1 s2.0 S1571064523000076 Main

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ScienceDirect
Physics of Life Reviews 44 (2023) 207–266
www.elsevier.com/locate/plrev
Review
Advances in pulsed electric stimuli as a physical method for treating

liquid foods
Farzan Zare a,b , Negareh Ghasemi a , Nidhi Bansal b , Hamid Hosano c,∗
a School of Information Technology and Electrical Engineering, The University of Queensland, Brisbane, St Lucia QLD 4072, Australia
b School of Agriculture and Food Sciences, The University of Queensland, St Lucia QLD 4072, Australia
c Biomaterials and Bioelectrics Department, Institute of Industrial Nanomaterials, Kumamoto University, Kumamoto 860-8555, Japan
Received 24 January 2023; accepted 28 January 2023

Available online 2 February 2023
Communicated by M. Frank-Kamenetskii
Abstract
There is a need for alternative technologies that can deliver safe and nutritious foods at lower costs as compared to conventional
processes. Pulsed electric field (PEF) technology has been utilised for a plethora of different applications in the life and physical
sciences, such as gene/drug delivery in medicine and extraction of bioactive compounds in food science and technology. PEF
technology for treating liquid foods involves engineering principles to develop the equipment, and quantitative biochemistry and
microbiology techniques to validate the process. There are numerous challenges to address for its application in liquid foods such
as the 5-log pathogen reduction target in food safety, maintaining the food quality, and scale up of this physical approach for
industrial integration. Here, we present the engineering principles associated with pulsed electric fields, related inactivation models
of microorganisms, electroporation and electropermeabilization theory, to increase the quality and safety of liquid foods; including
water, milk, beer, wine, fruit juices, cider, and liquid eggs. Ultimately, we discuss the outlook of the field and emphasise research
gaps.
© 2023 Elsevier B.V. All rights reserved.
Keywords: Pulsed electric field; Liquid foods; Safety; Quality
Contents
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
2. Pulse waveform, generation, and delivery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
3. Inactivation kinetics of microorganisms by PEF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
3.1. Bigelow model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
3.2. Peleg model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
3.3. Hülsheger and Niemann model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
* Corresponding author.
E-mail address: hamid@kumamoto-u.ac.jp (H. Hosano).
https://doi.org/10.1016/j.plrev.2023.01.007
1571-0645/© 2023 Elsevier B.V. All rights reserved.
F. Zare, N. Ghasemi, N. Bansal et al. Physics of Life Reviews 44 (2023) 207–266
3.4. Weibull distribution model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216

4. Electroporation, electropermeabilization, and associated mechanisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
5. Quality and safety of liquid foods treated with PEFs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
5.1. Water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
5.2. Milk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
5.3. Beer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
5.4. Wine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
5.5. Fruit juices and cider . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
5.6. Liquid egg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
6. Outlook . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Declaration of competing interest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240
1. Introduction
Bioelectrics encompasses both biological science and engineering principles to apply electrical stimuli to biologi-
cal systems [1]. Pulsed power technology is used to apply energy in a very short time (micro-to-nano second) in the
forms of pulsed electric fields (PEFs), plasmas, shock waves, pulsed electromagnetic waves, and pulsed light [1]. PEF
technology is a physical method of interest in the food, agricultural, and biomedical industries for its numerous and
continuously growing applications such as liquid food treatment [2,3]; growth stimulation and increasing the produc-
tion yield of plants and filamentous fungi [4–7]; modulating fermentation [8–10] and chemical reaction speeds [11];
increasing biomass and drying of insects [12]; enhanced extraction yields of biofuels and/or bioactive compounds
from microorganisms (i.e. algae and yeast) [13–17], fruits and vegetable waste [18–22], and meat products [23]; as-
sisting food preservation methods by reducing drying and freezing times corresponding to energy efficiency [24–27];
modification of biomacromolecules [28,29]; pre-treatment of fruits and vegetables with PEF for increasing physical
properties (e.g., smoothening [30], hardness and crispness [31]) and subsequent increased extraction of juices [32];
electrochemotherapy or drug delivery [33–35], transfection and transformation [36], electrofusion [37], or tumour
ablation [38–40]. The application of bioelectrics for cancer treatment, either as irreversible or reversible cell damage
modalities, has improved significantly. Albeit having its own limitations, such as minimising damage to surrounding
healthy tissue, the application in the medical fields can still be extended to assist food and agricultural science indi-
rectly. The mechanisms associated with optimising permeabilization, and hence mass transfer, has numerous benefits
for modulation of fermentation or kinetics of chemical reactions, extraction, and cell-death mechanisms.
PEF technology allows for the modulation of the resting membrane potential of cells, where transient pores of
various sizes (up to tens of nanometres in radius [41]) are created which allow larger or previously impermeable com-
pounds to cross the cell membrane or envelope. The application of electrostatic and/or time-varying fields to liquids
can cause numerous electrokinetic and dielectric phenomena (electromigration, electrophoresis, dielectrophoresis,
electroosmosis, electrorotation, electro-diffusion) [42].
PEF-induced changes to biological membranes can cause four states as outlined by Alan Barnett and James C.
Weaver [43,44]; (1) increase in membrane conductance, (2) mechanical rupture [45], (3) incomplete reversible break-
down, (4) and reversible breakdown. The formation of aqueous pores across the lipid bilayer under the influence of
electric fields has been extensively studied in passive and active models and is widely accepted as electroporation,
however the kinetics of reversible and irreversible pore-formation and -closure and associated lifetimes of such pores
are highly debated as they are intrinsically dependent on the compositions of the membrane(s) and solution(s) (intra-
and extracellular), as well as the simulation parameters and accuracy of the model [46–66]. Passive models refer to
simulation models where the electrical properties are ‘static’ in the time-domain. During the electroporation event
of the membrane, there will be a change in conductivity which is not accounted for in the passive (linear and time-
invariant) model [67]. However, the active model (non-linear) accounts for this difference and allows the electrical
properties to vary in the time-domain (e.g., conductivity increases during pore formation which consequently causes a
drop in the transmembrane voltage) [68]. The formation of aqueous pores across the lipid bilayer under the influence
of electric fields has been extensively studied in passive and active models and is widely accepted as electroporation.
208
The kinetics of reversible and irreversible pore-formation and -closure and associated lifetimes of such pores are in-
trinsically dependent on the compositions of the membrane(s) and solution(s) (intra- and extracellular). Moreover,
the simulation parameters and accuracy of the model are highly debated as they must subsequently be related to ex-
perimental investigations for validation [46,47,50,55,63]. Many authors have attempted to model electroporation and
“soft poration”, which is pore creation without the addition of an external electric field like in electroporation, using
the Smoluchowski equation [54,56,57,59–61]. The authors in [60] have interestingly derived pore parameters (such
as pore edge tension and initial distribution density of the number of pores) that are independent of the membrane
voltage when considering “soft poration” utilising the Smoluchowski equation. The Smoluchowski equation for the
purposes of analysing electroporation has also been coupled to the Laplace equations to relate the conductivity and
transmembrane voltage in a single shell [52] and double shell cell models [53]. It has been shown numerically [57,62]
that the asymmetry in the pore creation can be contributed from the resting membrane potential when operating near
the critical transmembrane potential required to induce electroporation [57]. An increase in permeability of the mem-
brane that cannot be explained exclusively by the electroporation model is referred to as electropermeabilization [69].
PEF-induced changes to the membrane of cells is a multi-phase process and may be solved using Poisson’s equations
rather through Laplace’s, as there is a non-zero charge density associated with the internal and external leaflets of the
membrane [70]. However, both the Laplace and Poisson equations treat ions as point charges and the dielectric mem-
brane as a continuous medium which may not be an appropriate assumption when analysing pores on the nanometre
scale [71].
Cell viability is reduced after PEF processing predominantly thought to be through irreversible permeabilization
of the membrane or cell viability is maintained via reversible permeabilization to extract or deliver a compound of
interest. The reversibility of a PEF protocol is dependent on the biophysical structure and composition of the cells
of interest, the response of the organism(s) to the physical stimuli, the matrix composition, and the PEF process
parameters (i.e., the pulse width, shape and polarity, electric field intensity, and specific energy input) [72].
The key advantage of PEF technology for the liquid food industry is that the electroporation/permeabilization
theory is a non-thermal method to facilitate the inactivation of microorganisms [69], and if optimised appropriately
it can operate at lower processing temperatures than thermal treatment (e.g., holder pasteurization) [73]. Therefore,
the food product has higher bioactive retention while also reducing the bacterial count of the liquid food sample (i.e.,
longer shelf life). Simulation packages that use the finite element method (FEM) are necessary to determine the spatial,
temporal, and frequency dependent interactions of the electric field and PEF-induced temperature increase as the
complexity of electrode geometries increases and simple approximations are not sufficient. These simulations coupled
with molecular dynamics (MD) models are powerful tools to evaluate the efficiency of the treatment process for
the target application [47,65,66,74–77]. Furthermore, through optimisation of the pulse protocol, treatment chamber
design, and with recent advances in wide-band gap (WBG) high powered electronic switches (i.e., silicon carbide and
gallium nitride [78]) the energy consumption may be lower than thermal treatment [79]. These WBG switches can be
used to develop pulsed power generators (PPGs) with ultrafast rise time (i.e., picoseconds) that can deliver hundreds
of kilovolts.
Within the scope of this review, pulsed electric field processing of liquid food products and the related process
parameters and mechanisms are discussed based on recent advances in the field.
2. Pulse waveform, generation, and delivery
Pulsed power generators (PPGs) store energy in the form of electric and/or magnetic fields through a capacitive
or inductive energy storage circuit, respectively. Although the instantaneous power of the delivered pulse can be very
large (i.e., gigawatt), the energy of each pulse is low (i.e., milli Joule to a few Joules) because the duration of the
pulses is usually in the microseconds to nanoseconds (µs – ns) range. This is shown in Eq. (1).
t2 t2
Eenergy = P (t) dt = V (t) i (t) dt (1)
t1 t1
where, Eenergy represents the total energy of the pulse (Joule), P (t), V (t) and i(t) are the instantaneous power (Watt),
voltage (V), and current (Ampere) delivered to the load (i.e., the sample of interest) from time t1 to t2 , respectively.
209
Key process parameters for PEF applications include the electric field intensity (kV.cm−1 ), treatment time (s),
specific energy input (J.kg−1 ), number of pulses per second (Hz), pulse waveform, pulse width, and temperature.
The pulse waveform can be of a unipolar or bipolar nature, where typical pulse shapes are chosen to be an expo-
nentially decaying or square wave for PEF treatment of liquid foods.
For a parallel-plate electrode geometry, the average electric field is characterised by Eq. (2a), where, Ef ield is the
electric field intensity, V is the applied voltage, and d is the distance between the two parallel-plate electrodes. For
a coaxial treatment chamber, the peak electric field strength can be determined by Eq. (2b), where ‘r’ is the radius
where the voltage is measured, and R1 and R2 are the inner and outer radii, respectively [80]. For complex electrode
geometries it is necessary to use numerical methods such as the finite element method to evaluate the electric field
distribution in the treatment chamber.
V
Ef ield = (2a)
d
V
Ef ield = (2b)
r. ln R 2
R1
The treatment time, t, is given by Eq. (3). Where, n is the number of pulses and τpulse is the pulse width of the applied
voltage waveform. For square wave pulses with fast rise times equation (3) is always valid, however for exponentially
decaying pulses the effective pulse width (36.8% of the peak voltage) may be used to approximate [81].
t = n.τpulse (3)
The specific energy input, Ws or Q (J.kg−1 ),
is used to evaluate the efficacy of the treatment and is given by Eq. (4).
Where, m is the mass of the liquid, Eenergy,k is the energy of the kth pulse from Eq. (1), and n is the applied number
of pulses.
t2
1 1
n n
Ws = Q = . Vk (t) ik (t) dt = . Eenergy,k (4)
m m
k=1 t1 k=1
The energy delivered to a sample or load (i.e., cuvette containing liquid sample) depends on the output impedance of
the PPG unit (i.e., switch impedance) and the impedance of the sample (cuvette and liquid). The impedance of the
sample will deviate during the application of a pulse train, such as from joule heating (i.e., the conductivity of the
liquid may increase which reduces the effective pulse width and temperature of electrode changes [82]) or changes in
the impedance of the electric double layer. Therefore, each pulse may have a different energy profile.
For continuous PEF mode, the total energy input can be calculated by Eq. (4a), where Eenergy is the energy per
pulse (assuming it remains constant for the entire duration of the treatment), f is frequency of excitation or pulse
repetition rate (Hz), and ṁ is the mass flow rate (kg.s−1 ) [79].
f.Eenergy
Ws = Q = (4a)
ṁ
The number of applied pulses per second (pps), or the pulse repetition rate is measured in Hertz and does not exclu-
sively correlate to the spectral content of the pulses. The spectral content refers to the frequency-domain representation
of the time-domain pulses through the Fourier Transformation (FT). The Fourier series of an ideal square wave (i.e.,
infinite dV.dt−1 or instantaneous rise/fall time) is characterised by the sum of all odd-integer sinusoidal harmonics,
e.g., a periodic bipolar square wave signal at 100 Hz will have 1st and 2nd harmonics at 300 Hz and 500 Hz, respec-
tively. For a short (µs – ns) square-wave in time-domain, the spectral content is represented as a sinc function over a
wide range of frequencies. This will be different depending on the shape and periodicity of the applied signal, and the
impact of the first and second harmonics may be significant due to the amplitude of the applied voltage (typically in
the kV range) [83].
The pulse delay time is characterised by Eq. (5), where ‘f ’ is the frequency of the applied pulse protocol which is
sometimes referred to as the pulse repetition rate (Hz).
1
tdelay = − τpulse (5)
f
210
The pulse width of the applied pulses, τpulse , can impact the biophysical responses, the energy delivery, and can con-
tribute to temperature increase of the load (i.e., liquid sample). The temperature of a solution is inversely proportional
to resistivity, so as the temperature of the solution increases the conductivity also increases. As the total number of
pulses, the pulse width, pulse amplitude or the pulse repetition rate is increased, the temperature rise can become
significant depending on the application and volume of the liquid sample. This rise in temperature will cause a drop
in the electric field strength and a consequent increase in the current consumption per pulse.
Emphasis needs to be placed on evaluating the influence of temperature during PEF excitation to isolate the effect
of temperature itself, and the combinatory mechanism. To measure the rising temperature, it is recommended to use
a fibre optic temperature sensor as the electromagnetic coupling will not interfere with the measurement system as
compared to thermocouples [84]. Assuming that no cooling is applied to the sample, the theoretical temperature
change (◦ C) in the liquid sample can be calculated from Eq. (6) for batch treatment [85], where Qv or Wv (J.m−3 ),
ρf (kg.m−3 ) and Cf (J.kg−1 .◦ C−1 ) are the specific energy input, fluid density, and specific heat capacity of the liquid
sample under investigation, respectively.
Qv
T = (6)
ρf .Cf
Equation (7) can be used to calculate the theoretical temperature increase for PEF treatment in continuous mode [86].
Where Eenergy is the energy per pulse calculated as per Eq. (1) (sometimes approximated as Eenergy = V I τ , although
this approximation is only appropriate for square wave pulses), f is frequency, tr is the resident time of the liquid
media within the treatment zone (s), and v is the volume of the PEF chamber.
E.f.tr
T = (7)
ρf .Cf .v
A transient thermal increase faster than the thermal expansion of the system can induce a thermal shock and strain in
the system [87].
The electrode configuration used within the treatment chamber, such as point-to-point (needle-to-needle) or plate-
to-plate, will have a significant influence on the homogeneity of the electric field and affect the current density
distribution within the media. The current density distribution contributes to joule heating (i.e., losses) and the gener-
ation of a magnetic field tangential to the flow.
The energy delivered to the treatment chamber will be absorbed by the liquid (including microorganisms) and the
electrodes. The energy balance is shown in Eq. (8) [88].
Qt = ρe .ve .Ce .Te + ρf .vf .Cf .Tf (8)
where Qt is the total energy absorbed by the load (J), ρe and ρf are the density of the electrode and fluid (kg.m−3 ),ve
and vf are the volumes of the electrode and fluid (m3 ), and Ce and Cf are the specific heat capacities of the electrode
and fluid (J.kg−1 .◦ C−1 ), and Te and Tf are the change in temperature for the electrode and fluid (◦ C), respectively.
The specific energy input delivered to the load is commonly used as an indicator of PEF process efficiency. How-
ever, this parameter should not be exclusively used to indicate the inactivation efficiency of the protocol. The energy
profile alone provides no information about the shape, duration, rise/fall time of the instantaneous voltage and current
waveforms or the pulse protocol (method of delivery, e.g., 100 unipolar square wave pulses with 10 µs pulse width at
1 kHz with an electric field strength of 20 kV.cm−1 ).
For PEF excitation the rise/fall time of the voltage waveform is a parameter which can influence an ‘effective
energy delivery’ to the load. The rise and fall time of a square wave pulse contains the dominant portion of the high
frequency content of the pulse [89]. Therefore, by reducing the rise/fall time (i.e., increasing dV.dt−1 ) we can increase
the high frequency spectra of the protocol and the temperature gradient of each pulse (i.e., dT.dt−1 ) [89].
The following two pulses in Figs. 1 (a, b) by Góngora-Nieto et al. [81] are an example of how the rise and fall
time of a pulse can influence the ‘effective treatment energy’. This definition of effective treatment energy may not be
true for all applications. Lower electric field strengths with longer treatment times, such as the long fall time of the
exponentially decaying pulse of Fig. 1a, were shown to enhance the permeabilization of tissues and microorganisms
in different media [90], and affect protein conformational changes [91].
Furthermore, the rapid changes in voltage (rise time of Fig. 1a and rise/fall time of Fig. 1b) corresponds to high
frequency spectra which will affect the load differently as compared to the mean amplitude (the DC component) of
211
Fig. 1. Effective treatment energy and non-effective treatment energy for (a) slightly overdamped unipolar exponentially decaying pulse, and (b) a
pseudo square pulse with slow rise and fall time. Reproduced from [81], with permission from Elsevier.
Fig. 2. Equivalent electrical model of a solution or bulk electrolyte between two parallel plate electrodes, where PPG represents the pulsed power
generator, Cdl is the electric double layer formed at the liquid-electrode interface, Rct and ZW are the charge transfer resistance and the Warburg
impedance.
the pulse. The pulse rise time was also found to contribute to the physicochemical and sensory properties of apple
juice treated with PEF [92]. PEFs were found to better maintain the antioxidant capacity, phenolic substance content,
organic acids, and anthocyanin compounds as compared to thermal pasteurization [92–94]. Short pulse rise times were
found to maintain the enzymatic activity as compared to longer pulse rise times [92].
PEF excitation has been ubiquitously shown to influence biological membranes when operating above a threshold
electric field strength and has little to no effects when below this threshold [90,95–97]. This threshold varies depend-
ing on numerous factors such as membrane composition, medium and pulse properties; thus, there is no universal
threshold electric field which will hold for all applications. In applications where pulses operate near the threshold
electric field the ‘effective treatment energy’ definition is more appropriate as it also correlates to portions of the pulse
which maximise the exposure time of the peak voltage that is induced on the plasma membrane by the pulse waveform
[98].
The equivalent electrical model of a typical two-electrode cuvette or treatment chamber can be described as shown
in Fig. 2, assuming the double layer capacitor, Cdl , is fully charged [99,100]. Rs and Cs are the resistance and capac-
itance of the solution, Cdl is the capacitance of the electrical double layer formed at the liquid-electrode interface for
each half-cell reaction [99,100]. Rct and ZW are the charge transfer resistance and the Warburg impedance, together
they represent the faradaic impedance which models the faradaic reactions at the electrode-electrolyte interface [100].
The faradaic impedance and the double layer capacitor behave non-linearly as they are intrinsically dependent on the
applied voltage [100].
In Fig. 3, Ohshima et al. [101] showed that when the experimental conditions except the liquid media were kept
constant (i.e., the PPG, the setting on the PPG which set the energy and amplitude of each pulse, and the treatment
chamber) the instantaneous voltage and current waveforms changed depending on the liquid media type. The effective
pulse width of the voltage waveform applied to the distilled water sample in Fig. 3a is approximately 500 ns with a
212
Fig. 3. Instantaneous voltage and current waveforms applied to distilled water (a, b), and full cream milk samples (c, d). Reproduced from [101],
with permission from Elsevier.
peak voltage of greater than 50 kV, whereas for milk in Fig. 3c the effective pulse width is less than 200 ns and has a
peak voltage of less than 40 kV [101]. Furthermore, the instantaneous current waveforms are drastically different in
shape, peak amplitudes, and relative time constants; as shown in Figs. 3 (b, d) [101].
The reason why this change in pulse characteristic occurs in different media types is due to the equivalent electrical
model of the system (i.e., source to load). Distilled water has low electrical conductivity as compared to the highly
conductive milk medium (predominantly due to its mineral content), as shown by Eq. (9). Resistance is described
by the conductivity of the solution (σ ), length of sample between electrodes (d) and surface area in contact with
electrodes (A).
d
R= (9)
σ.A
Furthermore, since both medias have different dielectric constants, εr , this will also influence the capacitance as shown
in Eq. (10).
ε0 .εr .A q
C= = (10)
d V
where the capacitance is affected by the surface area of the medium (A), relative permittivity of the medium, εr , and
the distance, d, between the charged surfaces. Distilled water has a larger impedance than milk because of its lower
conductivity and higher capacitance, therefore the pulse applied to the distilled water is expected to have a higher
amplitude and wider pulse width.
The conductivity of the liquid sample can influence the type of discharge occurring between the electrode inter-
faces. The lower the impedance of the liquid sample, the higher the probability of developing discharges at sufficient
electric field strengths, (such as 20-30 kV.cm−1 or greater), depending on the liquid sample, dissolved gases, presence
of bubbles, energy of the pulse, frequency, electrode geometry and material, temperature, and pressure [102–104]. If
it is a high impedance liquid sample, it typically will form an arc. The lower the impedance, it approaches a streamer-
like discharge. These discharges can cause secondary effects such as shock waves, UV radiation, and the production of
reactive oxygen species (ROS) and reactive nitrogen species (RNS). The interaction of oxidizing agents with stainless
steel alloys, commonly used in industrial food applications (i.e., SS316 or SS304), can corrode the electrodes which
consequently decreases their lifetime. Furthermore, it can negatively impact the nutritional or organoleptic quality of
the liquid food products due to release of ions from the electrode to the solution (i.e., in beer) [105]. Oxidation, loss
213
Fig. 4. An example pulse protocol showing a decrease in voltage due to an increase in conductivity, where correspondingly the current increases to
maintain the energy per pulse.
of an electron, occurs through the electrochemical reaction when the anode has lower potential than the anions in the
liquid and is dependent on the electrode material, temperature, current, and frequency [106]. If the impedance of the
pulse forming network remains the same and the energy of the charging unit of the PPG is also kept constant, then
the energy per pulse will be maintained constant irrespective of temperature. An example pulse protocol is shown
in Fig. 4 to illustrate this. If adequate resting time is not provided between each subsequent pulse, then the electric
field intensity will differ due to a change in liquid conductivity. This effect is more prevalent in batch treatment as
compared to continuous-mode treatment [64].
Turbulent flow has been recommended for continuous-mode PEF treatment of low-viscosity media such as juices,
beer and milk as the residence time distribution (the time during which the liquid is between the electrodes or within
the treatment chamber) is more homogeneous than in laminar flow [107]. Highly viscous fluids cause an increase in
the Prandtl number (ratio of momentum transport vs heat transport) which correspondingly minimises heat conduction
[108]. Therefore, continuous-mode PEF treatment with fluids that have high Prandtl numbers (i.e., 44.6 for tomato
juice and 38.4 for liquid egg at 20 ◦ C [107]) may have problems dissipating heat [107,108]. The change in electrical
conductivity due to Joule heating all follow a linear relationship, although the gradient will differ depending on the
liquid. This was shown by Heinz et al. [107], as illustrated in Fig. 5. If the generator voltage is compensated to maintain
the electric field intensity at the desired level (i.e., to avoid a decrease in voltage due to a rise in temperature and hence
the conductivity of the solution), then the energy per pulse will be different for all pulses. Therefore, PEF treatment
chambers for liquid food application may require a temperature compensation function to maintain the electric field
intensity as the sample temperature increases during treatment.
3. Inactivation kinetics of microorganisms by PEF
Microorganisms such as bacteria, fungi (yeasts and molds) [109], protozoa [110], archaea [111], algae [112],
viruses [113,114], and helminths [109,115,116] are ubiquitously present in food products. Their presence is either by
design to produce other foods (i.e., starter culture) or due to unwanted pre- or post-processing contamination sources
[117]. The unwanted microorganisms can cause food spoilage due to their cell density within the medium (i.e., high
numbers) and/or secretion of enzymes and toxins which can negatively degrade the nutritional profile of the food
product and may consequently pose a health risk to the consumer [117].
214
Fig. 5. Electrical conductivity of various liquid foods as a function of temperature. Reproduced from [107], with permission from Elsevier.
Table 1
Minimum detection limits and volumes of media used for the drop, spread, pour, and spiral plate methods [124,125].
Method Drop plate Spread plate Pour plate Spiral plate
Volume used for analysis (mL) 0.01-0.2 0.1 1.0 0.05 – 0.2
Minimum detection limit (CFU.mL−1 ) 10 or 1 log 100 or 2 logs 10 or 1 log 30 or 1 log
PEF technology has been used to decontaminate liquids in applications such as wastewater treatment or for inac-
tivating microorganisms in liquid foods [118,119]. Other technologies currently being developed and/or being used
in industry include thermal treatment, ohmic heating, ozone, microwave and radio frequency, pulsed light, oscillating
magnetic fields, gamma irradiation, pulsed x-rays, high pressure processing, ultrasound (i.e., sonoporation), UV ir-
radiation, shock waves, non-equilibrium atmospheric plasmas, and dielectric barrier discharge [120]. The method of
choice implemented ubiquitously in the food industry today for the processing of liquid foods is thermal treatment
[120]. A few forms of heating methodologies are utilised for the processing of liquid foods; (a) high temperature short
time (i.e., 73 ◦ C for 15-30 seconds); (b) low temperature long time (63 ◦ C for 30 minutes); (c), ultra-high temperature
(123 ◦ C for 15-30 seconds) pasteurization [120].
Thermal treatment at high temperatures can cause denaturing of enzymes and proteins which can degrade the
nutritional quality, colour, and volatile profile of the product. A key advantage of PEF technology over thermal pas-
teurization methods is the potential preservation of the natural bioactive compounds. PEF processing for the liquid
food industry requires the development of irreversible electropermeabilization protocols which irreversibly reduce
the cell count of the target contaminant(s) of interest to a safe level, while maintaining the nutritional profile of the
liquid food through careful consideration of the thermal gradient. The inactivation efficiency of PEF technology can
be validated based on the reduction of indicator microorganisms (i.e., coliform bacteria) through a cell count and is
represented as number of colony forming units per unit volume (e.g., CFU.mL−1 ). Four commonly used methods
for microbial count are the drop, spread, pour, and spiral plate methods [121–123]. Each of these methods require
different volumes for analysis and may have different minimum detection limits as noted in Table 1 [122].
Thermal inactivation of microorganisms typically follows a first-order rate reaction kinetics, and the D- and z-
values are used to describe these kinetics [120]. D-value is defined as the time required to reduce the specific
microorganisms cell count by 10-fold (i.e., 90% or 1 log) for a fixed temperature and medium [120]. Whereas the
z-value of a microorganism is the temperature increase required to reduce the D-value by 10-fold. These parameters
are appropriate for treatment methods which use temperature as a critical factor for reducing the microbial count, such
as microwave energy, ohmic heating, or thermal pasteurization [120]. In high pressure processing the time, tempera-
ture, and pressure are used to define new D- and z-values [120].
215
For PEF-induced inactivation, other processing parameters must also be chosen to determine the D- and z-values
[120]. For example, Puligundla et al. [126] used the electric field strength, frequency, and total treatment time as
processing parameters to define the DP EF - and zP EF -values. Other authors note the electric field strength and the
treatment time as dominant parameters. The inactivation kinetics of microorganisms by PEFs in liquids have been
modelled with the Bigelow model, Peleg model, Hülsheger and Niemann model [127], and the Weibull distribution
model [128]. Models for inactivation kinetics in-vitro most likely aren’t transferrable to in-vivo applications.
3.1. Bigelow model
First-order equation which uses treatment time [128,129].

