., 2013). There are several reports
suggesting increased susceptibility among Gram-negative species such
as pathogenic E. coli, Salmonella Typhimurium, and P. aeruginosa to
UV-LEDs than Gram-positive bacteria such as S. aureus and
L. monocytogenes, however, reported results have been inconsistent
(McSharry et al., 2022; Yu et al., 2022). A higher resistance of
Gram-positive pathogens to UV-LEDs irradiation has been proposed due
to the thick peptidoglycan layer in the cell wall (Hinds et al., 2019). In
contrast, McSharry et al. (2022) reported a higher susceptibility of
L. monocytogenes to UV-LEDs compared to S. Typhimurium which was
explained by the ability of Gram-negative bacteria to produce higher
amounts of porphyrins which makes them resistant to photoexcitation
(Endarko et al., 2012; Maclean et al., 2009; Nitzan et al., 2004). Another
explanation for this observed higher resistance in S. Typhimurium was
its efficient repair system (Gayán et al., 2012). Other studies have shown
the cell wall structure of bacteria may not be a key indicator of resistance
to UV-LEDs and suggest variation is dependent on strains within species
rather than their Gram status (Yu et al., 2022). It has been shown that
different strains or serotypes of Salmonella spp., Listeria spp., E. coli may
have different susceptibility when exposed to UV light (Kim & Yuk,
2017), because of a variation in the amounts and types of endogenous
porphyrins (Endarko et al., 2012; Nitzan et al., 2004).
UV-LEDs’ effectiveness against foodborne pathogens also can vary
depending on the wavelength applied; generally, a shorter wavelength
corresponds to a higher energy of photons and consequently penetra­
tion depth. McSharry et al. (2022) studied the inactivation efficiency of
UV-LED (255, 265, 285 nm) against L. monocytogenes and S. Typhimu­
rium in beef broth and on diced beef and reported 255 nm as the least
effective single wavelength even though it is within the UVC-LED
spectrum, whilst 285 nm was the most effective treatment. Akgün and
Ünlütürk (2017) also reported 280 nm as the most effective wavelength
to inactivate E. coli K12 in both cloudy and clear apple juice, with a 4.40
log reduction after 40 min exposure compared to treatments at 254, 365,
405 nm. UV-LEDs emitting at around 280 nm show a particularly high
inactivation efficiency for some specific bacterial species, probably
because they are close to the maximum absorption of UV light by DNA,
resulting in more direct damage to the microorganism (Forney et al.,
2009; Singh et al., 2021). Therefore, to ensure maximum efficacy of UV
LEDs against specific foodborne pathogens, the wavelengths should be
optimised for the target organisms. Due to different germicidal modes of
action of different UV-LED wavelengths; simultaneous/sequential
application of multiple wavelengths can exhibit additive/synergic
microbial inactivation. This can be due to the repression of photoreac­
tivation and dark repair by damaging of DNA repair enzymes induced by
one wavelength i.e. UVC and thus avoiding repair of DNA damage
caused by another wavelength such as UVB, resulting in a synergistic
inactivation effect (Li et al., 2017; Poepping et al., 2014). A synergistic
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Table 1
Recent studies applying UV-LED treatments on foodborne pathogens and main outcomes.
UV-LED conditions Sample Microorganism Observations Reference

Wavelength Intensity or
(fluency)

265, 285 nm and (0–301.3 Beef broth and Listeria monocytogenes isolated Reductions of 7.5 log reductions in beef broth and 0.7–0.9 McSharry et al.
combinations of 255, mJ/cm2) diced beef from beef log reductions in diced beef after 301.3 mJ/cm2 (2022)
265 and 285 nm Salmonella Typhimurium Reductions of 7.4 log reductions in beef broth and 0.2–0.6
isolated from beef log reductions in diced beef after 301.3 mJ/cm2
405 nm (0–150 mJ/ Phosphate- 18 Salmonella enterica strains Reductions of 1–2 log reduction with doses of 288 J/cm2 Kim and Yuk
cm2) buffered saline and 1.7–5.6 log reduction with doses of 576 J/cm2. Two (2017)
strains of S. enterica were more resistant to UV than the
others
285 and 280/365 nm (0–771.6 Cloudy and clear Escherichia coli K12 (ATCC Bacterial reduction of 2 and 3.9–4.4 log reductions in Akgün and
mJ/cm2) apple juice 25253), an indicator organism cloudy and clear apple juice, respectively. Ünlütürk
of E. coli O157:H7 (2017)
259, 268, 275, 289, and (0–14 mJ/ 0.9% saline Escherichia coli O157:H7, Synergistic effect during simultaneous exposure of E. coli to Green et al.
370 nm cm2) solution E. coli, Salmonella enterica and combination of 259/289 nm (14 mJ/cm2) with a 2.42 log (2018)
Listeria strains reduction.
265 and 365 nm (0–7.5 × Sterile distilled Escherichia coli ATCC 11229, E. Sequential application of 365 and 265 nm increased E. coli Xiao et al.
105 J/cm2) water coli ATCC 15597, and E. coli ATCC 11229, E. coli ATCC 15597, and E. coli ATCC 700891 (2018)
ATCC 700891 reductions by 1.3, 0.9, and 0.5 logs CFU/mL, respectively.