t
log (S) = − (11)
D
where, t is treatment time, and D is the time required to reduce the specific microorganisms cell count by 10-fold
at a specific temperature, S is the survival fraction, defined as Nt /N0 , ratio of number of colony forming units after
treatment Nt to the initial number of colony forming units N0 .
3.2. Peleg model
A sigmoid function which uses the electric field strength [128,130,131].

100
S= Ef ield −Ef ield ,c
(12)
1+e a
where, Ef ield is the applied electric field, Ef ield,c is the critical electric field strength to achieve 50% relative survival,
and a represents the gradient of the curve near Ef ield,c .

1−S
Ef ield,c = Ef ield − a ln (13)
S
3.3. Hülsheger and Niemann model
Uses electric field strength to characterise model [120].

ln (S) = −bE · Ef ield − Ef ield,c (14)
where, bE is the coefficient of regression (cm.kV−1 ) and the gradient of the survival curve.
ln (S)
Ef ield,c = 1 − (15)
bE
The critical field strength is solved by assuming the value of Ef ield for 100% survival [120].
3.4. Weibull distribution model

n
t
log10 (S) = − (16)
a
where, t is treatment time, n and a are parameters of shape and scale. If n = 1, then the model is equivalent to the
Bigelow model [128].
We believe it is more appropriate to use the energy density rather than treatment time, as this allows the comparison
between different electric field strengths (i.e., because the energy of a 20 kV.cm−1 pulse is different from the energy of
a 30 kV.cm−1 pulse if the treatment chamber dimensions and electrode geometries are kept constant). Treatment time
does not include the current waveform and hence its contribution to the energy density per pulse is not considered.
Treatment time is exclusively useful as a comparison for analysing PEF processing for the same electric field strengths
216
or for pulses with square waves, assuming the same pulse generator topology, pulse forming network, and load are
used.
Surprisingly, no model incorporates the pulse-width as a parameter. It is expected that for each range of pulse-width,
τpulse , and membrane charging time constant, τm , (e.g., τpulse > τm , τpulse < τm , or τpulse = τm ) the DP EF - and
zP EF -values would also change as the charging and discharging phenomena associated with the membrane voltage,
Vm , of both the plasma membrane and intracellular organelles are directly influenced.
If the membrane voltage, Vm , is dominant for triggering the electroporation phenomena, and assuming that it is
also the primary reason for cell death, the applied electric field strength required to induce a certain membrane voltage
(e.g., 1 V) increases as the pulse-width is reduced, for any single pulse of the same shape. In other words, as the pulse
width increases the critical electric field strength, Ef ield,c , decreases [82].
Here we represent an applied pulse, p, with an electric field strength of Ef ield , a pulse width of τ1 , and energy
of Eenergy,1 , as: p1 Ef ield,1 , τ1 , Eenergy,1 . Assuming the pulse width of pulses p1 , p2 , and p3 are τ1 = 10 ns,
τ2 = 100 ns, and τ3 = 1,000 ns, then based on the electroporation mechanism (i.e., membrane voltage exceeding 0.3
∼ 1 V), for a single applied pulse the survival fractions (i.e., S1 = S2 = S3 ) are equal if Ef ield,1 > Ef ield,2 > Ef ield,3 ,
and the total energy input of all three pulses are equivalent, as in Eenergy,1 = Eenergy,2 = Eenergy,3 . The critical
electric field strength, Ef ield,c , to induce electroporation phenomena is inversely proportional to the pulse width (i.e.,
Ef ield,c1 > Ef ield,c2 > Ef ield,c3 ) [132]. When extending the model to include charging and discharging phenomena
associated with pulse trains, then including frequency of excitation or the pulse repetition rate becomes appropriate.
The frequency spectrums of the three pulses discussed above are different and their relative interactions (in time and
frequency domains) with the cellular membrane/envelope, solution, and cytoplasm may induce different kinetics.
The chosen pulse repetition rate for a given pulse train creates a fundamental frequency at that chosen repetition
rate. For example, a pulse train of ‘n’ pulses pn Ef ield,n = 100 kV.cm−1 , τn = 1 ns, Eenergy,n = 0.1 J at 1 Hz pulse
repetition rate would have fundamental frequencies ranging from 1 Hz to 1 GHz.
4. Electroporation, electropermeabilization, and associated mechanisms
Electroporation models of planar lipid membranes and biological membranes which attempt to describe the theory
of pore formation can be categorised as deterministic (non-pore) or stochastic (pore) models [133].
The deterministic models do not include the creation of pores and have not been able to validate experimental
observations of the stochastic nature of rupture and/or its dependence on the transmembrane potential [133–135].
Whereas the stochastic models of electroporation are consistent with key features of the electroporation mechanism
such as its dependence on the transmembrane voltage. Cell membrane pore formation has been widely accepted as
a stochastic process where the probability of pore formation and cell survival is influenced by the transmembrane
potential [133,136].
The stochastic model includes the surface tension of the lipid membrane and how the aqueous pore influences
the energy balance equation. In these models the surface tension (energy per unit area) is considered positive and
pore formation influences the free energy of the membrane in two ways; (1) there is an increase in free energy as the
perimeter of the pore surface adds an ‘edge energy’ denoted by γ , and (2) a decrease in free energy because of the
water-filled pore which reduces the membrane surface area [133]. The change in free energy for a membrane voltage
of 0 V and assuming a cylindrical pore of radius, r, is denoted by Eq. (17) [136]:
Epore (r) = 2γ r − πr 2 (17)

The applied electric field to the bilayer membrane will induce a non-zero voltage across the cell and will cause pore
formation. Pore formation (electroporation) has been shown through MD simulations to occur first by the generation
of a single column of water molecules, which then expands depending on the change in free energy [137–139]. These
water-filled columns will cause an increase in the membrane capacitance as lipids have lower relative permittivity
(εr = 2.0 [138], 2.1 [137], 4.98 [140]) than pure water (εr = 80 [138]) [49,58,67,76,138]. Changes to the membrane
capacitance, Cm , due to the transient induced membrane voltage have been shown experimentally to be less than 2%
for a U O22+ -modified azolectin membranes [141,142]. However, this change in membrane capacitance will modify
the free energy Eq. (17) to Eq. (18) by including its relation to the transmembrane voltage [136]:
Epore energy (r, Vm ) = 2γ r − πr 2 − 0.5CLW Vm2 πr 2 (18)
217
Where, CLW denotes the new membrane capacitance as a result of pore formation and is shown in Eq. (19):

εw
CLW = − 1 Cm (19)
εm
The reversibility of the electroporation process has been linked to the size of the pores [54,143]. If the radius of the
pore, r, is greater than rcritical irreversible recovery occurs which may lead to cell death [143]. Alternatively, if the
radius of the pore is less than rcritical this will lead to a reversible process [143]. Stronger electric fields have been
shown to cause reduction in the radius of these pores and a rapid increase in membrane conductivity, whereas weaker
electric fields maintain the radius of the pores at a near constant value [143].
The electropermeabilization phenomenon (increased permeability due to the induced electric field) associated with
PEF technology is a non-thermal process which occurs when the membrane voltage is increased past a critical thresh-
old (commonly between 0.3 ∼ 1 V [87,144,145]), irrespective of temperature [69,146–149].
The transient transmembrane voltage, Vm , for a spherical cell is distributed asymmetrically, and its absolute value
is highest at the poles (i.e., θ = 0) given from Eq. (20) [5,14,41]:
Vm (t) = −fs .Ef ield (t) .R. cos θ (1 − e−t/τm ) + Vrest (20)
where fs , Ef ield (t), R, τm , θ , and Vrest , are the shape factor of the cell, time-dependent electric field strength, cell
radius, membrane time constant, angle between the field direction and the normal of the cell surface, and the voltage
across the membrane at rest, respectively [150]. The shape factor of a cell is given from Eq. (21), where fs is approx-
imately 1.5 for a spherical cell at physiological conditions and assuming that the membrane is absolutely insulating
(i.e., σm = 0) [151,152]:
3σe 3dR 2 σi + 3d 2 R − d 3 (σm − σi )]
fs = (21)
2R 3 (σm + 2σe ) σm + 12 σi − 2 (R − d)3 (σe − σm )(σi − σm )
where, ‘d’ is the membrane thickness and σe , σi , σm , are the extracellular, intracellular and membrane conductivities,
respectively. For a rod-shaped bacterium, the shape factor is given by Eq. (22), where l is length of bacterium, and r
is the radius of the hemisphere, assuming that the cellular envelope is absolutely insulating [120,152,153]:
l.(l − 2r)
fs = (22)
3
The time constant of membrane charging is given by Eq. (23):
Rεm
τm = (23)
2d(σi σe /(σi + 2σe )) + Rσm
The membrane charging time constant, τm , is between 100 ns and a few microseconds for most cells in physiological
conditions [89,154–158]. Pulses with a pulse width longer than τm may cause higher physical stresses on the outer
membrane layer, whereas pulses that have shorter pulse widths than τm may have a stronger influence on intracellular
organelles (depending on the relative permittivity and conductivity of solutions and membranes) [159,160].
From Eq. (23), the membrane charging time can be modulated by controlling the extracellular conductivity (i.e.,
sample conductivity). The peak induced membrane voltage in low conductivity media is lower than at higher extra-
cellular conductivities. However, as the conductivity of the extracellular medium is reduced, the membrane voltage
discharges at a slower rate which can allow for cumulative charging of the membrane as governed by the resistive
and capacitive model of the lipid bilayer [58,98]. The resistance is defined by Eq. (9), where ‘d’ is the length of the
resistor (in this case will be approximately equal to thickness of the membrane), σ is the conductivity of the pore,
and ‘A’ is the surface area of pores induced on the membrane. Capacitance is defined by Eq. (10), where ε0 is the
permittivity of vacuum (∼8.86 × 10−12 F.m−1 ), εm is the permittivity of the portion of membrane under analysis
(i.e., lipids, cholesterol, proteins, or aqueous pore), ‘A’ is the surface area, and ‘d’ is the thickness of the membrane or
pore. Typical measured values for specific membrane capacitance in neuronal cells with voltage-clamp experiments
are 0.5 ∼ 1.0 µF.cm−2 [161].
As the membrane of cells is a leaky bilayer (i.e., not fully impermeable to water, ions, and molecules) and the
bilayer is influenced by PEF excitation (i.e., changes in membrane thickness, total surface area, etc.) the capacitive
current, Ic , is defined by Eq. (24) [48,162,163].
218
dq d dVm dCm
Ic = = (Cm · Vm ) = Cm + Vm (24)
dt dt dt dt
The charges formed on opposing sides of the plasma membrane can induce a voltage-dependent mechanical force on
the bilayer which can cause deformation, reduction in its thickness and an increase in area, this phenomenon is called
electrostriction [162]. Assuming that the membrane and its properties are uniform, the force, F, induced on the bilayer
is expressed by Eq. (25) [162].
1 1 Vm 1 Cm Vm2
F = Ef ield .q = .q = (25)
2 2 d 2 d
Thomas Heimburg [162] noted that as the surface area of the membrane increases due to the increase in transmembrane
voltage, this may potentially reduce the melting point of the bilayer. The effect of electrostriction on the melting
temperature of a large unilamellar vesicle composed of 1,2-dipalmitoyl-sn-3-phosphatidylcholine (DPPC) is shown
in Fig. 6 [162]. For steady-state transmembrane voltages of approximately −70 mV, the influence of electrostriction
is negligible [162]. However, for PEF excitation of cells where the transmembrane voltage is known to exceed 1 V
the effect of electrostriction is significant. This seems to be a plausible explanation for the combinatory inactivation
mechanism of microorganisms through ‘mild’ heating combined with PEF treatment.
Modulation of the membrane voltage is also known to affect cell division, mechanosensation, membrane fluidity,
spore formation, cell-cell communications, and biofilm dynamics [164–171].
For most bioelectric applications (i.e., electrochemotherapy) there will not be a large deviation in conductance
as physiological conditions of the body (i.e., tissues) are very static at a macroscale (approx. 15 mS.cm−1 ) [172].
Although for liquid food applications, the matrix constituents and sample conductivity of even the same liquid food
are subject to huge variations between different countries, local farms, or even time of year. Furthermore, the PEF
processing itself can cause transient changes in sample temperature or impedance (i.e., will affect energy delivery)
and should be closely monitored and regulated to ensure both, the safety and quality of the liquid food.
For pulses which are much longer than the membrane charging time, the steady-state form of Eq. (20) can be used
for the transmembrane potential as shown in Eq. (26):
Vm (t) = −fs .Ef ield (t) .R. cos θ + Vrest (26)

There is a second-order model which must be used for membrane voltage calculations at higher frequencies of ex-
citation, such as in pulses of nanosecond duration or modulated pulse trains utilising high frequency spectra [173].
For AC excitation, Tadej Kotnik and Damijan Miklavčič [148] extended the TMP equation for higher frequencies of
excitation for a purely sinusoidal AC source for frequencies until 100 MHz (relaxation of lipid bilayers is in the 100
MHz range which limits the model). Other pulse shapes such as square wave, trapezoidal, and exponential pulses were
also explored for frequencies up to 1 MHz [98].
Some research groups have reported certain biophysical phenomena after changing the pulse delay time or polarity
of two or more pulses, i.e. electro-sensitization or -desensitization of the plasma membrane [173–175], bipolar can-
cellation (BPC) [47,176–184]. The use of bipolar pulses or manually alternating the cathode and anode reduces the
effects of electrolysis and solid deposition on electrodes (i.e., protein aggregates or mineralisation) [85,185]. How-
ever, bipolar pulses may cause BPC and contribute to reversible electropermeabilization process which may inhibit
the inactivation efficiency of the treatment, and hence the safety of the liquid food.
The time delay between consecutive pulses can affect the cumulative charging of the plasma membrane by a
pulse train (like charging a capacitor). The closer the subsequent pulses (i.e., lower pulse delay time) the higher
the peak membrane voltage for a given set of pulse parameters. If the delay between pulses is long (i.e., more than
5 membrane charging time constants) and in a high conductivity medium, then the impact of each pulse must be
analysed individually as the increased membrane voltage will be dissipated and no cumulative charging would occur
from subsequent pulses [98,139,182].
BPC occurs when two unipolar pulses of opposite polarity are applied with an interphase delay time of 2 ns up to
100 µs (dependent on the membrane charging time, refer to Eq. (23)) [186]. The reversal of the membrane voltage is
controlled by the ratio of opposite polarity pulses [177]. If the first unipolar pulse has a sufficiently large amplitude, it
has been reported that the second opposite polarity pulse does not cause reversibility [186]. Authors have been utilising
various complex electrode geometries to trigger BPC phenomena in regions of interest to maintain cell viability,
while sufficient PEF conditions (i.e., unipolar high intensity pulses) in other regions for selective inactivation [186–
219
Fig. 6. (Top) Changes in excess heat capacity at three membrane voltages for large unilamellar vesicles composed of 1,2-dipalmitoyl-sn-3-
phosphatidylcholine (DPPC). (Bottom) Voltage-induced change in melting temperature, Tm , of DPPC. Reproduced from [162], with permission
from Elsevier.
190]. Nanopore occlusion has been proposed as a biophysical mechanism of bipolar cancellation [183]. Amphitropic
proteins may also be involved in the reversibility mechanism due to the electro-mechanical coupling induced by PEFs
[191].
Furthermore, as the membrane gets charged and responds with creating transient pores of complex structure it con-
tributes to the equivalent electrical model of the membrane through the addition of a variable resistor (a varistor) with
linear and/or non-linear time-dependent conductance (i.e., change in surface area of pore and pore lifetimes contribute
to the dynamics of the varistor) [43,58,192]. Moreover, this also reduces the capacitance of the plasma membrane
due to a reduction in its surface area, which is associated with the dynamics of transient pores, and contributes to the
discharging of the transmembrane voltage [58,67]. These pores allow for the transport of small charged and neutral
molecules (i.e., calcium) which dominantly happen post-pulse [67,193]. The transport of larger molecules requires
longer pulses, and the exposure time is proportional to the amount of transport [194]. Authors tested different lipids
through MD simulations and quantitatively approximated the number of water and lipid molecules diffusing through
different pores at ns intervals (6-27 water molecules, and up to 0.07 lipid per ns) [76,195].
If lipid peroxidation occurs, this will further affect the capacitive model of the membrane due to the decrease in
membrane thickness and changes to its relative permittivity [58,196]. Fig. 7 by Kotnik et al. [98] illustrates the cumu-
lative charging numerically by making some necessary simplifications, such as assuming a homogeneous membrane
(i.e. no deviations in membrane thickness). Notice that the conductivity of the extracellular media can have a pro-
nounced effect on the charging and discharging (i.e., sensitization and relaxation) phenomena of the leaky capacitor.
Lower conductivity media, as shown by the thinner line in Fig. 7, holds the membrane voltage for much longer than
the high conductivity medium as shown by the bolder line.
If the pulse delay time is increased, the membrane may discharge further, however this is highly dependent on
the dielectric and conductive properties of the membrane and solutions (extracellular, intracellular, periplasmic, etc.)
220
Fig. 7. The left column shows pulse trains of unipolar rectangular pulses and the right column shows the induced transmembrane voltage on a
spherical cell when the horizontally adjacent pulse train is applied. (a) τpulse = 200 ns and pulse period = 400 ns, (b) τpulse = 1 µs and pulse
period = 2 µs. The dashed line shows the transmembrane voltage using the steady-state Eq. (23), the thin line shows the transmembrane voltage at
low conductivity, whereas the bolder line is at high conductivity. Reproduced from [98], with permission from Elsevier.
[198] and should be considered on a case-by-case basis as the theoretical model provided in Fig. 7 is not applicable to
rod-shaped microorganisms.
Pulses shorter than the membrane charging time constant (i.e., nanosecond pulses) were found to increase the num-
ber (i.e., millions) but not the size of the nanopores (i.e., 0.4 to 4 nm), called supra-electroporation [199,200]. Nanosec-
ond pulsed electric fields also cause changes in the arrangement and externalization of phospholipids [96,201–205],
may cause intracellular ionic imbalances which may contribute to pore formation [146,206], influence voltage gated
channels [74,207], disassembly of actin structures [208], calcium dynamics and calcium-induced-calcium-release
(CICR) [209], destruction of transthyretin aggregates when the applied electric field strength was greater than 1
MV.cm−1 [210], mitochondrial stress [159,160], induce cytoprotective mechanisms such as integrated stress response
(ISR) [211], and apoptosis of mammalian cells [146,147,160,212–218]. Whereas longer pulses would generate a
smaller number of total pores, but were larger in radius (i.e., tens of nanometres) and had different lifetimes than
pores induced by pulses shorter than τm [57,69,219].
The dielectric constant of most materials decreases with increasing frequency (depends on temperature), therefore,
pulses with high-frequency spectra (i.e., sub-microsecond or modulated pulse trains with high frequency spectra) may
influence the plasma membrane and will not exclusively affect intracellular organelles [67].
A material is a dielectric when the displacement or capacitive current is larger than the in-phase or resistive current
(valid for f > σm .(2πεm )−1 ) [67,220]. Changes in membrane conductivity (membrane conductance = displacement
currents + conduction currents) due to PEF excitation have been numerically compared in passive and active simu-
lation models, where the effects of displacement current (valid for f > σm .(2πεm )−1 ) dominates in passive models.
In active models (transient time-dependent models), the displacement current initially dominates, however due to
the creation of pores and an associated increase in membrane conductance (σm ), the conduction current (valid for
f < σm .(2πεm )−1 ) dominates [67].
The dielectric relaxation time of the cytoplasm can be approximated using Eq. (27) as ranging from 14 µs ∼ 700
ps for a dielectric constant of 80, and conductivities ranging from 5 ∼ 1000 mS.cm−1 , respectively [132]. Where εcp
and σcp are the relative permittivity and conductivity of the cytoplasm, respectively.
ε0 .εcp
τ= (27)
σcp
The electric field distribution of picosecond pulses with effective pulse widths shorter than the dielectric relaxation
time of the cytoplasm will be dominated by the relative permittivity of the cytoplasm and membrane, rather than
conductivity [132]. Membrane currents will be limited to displacement currents as conduction currents (diffusion of
ions) are too slow to occur [132].
221
Fig. 8. PEF-induced changes in the transmembrane potential of a spherical cell with different membrane charging time constants for (A) square
wave with 10 µs pulse width and (B) an exponentially decaying pulse with a time constant of 10 µs. Reproduced from [139], with permission from
Springer.
From Eq. (26), it is quite clear that as the cells become larger the induced membrane voltage also increases.
Furthermore, as the size of the cell varies the intrinsic time constant of the cell also changes as governed by the leaky
resistive-capacitive model of the lipid bilayer. This illustrates that within a real cell population of various cell radii,
the characteristic time constant will be different for cells of different sizes [189]. Consequently, the onset and degree
of electroporation and/or electropermeabilization phenomena will also be unique for cells which are of the same
species and strain yet different size [221]. Fig. 8 by Gintautas Saulis [139] shows PEF-induced dynamic changes in
the transmembrane potential of a spherical cell’s plasma membrane with different membrane time constants.
Strain differences were also found to affect PEF resistance, indicating that PEF-induced inactivation is not exclu-
sively due to the biophysical structure [222]. Beatrice et al. [222] reported that PEF treatment decreased the expression
of three major heat shock proteins in Listeria Monocytogenes, which is inherently ambiguous as an increase in expres-
sion of these proteins is usually associated with a response to external stresses (i.e. heat, UV, and oxidative stress).
Bacteria in the stationary phase are highly resistive to external environmental stresses [223]. Researchers have iden-
tified that non-specific DNA-binding proteins such as Dps can protect microorganisms (e.g., E. coli and Salmonella)
from physical and chemical stresses such as UV and gamma irradiation, high temperature, metal toxicity, and oxida-
tive stress [224–227]. The same may be true for PEF excitation, however to our knowledge no such study has been
performed. The protection mechanism of Dps in E. coli has been associated with how it congregates DNA allowing
for a compact protective structure and its iron oxidation mechanism, which further protects DNA from damage [228].
PEF was also found to not induce stress-resistance in vitro in future generations of CHO cell line and Pseudomonas
putida [229,230].
The change in membrane temperature as outlined by Sale and Hamilton [231], is shown in Eq. (28). Where
Rm is the membrane resistance ( .cm2 ), dm is membrane thickness, and km is the membrane thermal conductiv-
ity (J.s−1 .cm−1 .◦ C−1 ).
Vm2 .dm
Tm = (28)
8.Rm .km
The change in temperature gradient of the membrane (from the medium and applied pulse) causes an osmotic pressure
difference due to the colligative effect which makes water flow out of the cell [87]. The thermal osmosis effect opposes
this colligative force [87].
PEF excitation does not generate reactive oxygen species (ROS) itself [69,232], however it can trigger the gen-
eration of ROS within cells (i.e., mitochondria) which can cause lipid peroxidation [232–236]. Oxidative damage to
lipids can increase permeability to water, reduce the thickness of the plasma membrane, and increases its suscepti-
bility to electroporation [69,196,237]. Recently it was reported that lipid oxidation is not a likely mechanism which
can explain electropermeabilization on longer timescales [238]. The authors measured lipid oxidation through C11-
BODIPY staining with total internal reflection fluorescence microscopy and noted that for microsecond pulses lipid
222
oxidation was inhibited in low conductivity media whereas the uptake of YO-PRO-1 (630 Da) was increased. This did
not occur for the applied nanosecond PEF treatment [238].
Transient single large pores (micrometre radius) have been observed in giant unilamellar vesicles (GUVs) under
special conditions (i.e., viscous solution) although the complex dynamics of pore initiation and closure are not fully
understood [74,195,206]. While the investigation of the kinetics of nanopores (up to several nanometres in diameter)
remains a challenge, the size of nanopores can be inferred through the selective entry of impermeable compounds
[69,202,239,240]. These reporter molecules may affect the bilayer and the pore composition during diffusion due to
hydrogen bonding or electrostatic interactions of the matrix (i.e. complex/dark pores [241]), some are cytotoxic (de-
pendent on the exposure time and may not be suitable for certain studies). Membrane-impermeable fluorescent dyes
(i.e., propidium iodide, or ethidium bromide and acridine orange staining) are useful indicators of permeabilization for
in vitro studies, however, they are not able to detect pores which are smaller than their size [67,242–245]. Propidium
iodide has been shown to be an inaccurate method for assaying the viability of prokaryotes [246]. Recently, authors
have shown that Ba2+ and Ca2+ can be useful nanoporation markers for in vitro electroporation [239]. Biolumines-
cence has also been recently shown to be a useful indicator in vivo for the onset of permeabilization/electroporation
phenomena [247]. Fluorescent voltage indicators (i.e. voltage-sensitive dyes or genetically encoded voltage indicators
(GEVIs)) have widespread use in neuroscience and electrophysiology studies to spatially and temporally visualise the
modulation of membrane voltage through neuronal circuits [164,248,249], but are also useful for electroporation and
electropermeabilization studies [154,250]. Although there remain numerous challenges during transformation such as
low voltage sensitivity, low signal to noise ratio, and poor localisation of voltage indicators [251–256].
Another method is to evaluate pore conductance through constant current experiments [257]. Pore initiation time,
as shown by current-controlled experiments, decreases with increasing electric field and membrane voltage [257]. Re-
cently, pulsed electric fields were found to induce changes (i.e., pore formation) in voltage-gated ion channels (VGICs)
which are ubiquitously present in prokaryotic and eukaryotic organisms [74]. This is inherently important as it can
affect signal transduction pathways and contribute to the electropermeabilization phenomena [74]. The generation of
pores also affects the surface tension at the membrane interface (and vice versa [49]), this can trigger one of the many
mechanosensitive gating mechanisms (such as shear stress) of proteins ranging from mammals to prokaryotes (i.e.,
PIEZO1, PIEZO2, and MscL) [258–262]. These proteins act as electromechanical transducers and accurate modelling
of their kinetics is of huge significance for drug delivery or evaluating cell-death mechanisms.
The equivalent electrical model of membrane proteins (linear and non-linear conductance) and ionic gradients
(represented as batteries) have been studied by Hodgkin and Huxley (H-H), however the contribution of cholesterol
and integral membrane proteins to the regulation of the transmembrane potential during PEF excitation and the charg-
ing and discharging kinetics associated with it has not been studied. Some proteins with mechanosensitive gating
mechanisms such as PIEZO1 and PIEZO2 may be modelled with an equivalent RLC model (resistive, inductive,
and capacitive model). Moreover, modelling the equivalent electrical model of cellular envelopes of monoderm and
diderm prokaryotes remains a major unsolved challenge [263].
Cholesterol was found to require a higher electric field strength to induce electroporation in experimentation and
in silico models, and lower amounts of cholesterol in the cell membrane increased the cell sensitivity to nsPEFs
[64,69,264]. The H-H model does not incorporate temperature dependence, and to further complicate; the extended
thermodynamic H-H models could also not validate the conductance of potassium channels at different temperatures
[265].
The polarization field associated with Maxwell’s equations can influence the analysis of protein channels and PEF
processing on biological membranes [266]. The relative permittivity of a material, εr (F.m−1 ), is used to approximate
the materials interactions with electric fields referenced to vacuum and is frequency dependent as in the case of a lossy
dielectric (i.e., membrane) [267]. As governed by Maxwell’s equations, the relative permittivity is treated independent
of time and as a single constant [266]. This over-approximation is not adequate to describe the transient equivalent
electrical model of membranes, and the mathematical model for εr (t) and contributing factors affecting polarization
(i.e., electro-diffusion) must be coupled and solved in conjunction with Maxwell’s equations as outlined recently by
Robert S. Eisenberg [266,268].
When conducting studies that involve the inactivation kinetics of microorganisms, it is imperative to report the
transient temperature gradient of the liquid due to the pulse protocol to analyse the influence of heating induced by
Joule heating [84,269]. For the decontamination of liquid foods, combining mild heating of the liquid sample (50 ◦ C)
with PEF treatment has been reported to cause higher inactivation than both treatments applied separately [270]. We
223
refer the reader to a comprehensive review paper by Raso, et al. [84] that lists the relevant PEF process parameters
that are required for the comparison of results between different research groups [73].
Garner et al. [269] discussed the influence of nsPEFs on cell membrane thermal gradients which can further in-
fluence the associated phenomena (i.e., electroporation/electropermeabilization) by contributing to the modulation of
the membrane voltage. By considering the bilayer as a thermoelectric, they note that higher pulse repetition rates at
the same pulse duration can maximise these thermal gradients through numerical techniques [269]. Recently, Das
et al. [271] showed numerically that under certain conditions bipolar pulsed-DC fields caused strong deformation of
non-spherical vesicles (e.g. applicable to rod-shaped microorganisms). Furthermore, they attribute the changes to the
switching behaviour of the field (i.e., reversal of polarity) which causes changes in the spatial dynamics of tensile and
compressive stresses [271,272].
When a unipolar pulse is applied to a cell where the relative conductivity is high (i.e., σr = 10) and the pulse
width is equal to the membrane charging time (i.e., τp = τm ) the deformation in the cell is low [271]. The relative
conductivity is defined by Eq. (29), where σi and σe are the intra- and extracellular conductivities, respectively [271].
σi
σr = (29)
σe
The authors state that when the medium has low relative conductivity (i.e., σr = 0.1), switching the electric field
polarity triggers tensile equatorial stresses [271].
Sub-nanosecond pulsed electric fields (58 kV.cm−1 , 303 ps and a FWHM of 773 ps, 200 Hz, up to 50,000 pulses)
have been shown to induce electropermeabilization as measured by YO-PRO−1 and SYTO 9 staining of E. coli
[146]. The authors report an increase in the fluorescence intensity as the number of pulses was increased from 5,000
to 50,000 pulses, furthermore they estimate an increase in temperature of less than 0.01 ◦ C after 50,000 consecutive
pulses through Eq. (8) [146]. However, electropermeabilization does not necessarily mean loss of cell viability and it
would be useful to investigate the inactivation kinetics associated with sub-nanosecond pulsed electric fields for liquid
food treatment and how to optimise it for cell-death mechanisms.
A review of the effects of nanosecond and microsecond high-voltage pulses from primary research articles ranging
from the year 1990 to 2014 is presented in [159]. The authors categorized original research articles by the duration
of the pulses; (A) 1-10 ns, (B) 11-100 ns, and (C) 100-1000 ns. Longer duration pulses (i.e., 100 ns – 1 µs) affect the
plasma membrane more than shorter pulse durations of (1-100 ns). Plasma membrane effects were denoted through
cellular uptake of large molecular dyes such as propidium iodide (PI), trypan blue (TB), ethidium homodimer (EtH),
or YOPRO [159]. A single nanosecond electric pulse was found to create plasma membrane pores of diameter ∼1 nm
or less [273–275].
Plasma membrane effects reported from experiments which used single pulse protocols were more frequent when
the duration of the pulse was longer [159]. The uptake of dyes was significantly dependent on the pulse duration [159].
The effect of pulse repetition rate on the plasma membrane is less explored than other electrical parameters [159].
Four reports suggest that higher repetition rates produced more effects [202,274,276,277]. Three authors reported that
higher repetition rates produced fewer effects [278–280]. Two groups reported a band-pass response to repetition rate,
where the mid-band ranges of the repetition rate corresponded to more effects on the plasma membrane [281,282].
In [280], the authors note a sensitization effect during PEF treatment, where lower repetition rates corresponded
to higher inactivation. They also found that the treatment time (i.e., longer exposure) were more efficient irrespective
of the repetition rate [280]. Furthermore, they found that by splitting the high repetition rate pulse train and adding
a delay of 1-5 minutes in between the split pulse train resulted in higher inactivation rates (through PI uptake) for
electric field strengths greater than 1.8 kV.cm−1 [280].
5. Quality and safety of liquid foods treated with PEFs
PEF treatment of liquid food samples must ensure that the functionally important bioactive compounds are not in-
activated by the thermal gradient or the electric field (quality), while harmful microorganisms are inactivated through
irreversible electropermeabilization (safety). This is a key advantage of PEF treatment technology over thermal pas-
teurization techniques [283,284].
Fig. 9 shows the cumulative peer-reviewed publication number of primary articles for the application of PEF
treatment for assessing the quality and/or safety of water, wine, milk, liquid whole egg (LWE), juice, cider, and beer.
224
Fig. 9. Cumulative peer-reviewed publication number of primary articles on PEF technology treatment of liquid foods for the application
of assessing quality and/or safety. Liquid foods include water [285–311], wine [126,312–342], milk [101,287–289,293,297,304,343–453],
liquid egg [285,374,376,416,428,454–495], juice [2,29,92,93,105,128,197,290,312,313,326,337,357,362–364,369,372,378,400,406,408,409,418,
422,435,442,444,445,459,496–662], cider [505,612,663,664,749], and beer [105,124,125,665,666]. References were procured from Web of Sci-
ence in June 2022.
Most of the literature has been focused on juice and milk as they include multiple sub-groups, for example in milk
we have human, bovine/cow, goat, and camel milk. Relative to other food products, a low number of studies has been
conducted on beer, cider, and wine.
The quality of liquid foods (e.g., colour, rheology, viscosity, appearance, mouth feel, aroma, and flavour) can be
negatively influenced by degenerative enzymes, microorganisms which secrete enzymes, and oxidative reactions [668,
669]. Some enzymes such as alkaline phosphatase (ALP) in milk are used as an indicator of thermal pasteurization
[669].
Nutritionally important compounds within liquid foods include proteins and free amino acids, lipids and free fatty
acids, vitamins, and polyphenols [670]. Some authors have reported changes in free amino acids [545,671]; structural
changes to proteins (vitamin modifications) including changes in the secondary (α-helix, β-sheets, etc.), tertiary (spa-
tial conformation), and quaternary (number and arrangement of protein subunits) structures [668,672]. No evidence of
PEFs modifying the primary structure of proteins (i.e., modifying polypeptide chains) has been identified [669,673].
Other authors have reported total fat changes, increase and decrease in free fatty acids (FFA) [545], conversion
of carotenoid molecules [674], changes in the secondary and tertiary structure of proteins (i.e., changes in the bond
vibrations within the sidechains of β-sheets) [670,675].
It has been reported that depending on the energy input and the electric field intensity, PEFs can inactivate or
stimulate enzymatic activity. Large specific energy input have been reported to inactivate enzymes [668], whereas
enzymatic activity may be stimulated at low electric field treatments (i.e., few kV.cm−1 ) [668]. Moreover, the total
phenolic compounds within the fluid were found to contribute to PEF-resistance [670].
Thermal pasteurization of liquid food products cannot effectively inactivate spores from pathogenic microorgan-
isms such as Clostridium botulinum, which produces a neurotoxin that causes botulism. These spores are able to
remain dormant in low oxygen (i.e., canned foods) or acidic conditions. Combining PEF with ultra-high tempera-
ture (UHT) treatment has been shown to reduce spores of heat resistant species of Bacillus subtilis and Geobacillus
stearothermophilus by 3-5 log [676]. Although the low-acid canned industry aims at 12 log reduction of Clostridium
botulinum spores as a thermal pasteurization target [120].
225
Microorganisms change their fatty acid composition in response to external stresses such as heat, which will con-
sequently affect membrane properties such as fluidity [234]. The culture temperature was shown to affect the PEF
resistance of S. aureus in [677]. The size of the microorganism was found to be more dominant than the gram staining
classification for higher PEF-induced inactivation [678]. Gram-negative bacteria contain numerous binding proteins
which break the continuity of the peptidoglycan layer (PG) and cause adherence between the membranes [73]. Al-
though, the gram-staining classification has been identified as an oversimplification because not all gram-negatives
have two membranes (i.e., diderm) and similarly not all gram-positive bacteria have a single membrane (i.e., mono-
derm) [263,679]. It is recommended that prokaryotes are classified by the number of membranes they contain and the
presence and/or absence of lipopolysaccharides, this also becomes relevant for modelling and classifying microor-
ganisms based on their equivalent electrical model [263,679]. The average thickness of the cellular envelope for a
diderm is 35 nm, membrane thickness of 5-8 nm, and the PG being 6.4 nm for E. coli and 2.4 nm for Pseudomonas
aeruginosa [263,680]. Comparatively, the average thickness of a monoderm is variable predominantly dependent on
the thickness of the PG. Membrane thickness is typically 6 nm, whereas PG thickness ranges from 20 nm in S. aureus
and 34 nm in B. subtillis [263,680].
Among some of the relevant microorganisms for food safety and stability [117], for spoilage non-spore form-
ing bacteria, Lactobacillus and E. coli are the most studied in combination with PEF technology for inactivation or
fermentative processes. Other spoilage non-spore forming bacteria with a few studies include Pediococcus, Strep-
tococcus, Lactococcus, and Leuconostoc. With no studies on Carnobacterium, Kurthia, nor Weissella. For spoilage
spore-forming bacteria Bacilli is one of the most studied, whereas Clostridia has no studies. For pathogenic non-spore
forming bacteria there are numerous studies on Salmonella, Listeria, and E. coli (i.e., O157:H7), with little attention
on Staphylococcus Aureus, Yersinia, Campylobacter and no studies on Brucella [681]. For pathogenic spore-forming
bacteria, the most studied is Bacillus cereus, whereas Clostridium has none.
Nearly all studies for Ascomycetous (yeasts) are done with model organism Saccharomyces cerevisiae, with only
one paper on Schizosaccharomyces pombe and Candida spp. For Basidiomycetous only three papers are published
on Rhodotorula spp. For molds there is a few papers on Aspergillus, Penicillium, and Alternaria, with no papers
on Mucor, Rhizopus, nor Bothrytis. Only four papers have evaluated the efficacy of PEF technology on viral loads
[291,682–684].
Prions were found to increase the PEF resistance of yeast, Saccharomyces Cerevisiae [685]. A PEF treatment of 5
pulses with 1 ms duration at 26 kV.cm−1 was found to reduce the cell viability of the yeasts and not contribute to the
generation of prions, called ‘prionization’ [685]. Research into the interactions of PEFs and with prions are limited
and there is a need to study prions in other smaller microorganisms such as bacteria [686,687] which have generally
lower susceptibilities to PEF treatment than yeasts due to their size.
The hurdle preservation approach for food processing, shown in Fig. 10 [688], includes using combinatory tech-
niques to preserve both the quality and safety of the food product. Hurdles can include chemical, physical, and
microbial processes. Some examples of commonly used hurdles include pH, PEF, high pressure, UV, shock waves, ul-
trasound, antimicrobials, temperature, acids, oils, lysozyme, salts, bacteriocins, antibiotics, and packaging [688,689].
The hurdle approach may be useful to target highly resistant species of microorganisms, spores, biofilms, and viral
loads. As compared to other non-thermal technologies, high pressure processing cannot be applied in continuous mode
and UV is affected by the opaqueness of the product and is limited to surface treatment [124]. The inactivation by UV
treatment is because of induced cross-linking between DNA strands, preventing cell replication [690].
Sub-nanosecond PEFs may further enhance the bioactive retention of the product as the increase in temperature
of the protocol is insignificant (a few degrees) and may also be more energy efficient than PEF combined with mild
heating. Other authors have suggested using the losses associated with PEF treatment (Joule heating) to pre-heat
the incoming liquid and for post-treatment holding time to maximise energy efficiency [79]. The exposure time to
different temperatures post-PEF treatment (i.e. holding time) has been shown to contribute to inactivation kinetics of
microorganisms [101].
The use of inert electrode materials (i.e. electrodes which do not contribute to electrochemical kinetics and release
of metals) such as carbon have been shown to have higher inactivation than treatment with stainless steel [691], which
is a commonly used material in the food industry (i.e., 316 stainless steel).
A combination of PEF and ultrasound (US) excitation was shown to enhance the lethality of a thermotolerant
microorganism, Streptococcus thermophilus, where the order of excitation (i.e. (PEF + US) or (US + PEF)) was
226
Fig. 10. Example of hurdle preservation methods where individual hurdles at low intensity cannot provide adequate protection against microbial
growth, whereas individually applied hurdles at high-intensity or a combination of low-intensity hurdles can be used to maintain the quality and
safety. Reproduced from [688], with permission from Springer.
also found to contribute [79]. Alternatively, mild heating and sub-nanosecond PEFs were also shown to increase the
lethality of the protocol for mammalian and prokaryotic cells [213].
PEF-treated media was also reported to contribute to the inactivation of certain microorganisms (E. coli, S. Ty-
phimurium, and L. innocua) in buffer media and water [295]. The authors attribute the mechanism of cell inactivation
to be due to electrochemical oxidation of Cl− ions during PEF treatment as the addition of 0.5% sodium thiosulfate
inhibited the inactivation [295].
Researchers discuss the feasibility of PEF-induced inactivation of microorganisms for liquid food treatment only
applicable for low conductivity samples and without microbubbles [28]. Microbubbles within the liquid media can
affect the treatment homogeneity of the pulse protocol and cause various unwanted effects. Bubbles have lower di-
electric breakdown strength than the liquid media, so sufficient electric field strengths can cause partial discharges of
these bubbles which can cause (1) a reduction in the electric field strength and losses due to heating, (2) formation of
free radicals and reactive species depending on the gas composition, (3) UV generation, (4) and micro shock waves.
The presence of air bubbles can cause non-homogeneous PEF treatment as well as operational and safety problems
[692]. The use of vacuum degassing or pressurizing the treatment media during processing can be used to minimise
the presence of air bubbles [692].
In order to reduce arc discharge in treatment chambers, the following methods have been suggested:
• Smooth electrodes which avoid sharp points [85].