Pre-treatment with 365 nm made them more sensitive to
265 nm achieved after 220 mJ/cm2.
369, 288 and 271 nm 0.75 and Solid agar surface Escherichia coli, Staphylococcus Highest reductions of 1.8–3.2 log were achieved in all the Lu et al. (2021)
and combinations 6.75 mJ/ epidermidis and Salmonella studied bacteria after application of 271 nm. A positive
cm2 typhimurium synergistic effect was obtained when a combination of 288
and 271 nm was conducted.
395 nm 1188 J/cm2 Pet food pellets Two S. enterica strains Bacterial reductions of 1.2 log were achieved after 1188 J/ Subedi and
cm2 Roopesh (2020)
280 nm 21.6 mJ/ Sliced deli meat Escherichia coli O157:H7, All the studied bacteria were reduced in 1–1.6 and 2.4–2.6 (Kim & Kang,
cm2 and spinach Salmonella Typhimurium and log reductions in sliced meat and spinach surfaces, 2020a)
surfaces Listeria monocytogenes respectively. Differences in the bacterial inactivation rates
were observed.
275 nm 4000 mJ/ Raw tuna fillets Salmonella Typhimurium, Reductions of 1.31, 1.86, and 1.77 logs were achieved in S. Fan et al.
cm2 Listeria monocytogenes, and Typhimurium, L. monocytogenes, and E. coli O157:H7 after (2021)
Escherichia coli O157:H7 4000 mJ/cm2, respectively.
275 nm 1200 mJ/ Fresh-cut white Escherichia coli O157:H7 A strain E. coli O157:H7 was reduced by 2.21 log Zhai, Tian,
cm2 pitaya reductions in white pitaya after 1200 mJ/cm2. Ping, Yu, et al.
(2021)

effect during simultaneous exposure of E. coli ATCC 8739 to a combi­
nation of 259/289 nm (14 mJ/cm2) was reported by Green et al. (2018),
who observed a 2.42 log reduction using a 259 and 289 nm combination.
Similarly, Xiao et al. (2018) showed that the sequential application of
UVA-LEDs and UVC-LEDs increased E. coli ATCC 11229, E. coli ATCC
15597, and E. coli ATCC 700891 reductions by 1.3, 0.9, and 0.5 logs
CFU/mL, respectively, where UVA pre-treatment made them more
sensitive to UVC-LEDs. Nevertheless, there are some conflicting results
in the literature such as Song et al. (2019) who reported 1.8 log and 2.8
log reductions for E. coli using 265 nm UV-LED (4.2 mJ/cm2) and 285
nm UVB-LED (15.3 mJ/cm2), respectively, whilst a combination of
these, either simultaneously or sequentially, resulted in up to 4.6 log
inactivation, indicating no additional effect from the 265 nm (UVC-LED)
and 285 nm (UVB-LED) combination. Another study also revealed no
significant synergistic effects using a UVA/UVB (369/288 nm) combi­
nation against five bacteria including E. coli, S. epidermidis, S. Typhi­
murium, Serratia marcescens, and P. alcaligenes compared to the sum of
the two respective UV-LEDs, operating alone (Lu et al., 2021). However,
a synergistic effect was observed for the application of UVB/UVC
(288/271 nm) in combination against these bacterial species. Such an
absence of a synergistic effect using multiple wavelengths has been
explained by referring to the second law of photochemistry, which states
that the photochemical effects of different wavelengths on a molecule
should be independent of each other. Thus, the photonic response
induced by a multiple wavelength combination should only produce an
additive inactivation effect (Beck et al., 2017). Another potential mode
of use of the UV-LED technology could be employing subsequent UV
treatments. This application may influence the response of bacteria and
its investigation could help us to understand better this technology.
The other challenge in developing UV-LED treatments for food safety
applications is the dependence of UVC irradiation on food matrix
characteristics i.e. relative humidity (Subedi et al., 2020), chemical
composition, density, colour, pH, turbidity, and optical viscosity (Koca
et al., 2018). Thus, the processing parameters of UVC should be opti­
mised for a specific food product. The particle size of the food sample is
one such characteristic, which can affect UV inactivation of microor­
ganisms. For example, penetration by UV light irradiation of powders is
difficult due to the shadowing effect of food particles, which protects the
bacterial cells from complete exposure, while bacterial cells can also
protect each other from UV light (Subedi & Roopesh, 2020). UV-LED
treatments are also more efficient for the decontamination of clear so­
lutions and/or smooth surfaces as the presence of UV-absorbing com­
ponents such as pigments, sugars, organic acids, suspended matter
present in juices may increase absorption and reduce the transmission of
UV light, consequently reducing the microbial inactivation efficiency of
UVC-LEDs (Akgün & Ünlütürk, 2017; Koutchma & Popović, 2019).
Differences in inactivation of E. coli O157:H7, S. Typhimurium, and
L. monocytogenes were found when treatments were performed in se­
lective media, deli meat, or spinach by Kim and Kang (2020b). McSharry
et al. (2022) also observed a greater susceptibility for L. monocytogenes
and S. Typhimurium in beef broth compared to the surface of solid beef
after UV-LED treatment. Differences in the UV-LED irradiation effect
according to food type is believed to be due to the different degrees of
absorption of UV light because the surface colour and surface structure
vary for different food materials (Hinds et al., 2019).