• Degassing of the sample (i.e. sparging) prior to PEF treatment [85].
◦ Degassing of liquids increases conductivity [693], [694]. Conductivity values of the degassed sample were
found to be dependent on temperature and the extent of degassing. The author explains that the dominant factor
was the extent of degassing. Thus, the higher the removal of the dissolved gases, the higher the conductivity of
the solution. Mathew J. Francis [694] compared how the conductivity of various concentrations of potassium
chloride solutions would be affected by degassing. At higher concentrations, the effect of degassing was negli-
gible as ionic effects dominated. For all potassium chloride solutions, a conductivity difference of 1 µS.cm−1
was reported after removal of dissolved gas [694].
227
Fig. 11. Comparison of (a) pulsed corona plasma and (b) PEF treatment on Legionella suspensions. Reproduced from [678], with permission from
Elsevier.
• Pressurising the sample to prevent bubble formation [85,695].

◦ At atmospheric pressure, the dielectric breakdown strength of bubbles is exceeded in PEF liquid food chambers
[696]. Góngora-Nieto et al. [696] recommend pressurising the chamber to 810,600 Pa (8 atm) to minimise the
risk of arcing by small bubbles (∼1 mm). The drop in the electric field strength predominantly occurs at the
boundary interface of the bubbles [696].
• Minimising temperature increases in liquid. Temperature rise reduces the solubility of gases in liquid and hence
causes bubble formation [85,695].
◦ If the pulse repetition rate is low, cooling electrodes may be adequate to minimise bubble formation. Alter-
natively, applying PEF treatment in stages with cooling steps in-between can be advantageous in continuous
systems.
If the electric field strength applied to the liquid sample exceeds the breakdown voltage of the media, then an arc
discharge may occur [85]. Gasses have lower dielectric breakdown strength than liquids (20-30 kV.cm−1 for air, versus
MV.cm−1 order for low conductivity water [695]), so other arc discharging phenomena more commonly experienced
than the breakdown of liquid includes surface discharges at gas-liquid or solid-liquid interfaces [85].
Soluble gasses in the liquids can influence the heterogeneity and amplitude of the PEF treatment. If the energy den-
sity delivered to the sample causes a net-rise in bulk liquid temperature, then the solubility of the gasses will decrease,
causing gas bubble formation and partial vaporisation [85]. Since vapour and gas bubbles have lower breakdowns than
most liquids, it may attribute to arc discharging phenomena at the gas-liquid interface or micro-discharges within the
bulk media [85].
5.1. Water
The availability of safe and clean drinking water is a major global challenge. A plethora of different microorgan-
isms have been independently studied in liquid water applications. The treatment of water for decontamination of
various microorganisms serves as a useful model to independently examine the inactivation efficacy of various mi-
croorganisms in various environmental conditions (i.e., conductivity and temperature). Buffers are more appropriate
to study inactivation efficiency at fixed pH values. The matrix constituents can also be modified to emulate a media of
interest (i.e., simulated milk ultrafiltrate).
Pulsed corona plasma and pulsed electric field treatment methods used to inactivate Legionella pneumophila in
water are compared in [678]. The dissipated energy and treatment time were kept constant for both methodologies.
As shown in Fig. 11, plasma was more efficient (in terms of energy) than PEF treatment for inactivation of Legionella
suspensions [678]. However, plasma was found to generate more hydrogen peroxide than PEF treatment and the au-
thors attributed the higher inactivation efficiency of pulsed corona plasma to be due to hydroxyl radicals and hydrogen
peroxide formed [678]. Scanning electron microscopy (SEM) results (Fig. 12) showed that after PEF treatment some
cells were intact, whereas for plasma treatment the cells were ruptured [678].
228
Fig. 12. Comparison of scanning electron microscopy pictures of L. pneumophila after pulsed corona plasma and PEF treatment on Legionella
suspensions. Reproduced from [678], with permission from Elsevier.
Pulsed corona plasma treatment can induce various chemical reactions (i.e., formation of hydroxyl radicals and
hydrogen peroxide), shock waves, ultraviolet irradiation, as well as the electric field generated [697]. The effects of
electric fields, UV light, and shockwaves on the inactivation of the gram-negative E. coli species were studied in [698].
They utilised needle-to-plate electrode configurations to generate pulsed discharges in water [698], whereas electric
fields were applied with a plate-to-plate setup [698]. The authors reported that the discharge location can significantly
affect the impacts of the shockwaves and electric fields, however, it did not affect the UV radiation [698]. As the
distance from the discharge centre axis was reduced, the bacterial inactivation was increased [698]. The inactivation
rates of 36.1%, 30.8%, and 12.7% were achieved using shock waves, UV light and electric fields, respectively [698].
The effects of PEFs on the cell wall architecture of gram-positive vegetative bacteria (B. pumilus species) were
studied in [185]. They used a 4.1 mM NaCl solution with a set conductivity of 0.5 mS.cm−1 to study reversible or
irreversible permeabilization rates through propidium iodide (PI) staining [185]. They applied 1000 pulses of 5 µs
with 2.0 - 7.5 kV.cm−1 with a frequency of 1 kHz, where the inactivation rate varied between 38% to 98% [185]. The
authors also tested spores and found that 1,000 pulses at 7.5 kV.cm−1 was insufficient for inactivation [185], whereas
10,000 pulses were enough to reach 67% inactivation [185]. To avoid temperature increase, a delay of 60 seconds
was applied between each series of 1,000 pulses. To limit the production of electrolytes, the polarity was manually
inverted every 500 pulses [185].
5.2. Milk
The use of PEFs as a non-thermal technology for milk treatment has seen rapid advancements. Most literature in
the field has examined cow/bovine milk, where a smaller number of papers evaluated PEF treatment for human, goat,
and camel milk. For grade “A” bovine milk, it has been recommended that pasteurized milk has a total microbial count
of 20,000 CFU.mL−1 , with only 10 coliform bacteria [699]. Recently, the pre- and post-pasteurization criterion for
human milk has been changed to ≤105 CFU.mL−1 and <1 CFU.mL−1 , respectively [700].
The compounds present in milk include water, fat, proteins, minerals, vitamins and sugars (i.e., lactose). From
these compounds, the most dominant effect on conductivity originates from the minerals [701,702]. Whereas lactose
and casein were found to have a small effect on conductivity [703]. Major salt fractions within milk are the cations
calcium, magnesium, potassium, and sodium and the anions citrate, phosphate, and chloride [703]. Multiple research
groups [701,702,704] reported that the conductivity of milk decreases as the percentage of fat increases. Milk fat
229
Fig. 13. Changes in properties of milk due to a change in MFG size. Reproduced from [708], with permission from Springer.
globules (MFG), from 0.5 µm to 8.0 µm in diameter [425,705,706] for raw milk, are contained in milk fat globule
membranes (MFGMs) which contains proteins, glycoproteins, polar lipids, phospholipids, enzymes, and cholesterol
[707].
MFG are homogenised (i.e., reducing diameter variance of MFG present in milk) for dairy applications to a size of
approximately 0.3 ∼ 0.8 µm for colour, texture, and to modulate whipping time or creaming at the surface [705,706].
Modifying the size of MFG can influence the physicochemical properties of the liquid as shown in Fig. 13 [708].
The membrane protects triglycerides from hydrolysis by native or bacterial lipases and prevents coalescence [707].
Thermal pasteurization and ultrafiltration of milk can modify the size and structure of the MFGM and cause dena-
turing of whey proteins such as β-lactoglobulin and α-lactalbumin [707]. Heating can also impact physicochemical
properties of milk, such as changes in secondary and tertiary structures of proteins [707]. As shown in Fig. 14 by
Sharma et al. [707], MFGM are susceptible to thermal, mechanical, and PEF-induced changes in membrane structure
and aggregation.
Two of the most economically viable compounds within milk are its fat and protein content [703]. The sustainability
of these constituents can increase yields in secondary production streams such as cheese and butter. Phosphoproteins
such as casein are important to produce cheese. The properties of the casein micelles can affect the processability of
the cheese products [703]. Specifically, the distribution of calcium phosphate between the casein micelle and serum
phase impacts the structural and physical properties of the micelles. Proteolytic activity is considered a contributor to
casein degradation and negatively affects cheese yields [709].
Bijl et al. [703] report that the historical increase in the protein content of cow milk is attributed to changes in
casein and micellar salt fractions of the milk. They state that the casein content and micellar fractions of calcium,
phosphorous, and magnesium have increased significantly in the past 75 years. However, the salt content of serum and
the salt composition of casein micelles have remained the same [703]. Moreover, that the nutritional quality of milk
has also improved due to an increase in calcium, magnesium, and phosphorus.
High flow rates in continuous PEF treatment chambers can induce changes to the casein micelle size, where this
process can be reversible [405]. The authors state that this is an indication of the hypersensitivity of the micelle
structures to the shear forces through the continuous system. Pulse electric field intensity, number of pulses, and
concentration of proteins can possibly alter the structure of proteins [405]. Ascorbic acid was found to be the most
sensitive vitamin to PEF treatment [405].
However, the impact of PEF on the bioactive compounds and enzymatic activity of milk is not well understood.
Moreover, milk samples from different mammals have different compositions (i.e., casein content) and bacterial pro-
files. There is also a drastic variation in the bacterial and nutritional composition of the milk sample depending on the
postpartum timing (i.e., colostrum, transitional or mature human milk) [710].
230
Fig. 14. Impact of heat, mechanical or PEF disruption on the structure and aggregation of MFGM. Reproduced from [707], with permission from
Elsevier.
A variety of gases are used in the dairy industry for a change in taste, texture, shelf-life, or manufacturing a
secondary product [711]. Carbon dioxide and argon are used in milk beverages and raw milk to reduce microbial
growth rate and hence the shelf life [711]. Oxygen content for milk beverages or dairy powders needs to be kept
to a minimum as it can affect the flavours of the product [711]. Furthermore, contact of air with the dairy beverage
can support the growth of aerobic gram-negative bacteria [711]. The solubility of CO2 is reported to increase with
pressure, whereas it reduces when temperature rises [711]. N2 is insoluble in water, comparatively to CO2 which is
highly soluble, is reported to improve sensory qualities of milk when flushed [711].
Chantry et al. [712] used high-temperature–short-time (HTST) on HIV-infected human milk and subsequently as-
sessed its impact on the antimicrobial properties of the milk. Lactoferrin was not significantly affected by the flash-heat
treatment pre-digestion, however, it was considerably dropped post-digestion in flash-heated milk [712]. Pre-digestion
lysozyme had a significant drop when flash-treated, however, post-digestion there was no significant difference be-
tween post- and pre-digested amounts; indicating that flash-treatment did not affect the digestion survivability of
lysozyme [712]. Inhibitors of bacterial growth (i.e., hurdles) include lactoferrin (present in raw milk), carbon dioxide,
lysozyme, or natural inhibitors formed by microorganisms (nisin).
Psychrotrophic bacteria can grow in refrigeration temperatures (i.e., 4 ◦ C) and pose a safety risk to the shelf-life
of dairy products. Psychrotrophic bacteria in milk (e.g., Pseudomonas) may produce enzymes that remain active after
thermal pasteurization and continue to deteriorate the milk quality. Coliform bacteria appear in processed milk as
post-pasteurized contaminates (PPC) and produce acid and gas through the fermentation of lactose.
Thermoduric bacteria can survive thermal pasteurization (e.g., Bacillus, Paenibacillus) and may be psychrotrophic.
Heat resistant spore-forming psychrotrophs can significantly impact milk shelf-life and represent a major safety chal-
lenge for the industry. Spore-forming bacteria are one of the dominant causes of the reduction of milk-shelf life,
specifically the Bacillus and Paenibacillus species [713,714]. The spores are quite tolerant of heat, pressure, antimi-
crobial, UV, and PEF treatment [713,715]. The spores from some Bacillus species have high adhesion properties to
electrode material choices used in treatment chambers such as stainless steel [716,717]. Paenibacillus species produce
class I and II bacteriocins (Lantibiotics and pediocins) [714].
Inactivation of spores with PEF (40 kV.cm−1 ) or a combination of PEF and nisin is reported to be not effective,
where inactivation of up to 2 logs was achieved after hours of germination [718]. A combination of pressurised flow
system and PEF treatment (110 kV.cm−1 , 0.5 MPa) caused up to 6.7 log reduction in spores [719].
231
Inactivation of E. coli and S. aureus was examined in simulated milk ultrafiltrate (SMUF) by Pothakamury et al.
[720]. They used up to 60 pulses of 200 - 300 µs unipolar exponentially decaying waveforms at peak electric field
strength of 16 kV.cm−1 , with an electrode gap of 0.1 cm, and a liquid volume of 100 µL [720]. They cooled their
cuvettes after every 10 pulses for 5 - 10 seconds to avoid heating above 30 ◦ C. The survival fraction of the gram-
negative E. coli and gram-positive S. aureus were decreased by nearly 5 logs and 4 logs respectively when applying
60 pulses at 16 kV.cm−1 with 200 µs pulse width [720].
Viruses are more resistant to chemical methods (e.g. ROS and chlorine) than vegetative cells [682] and remain
stable in cold storage commonly used for food products [721]. With the recent and still ongoing SARS-CoV-2 pan-
demic, researchers noted that holder pasteurization (63 ◦ C or 56 ◦ C for 30 min) can cause inactivation of the viral
SARS-CoV-2 load within the treated human milk samples [722]. Bidawid et al. [723] noted that an increase in matrix
constituents such as fat can affect the thermal inactivation of hepatitis A virus (HAV) in cream (18% fat), skim (0%
fat) and homogenised milk (3.5% fat).
Other researchers reported no protective effects of milk matrix constituents (i.e., fat) for the thermal inactivation
mechanism of HAV [724]. The inactivation mechanism of thermal treatment is thought to be due to capsid disintegra-
tion [725]. Parvovirus and non-lipid enveloped viruses (plum plox virus) was reported to have higher heat tolerance
than other viral loads (95 ◦ C treatment for 2 hours could not significantly reduce the infectivity) [726]. There is a
prevalent safety concern for PEF treatment of foods if the treatment process cannot guarantee the inactivation of
viruses [291,682–684,727]. If not, then this technology may not be ready for wide-scale integration and would require
a combinatory technique to address food safety concerns from viral loads.
Drees et al. [682] reported that E. coli and bacteriophage (virus) inactivation was inhibited when the media in-
cluded glutathione (GSH), which has antioxidant properties protecting the microorganisms from oxidative stress and
free radicals. The authors noted that the inhibition of direct-current-induced inactivation was much higher for E. coli
than that of the bacteriophage (MS2), and therefore attributed the cell death mechanism to be due to oxidation by
electrochemically generated oxidants [682]. They also noted that E. coli and the bacteriophage were able to tolerate
higher treatment doses at higher concentrations [682]. As an example, milk contains catalase which also has antioxi-
dant properties and may protect microorganisms from the free radicals and oxidants produced during PEF excitation.
Sources of free radicals and oxidants can include electrochemical reactions at the electrode-electrolyte interface, sur-
face discharge in the treatment chamber or partial discharge from microbubbles, and high-current waveforms with
rapid changes in current (i.e., high di.dt−1 ) [87,238].
5.3. Beer
The influence of PEFs on beer has been minimally investigated as compared to milk or juice applications. This
may be due to the carbonation process in beer (i.e., either through fermentation or via forced carbonation), as the mi-
crobubbles can cause a decrease in the electric field intensity and produce secondary effects such as micro-discharges.
Although, PEF can be applied prior to the carbonation and/or fermentation steps.
The effects of the electric field strength and total energy input on the inactivation of late-stationary phase Lacto-
bacillus plantarum, a beer spoilage microorganism, were investigated in model beer [665]. Untreated cells and heat
treatment at 80 ◦ C for 10 minutes were used as comparisons, negative and positive controls, respectively. Authors
note that the temperature of the model beer never exceeded 31 ◦ C although no mention on the method of temperature
measurement, nor the pulse shape and effective pulse width used to excite the liquid sample was made. The authors
noted little to no effects on inactivation of L. plantarum for electric field strengths of up to 18.7 kV.cm−1 and corre-
sponding energy density of 42.1 kJ.kg−1 [665]. However, at 13.8 kV.cm−1 and at an energy density of 98 kJ.kg−1 ,
there was a 1.0 log decrease in L. plantarum [665]. The authors used the positive control (heat treatment of 80 ◦ C for
10 minutes) as an indicator of 0% membrane integrity and metabolic activity, whereas untreated samples (negative
control) were 100% [665]. The authors reported that loss of cell viability was noticeable when the relative metabolic
activity and membrane integrity was below 15% [665]. Furthermore, the metabolic activity and membrane integrity
at 13.8 kV.cm−1 and 98 kJ.kg−1 was equivalent to heat treatment of 80 ◦ C for 10 minutes which caused a 95% loss
relative to the untreated sample. Loss of membrane integrity and metabolic activity varied linearly as the energy input
was increased [665].
At an electric field strength of approximately 30 kV.cm−1 , the authors report 5-6 log reduction of L. plantarum at a
similar energy density of up to approximately 70 kJ.kg−1 in beer at a pH of 3.6 [665]. The addition of nisin (1 mg.L−1 )
232
or hop extract (100 p.p.m.) during PEF treatment caused additional 1.5 log reduction when the electric field strength
was approximately 30 kV.cm−1 and energy densities greater than 30 kJ.kg−1 [665]. The use of hop extract was also
found to cause a 2-log increase in sub-lethal injury, where the difference between cell counts in MRS and MRS-NaCl
agar were used as a differentiator for sublethal injury [665]. The authors also report that 15 hours of storage of the
PEF-treated model beer at 10 ◦ C containing 100 p.p.m. of hops killed surviving cells, whereas untreated cells were
reduced by 0.5 log after 25 hours of storage [665].
Synergistic effects of iso-α-acids of hops in beer with PEF have been suggested to potentially improve the lifetime
of the product during storage [665]. When utilising hops bitter acids in combination with PEF for beer preservation, as
outlined by Ulmer et al. [665], the amount of hop bitter acids and the pH of the medium need to be considered further
[665]. Although the medium pH of 4.0 and 3.6 will not cause a loss of viability of L. plantarum, the antimicrobial
activity of hop extract is increased by a factor of 2 when the pH is reduced from 4.0 to 3.6 [665].
Walkling-Ribeiro et al. [124] utilised a continuous co-axial treatment chamber to study the inactivation of four
inoculants in lager beer: yeast (Saccharomyces cerevisiae), gram-negative pathogenic bacteria (Salmonella Cholerae-
suis), gram-positive spoilage/probiotic (Lactobacillus plantarum) and spore-forming pathogenic bacterium (Bacillus
subtilis). For S. cerevisiae, the same reduction of greater than 6.8 log was reported for thermal pasteurization (76 ◦ C
for 30 seconds) and a treatment of 45 kV.cm−1 , energy density of 585 kJ.L−1 , and Toutlet of 43.7 ◦ C, at an inlet temp
of approximately 14.0 ◦ C [124]. The same treatment conditions only gave a 3-4 log reduction of L. plantarum, B. sub-
tilis, and S. Cholerasesuis [124]. When the energy was doubled to ∼1100 kJ.L−1 , and the Toutlet was approximately
55 ◦ C, L. plantarum had greater than 7.5 log reduction, B. subtilis had 8.1 log reduction, and S. Cholerasesuis had
5.9 log reduction [124]. The pH of the lager beer changed after PEF excitation, from 4.21 to 4.35, whereas thermal
pasteurization caused a change from 4.21 to 4.39 [124]. The ◦ Brix of untreated, PEF-treated and thermally pasteurized
beer showed no statistically significant change [124].
At Toutlet of below 44 ◦ C and an energy density of approximately 500 kJ.L−1 , when the electric field strength was
increased from 35 to 45 kV.cm−1 , the reduction of S. cerevisiae and S. Cholerasesuis was increased by 3 log and
2 log, respectively [124]. Whereas, for L. plantarum and B. subtilis there was only an increase of 1.0 – 1.5 log due to
the increase of the electric field strength [124].
In terms of the effect of the alcohol content, L. plantarum had 1.5 log higher reduction in beer with 3.5% alcohol
content as compared with 0.5% alcohol content [124]. Whereas, the other 3 inoculants did not show any statistically
significant change in reduction [124].
In carbonated beer inoculated with S. Cholerasesuis, when the electric field intensity was 40 kV.cm−1 with an
energy density of approximately 700 kJ.L−1 , the authors report approximately 4 log reduction whereas in degassed
beer it only had 2 log reduction [124]. This difference in reduction was not noticed at 35 kV.cm−1 nor for any of the
other three inoculants [124]. Furthermore, the authors report that no change in the carbon dioxide content between
untreated and PEF-treated beer [124]. The authors attribute the spore-forming ability and virulence gene expression
of B. subtilis and S. Choleraesuis to be possible explanations for their higher resistance to the PEF treatment [124].
Evrendilek et al. [105] utilised a continuous treatment chamber to study the inactivation of five inoculants in
beer: yeast (Rhodotorula rubra and Saccaromyces uvarum), gram-positive spoilage/probiotic (Lactobacillus plan-
tarum and Pediococcus damnosus) and spore-forming pathogenic bacterium (Bacillus subtilis). The authors applied
bipolar square waves to beer inoculated with yeast cells with 10.5 mL.s−1 flowrate, 22 kV.cm−1 , 10 µs, 800 Hz, 413.7
kPa (60 psi), 154 µs total treatment time and achieved 4-5 log reduction of the yeast inoculants [105]. They applied
the same conditions to the bacterial inoculants and did not notice a significant reduction, therefore they changed the
PEF excitation to 1.0 mL.s−1 flow rate, 41 kV.cm−1 , 600 Hz, 4 µs pulse duration, 158.6 kPa (23 psi), and 175 µs total
treatment time [105]. At these conditions, the authors report a 4-6 log reduction of the bacterial inoculants in beer
with a peak temperature of ∼44 ◦ C as measured with K-type thermocouples on the outer surface of the SS316 tubing
[105]. Although it is not clear if similar energy densities were applied as the pulse width, electric field strength, and
total treatment times were different between the two treatment protocols investigated.
Cr, Zn, Fe, Mn, Mg, Al, Cu, Na, K, and Ca were analysed by inductively couple plasma-atomic emission spec-
troscopy (ICP-AES), and the authors reported a significant increase of Fe, Cr, Zn, and Mn in PEF-treated beer [105].
Based on the review of panellists, a significant difference in flavour and mouth feeling for PEF-treated beer, where
a metallic taste was reported [105]. No significant difference was reported in foam condition, colour, and overall ac-
ceptability of the control and PEF-treated samples [105]. Authors report that the release of metals in solution from the
electrodes was dependent on being below a critical frequency and above a certain pulse width [105,728].
233
Continuous PEF was used by Milani et al. [125] to investigate the inactivation of S. cerevisiae ascospores in nine
types of beer (i.e. ale, lager, dark) with varying alcoholic contents ranging from approximately 0% up to 7%. Thermal
pasteurization of 50 ◦ C for 20 min in a water bath was used as a positive control [125]. They applied an electric field
intensity of 45 kV.cm−1 , 46.3 pulses with bipolar square waveforms and a pulse width of 1.5 µs [125]. The initial
temperature of the inlet was slightly heated to 33 ◦ C, with peak processing temperatures reported to range between
37.4 to 53.1 ◦ C as measured with fibre optic temperature sensors [125].
When the alcohol content in the beer was increased from 0% to 4% or 7%, there was a 1-2 log increase in in-
activation [125]. Furthermore, when the temperature was increased from 43 to 53 ◦ C, there was an additional 2 log
reduction (maximum of 4 log) when the alcohol content was 4 or 7%, and an additional 1 log when the beer contained
0% alcohol [125].
The carbonation, alcohol content, and the addition of hops all serve as antimicrobial preservatives (i.e., hurdles)
[125]. There have been reports of safety concerns in alcohol-free beer as lactic acid bacteria (LAB) showed a 4-
to 7-fold increase in heat resistance, and pathogenic microorganisms (Escherichia coli O157:H7 and Salmonella
typhimurium) showed a 3- to 17-fold increase to heat resistance as compared to beer with 5% alcohol content [729].
Gad and Jayaram [666] treated carbonated ale beer with continuous-mode PEF, and evaluated the release of metal
ions from the SS316 electrodes and its subsequent effect on the quality of beer (i.e. taste and flavour) and the in-
activation of stationary-phase Saccharomyces cerevisiae, Lactobacillus plantarum, Bacillus subtilis, and Salmonella
enterica. SS316 contains 66% iron, 17% chromium, and 12% nickel [666]. Thermal pasteurization at 60 ◦ C for 20
minutes was used as a positive control [666]. Unipolar exponentially decaying pulses with an electric field strength of
40 kV.cm−1 at 30 Hz was applied with a pulse-width of 1.3 µs [666]. As the energy was increased the beer became
darker, and for every 86.4 kJ.L−1 energy applied there was approximately 100 part per billion (p.p.b.) increase in the
iron content [666]. The authors report up to 1 log reduction with PEF treatment for all four inoculants (originally
inoculated at 8-log), although with a peak processing temperature of 5 ◦ C [666]. The thermally pasteurized beer in
comparison had 2-3 log reduction of all 4 inoculants [666]. Iron released from the SS316 electrode material may
accelerate the aging of beer, where the aging of beer is also affected by exposure to air [666]. An increase in the
temperature of beer and/or if treated with mechanical shock (i.e., ultrasound) may cause a reduction in carbonation,
called foaming [666]. This foaming of beer is attributed to its carbonation, sugar, and alcohol content [666].
The same research group applied continuous mode PEF to beer but instead with titanium electrodes, and sub-
sequently evaluated the changes in trans-iso-α-acid and metal ions with high-performance liquid chromatography
(HPLC) and ICP-AES, respectively [667]. The authors report no significant difference in the total iron content between
PEF-treated and untreated beer due to the use of titanium electrodes [667]. Furthermore, no significant difference in
the trans-iso-α-acids (isocohumulone, isohumulone, and isoadhumulone) were detected between PEF-treated and
untreated beer samples [667]. Degradation of trans-iso-α-acids can limit the shelf-life and affect the flavour (i.e.
bitterness) of the beer [667].
Zufall and Tyrell [730] investigated the role of heavy metal ions such as iron and manganese and found that they
contribute to changes in the quality of foaming and the organoleptic stability of beer. Iron and manganese ions can
contribute to the generation of ROS such as hydroxyl radicals and superoxide anions which may oxidise organic
compounds present in the liquid media [666,730].
5.4. Wine
Recently at the end of 2020, “The International Organisation of Vine and Wine” approved PEF technology to be
used for destemmed and crushed grapes [731] to reduce maceration time and increase the extraction of bioactive
compounds (i.e. polyphenols) [235,732,733].
Inactivation of acetic acid bacteria (AAB) or lactic acid bacteria (LAB) for the wine industry is important as these
bacteria can utilise ethanol and glucose to generate acetic/lactic acid which may influence the volatile acidity of the
product [333]. AAB are aerobic microorganisms so sealing the containers can minimise growth. Most of the volatile
acidity in wines are produced from microbial metabolism and includes acetic, D-lactic, formic, butyric, and propionic
acids [734]. There are legal limits on wine products with high volatile acidity such as 0.24% (w.v−1 ) in Canada [735].
Important spoilage microorganisms include Brettanomyces and its sporulating form Dekkera (yeasts), LAB, and
AAB [325]. These yeasts produce ‘leather’, ‘animal’, and ‘horse sweat’ odours in wines, which are attributed to the
production of ethylphenols and from grape hydroxycinnamic acids [325,736,737]. Some lactic acid bacteria produce
234
acetic acid from glucose if high concentrations of sugars exist. Wine yeast such as Brettanomyces produce high
concentrations of acetic acid and were also found to spoil olive oil [738,739]. Spoilage bacteria and yeasts come from
surfaces of grape skins [325,740,741].
Sulphur dioxide is used as a ‘natural’ additive for wines as it inhibits the growth of the spoilage microbiota and acts
as an antioxidant minimising changes to the organoleptic pattern of the wine [742,743]. Sulphur dioxide is considered
a ‘natural’ compound in wine as most yeasts produce 10-40 mg.L−1 as part of their metabolism, even though the wine
does not naturally contain the compound [742]. Sulphur dioxide may also influence the anthocyanins within the wine
(i.e., red wine) and change its organoleptic pattern [742,744,745]. The use of PEF for wine can help minimise the use
of sulphur dioxide to maintain the safety and shelf-life of the product [746]. Low concentrations of acetic and formic
acid were found to contribute to higher inactivation rates for S. aureus and P. aeruginosa [747].
Ethanol with Acetobacter sp. in the stationary phase caused higher inactivation than without ethanol [333]. Authors
attribute the enhanced PEF inactivation to changes in the membrane fluidity (changed in membrane fatty acid compo-
sition and membrane lipid conformation) [333]. Alternatively, vegetative Bacillus subtilis cells were also studied with
and without ethanol at pH of 5.5 and 7.0 in combination with PEF treatment [748]. Heinz and Knorr report that the
addition of ethanol at a pH of 5.5 was synergistic for increased inactivation of Bacillus subtilis cells. However, at pH
of 7.0 the addition of ethanol caused a higher resistance to the PEF protocol. The authors suggest to apply PEF prior
to alcohol fermentation to maximise inactivation [748].
5.5. Fruit juices and cider
Most inactivation studies with juices involve store-bought thermally pasteurised juices which are subsequently
inoculated with a microorganism of interest. Studies with raw juices, such as apple cider, is a more appropriate
media to investigate the feasibility of PEF for industry. Apple cider, as stated in literature, is considered as non-
alcoholic freshly squeezed apples with no further clarification (i.e., including fruit pulp), sometimes referred to as
cloudy apple juice [664,749]. Yeasts, moulds, E. coli, coliforms, and LAB include microorganisms which commonly
influence the safety of apple cider or juice [749]. Thermal pasteurisation of apple cider was found to decrease the ester
concentrations [750], whereas for carrot juice there was a decrease in α-carotene, β-carotene and colour [751].
Buckow et al. [752] reviewed PEF techniques being used in orange juice for microbial inactivation and increasing
shelf life. PEF treatment of <10 kV.cm−1 for <20 µs (at room temperature) is reported to reduce yeast and mould
counts in orange juice [752]. PEF-resistant bacteria such as lactic acid bacteria and pathogenic E. coli become harder
to inactivate [752].
From the review in [752], it is indicated that a combinatory treatment of PEF and heat (up to 50 ◦ C) can be beneficial
as it reduces the total treatment time and specific energy requirements to achieve a 5-log reduction for E. coli. The
authors also state that the combinatory effect of antimicrobials such as nisin, benzoate, and sorbate allow for energy
efficient treatment [752].
The effect of PEF on enzyme inactivation is minimal compared to thermal treatment. PEF at room temperatures has
minimal impact on enzyme inactivation. Some studies have reported PEF inactivation of orange pectin methylesterase
(PME) and peroxidase (POD) at specific PEF conditions [752]. PEF was also found to degrade mycotoxins in juice
[753].
PEF treatment requires 4 to 5 times the specific energy input for enzyme inactivation levels like thermal treatment
[752]. PEF treatment at low to moderate treatment conditions caused minimal effects on the sensorial and nutritional
quality of orange juice. Furthermore, the impact of PEF treatment on vitamin C, carotenoids, polyphenols, and volatile
aroma compounds in orange juice was less than in thermal treatment (e.g., 95 ◦ C for 30 s).
Dziadek et al. [524] assessed the nutritional quality and shelf life of apple juice after PEF treatment. The authors
found that PEF treatment did not affect the bioactive compounds within apple juice (vitamin C and total polyphenols
during the storage for 72 h under refrigeration) [524].
Iu et al. [749] investigated the inactivation of non-pathogenic strains of E. coli O157:H7, yeasts, moulds, LAB,
and coliforms in apple cider with PEF treatment ranging up to 90 kV.cm−1 . Difference in cell count by tryptic soy
agar (TSA) and sorbitol MacConkey agar (SMA) to investigate sublethal injury of E. coli O157:H7 were found to
be insignificant (i.e. P > 0.05) [749]. At 90 kV.cm−1 and 10 pulses with a peak temperature of 42 ◦ C there was
approximately 5 log reduction, whereas at 70 kV.cm−1 and 10 pulses there was approximately 4 log reduction [749].
The authors used pulse number rather than energy density and therefore cannot compare the treatment protocols, as
235
the energy per pulse for a 70 kV.cm−1 pulse is not equivalent to a 90 kV.cm−1 pulse if the same electrode geometry
is used. Interestingly, for the same electric field intensity, increasing the number of pulses from 10 to 30 did not cause
any increase in the inactivation of E. coli O157:H7 for electric field intensities ranging from 60 to 80 kV.cm−1 nor for
peak temperatures ranging from 20 to 42 ◦ C [749].
With a treatment of 80 kV.cm−1 with 10 pulses and a peak temperature of 42 ◦ C, the authors report approximately
1 log decrease in yeasts and moulds (initial count of 4 log), and 1-2 log decrease of E. coli, LAB, and coliforms (lower
than the detection limit of the spread-plate method). The authors attribute the higher PEF resistance of the yeasts and
moulds to the highly acidic apple cider medium [749]. The addition of 2% (w.v−1 ) cinnamon or nisin increased the
inactivation of E. coli O157:H7 by 1-2 or 3-4 log, respectively [749].
Sources of E. coli O157:H7 outbreaks include raw milk, water, and highly acidic foods (pH of 4.0-4.5) such as
apple cider, mayonnaise and yoghurt [749,754,755]. Temperature treatment above 42 ◦ C was found to contribute to
rapid inactivation of E. coli O157:H7 [749]. Other sources of outbreaks include Salmonella sp. and Cryptosporidium
parvum [664].
Liang et al. [612] utilised continuous-mode PEF treatment to investigate the inactivation of yeasts and molds in
freshly squeezed apple cider. Nisin and lysozyme (0.4 mg.kg−1 ) caused an approximate 1-2 log increase in inactiva-
tion, whereas clove oil (0.3 or 0.5 mL.L−1 ) caused a 2-log increase in inactivation [612]. The effect of PEF treatment
on the inactivation of polyphenol oxidase (PPO) was investigated. PPO cause enzymatic browning in various fruits
and vegetables such as apples, avocados, and lettuce, which leads to nutritional loss [756]. Apple cider was heated to
50 ◦ C for 20 minutes as a positive control and the PPO activity did not change up to 20 minutes reaction time [612].
However, after continuous PEF treatment of 27 kV.cm−1 , 3 L.h−1 (58.7 pulses), and 200 Hz; the authors report 33%
reduction in PPO activity at 10 min reaction time [612]. Inactivation increased when the flow rate was decreased, as
the residence time was subsequently increased [612].
Valappil et al. [664] compared how three different treatments, PEF, UV and thermal processing, affected the quality
and safety of apple cider which was subsequently stored at 4 ◦ C and compared at 0 and 4 weeks. Samples treated with
UV fermented after 2 weeks, whereas neither PEF nor thermal treatments did [664]. All three treatments achieved 5
log reduction of E. coli, the PEF treatment was continuous mode with square wave pulses of 2.5 µs duration, electric
field intensity of 23 kV.cm−1 , and peak temperature of 51 ◦ C, although no mention of the energy consumption of the
PEF treatment was made [664]. Apple cider treated with PEF retained ∼98% of its total ester and aldehydes after 4
weeks of storage, except for hexyl acetate where it retained 35% (thermal: 33%, UV: 6%) [664].
5.6. Liquid egg
LWE is commonly pasteurized at 60-68 ◦ C for 2-5 min [474,757,758], which also reduces its viscosity and foam-
ing capabilities [758]. Salts and organic compounds such as lysine and glycerol are synthesised or accumulated via
transport by microorganisms to negate the effects of heat, osmolarity, turgor pressure, and provide ribosomal stability
[759–764]. These are called compatible solutes, where high concentrations of compatible solutes increase the dielec-
tric constant of the solution (i.e., solution capacitance will be altered – refer to Eq. (10)) [760]. LWE is rich with
compatible solutes that may be used by microorganisms to reduce the PEF-induced stresses, although the influence
and protective mechanisms of each compatible solute (if any) are yet to be elucidated. Lysozyme inhibits the growth
of gram-positive bacteria such as S. aureus in LWE as it contributes to the breakdown of the peptidoglycan layer of
the cellular envelope [765]. Antimicrobials such as nisin (10-100 IU nisin.mL−1 ) added to LWE after PEF exposure
increased reduction of L. innocua by 1-3 log depending on the concentration of nisin added [474].
LWE is rich in nutrients such as protein, fat, and cholesterol; minerals such as calcium, phosphorus, potassium, and
sodium; vitamins such as vitamin E, niacin, vitamin A, B6, B12, and riboflavin; amino acids such as alanine, lysine,
aspartic acid, leucine, serine, and histidine [766]. Microorganisms relevant in LWE include Listeria monocytogenes,
Salmonella enterica, Enterococcus, Escherichia, Enterobacter, Serratia, Shigella, Pseudomonas, and Bacillus cereus
[767,768].
PEF processing of LWE can lead to ohmic heating due to the high conductivity of the sample [692]. Therefore,
integration of secondary operations/considerations such as a cooling system or maybe lower treatment time is re-
quired. The impact of these secondary integrations on the PEF efficiency (log reduction) on microorganisms, and the
economics of the system needs to be evaluated [692].
236
Combinatory treatment of LWE with PEF and heat treatment has shown higher pasteurization levels than indi-
vidually treating with PEF or heating [692]. Therefore, optimization processes need to be identified to increase the
efficiency of PEF treatment at low electric field strengths if minimal heating is to be delivered. The high Prandtl num-
ber of LWE (see Table 2) is also problematic as it influences the heat dissipation capability of the liquid when PEF
treatment is applied in continuous mode [107].
6. Outlook
High impedance liquid foods, such as olive oil, represent a unique group of food products that are highly advan-
tageous to treat with PEFs. Oil products have a large impedance which is significant from an engineering perspective
as it allows for higher electric field strengths (and wider pulse-widths [82]) to be achieved as compared with low
impedance media (i.e., high conductivity media such as tomato juice). The high impedance media can also limit
breakdown phenomena occurring between the electrodes. Furthermore, the discharging transients of the accumulated
plasma membrane charge after PEF excitation may be slowed because of the high impedance nature of the medium.
This consequently decreases the threshold electric field for subsequent pulses to reach the same membrane voltage
[82].
Most studies involving oil products involve pre-treatment of the fruit with PEFs for the purpose of increasing the
extraction yield of valuable compounds such as colourants, sucrose, or polyphenols [769]. The topic of PEF treatment
for the application of extraction from fruits and vegetables, or extraction from microorganisms such as yeasts and
algae, or the application on solids (i.e., meats) are out of scope of this review paper as these fields are extensive
enough to merit their own reviews.
The following are further prospects for bioelectricity and PEF technology in the liquid food and beverage industry.
• There is a need for novel pulsed power generators which can deliver large amplitudes (e.g., >50 kV) with variable
pulse widths at high repetition rates (e.g., >1 kHz) and minimal distortion in the pulse shape.
• For studies concerning the safety or quality of the liquid food, such as the inactivation of microorganisms or
enzymes, the contribution of thermal- and PEF- induced inactivation can be better understood by evaluating
transient thermal gradients.
• The hurdle approach may be extended to inactivate dormant spores from heat resistant spore-forming psy-
chrotrophs, viral loads, and to minimise the recovery of viable but non-culturable cells during storage.
• There have been numerous studies on the high-frequency dielectric permittivity of liquid foods ranging from a few
MHz to several GHz for the applications of microwaves. The dielectric permittivity at high frequencies will be
relevant to the frequency domain spectra of the pulse applied in PEF technology. PEF technology typically applies
narrow pulses (microsecond to nanoseconds) with pulse repetition rates in the low frequency bands ranging from
1 Hz to several kHz. The pulse repetition rate will also create a fundamental frequency; therefore, it is necessary
to explore the dielectric permittivity of liquid foods in the low frequency bands ranging from 1 Hz to several kHz
to accurately model the response to PEF excitation.
• Modelling the inactivation kinetics of prokaryotes in different media and at controlled temperatures is essential in
order to subsequently compare them with those of eukaryotes.
• Simulation models in both the time and frequency domains are required to accurately predict the kinetics of
physical, chemical, and thermal processes. These simulation models need to be designed in a manner which can
be subsequently validated with experimental investigations.
• For industrial integration, processing liquid foods is highly dependent upon energy efficiency and economics
(i.e., feasibility studies). To be able to integrate or choose a suitable technology for the processing, an in-depth
analysis of competing techniques (positive controls) such as heat pasteurisation (low-temperature-long-time, high-
temperature-short-time, and ultra-high-temperature treatments), and high-pressure processing is required.
Declaration of competing interest
The authors declare that they have no known competing financial interests or personal relationships that could have
appeared to influence the work reported in this paper.
237
F. Zare, N. Ghasemi, N. Bansal et al.
Table 2
Physical and chemical properties of liquid foods.
Sample pH Conductivity Specific gravity Specific heat Conductivity changes due to Prandtl Number
(mS.cm−1 ) capacity, Cp adiabatic Joule heating
(kJ.kg−1 K) (∂σ /∂W ) = ((S/m)/kJ/kg)
Pure water 7 0.06 × 10−3 1 4.186 - 6.9
Drinking water 7 0.5 – 1.0 at 23 ◦ C 1 4.18 at 20 ◦ C [107] 0.301 × 10−3 at 20 ◦ C [107] 7.0 at 20 ◦ C
0.6 at 20 ◦ C [107] [125] [107]
Milk Raw milk 6.6 ± 0.2 at 4 ± 2 ◦ C 4.75 ± 0.25 at 4 ± 1.03 ± 0.005 at 4 - - -
[88] 2 ◦ C [88] ± 2 ◦ C [88]
Skim milk 6.6 ± 0.2 at 4 ± 2 ◦ C 5.25 ± 0.25 at 4 ± 1.04 ± 0.005 at 4 Milk (1.5% fat) Milk (1.5% fat) 3.019 × Milk (1.5% fat)
[88] 2 ◦ C [88] ± 2 ◦ C [88] 4.14 at 20 ◦ C [107] 10−3 at 20 ◦ C [107] 9.6 at 20 ◦ C
5.7 at 20 ◦ C (1.5% [107]
fat) [107]
4.5 for UHT milk
with 1.5% and 3.5%
fat [770]
4.1 at 5 ◦ C [480] 3.2 at 20 ◦ C [107] 3.906 × 10−3 at 20 ◦ C [107] 38.4 at 20 ◦ C
238
Liquid Egg Whole egg 8.1-8.3 [480] -