3.2. UV-LED effects on foodborne spore forming bacteria
Endospore formation is a mechanism of survival for some species of
microorganisms in harsh environmental conditions. Endospores are

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coated by a multi-layered protective structure which enables them to
resist normal physicochemical treatments, commonly used for inacti­
vation of foodborne pathogens. Bacterial spores are much more resistant
than their vegetative counterparts, and sensitivity to UV-LED treatment
is influenced by this (Brown et al., 2000). In Table 2, recent studies
focused on the application of UV-LED treatments on spore-forming
bacteria are presented. For instance, as highlighted by Zhai, Tian,
Ping, Yu, et al. (2021), at a fluence of 220 mJ/cm2 applied to Alicyclo­
bacillus acidoterrestris cells resulted in a 6.04-log reduction for vegetative
cells, and only a 2.49-log reduction for spores. This is due to the fact that
DNA is located in the core, the inner compartment of the spore, so it is
protected by surface layers and by UV light-absorbing compounds sur­
rounding it (Nicholson et al., 2005). Another reason for increased
resistance is due to the presence of higher levels of the DNA repair
proteins, α/β-type small, acid-soluble spore proteins, and the Ca-DPA
(Lee et al., 2021). A solution proposed for this could include a
pre-treatment of the product to germinate bacterial spores and apply UV
light to their vegetative form. One or two pre-treatments can stimulate
germination of spores could improve the effectiveness of the decon­
tamination technology. According to Buhr et al., 2020, Bacillus anthracis
spores were stimulated with a L-alanine + inosine + calcium dipicolinate
(CaDPA) solution which favoured the germination efficiency at 0–40 ◦ C
with >99% at <8 log spores per mL. These authors observed a reduction
of >99% of the spore population after thermal treatment at 60 ◦ C due to
a higher susceptibility of germinated cells compared to spores. The
lower resistance of germinated cells compared to their spores could also
be an advantage for UV-LED treatment. Moreover, in another study,
high pressure processing (HPP) technology was assessed as a potential
strategy to induce germination of A. acidoterrestris spores. At low pH,
spore germination of up to 3.59–3.75 log and inactivation of 1.85–2.04
log was observed in a low pressure window (200–300 MPa) applied at
50 ◦ C for 20 min. The degree of germination in apple juice after pres­
surization for 30 min with 200 MPa at 20 ◦ C was 2.04 log, with only 0.61
log of spores being inactivated, while at 70 ◦ C spore germination was
5.94 log and inactivation 4.72 log (Sokołowska et al., 2015). These data
represent between 1.22 and 1.74 log cells that have been pre-treated but
not inactivated, germinating and being more susceptible for further
treatments, such as UV-LED. Despite this, some spore-forming bacteria
may germinate more effectively than others, so the effectiveness of a
germination step should be taken into account. This phenomenon could
be affected by the selected pre-treatment and therefore, an effective
combination of pre-germination techniques and further strategies for
decontamination with UV-LED would require optimisation (Krawczyk
et al., 2017). Other studies like Taylor et al. (2020) and Narita et al.
(2020) observed that spores of Bacillus cereus, Bacillus subtilis, Bacillus
thuringiensis, S. aureus, and Clostridioides difficile were sensitive to UV
light at 222 nm and therefore, it should be taken into consideration the
variability in resistance to UV light among spore-forming bacteria.
3.3. UV-LED effect on food spoilage microorganisms
Spoilage microorganisms, which are commonly present in foods, can
influence food quality by their deleterious metabolic activities.
Although the focus on the use of light-based treatment methods have
primarily been targeted to inactivate pathogenic microorganisms in
food, there are some reports indicating their potential to reduce spoilage
microorganisms as well. Thus, these technologies could contribute to
food preservation and extension of the shelf life. Recent publications on
this topic are summarized in Table 3. Bacteria such as A. acidoterrestris
(Zhai, Tian, Ping, Yu, et al., 2021) or S. marcensens (Lu et al., 2021),
yeasts such as Zygossaccharomyces rouxii (Xiang et al., 2020) and molds
including Aspergillus niger (Wan et al., 2020) and Penicillium expansum
(de Souza et al., 2020) have been studied. However, as for pathogenic
bacteria, the different UV equipment configurations and information on
irradiation intensity, and the different ways to express treatment con­
ditions (fluence or time, different units) make comparison between
studies a challenging task. Controlling spoilage microbiota and under­
standing the effect of UV-LED treatments have on food quality attributes
(section 4), is key to increase the shelf life of food products. Having a
collection of comparable studies on the effect of UV-LED on various
microorganisms and complete information on the necessary inactivation
conditions could help to find common strategies to control their growth
and prevent food deterioration.
Some challenges have been already highlighted in section 3.1, that
are also relevant for spoilage microorganisms. A further challenge is
posed by the protective pigments present in some mould species. For

Table 2
Recent studies applying UV-LED treatments on spore-forming bacteria and main outcomes.
UV-LED conditions Sample Microorganism Observations Reference

Wavelength Intensity or
(fluency)

405 nm (0–1.8 kJ/ Bacterial suspension Bacillus cereus NCTC 11143 Reductions of 3.6 log reductions after 1.73 kJ/cm2 Maclean et al.
cm2) Clostridioides difficile NCTC Reductions of 3.2 log reductions after 1.15 kJ/cm2 (2013)
11204
260 and 280 nm (0–150 mJ/ Wastewater from a Bacillus subtilis ATCC 6633 Combination with chlorine (4.0 mg/L) with 265 and Li et al. (2018)
cm2) treatment plant 280 nm LEDs causes up to about 5 log reduction in
B. subtilis spores population(0.4–0.5-log reduction
enhanced with UV alone) in water and significantly
extended the lag phase of the survived spores
285, 365, 405 nm (a) Phosphate buffered Baciluus subtilis DSM 618 Complete inactivation (6.82 log CFU/mL) was achieved (Hinds et al.,
(individually or solution after 285/365 and 285/365/405 nm wavelength 2020)
combined) (b) Nutrient broth combination (5 min treatment, PBS solution). Lower
efficacy observed in NB. Other wavelength
combinations led to reductions ranging from 0.05 log
CFU/mL (365/405 nm, 10 min) to 6.25 log CFU/mL
(285/405 nm, 10 min)
270 nm (0–128 mJ/ (a) Solid agar surface Bacillus subtilis subsp. Complete inactivation on agar (4-log decrease) after 5 s (Nyhan et al.,
cm2) (b) Seasoning powders: Spizizenii ATCC 6633 treatment (128 mJ/cm2). 2021)
garlic, onion, cheese Different inactivation depending on powder: 1-log
and onion, chilli) (chilli), 2.2-log (onion), 2.5-log (garlic, cheese and
onion) after 40 s treatment ((128 mJ/cm2).
275 nm (0–220 mJ/ Orange juice (6 log CFU/ Alicyclobacillus 1-D value equal to 55.98 mJ/cm2 (vegetative cells) and Zhai, Tian,
cm2) mL) acidoterrestris DSM 3922 60.20 mJ cm2 (spores). Complete inactivation of Ping, Yu, et al.
(0–367 mJ/ (vegetative cells and spores) vegetative cells was achieved after 220 mJ/cm2. (2021)
cm2) However, extending the treatment to 367 mJ/cm2 did
not led to an enhanced inactivation on spores.

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Table 3
Recent studies applying UV-LED treatments on spoilage microorganisms and main outcomes.
UV-LED conditions Sample Microorganism Observations Reference

Wavelength Intensity or (fluency)

266, 270, 275, (0.1, 0.2, 0.3, 0.4, 0.5, 0.6 Potato dextrose agar Pichia membranaefaciens Inactivation reached 4-log units after 0.6 mJ/cm2 (Kim, Kim, &
279 nm mJ/cm2) surfaces KCCM 12470 Kang 2017)
Saccharomyces pastorianus Inactivation reached 1-log unit after 0.6 mJ/cm2
KCCM 11523
(a) 255 nm (a) 53.66 mW/cm2 Water Aspergillus niger Less than 1-log reduction was obtained even for the Oliveira et al.
(b) 265 nm (b) 250.3 mW/cm2 highest UV fluence (60 min) for both wavelengths. (2020)
277 nm 2.33 mW cm2 Apple discs (inoculated Penicillium expansum 1-D value equal to 15.27 ± 0.69 mJ/cm2 (25 ◦ C) and (Rios de Souza
on the surface) 16.60 ± 0.21 mJ/cm2 (4 ◦ C) et al., 2020)
2
(a) 265 nm (a) 0.215 mW/cm Phosphate buffered Aspergillus niger 2-D value was 60, 59 and 58 mJ/cm2 for 265, 280 and Wan et al.
(b) 280 nm (b) 0.214 mW/cm2 solution 265/280 nm LEDs, respectively. (2020)
275 nm (200, 400, 800, 1200 mJ/ (a) Saline solution Zigosaccharomyces rouxii Reductions of 4.87-log values after irradiation at 40 Xiang et al.
cm2) (0.85% NaCl) mJ/cm2 in saline solution and of 4.86-log values after (2020)
(b) Apple juice irradiation at 800 mJ/cm2.
(a) 280 (a) (7.71–77.160 mJ/ Mixed beverage (carrot, Mesophilic bacteria and Mesophilic bacteria, and yeast and molds were reduced (Baykus et al.,
(b) 365 cm2) carob, ginger, grape and yeast and molds by 2.59 log CFU/mL (from 5.69 log CFU/mL of initial 2021)
(b) (21.72–217.2 mJ/ lemon juice) load), and 0.17 log CFU/mL (from 3.28 log CFU/mL of
cm2) the initial load), respectively. Although,
(a) 255 nm (a) 3 mW/cm2 Chicken breast fillets Total viable counts Reductions between 1.17 and 1.67 log CFU/g for total Soro et al.