7.96 at 25 ◦ C [771] 5.7 at 20 ◦ C [107] [107]
7.74 at 60 ◦ C [771] 5.9 at 20 ◦ C [480]
9.4 at 50 ◦ C [480]
Egg white 7.8 [480] 4.4 at 5 ◦ C [480] - 3.81 at 20 ◦ C [107] - 27.8 at 20 ◦ C
9.43 at 4 ◦ C [771] 6.5 at 20 ◦ C [107] 3.8 [772] [107]
9.14 at 25 ◦ C [771] 6.4 at 20 ◦ C [480]
8.61 at 55.6 ◦ C [771] 10.4 at 50 ◦ C [480]
Egg yolk 6.4 [480] 2.1 at 5 ◦ C [480] - 3.12 [772] - -
6.25 at 4 ◦ C [771] 3.3 at 20 ◦ C [480]

6.13 at 25 ◦ C [771] 5.7 at 50 ◦ C [480]
5.92 at 60 ◦ C [771]
Wine Red wines 3.4 [773] 197 to 252 [774] - 4.18 [776] 4.15 - -
3.15 – 3.96 [774] 0.88 – 2.57 at 25 ◦ C [776] 4.5 for wines
2.80 – 3.54 [775] [775] with a density of
995 kg.m−3 [776]
White wines 2.78 – 2.93 [774] 1.74 – 1.86 at 25 ◦ C - - -
3.10 – 3.15 [775] [775]
F. Zare, N. Ghasemi, N. Bansal et al.
Table 2 (continued)
Sample pH Conductivity Specific gravity Specific heat Conductivity changes due to Prandtl Number
(mS.cm−1 ) capacity, Cp adiabatic Joule heating
(kJ.kg−1 K) (∂σ /∂W ) = ((S/m)/kJ/kg)
Beer 3.6 – 4.0 [665] 1.20–1.78 [665] - 3.18 at 20 ◦ C [107] 1.211 × 10−3 at 20 ◦ C [107] 10.6 at 20 ◦ C
4.21 [124] 1.38–1.41 [124] [107]
4.4 ± 0.1 [667] 1.58 at 4 ◦ C [105]
1.37–2.76 at 23 ◦ C
[125]
2.4 ± 0.05 [667]
2 at 20 ◦ C [107]
Juice Orange 3.7 ± 0.2 at 4 ± 2 ◦ C 3.5 ± 0.1 at 4 ± 2 ◦ C 1.03 ± 0.005 at 4 3.77 at 20 ◦ C [107] 2.801 × 10−3 at 20 ◦ C [107] 12.2 at 20 ◦ C
[88] [88] ± 2 ◦ C [88] [107]
3.8 [480] 2.3 at 5 ◦ C [480]
4.4 at 20 ◦ C [107]
3.5 at 20 ◦ C [480]
239
5.9 at 50 ◦ C [480]
Apple 3.6 [480] 1.3 at 5 ◦ C [480] - 3.85 at 20 ◦ C [107] 1.958 × 10−3 at 20 ◦ C [107] 13.3 at 20 ◦ C
2.9 at 20 ◦ C [107] [107]
2 at 20 ◦ C [480]
3.4 at 50 ◦ C [480]
Tomato 4.05 – 4.65 [777] 15 at 20 ◦ C [107] - 3.98 at 20 ◦ C [107] 7.915 × 10−3 at 20 ◦ C [107] 44.6 at 20 ◦ C
[107]
Apple cider 4.22 ± 0.03 [749] 2.5 ± 0.1 at 4 ± 2 ◦ C 1.03 ± 0.005 at 4 - - -

4.0 ± 0.01 at 5 ◦ C [88] ± 2 ◦ C [88]
[749]
3.7 ± 0.2 at 4 ± 2 ◦ C
[88]
Waste brine 7.0 ± 0.1 at −7 ± 300 ± 20 at −7 ± 1.25 ± 0.01 at - - -
2 ◦ C [88] 2 ◦ C [88] −7 ± 2 ◦ C [88]
- Cannot find.
Acknowledgements
The authors would like to thank Dr. Nushin Hosano for her advice regarding inactivation mechanisms and Dr.
Elisabeth Prabawati for her comments concerning food safety and methods. This work was supported in part by
Grant-in-Aid for Scientific Research from the Ministry of Education, Culture, Sports, Science and Technology of
Japan (21H01253).
References
[1] Akiyama H, Heller R. Bioelectrics. 1st ed. Tokyo: Springer Japan: Imprint: Springer; 2017.
[2] Zhao W, Yang R, Lu R, Wang M, Qian P, Yang W. Effect of PEF on microbial inactivation and physical–chemical properties of green tea
extracts. Lebensm-Wiss Technol 2008;41(3):425–31. https://doi.org/10.1016/j.lwt.2007.03.020.
[3] Wan J, Coventry J, Swiergon P, Sanguansri P, Versteeg C. Advances in innovative processing technologies for microbial inactivation and
enhancement of food safety – pulsed electric field and low-temperature plasma. Trends Food Sci Technol 2009;20(9):414–24. https://doi.org/
10.1016/j.tifs.2009.01.050.
[4] Zare F, Ghasemi N, Bansal N, Garg A, Hosano H. Increasing the production yield of white oyster mushrooms with pulsed electric fields.
IEEE Trans Plasma Sci 2021;49(2):805–12. https://doi.org/10.1109/tps.2021.3053071.
[5] Gusbeth CA, Eing C, Gottel M, Frey W. Boost of algae growth by ultra short pulsed electric field treatment. Presented at the 2013 Abstracts
IEEE International Conference on Plasma Science (ICOPS). https://doi.org/10.1109/plasma.2013.6633325, 2013.
[6] Ahmed Z, et al. Impact of pulsed electric field treatments on the growth parameters of wheat seeds and nutritional properties of their wheat
plantlets juice. Food Sci Nutr May 2020;8(5):2490–500. https://doi.org/10.1002/fsn3.1540.
[7] Takaki K, Hayashi N, Wang D, Ohshima T. High-voltage technologies for agriculture and food processing. J Phys D, Appl Phys
2019;52(47):473001. https://doi.org/10.1088/1361-6463/ab2e2d.
[8] Al Daccache M, Koubaa M, Maroun RG, Salameh D, Louka N, Vorobiev E. Pulsed electric field-assisted fermentation of Hanseniaspora sp.
yeast isolated from Lebanese apples. Food Res Int Mar 2020;129:108840. https://doi.org/10.1016/j.foodres.2019.108840.
[9] Chanos P, Warncke MC, Ehrmann MA, Hertel C. Application of mild pulsed electric fields on starter culture accelerates yogurt fermentation.
Eur Food Res Technol 2020;246(3):621–30. https://doi.org/10.1007/s00217-020-03428-9.
[10] Kanafusa S, Uhlig E, Uemura K, Gómez Galindo F, Håkansson Å. The effect of nanosecond pulsed electric field on the production of
metabolites from lactic acid bacteria in fermented watermelon juice. Innov Food Sci Emerg Technol 2021;72:102749. https://doi.org/10.
1016/j.ifset.2021.102749.
[11] Zhang ZH, Han Z, Zeng XA, Wang MS. The preparation of Fe-glycine complexes by a novel method (pulsed electric fields). Food Chem
Mar 15 2017;219:468–76. https://doi.org/10.1016/j.foodchem.2016.09.129.
[12] Alles MC, et al. Bio-refinery of insects with pulsed electric field pre-treatment. Innov Food Sci Emerg Technol 2020;64:102403. https://
doi.org/10.1016/j.ifset.2020.102403.
[13] Martinez JM, Delso C, Alvarez I, Raso J. Pulsed electric field-assisted extraction of valuable compounds from microorganisms. Compr Rev
Food Sci Food Saf Mar 2020;19(2):530–52. https://doi.org/10.1111/1541-4337.12512.
[14] Zbinden MD, et al. Pulsed electric field (PEF) as an intensification pretreatment for greener solvent lipid extraction from microalgae. Biotech-
nol Bioeng Jun 2013;110(6):1605–15. https://doi.org/10.1002/bit.24829.
[15] Guionet A, Hosseini B, Teissie J, Akiyama H, Hosseini H. A new mechanism for efficient hydrocarbon electro-extraction from Botryococcus
braunii. Biotechnol Biofuels 2017;10(1):39. https://doi.org/10.1186/s13068-017-0724-1.
[16] Goettel M, Eing C, Gusbeth C, Straessner R, Frey W. Pulsed electric field assisted extraction of intracellular valuables from microalgae.
Algal Res 2013;2(4):401–8. https://doi.org/10.1016/j.algal.2013.07.004.
[17] Gorte O, et al. Pulsed electric field treatment promotes lipid extraction on fresh oleaginous yeast Saitozyma podzolica DSM 27192. Front
Bioeng Biotechnol 2020;8:575379. https://doi.org/10.3389/fbioe.2020.575379.
[18] Nowosad K, Sujka M, Pankiewicz U, Kowalski R. The application of PEF technology in food processing and human nutrition. J Food Sci
Technol Feb 2021;58(2):397–411. https://doi.org/10.1007/s13197-020-04512-4.
[19] Parniakov O, Barba FJ, Grimi N, Lebovka N, Vorobiev E. 1st world congress on electroporation and pulsed electric fields in biology, medicine
and food & environmental technologies. Singapore: Springer Singapore; 2015. p. 418–21.
[20] Frontuto D, et al. Optimization of pulsed electric fields-assisted extraction of polyphenols from potato peels using response surface method-
ology. Food Bioprocess Technol 2019;12(10):1708–20. https://doi.org/10.1007/s11947-019-02320-z.
[21] Peiró S, Luengo E, Segovia F, Raso J, Almajano MP. Improving polyphenol extraction from lemon residues by pulsed electric fields. Waste
Biomass Valoriz 2017;10(4):889–97. https://doi.org/10.1007/s12649-017-0116-6.
[22] Andreou V, Dimopoulos G, Dermesonlouoglou E, Taoukis P. Application of pulsed electric fields to improve product yield and waste val-
orization in industrial tomato processing. J Food Eng 2020;270:109778. https://doi.org/10.1016/j.jfoodeng.2019.109778.
[23] Ghosh S, Gillis A, Sheviryov J, Levkov K, Golberg A. Towards waste meat biorefinery: extraction of proteins from waste chicken meat with
non-thermal pulsed electric fields and mechanical pressing. J Clean Prod 2019;208:220–31. https://doi.org/10.1016/j.jclepro.2018.10.037.
[24] Barba FJ, et al. Current applications and new opportunities for the use of pulsed electric fields in food science and industry. Food Res Int
2015;77:773–98. https://doi.org/10.1016/j.foodres.2015.09.015.
[25] Golberg A, et al. Energy-efficient biomass processing with pulsed electric fields for bioeconomy and sustainable development. Biotechnol
Biofuels 2016;9(94):94. https://doi.org/10.1186/s13068-016-0508-z.
[26] Frey W, et al. Bioelectrics. Tokyo: Springer Japan; 2016. p. 389–476.
240
[27] Zhang C, Ye J, Lyu X, Zhao W, Mao J, Yang R. Effects of pulse electric field pretreatment on the frying quality and pore characteristics of
potato chips. Food Chem Feb 1 2022;369:130516. https://doi.org/10.1016/j.foodchem.2021.130516.
[28] Niu D, et al. Review of the application of pulsed electric fields (PEF) technology for food processing in China. Food Res Int Nov
2020;137:109715. https://doi.org/10.1016/j.foodres.2020.109715.
[29] Zhang F, Tian M, Du M, Fang T. Enhancing the activity of pectinase using pulsed electric field (PEF) treatment. J Food Eng 2017;205:56–63.
https://doi.org/10.1016/j.jfoodeng.2017.02.023.
[30] Fauster T, et al. Impact of pulsed electric field (PEF) pretreatment on process performance of industrial French fries production. J Food Eng
2018;235:16–22. https://doi.org/10.1016/j.jfoodeng.2018.04.023.
[31] Zhang C, et al. Effect of pulsed electric field pretreatment on oil content of potato chips. Lebensm-Wiss Technol 2021;135:110198. https://
doi.org/10.1016/j.lwt.2020.110198.
[32] Toepfl S, Heinz V. Pulsed electric fields (PEF) applications in food processing - process and equipment design and cost analysis. Presented
at the 2008 IEEE 35th International Conference on Plasma Science. https://doi.org/10.1109/plasma.2008.4590818, 2008.
[33] Stewart MP, Langer R, Jensen KF. Intracellular delivery by membrane disruption: mechanisms, strategies, and concepts. Chem Rev Aug 22
2018;118(16):7409–531. https://doi.org/10.1021/acs.chemrev.7b00678.
[34] Weaver JC, Vanbever R, Vaughan TE, Prausnitz MR. Heparin alters transdermal transport associated with electroporation. Biochem Biophys
Res Commun May 29 1997;234(3):637–40. https://doi.org/10.1006/bbrc.1997.6701.
[35] Šatkauskas S, Jakštys B, Ruzgys P, Jakutavičiūtė M. Different cell viability assays following electroporation in vitro. In: Miklavcic D, editor.
Handbook of electroporation. Cham: Springer International Publishing; 2016. p. 1–14, Chapter 140-1.
[36] Kotnik T. Lightning-triggered electroporation as a mechanism for horizontal gene transfer. In: Miklavcic D, editor. Handbook of electropo-
ration. Cham: Springer International Publishing; 2016. p. 1–18, Chapter 25-1.
[37] Bu B, Tian Z, Li D, Ji B. High transmembrane voltage raised by close contact initiates fusion pore. Front Mol Neurosci 2016;9:136. https://
doi.org/10.3389/fnmol.2016.00136.
[38] Wolff CM, et al. Combination of cold plasma and pulsed electric fields – a rationale for cancer patients in palliative care. Clin Plasma Med
2019;16:100096. https://doi.org/10.1016/j.cpme.2020.100096.
[39] Nuccitelli R, Tran K, Sheikh S, Athos B, Kreis M, Nuccitelli P. Optimized nanosecond pulsed electric field therapy can cause murine
malignant melanomas to self-destruct with a single treatment. Int J Cancer Oct 1 2010;127(7):1727–36. https://doi.org/10.1002/ijc.25364.
[40] Pakhomova O, Gianulis E, Pakhomov AG. Different cell sensitivity to pulsed electric field. In: Miklavcic D, editor. Handbook of electropo-
[41] Tieleman DP. The molecular basis of electroporation. BMC Biochem Jul 19 2004;5(1):10. https://doi.org/10.1186/1471-2091-5-10.
[42] Beretta G, Mastorgio AF, Pedrali L, Saponaro S, Sezenna E. The effects of electric, magnetic and electromagnetic fields on microorganisms
in the perspective of bioremediation. Rev Environ Sci Bio/Technol 2019;18(1):29–75. https://doi.org/10.1007/s11157-018-09491-9.
[43] Barnett A, Weaver JC. Electroporation: a unified, quantitative theory of reversible electrical breakdown and mechanical rupture in artificial
planar bilayer membranes. J Electroanal Chem Interfacial Electrochem 1991;320(2):163–82. https://doi.org/10.1016/0022-0728(91)85625-y.
[44] Weaver JC, Barnett A. Progress toward a theoretical model for electroporation mechanism: membrane electrical behavior and molecular
transport. In: Chang DC, Chassy BM, Saunders JA, Sowers AE, editors. Guide to electroporation and electrofusion. San Diego: Academic
Press; 1992. p. 91–117.
[45] Karal MAS, et al. Electrostatic effects on the electrical tension-induced irreversible pore formation in giant unilamellar vesicles. Chem Phys
Lipids Sep 2020;231:104935. https://doi.org/10.1016/j.chemphyslip.2020.104935.
[46] Vernier PT, Ziegler MJ. Nanosecond field alignment of head group and water dipoles in electroporating phospholipid bilayers. J Phys Chem
B Nov 15 2007;111(45):12993–6. https://doi.org/10.1021/jp077148q.
[47] Tarek M. Membrane electroporation: a molecular dynamics simulation. Biophys J Jun 2005;88(6):4045–53. https://doi.org/10.1529/biophysj.
104.050617.
[48] Ziegler MJ, Vernier PT. Interface water dynamics and porating electric fields for phospholipid bilayers. J Phys Chem B Oct 30
2008;112(43):13588–96. https://doi.org/10.1021/jp8027726.
[49] Marrink SJ, de Vries AH, Tieleman DP. Lipids on the move: simulations of membrane pores, domains, stalks and curves. Biochim Biophys
Acta Jan 2009;1788(1):149–68. https://doi.org/10.1016/j.bbamem.2008.10.006.
[50] Bockmann RA, de Groot BL, Kakorin S, Neumann E, Grubmuller H. Kinetics, statistics, and energetics of lipid membrane electroporation
studied by molecular dynamics simulations. Biophys J Aug 2008;95(4):1837–50. https://doi.org/10.1529/biophysj.108.129437.
[51] Zheng X, Gallot G. Dynamics of cell membrane permeabilization by saponins using terahertz attenuated total reflection. Biophys J Aug 18
2020;119(4):749–55. https://doi.org/10.1016/j.bpj.2020.05.040.
[52] Hu Q, Joshi RP. Transmembrane voltage analyses in spheroidal cells in response to an intense ultrashort electrical pulse. Phys Rev E, Stat
Nonlinear Soft Matter Phys Jan 2009;79(1 Pt 1):011901. https://doi.org/10.1103/PhysRevE.79.011901.
[53] Hu Q, Joshi RP. Analysis of intense, subnanosecond electrical pulse-induced transmembrane voltage in spheroidal cells with arbitrary orien-
tation. IEEE Trans Biomed Eng Jun 2009;56(6):1617–26. https://doi.org/10.1109/TBME.2009.2015459.
[54] Anosov AA, Sharakshane AA, Smirnova EY, Nemchenko OY. Application of the Smoluchowski equation with a source term to the model
of lipid pore formation during a phase transition. Biophysics 2017;61(6):936–41. https://doi.org/10.1134/s0006350916060051.
[55] Weaver JC. Transient aqueous pores: a mechanism for coupling electric fields to bilayer and cell membranes. In: Blank M, Findl E, editors.
Mechanistic approaches to interactions of electric and electromagnetic fields with living systems. Boston, MA: Springer US; 1987. p. 249–70,
Chapter 15.
[56] Smith KC, Neu JC, Krassowska W. Model of creation and evolution of stable electropores for DNA delivery. Biophys J May
2004;86(5):2813–26. https://doi.org/10.1016/S0006-3495(04)74334-9.
[57] Krassowska W, Filev PD. Modeling electroporation in a single cell. Biophys J Jan 15 2007;92(2):404–17. https://doi.org/10.1529/biophysj.
106.094235.
241
[58] Freeman SA, Wang MA, Weaver JC. Theory of electroporation of planar bilayer membranes: predictions of the aqueous area, change in
capacitance, and pore-pore separation. Biophys J Jul 1994;67(1):42–56. https://doi.org/10.1016/S0006-3495(94)80453-9.
[59] Stewart DA, Gowrishankar TR, Weaver JC. Transport lattice approach to describing cell electroporation: use of a local asymptotic model.
IEEE Trans Plasma Sci 2004;32(4):1696–708. https://doi.org/10.1109/tps.2004.832639.
[60] Anosov AA, Smirnova EY, Ryleeva ED, Gligonov IA, Korepanova EA, Sharakshane AA. Estimation of the parameters of the Smoluchowski
equation describing the occurrence of pores in a bilayer lipid membrane under soft poration. Eur Phys J E, Soft Matter Oct 6 2020;43(10):66.
https://doi.org/10.1140/epje/i2020-11989-0.
[61] Anosov AA, Smirnova EY, Sharakshane AA, Nikolayeva EA, Zhdankina YS. Increase in the current variance in bilayer lipid membranes
near phase transition as a result of the occurrence of hydrophobic defects. Biochim Biophys Acta, Biomembr Feb 1 2020;1862(2):183147.
https://doi.org/10.1016/j.bbamem.2019.183147.
[62] DeBruin KA, Krassowska W. Modeling electroporation in a single cell. I. Effects of field strength and rest potential. Biophys J Sep
1999;77(3):1213–24. https://doi.org/10.1016/S0006-3495(99)76973-0.
[63] Levine ZA. Effects of heterogeneous membranes and electrolytes on electropore formation. In: Miklavcic D, editor. Handbook of electropo-
[64] Fernández ML, Risk MR. Lipid electropore stabilization. In: Miklavcic D, editor. Handbook of electroporation. Cham: Springer International
Publishing; 2016. p. 1–17, Chapter 83-1.
[65] Levine ZA. Lipid electropore lifetime in molecular models. In: Miklavcic D, editor. Handbook of electroporation. Cham: Springer Interna-
tional Publishing; 2016. p. 1–19, Chapter 86-1.
[66] Marracino P, Vernier PT, Liberti M, Apollonio F. Lipid electropore geometry in molecular models. In: Miklavcic D, editor. Handbook of
electroporation. Cham: Springer International Publishing; 2016. p. 1–16, Chapter 88-1.
[67] Smith KC, Weaver JC. Active mechanisms are needed to describe cell responses to submicrosecond, megavolt-per-meter pulses: cell models
for ultrashort pulses. Biophys J Aug 2008;95(4):1547–63. https://doi.org/10.1529/biophysj.107.121921.
[68] Son RS, Smith KC, Gowrishankar TR, Vernier PT, Weaver JC. Basic features of a cell electroporation model: illustrative behavior for two
very different pulses. J Membr Biol 2014;247(12):1209–28. https://doi.org/10.1007/s00232-014-9699-z.
[69] Kotnik T, Rems L, Tarek M, Miklavcic D. Membrane electroporation and electropermeabilization: mechanisms and models. Annu Rev
Biophys May 6 2019;48(1):63–91. https://doi.org/10.1146/annurev-biophys-052118-115451.
[70] Timoshkin IV, MacGregor SJ, Fouracre RA, Crichton BH, Anderson JG. Transient electrical field across cellular membranes: pulsed electric
field treatment of microbial cells. J Phys D, Appl Phys 2006;39(3):596–603. https://doi.org/10.1088/0022-3727/39/3/026.
[71] Kuyucak S, Hoyles M, Chung S-H. Analytical solutions of Poisson’s equation for realistic geometrical shapes of membrane ion channels.
Biophys J 1998;74(1):22–36. https://doi.org/10.1016/S0006-3495(98)77763-X.
[72] Garcia D, Gomez N, Manas P, Condon S, Raso J, Pagan R. Occurrence of sublethal injury after pulsed electric fields depending on the
micro-organism, the treatment medium ph and the intensity of the treatment investigated. J Appl Microbiol 2005;99(1):94–104. https://
doi.org/10.1111/j.1365-2672.2005.02611.x.
[73] Buchmann L, Mathys A. Perspective on pulsed electric field treatment in the bio-based industry. Front Bioeng Biotechnol 2019;7:265. https://
doi.org/10.3389/fbioe.2019.00265.
[74] Rems L, Kasimova MA, Testa I, Delemotte L. Pulsed electric fields can create pores in the voltage sensors of voltage-gated ion channels.
Biophys J Jul 7 2020;119(1):190–205. https://doi.org/10.1016/j.bpj.2020.05.030.
[75] Sozer EB, Haldar S, Blank PS, Castellani F, Vernier PT, Zimmerberg J. Dye transport through bilayers agrees with lipid electropore molecular
dynamics. Biophys J Nov 3 2020;119(9):1724–34. https://doi.org/10.1016/j.bpj.2020.09.028.
[76] Tieleman DP, Leontiadou H, Mark AE, Marrink SJ. Simulation of pore formation in lipid bilayers by mechanical stress and electric fields. J
Am Chem Soc May 28 2003;125(21):6382–3. https://doi.org/10.1021/ja029504i.
[77] Chen G, Huang K, Miao M, Feng B, Campanella OH. Molecular dynamics simulation for mechanism elucidation of food processing and
safety: state of the art. Compr Rev Food Sci Food Saf Jan 2019;18(1):243–63. https://doi.org/10.1111/1541-4337.12406.
[78] Xu J, Gu L, Ye Z, Kargarrazi S, Rivas-Davila JM. Cascode GaN/SiC: a wide-bandgap heterogenous power device for high-frequency
applications. IEEE Trans Power Electron 2020;35(6):6340–9. https://doi.org/10.1109/tpel.2019.2954322.
[79] Jäger H. Process performance analysis of pulsed electric field (PEF) food applications. Technische Universität Berlin; 2011.