(b) 280 nm (b) 68 mW/cm2 (mesophilic or viable counts of mesophilic, psychrophilic bacteria and (2021)
(c) 300 nm (c) 49 mW/cm2 psychrophilic) total Enterobacteriaceae counts were observed,
(d) 365 nm (d) 45 mW/cm2 Enterobacteriaceae whereas, up to 2 log CFU/g was obtained for
Pseudomonas Pseudomonas and lactic acid bacteria groups after
Lactic acid bacteria treatment with wavelengths of 280, 300 and 365 nm
265 nm 0.34 mW/cm2 (20.4 ± Transport crate Total aerobic bacteria Significant reductions of 1.4 ± 0.4 log CFU/mL (20.4 ± Moazzami
3.36 and 61.2 ± 10.8 mJ/ 3.36 mJ/cm2) and 1.6 ± 0.8 log CFU/mL (61.2 ± 10.8 et al. (2021)
cm2) mJ/cm2)
(a) 265 nm 5.5 mW/cm2 Polyethylene Hyphopichia burtonii Complete inactivation with 5 s in single layer. Belloli et al.
(b) 275 nm 9.0 mW/cm2 terephthalate lids SSICA 175717 (conidia) 1D-value equal to 2–4 s. (2022)
- a single layer (4–5 log Penicillium Complete inactivation with 5 s in single layer.
CFU/lid) brevicompactum SSICA 1D-value equal to 3–13 s.
- multiple layer of cells 14220 (conidia)
(5–6 log CFU/lid) Aspergillus brasiliensis 1-D value equal to 23–58 s (9.0 mW/cm2) and 46–58 s
ATCC 16404 (conidia) (5.5 mW/cm2)
265 nm (420, 840, 1260, and Kimchi seasoning Candida sake MGB0659 Treatments of 1680 mJ/cm2 reduced the number of Song et al.
1680 mJ/cm2) Kazachstania servazzi C. sake cells in kimchi seasoning by 2.29–2.79 log CFU/ (2022)
MGB0660 g, whereas the reduction in the number of K. servazzii
cells was observed to be < 2 log10 CFU/g.

instance, as shown in Table 2, Belloli et al. (2022), found that hyaline
and green-pigmented cells were totally inactivated in times compatible
with industrial practice, whereas melanin-containing fungal spores were
not completely inactivated even after extended treatment times, which
could represent a challenge for the food industry. Melanin and
melanin-related compounds may be responsible for this increased
resistance, as they absorb light in a wide range of wavelengths, including
UV light (Gessler et al., 2014). Melanized fungi such as Cladosporium,
Alternaria and Aspergillus niger will therefore need higher UV fluence to
be controlled, so equipment configurations and treatment times would
need to be carefully considered for use in the food industry (Grishkan,
2011).
4. UV-LED combined with other preservative methods in a
hurdle approach
While there are promising reports indicating the potential of UV-
LEDs for food safety, its application has some limitations such as low
penetration depth, consumer acceptance, and marginal germicidal po­
tency compared to those of thermal treatments which make it chal­
lenging to employ UV-LED for adequate inactivation of foodborne
pathogens (in various food products). Some of these challenges can be
overcome by combining UV-LED treatment with other preservation
methods to cause a hurdle effect. For instance, Shin et al. (2020) com­
bined 222-nm krypton-chlorine (KrCl) excilamp (422.15 μW/cm2) with
UVC-LED (280 nm, 104.31 μW/cm2) to enhance the anti-pathogenic
potential of UV-LEDs against S. Typhimurium and L. monocytogenes in
water. They observed an additive and synergistic inactivation effect
using this approach which was accompanied by reduced populations of
surviving bacteria mainly due to causing irreversible cell membrane
damage. Such a synergic effect may be a result of multiple modes of
action of these treatments, such as inactivation due to membrane
damage caused by the excilamp (Ha et al., 2017), and LED-UVC causing
DNA damage (Kim et al., 2017), resulting in more efficient inactivation.
The germicidal efficiency of UV-LED can also be enhanced by its com­
bination with antimicrobial compounds. A study by Lee et al. (2022)
revealed that the combination of UV-LEDs (365, 385, and 405 nm) with
100 μM folic acid enhanced increases reduction levels of E. coli O157:H7
ATCC 35150 and methicillin-resistant S. aureus ATCC 33591 with up to
>5 log CFU/mL in PBS and apple juice while <1.5 log CFU/mL re­
ductions were achieved for individual LED treatments. Washing with
functionalised liquids before and following UV-LED treatment has also
been shown to give a synergic effect on foodborne pathogens inactiva­
tion by UV-LED. Jiang et al. (2020), reported enhanced decontamination
efficacy when UVC-LED (80, 160, 240 μW/cm2 for 5, 15, 30 min) was
applied to Salmonella ssp. and E. coli inoculated coriander that had
previously been washed in slightly acidic electrolyzed water (SAEW)
reporting reductions of 2.72 log CFU/g and a 2.42 log CFU/g reductions
in Salmonella and E. coli populations, respectively. A 2.97 log CFU/g
reduction of Salmonella spp. in lettuce was also reported by Han et al.
(2021) using a combined treatment of dipping for 2 min in 80 ppm
SAEW followed by UVC-LED treatment at 100 W/cm2. Similar results
have been reported for E. coli O157:H7 and S. Typhimurium in romaine
lettuce and kale by dipping for 5 min in 50 ppm ClO2 and treatment with
UVC at 254 nm (Lee et al., 2012), indicating an improved UV-LED effects
when used in combined with a washing step containing antimicrobials.
Kim et al. (2021) and Lee et al. (2021) also found a similar phenomenon
in which antibacterial activity was synergistically increased by

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sequential treatment of weakly acidic electrolyzed water and UVC-LED
irradiation in fresh-cut vegetables compared with the individual treat­
ments alone. Combinations of UV-LED and mild thermal treatment have
also successfully been used for adequate decontamination of foodborne
pathogenic microorganisms from foods. For instance, Kim and Kang
(2020a) reported that UVC-LED treatment at doses ranging from 0.5 to 3
mJ/cm2 combined with 60 ◦ C mild heat, represents a synergistic
bactericidal effect against L. monocytogenes (~2 log reductions), E. coli
O157: H7 and S. Typhimurium on different contact surfaces during food
processing. In another study, Subedi and Roopesh (2020) developed a
pulsed LED unit to inactivate Salmonella on pet food pellets and reported
that 30 min treatments of the samples with LED alone (395 nm), vi­
bration + mild hot air (50 ◦ C) fluidization, and 395 nm LED treatment +
vibration + mild hot air (50 ◦ C) fluidization led to 1.2, 0.33, and 2.26 log
CFU/g reductions, respectively. They stated that vibration and air
fluidization could increase the exposure of Salmonella to light pulses by
decreasing the shadowing effect. Similar results were also confirmed by
Song et al. (2022), who reported improved inactivation for Candida sake
and Kazachstania servazzii in kimchi seasoning by applying a combina­
tion of an impeller system (70 and 140 rpm) with UVC-LED processing.
Ultrasound pre-treatment or simultaneous application of ultrasound
and UV light has also been considered as an approach to enhance the
disinfection efficiency of UV-LED treatment and reduce photoreactiva­
tion in the effluent, which offers a promising practical application of the
technology. Zhou et al. (2017) reported that the US pre-treatment
positively affects UV-LED inactivation of E. coli in deionized water
containing a kaoline suspension and secondary effluent from a munic­
ipal wastewater treatment plant with enhanced inactivations from 0.24
to 1.07 log CFU/mL and 0.07 to 0.55 log CFU/mL observed, respec­
tively. It was suggested by the authors that by attenuating the protective
effects of suspended particles and decreasing photoreactivation as a
consequence of sonication resulted in improved inactivation efficiency
of UV-LED treatments.
UV-LED combinations with antimicrobial packaging is another pro­
posed approach to enhance inactivation of foodborne pathogenic mi­
croorganisms. In a study conducted by Kim et al. (2022), UVC-LED
treatment led to a reduction of 0.47, 0.55, and 0.61 log CFU/g for
E. coli (EHEC) and 0.46, 0.39, and 0.33 log CFU/g for S. aureus in
fresh-cut onion, cabbage, and carrot, respectively. However, when
combined with antimicrobial packaging containing grapefruit seed
extract and zinc oxide nanoparticles added to a blend film of poly(lac­
tide) and poly (butylene adipate-co-terephthalate)) resulted in a signif­
icant synergistic effect on reducing these bacteria 7 days of storage at
10 ◦ C.
5. Challenges to UV-LED application related with food quality
In order to evaluate the feasibility of light-based treatments for food
safety or shelf life applications, it is important to consider possible
quality changes following UV-LED treatment. A number of studies have
demonstrated positive effects of UV-LED on quality attributes of foods
such as inactivation of food spoilage microorganisms, improving nutri­
tional quality by increasing phytochemical compounds, reduction of
weight loss, and improving techno-functional properties of food mate­
rials (Fan et al., 2021; Soro et al., 2021; Subedi et al., 2020). However,
the main concern following UV-LED treatment is induced photochemical
reactions of food ingredients, especially in food pigments and lipid
peroxidation. The following section will give an overview of potential
quality changes following UV-LED treatment.
Visual appearance is the first quality feature of food which de­
termines consumers’ direct judgment of quality and safety. Several
studies have shown that UV-LED treatment does not induce significant
changes in colour parameters of food. For example, Akgün and Ünlütürk
(2017) reported that moderate multiple LED (UVA and UVC) treatments
even exhibit a better preservation of the colour of cloudy apple juice
compared to thermal treatments, although adverse effects were
272
Trends in Food Science & Technology 131 (2023) 264–276
observed in colour compounds after long exposure times. Subedi et al.
(2020) also reported no colour changes in wheat flour following 275-nm
pulsed LED treatment at a dose of 60.1 mJ/cm2 and an intensity of 16.7
μW/cm2 for 60 min, although, treatment at 365, 395, and 455 nm
caused a significant change in wheat flour colour. Similarly, negligible
changes in the colour of mixed beverages were observed after exposure
to 280, 365 nm and a wavelength combination of 280/365 nm for 10,
20, 30, 40, 60, 80, and 100 min at room temperature (25 ◦ C) (Baykuş
et al., 2021) and diced chicken breast fillets exposed to UV light wave­
lengths of 255, 280, 300 and 365 nm for 2, 4, 6, 8 and 10 min (Soro et al.,
2021). Soro et al. (2021) reported that treated samples had a better
colour quality after 7 days storage at 4 ◦ C, for samples treated at 280 nm,
with no significant changes during storage in terms of L*, a* and b*
values for treated samples. Fan et al. (2021) also noted no remarkable
changes in L*, a*, and b* and whiteness index, and the percentage of
metmyoglobin in raw tuna fillets following UVC-LED irradiation
(500–4000 mJ/cm2). In contrast, Zhai, Tian, Ping, Yu, et al. (2021)
observed that UVC-LED (275 nm) treatment caused significant changes
in the colour attributes (a*, b*, chroma, hue angle, and total colour
difference) and browning index of orange juice. Similar results were
reported by Xiang et al. (2020), who also observed remarkable changes
in some colorimetric parameters of apple juice, exposed to UVC-LED
irradiation; significant decrease in a* value (greenness) and increases
in b* value (yellowness) of apple juices which were irradiation dosage
dependent. An increase in C* (chroma) value and hue angles could be
due to colour saturation and oxidation reactions during UVC-LED
treatment, respectively. Another possible explanation for the changes
in colour of UVC-LED-treated juices could be due to accelerated isom­
erization of carotenoids and their interactions with free radicals pro­
duced during light treatment (Bhat, 2016). A significant increase in the
browning index was also observed after UVC-LED treatment at 1200
mJ/cm2, and was explained by the Maillard reaction and oxidation of
certain phytochemicals such as ascorbic acids and some polyphenols
during UVC-LED irradiation (Riganakos et al., 2017).
It has been well documented that UV-LED treatment does not influ­
ence physicochemical properties, such as pH, titratable acidity, total
soluble solids, dissolved solids (Brix), clarity of exposed food materials,
including apple juice (Xiang et al., 2020), mixed beverage (Baykuş et al.,
2021), fresh-cut white pitaya (Zhai, Tian, Ping, Yu, et al., 2021), kimchi
seasoning (Song et al., 2022), chicken breast fillets (Soro et al., 2021), or
raw tuna fillets (Fan et al., 2021).
A challenge of UV-LED application for food safety purposes are also
photochemical reactions (oxidation) of food ingredients, which can
affect the quality of the treated food materials. However, limited studies
are available which address photochemical reactions of food ingredients
under UV-LED treatment. Fan et al. (2021) reported that UVC-LED
irradiation led to increased levels of lipid oxidation in tuna fillets. This
was confirmed by a significant increase in the TBARS values following
UVC-LED irradiation at 1000, 2000, and 4000 mJ/cm2. The TBARS
values of raw tuna fillets were increased to 0.50, 0.58, and 0.73 mg
MDA/100 g, respectively, which were significantly higher than that of
the control samples (0.43 mg MDA/100 g). This can be explained by
induced oxidation by various reactive species such as superoxide anion
(O2•-), hydrogen peroxide (H2O2), and the hydroxyl radical (•OH)
(D’Orazio et al., 2013) generated during UV-light exposure. Indeed,
some researchers have used UV-LED oxidative effects to improve the
techno-functional properties of some food products. For instance, Subedi
et al. (2020) reported improved functional properties of UV-LED treated
wheat flour at 275, 365, 395, and 455 nm. Briefly, wheat flour showed a
reduction in yellowness, water holding capacity (by increasing free –SH
groups, thus affecting surface hydrophobicity), polymerization,
improved drying process, and an altered secondary structure of gluten
following UV-LED treatment in all the studied wavelengths, except for
275 nm which had no impact on the colour. The authors of this study
suggested that colour changes after treatment with UV-LED at 365, 395
and 455 nm may be due to the oxidation and thermal degradation of
A.B. Soro et al.
carotenoids in wheat flour (Subedi et al., 2020).
UV-LED treatment does not have an immediate effect on texture
parameters of treated food. In a study by Fan et al. (2021), there were
no significant differences in texture parameters between raw tuna fillets
exposed to UVC-LED (up to 4000 mJ/cm2) and control groups, including
hardness, springiness, cohesiveness, gumminess, chewiness, and resil­
ience were observed. Similarly, Soro et al. (2021) reported no significant
changes in the texture of chicken breast fillets, including hardness,
springiness, cohesiveness, gumminess, and chewiness after UV-LED
treatment. Furthermore, tapioca starch samples subjected to various
UV-LED treatments (280, 300, and 365 nm for 5, 10, and 20 min)
showed no significant changes in pasting properties (peak viscosity,
breakdown viscosity, final viscosity, setback viscosity, and pasting
temperature) or surface morphology for any treatments investigated
(Hinds et al., 2022).
Regarding the effects of UV-LED on sensory attributes of food
products, conflicting results have been reported. While Jiang et al.
(2020) described preserved sensory properties and an extended shelf life
of coriander after UVC-LED and slightly acidic electrolysed water
treatment, Martin et al. (2016) observed a negative impact of UV-LED
treatment on the sensory quality of liquid milk.
Various studies have shown both increases and decreases in phyto­
chemical content and antioxidative properties of food products after
exposure to UV-LED irradiation, depending on the treatment conditions
and the food matrices, which can affect nutritional quality of the treated
food. Baykuş et al. (2021) reported that a mixed beverage subjected to
UV-LED treatment [at 280 and 365 nm, and their combination for 10,
20, 30, 40, 60, 80, and 100 min at room temperature (25 ◦ C)] had a
significantly higher total phenolic compounds (TPC) content and anti­
oxidant capacity compared to untreated and heat-treated samples
(pasteurized at 70 ◦ C for 120 s). TPC and antioxidant capacity in UV-LED
treated samples were 1.75-fold and 4.60 fold higher compared to
heat-treated samples, respectively. Nevertheless, UV-LED treatment
caused a significant decrease of up to 71.3% in carotenoid content,
however this decrease was still lower than that of heat-treated samples
(88.9%), indicating that UV-LED irradiation preserved the total carot­
enoid content better than the heat treatment. Similar results were re­
ported by Hinds et al. (2021) for black peppercorns subjected to UV LED
at 280, 300 and 365 nm for 5, 10, and 20 min. Others have reported
sizeable reductions in TPC and antioxidant activity of food materials
following UV-LED irradiation. For example, Xiang et al. (2020) reported
a significant decrease of 3.03, 5.40, 8.97, and 10.45% in TPC of apple
juice after exposure to UVC-LED at 200, 400, 800, and 1200 mJ/cm2.
Such a decrease in TPC was explained by oxidative degradation of
phenolic compounds induced by UVC-LED irradiation (Islam et al.,
2016). This was further confirmed by observing a lower antioxidant
capacity in apple juice treated with UVC-LED. Another reason for a
lower antioxidant capacity was the degrading effects of UVC-LED irra­
diation on ascorbic acid, a water-soluble antioxidant in fruit and vege­
table juices (Bhat, 2016). Zhai, Tian, Ping, Yu, et al. (2021) also reported
that UV-LED treatment (275 nm) of fresh-cut pitaya at high fluence
(800–1200 mJ/cm2) leads to a small but statistically significant reduc­
tion in antioxidant properties of fresh-cut pitaya, while TPC remained
unchanged after UVC-LED treatment at a range of 200–1200 mJ/cm2.
Similar fluence dependent decrease in the TPC of orange juice exposed
to UV-LED was also reported by Zhai, Tian, Ping, Yu, et al. (2021).
6. Conclusion and future directions
In recent years UV-LED technology has attracted considerable in­
terest as an alternative method for inactivation of foodborne pathogens.
UV-LED possesses several advantages such as low radiant heat emis­
sions, high emissions of monochromatic light, and low energy con­
sumption compared to conventional UV low-pressure mercury lamps.
Although, the effectiveness of UV-LED to inactivate a wide range of
foodborne pathogenic microorganisms has been proven, various stress
273
Trends in Food Science & Technology 131 (2023) 264–276
adaptive responses may allow pathogens to survive UV light exposure.
These adaptive responses can lead to modifications in cell physiology
due to DNA repair mechanisms, such as photoreactivation, recombina­
tional and excision repair (dark repair), and SOS response. Further
studies are therefore required to explore DNA repair mechanisms of
different microbial species exposed to UV light at different wavelengths
and their combinations either simultaneously and/or sequentially. For
these purposes employing techniques, such as whole genome
sequencing, transcriptomics, proteomics, and metabolomics should be
considered for future studies. Adapted cells may exhibit alterations in
their cellular structures and functions, such as biofilm formation,
turning to viable but non-culturable states, AMR, transcriptional regu­
lation, and changes in protein expression. Such adaptations need to be
investigated in the future, particularly in the context of how it may in­
fluence tolerance to other stressors in the food production chain. Some
limitations of UV-LED application, such as low penetration depths and
marginal germicidal potency compared to those of thermal treatments
may increase the changes of cell survival. To overcome microbial
adaptive responses, a combination of UV-LED treatment with other
preservation methods such as visible and blue light LEDs, mild heat
treatment, washing with electrolyzed water and ultrasound may need to
be considered as a hurdle approach to enhance UV-LED germicidal ef­
fects. However, there are limited studies, some with contradicting re­
sults, on UV-LED and their application for food safety purposes.
Therefore, additional research is required to reach conclusive results and
recommendations that permit the employment of UV-LED to inactivate
foodborne pathogens. Furthermore, there is some evidence showing UV-
LEDs can negatively affect food quality properties by altering the
physicochemical attributes of food materials such as colour, lipid
oxidation, antioxidant capacity, and texture. Thus, more attention
should be paid to the underlying mechanism of quality changes induced
by UV-LED, when used for pathogen inactivation. To the best of our
knowledge there is also a lack of information on consumer acceptability
of UV-LED-treated foods, which would also need to be addressed in
future studies.
Author contributions
Conceptualization, writing—original draft preparation, wri­
ting—review and editing, Soro, A.B.; conceptualization, writing-original
draft preparation, Shokri, S.; conceptualization, writing-original draft
preparation, Nicolau-Lapeña, I.; writing—review and editing, graphical
design of figure, Ekhlas, D.; writing—review and editing, supervision,
Burgess C.M; writing—review and editing, supervision, Whyte, P.; wri­
ting—review and editing, supervision, Bolton, D.J.; writing—review
and editing, supervision, Bourke, P.; writing—review and editing, su­
pervision, project administration, funding acquisition, Tiwari, B.K.
Declaration of competing interest
The authors declare no conflict of interest.
Data availability
No data was used for the research described in the article.
Acknowledgments
The present work was funded by the Teagasc Walsh Scholarship
program, Department of Agriculture, Food, and Marine (DAFM) under
the Food Institutional Research Measure (FIRM) program [Grant num­
ber: DAFM/17/F/275].
A.B. Soro et al. Trends in Food Science & Technology 131 (2023) 264–276

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