[80] Jin TZ. Pulsed electric fields for pasteurization: defining processing conditions. In: Miklavcic D, editor. Handbook of electroporation. Cham:
Springer International Publishing; 2017. p. 1–25, Chapter 132-1.
[81] Góngora-Nieto MM, Sepúlveda DR, Pedrow P, Barbosa-Cánovas GV, Swanson BG. Food processing by pulsed electric fields: treatment
delivery, inactivation level, and regulatory aspects. Lebensm-Wiss Technol 2002;35(5):375–88. https://doi.org/10.1006/fstl.2001.0880.
[82] Hulsheger H, Potel J, Niemann EG. Killing of bacteria with electric pulses of high field strength. Radiat Environ Biophys 1981;20(1):53–65.
https://doi.org/10.1007/BF01323926.
[83] Lu W, et al. Theoretical analysis of transmembrane potential of cells exposed to nanosecond pulsed electric field. Int J Radiat Biol Feb
2017;93(2):231–9. https://doi.org/10.1080/09553002.2017.1230244.
[84] Raso J, et al. Recommendations guidelines on the key information to be reported in studies of application of PEF technology in food and
biotechnological processes. Innov Food Sci Emerg Technol 2016;37:312–21. https://doi.org/10.1016/j.ifset.2016.08.003.
[85] Zhang Q, Barbosa-Cánovas GV, Swanson BG. Engineering aspects of pulsed electric field pasteurization. J Food Eng 1995;25(2):261–81.
https://doi.org/10.1016/0260-8774(94)00030-d.
[86] Yang RJ, Li SQ, Zhang QH. Effects of pulsed electric fields on the activity of enzymes in aqueous solution. J Food Sci 2004;69(4):FCT241–8.
https://doi.org/10.1111/j.1365-2621.2004.tb06323.x.
[87] Kinosita Jr K, Tsong TT. Hemolysis of human erythrocytes by transient electric field. Proc Natl Acad Sci USA May 1977;74(5):1923–7.
https://doi.org/10.1073/pnas.74.5.1923.
242
[88] Ho SYW. Analysis of two high voltage electric pulse systems for batch and continuous pasteurization of selected fluid products. ProQuest
Dissertations Publishing. 1997 [Online]. Available: https://hdl.handle.net/10214/21965.
[89] Beebe SJ, Chen YJ, Sain NM, Schoenbach KH, Xiao S. Transient features in nanosecond pulsed electric fields differentially modulate
mitochondria and viability. PLoS ONE 2012;7(12):e51349. https://doi.org/10.1371/journal.pone.0051349.
[90] Vaessen EMJ, Timmermans RAH, Tempelaars MH, Schutyser MAI, den Besten HMW. Reversibility of membrane permeabilization upon
pulsed electric field treatment in Lactobacillus plantarum WCFS1. Sci Rep Dec 27 2019;9(1):19990. https://doi.org/10.1038/s41598-019-
56299-w.
[91] Takaki K, Takahashi K, Guionet A, Ohshima T. Pulsed power applications for protein conformational change and the permeabilization of
agricultural products. Molecules Oct 18 2021;26(20):6288. https://doi.org/10.3390/molecules26206288.
[92] Bi X, et al. Effects of electric field strength and pulse rise time on physicochemical and sensory properties of apple juice by pulsed electric
field. Innov Food Sci Emerg Technol 2013;17:85–92. https://doi.org/10.1016/j.ifset.2012.10.008.
[93] Aadil RM, et al. Influence of different pulsed electric field strengths on the quality of the grapefruit juice. Int J Food Sci Technol
2015;50(10):2290–6. https://doi.org/10.1111/ijfs.12891.
[94] Evrendilek GA. Pulsed electric field treatment for beverage production and preservation. In: Miklavcic D, editor. Handbook of electropora-
tion. Cham: Springer International Publishing; 2016. p. 1–17, Chapter 180-1.
[95] Trainito C. Study of cell membrane permeabilization induced by pulsed electric field – electrical modeling and characterization on biochip.
https://tel.archives-ouvertes.fr/tel-01254036, 2015.
[96] Vincelette RL, Roth CC, McConnell MP, Payne JA, Beier HT, Ibey BL. Thresholds for phosphatidylserine externalization in Chinese hamster
ovarian cells following exposure to nanosecond pulsed electrical fields (nsPEF). PLoS ONE 2013;8(4):e63122. https://doi.org/10.1371/
journal.pone.0063122.
[97] Yang E, Li J, Cho M, Xiao S. Cell fragmentation and permeabilization by a 1 ns pulse driven triple-point electrode. BioMed Res Int
2018/03/18;2018:4072983. https://doi.org/10.1155/2018/4072983.
[98] Kotnik T, Miklavčič D, Slivnik T. Time course of transmembrane voltage induced by time-varying electric fields—a method for theoretical
analysis and its application. Bioelectrochem Bioenerg 1998;45(1):3–16. https://doi.org/10.1016/s0302-4598(97)00093-7.
[99] Grossi M, Riccò B. Electrical impedance spectroscopy (EIS) for biological analysis and food characterization: a review. J Sens Sens Syst
2017;6(2):303–25. https://doi.org/10.5194/jsss-6-303-2017.
[100] Pataro G, Donsì G, Ferrari G. Modeling of electrochemical reactions during pulsed electric field treatment. In: Miklavcic D, editor. Handbook
of electroporation. Cham: Springer International Publishing; 2016. p. 1–30, Chapter 5-1.
[101] Ohshima T, Tanino T, Kameda T, Harashima H. Engineering of operation condition in milk pasteurization with PEF treatment. Food Control
2016;68:297–302. https://doi.org/10.1016/j.foodcont.2016.03.047.
[102] Akiyama H. Streamer discharges in liquids and their applications. IEEE Trans Dielectr Electr Insul 2000;7(5):646–53. https://doi.org/10.
1109/94.879360.
[103] Sakamoto S, Yamada H. Optical study of conduction and breakdown in dielectric liquids. IEEE Trans Electr Insul 1980;EI-15(3):171–81.
https://doi.org/10.1109/TEI.1980.298310.
[104] Zhu L, He Z-H, Gao Z-W, Tan F-L, Yue X-G, Chang J-S. Research on the influence of conductivity to pulsed arc electrohydraulic discharge
in water. J Electrost 2014;72(1):53–8. https://doi.org/10.1016/j.elstat.2013.11.004.
[105] Evrendilek GA, Li S, Dantzer WR, Zhang QH. Pulsed electric field processing of beer: microbial, sensory, and quality analyses. J Food Sci
2004;69(8):M228–32. https://doi.org/10.1111/j.1365-2621.2004.tb09892.x.
[106] Hung PSK, Lee OW, Yeung KL, Kwan SM, Kwan JKC. Pulsed electric field for drinking water disinfection. Patent US10941060 (B2). 2021.
[107] Heinz V, Alvarez I, Angersbach A, Knorr D. Preservation of liquid foods by high intensity pulsed electric fields—basic concepts for process
design. Trends Food Sci Technol 2001/03/04;12(3–4):103–11. https://doi.org/10.1016/s0924-2244(01)00064-4.
[108] Rapp BE. Fluids. In: Rapp BE, editor. Microfluidics: modelling, mechanics and mathematics. Oxford: Elsevier; 2017. p. 243–63.
[109] Snyder AB, Churey JJ, Worobo RW. Association of fungal genera from spoiled processed foods with physicochemical food properties and
processing conditions. Food Microbiol Oct 2019;83:211–8. https://doi.org/10.1016/j.fm.2019.05.012.
[110] Dawson D. Foodborne protozoan parasites. Int J Food Microbiol Aug 25 2005;103(2):207–27. https://doi.org/10.1016/j.ijfoodmicro.2004.
12.032.
[111] Lee HS. Diversity of halophilic archaea in fermented foods and human intestines and their application. J Microbiol Biotechnol Dec
2013;23(12):1645–53. https://doi.org/10.4014/jmb.1308.08015.
[112] Abeysiriwardena NM, Gascoigne SJL, Anandappa A. Algal bloom expansion increases cyanotoxin risk in food. Yale J Biol Med Jun
2018;91(2):129–42 [Online]. Available: https://www.ncbi.nlm.nih.gov/pubmed/29955218.
[113] Bosch A, et al. Foodborne viruses: detection, risk assessment, and control options in food processing. Int J Food Microbiol Nov 20
2018;285:110–28. https://doi.org/10.1016/j.ijfoodmicro.2018.06.001.
[114] Rodriguez-Lazaro D, et al. Virus hazards from food, water and other contaminated environments. FEMS Microbiol Rev Jul
2012;36(4):786–814. https://doi.org/10.1111/j.1574-6976.2011.00306.x.
[115] Ryan U, Hijjawi N, Xiao L. Foodborne cryptosporidiosis. Int J Parasitol Jan 2018;48(1):1–12. https://doi.org/10.1016/j.ijpara.2017.09.004.
[116] Hazards EPoB, et al. Public health risks associated with food-borne parasites. EFSA J Dec 2018;16(12):e05495. https://doi.org/10.2903/j.
efsa.2018.5495.
[117] Lorenzo JM, Munekata PE, Dominguez R, Pateiro M, Saraiva JA, Franco D. Main groups of microorganisms of relevance for food safety
and stability. In: Innovative technologies for food preservation; 2018. p. 53–107.
[118] Hamilton W, Sale A. Effects of high electric fields on microorganisms II. Mechanism of action of the lethal effect. Biochim Biophys Acta G,
Gen Subj 1967/12/27;148(3):789–800. https://doi.org/10.1016/0304-4165(67)90053-0.
[119] Sale A, Hamilton W. Effects of high electric fields on microorganisms I. Killing of bacteria and yeasts. Biochim Biophys Acta G, Gen Subj
1967/12/27;148(3):781–8. https://doi.org/10.1016/0304-4165(67)90052-9.
243
[120] Special supplement: kinetics of microbial inactivation for alternative food processing technologies - IFT’s response to task order #1, US Food
and Drug Administration: how to quantify the destruction kinetics of alternative processing technologies. J Food Sci 2000:4–108 [Online].
Available: https://books.google.com.au/books?id=On6HzgEACAAJ.
[121] Richmond D, Chang TS. A comparison of drop-plate and pour-plate methods for bacterial population counts of poultry anaphage (dehydrated
caged layer excreta). Poult Sci Jan 1978;57(1):293–5. https://doi.org/10.3382/ps.0570293.
[122] Hoben HJ, Somasegaran P. Comparison of the pour, spread, and drop plate methods for enumeration of Rhizobium spp. in inoculants made
from presterilized peat. Appl Environ Microbiol Nov 1982;44(5):1246–7. https://doi.org/10.1128/aem.44.5.1246-1247.1982.
[123] Herigstad B, Hamilton M, Heersink J. How to optimize the drop plate method for enumerating bacteria. J Microbiol Methods
2001;44(2):121–9. https://doi.org/10.1016/s0167-7012(00)00241-4.
[124] Walkling-Ribeiro M, Rodríguez-González O, Jayaram SH, Griffiths MW. Processing temperature, alcohol and carbonation levels and their
impact on pulsed electric fields (PEF) mitigation of selected characteristic microorganisms in beer. Food Res Int 2011;44(8):2524–33. https://
doi.org/10.1016/j.foodres.2011.01.046.
[125] Milani EA, Alkhafaji S, Silva FVM. Pulsed electric field continuous pasteurization of different types of beers. Food Control 2015;50:223–9.
https://doi.org/10.1016/j.foodcont.2014.08.033.
[126] Puligundla P, Pyun YR, Mok C. Pulsed electric field (PEF) technology for microbial inactivation in low-alcohol red wine. Food Sci Biotechnol
Dec 2018;27(6):1691–6. https://doi.org/10.1007/s10068-018-0422-1.
[127] Hulsheger H, Niemann EG. Lethal effects of high-voltage pulses on E. coli K12. Radiat Environ Biophys 1980;18(4):281–8. https://doi.org/
10.1007/BF01324271.
[128] Singh J, Singh M, Singh B, Nayak M, Ghanshyam C. Comparative analyses of prediction models for inactivation of Escherichia coli in carrot
juice by means of pulsed electric fields. J Food Sci Technol May 2017;54(6):1538–44. https://doi.org/10.1007/s13197-017-2585-9.
[129] Bigelow WD. The logarithmic nature of thermal death time curves. J Infect Dis 1921;29(5):528–36. https://doi.org/10.1093/infdis/29.5.528.
[130] Peleg M. A model of microbial survival after exposure to pulsed electric fields. J Sci Food Agric 1995;67(1):93–9. https://doi.org/10.1002/
jsfa.2740670115.
[131] Peleg M, Cole MB. Reinterpretation of microbial survival curves. Crit Rev Food Sci Nutr Jul 1998;38(5):353–80. https://doi.org/10.1080/
10408699891274246.
[132] Semenov I, Xiao S. Electropermeabilization and electrostimulation by picosecond pulses. In: Miklavcic D, editor. Handbook of electropora-
[133] Chen C, Smye SW, Robinson MP, Evans JA. Membrane electroporation theories: a review. Med Biol Eng Comput 2006;44(1–2):5–14.
https://doi.org/10.1007/s11517-005-0020-2.
[134] Crowley JM. Electrical breakdown of biomolecular lipid membranes as an electromechanical instability. Biophys J 1973;13(7):711–24.
https://doi.org/10.1016/S0006-3495(73)86017-5.
[135] Michael DH, O’Neill ME. Electrohydrodynamic instability in plane layers of fluid. J Fluid Mech 1970;41(3):571–80. https://doi.org/10.
1017/S0022112070000757.
[136] Abidor IG, Arakelyan VB, Chernomordik LV, Chizmadzhev YA, Pastushenko VF, Tarasevich MP. Electric breakdown of bilayer lipid mem-
branes. I. The main experimental facts and their qualitative discussion. J Electroanal Chem Interfacial Electrochem 1979;104(C):37–52.
https://doi.org/10.1016/S0022-0728(79)81006-2.
[137] Powell KT, Weaver JC. Transient aqueous pores in bilayer membranes: a statistical theory. Bioelectrochem Bioenerg 1986;15(2):211–27.
https://doi.org/10.1016/0302-4598(86)80029-0.
[138] Cunill-Semanat E, Salgado J. Spontaneous and stress-induced pore formation in membranes: theory, experiments and simulations. J Membr
Biol Oct 2019;252(4–5):241–60. https://doi.org/10.1007/s00232-019-00083-4.
[139] Saulis G. Electroporation of cell membranes: the fundamental effects of pulsed electric fields in food processing. Food Eng Rev
2010;2(2):52–73. https://doi.org/10.1007/s12393-010-9023-3.
[140] Stewart Jr DA, Gowrishankar TR, Smith KC, Weaver JC. Cylindrical cell membranes in uniform applied electric fields: validation of a
transport lattice method. IEEE Trans Biomed Eng Oct 2005;52(10):1643–53. https://doi.org/10.1109/TBME.2005.856030.
[141] Chernomordik LV, Sukharev SI, Abidor IG, Chizmadzhev YA. 468—the study of the BLM reversible electrical breakdown mechanism in
the presence of UO22+. Bioelectrochem Bioenerg 1982;9(2):149–55. https://doi.org/10.1016/0302-4598(82)80171-2.
[142] Chernomordik LV, Sukharev SI, Abidor IG, Chizmadzhev YA. Breakdown of lipid bilayer membranes in an electric field. Biochim Biophys
Acta, Biomembr 1983;736(2):203–13. https://doi.org/10.1016/0005-2736(83)90285-7.
[143] Abidor IG, Arakelyan VB, Chernomordik LV, Chizmadzhev YA, Pastushenko VF, Tarasevich MP. Electric breakdown of bilayer lipid mem-
branes. J Electroanal Chem Interfacial Electrochem 1979;104(C):37–52. https://doi.org/10.1016/s0022-0728(79)81006-2.
[144] Tovar O, Tung L. Electroporation and recovery of cardiac cell membrane with rectangular voltage pulses. Am J Physiol Oct 1992;263(4 Pt
2):H1128–36. https://doi.org/10.1152/ajpheart.1992.263.4.H1128.
[145] Bae C, Butler PJ. Automated single-cell electroporation. BioTechniques Oct 2006;41(4):399–400, 402. https://doi.org/10.2144/000112261.
[146] Ibrahimi N, et al. A subnanosecond pulsed electric field system for studying cells electropermeabilization. IEEE Trans Plasma Sci
2020;48(12):4242–9. https://doi.org/10.1109/tps.2020.3034286.
[147] Xiao S, Guo S, Nesin V, Heller R, Schoenbach KH. Subnanosecond electric pulses cause membrane permeabilization and cell death. IEEE
Trans Biomed Eng May 2011;58(5):1239–45. https://doi.org/10.1109/TBME.2011.2112360.
[148] Kotnik T, Miklavcic D. Second-order model of membrane electric field induced by alternating external electric fields. IEEE Trans Biomed
Eng Aug 2000;47(8):1074–81. https://doi.org/10.1109/10.855935.
[149] Teissie J. Mechanistic description of membrane electropermeabilization. In: Miklavcic D, editor. Handbook of electroporation. Cham:
[150] Teissie J. Critical electric field and transmembrane voltage for lipid pore formation in experiments. In: Miklavcic D, editor. Handbook of
244
[151] Pakhomov AG, Miklavcic D, Markov MS. Advanced electroporation techniques in biology and medicine. CRC Press; 2010 [Online]. Avail-
able: https://books.google.com.au/books?id=sTHNBQAAQBAJ.
[152] Kotnik T, Bobanović F, Miklavcˇicˇ D. Sensitivity of transmembrane voltage induced by applied electric fields—a theoretical analysis.
Bioelectrochem Bioenerg 1997/08/01;43(2):285–91. https://doi.org/10.1016/s0302-4598(97)00023-8.
[153] Zimmermann U, Pilwat G, Riemann F. Dielectric breakdown of cell membranes. Biophys J 1974/11/01;14(11):881–99. https://doi.org/10.
1016/s0006-3495(74)85956-4.
[154] Frey W, et al. Plasma membrane voltage changes during nanosecond pulsed electric field exposure. Biophys J May 15 2006;90(10):3608–15.
https://doi.org/10.1529/biophysj.105.072777.
[155] Schoenbach K, et al. Bioelectric effects of intense nanosecond pulses. IEEE Trans Dielectr Electr Insul 2007;14(5):1088–109. https://doi.
org/10.1109/tdei.2007.4339468.
[156] Flickinger B, Berghofer T, Hohenberger P, Eing C, Frey W. Transmembrane potential measurements on plant cells using the voltage-sensitive
dye ANNINE-6. Protoplasma Nov 2010;247(1–2):3–12. https://doi.org/10.1007/s00709-010-0131-y.
[157] Deng J, Schoenbach KH, Buescher ES, Hair PS, Fox PM, Beebe SJ. The effects of intense submicrosecond electrical pulses on cells. Biophys
J Apr 2003;84(4):2709–14. https://doi.org/10.1016/s0006-3495(03)75076-0.
[158] Tekle E, Oubrahim H, Dzekunov SM, Kolb JF, Schoenbach KH, Chock PB. Selective field effects on intracellular vacuoles and vesicle
membranes with nanosecond electric pulses. Biophys J Jul 2005;89(1):274–84. https://doi.org/10.1529/biophysj.104.054494.
[159] Batista Napotnik T, Rebersek M, Vernier PT, Mali B, Miklavcic D. Effects of high voltage nanosecond electric pulses on eukaryotic cells (in
vitro): a systematic review. Bioelectrochemistry Aug 2016;110:1–12. https://doi.org/10.1016/j.bioelechem.2016.02.011.
[160] Beebe SJ, Fox PM, Rec LJ, Willis EL, Schoenbach KH. Nanosecond, high-intensity pulsed electric fields induce apoptosis in human cells.
FASEB J Aug 2003;17(11):1493–5. https://doi.org/10.1096/fj.02-0859fje.
[161] Gentet LJ, Stuart GJ, Clements JD. Direct measurement of specific membrane capacitance in neurons. Biophys J Jul 2000;79(1):314–20.
https://doi.org/10.1016/S0006-3495(00)76293-X.
[162] Heimburg T. The capacitance and electromechanical coupling of lipid membranes close to transitions: the effect of electrostriction. Biophys
J Sep 5 2012;103(5):918–29. https://doi.org/10.1016/j.bpj.2012.07.010.
[163] Carius W. Voltage dependence of bilayer membrane capacitance. J Colloid Interface Sci 1976;57(2):301–7. https://doi.org/10.1016/0021-
9797(76)90205-8.
[164] Strahl H, Hamoen LW. Membrane potential is important for bacterial cell division. Proc Natl Acad Sci USA Jul 6 2010;107(27):12281–6.
https://doi.org/10.1073/pnas.1005485107.
[165] Bruni GN, Weekley RA, Dodd BJT, Kralj JM. Voltage-gated calcium flux mediates Escherichia coli mechanosensation. Proc Natl Acad Sci
USA Aug 29 2017;114(35):9445–50. https://doi.org/10.1073/pnas.1703084114.
[166] Sirec T, Benarroch JM, Buffard P, Garcia-Ojalvo J, Asally M. Electrical polarization enables integrative quality control during bacterial
differentiation into spores. iScience Jun 28 2019;16:378–89. https://doi.org/10.1016/j.isci.2019.05.044.
[167] Prindle A, Liu J, Asally M, Ly S, Garcia-Ojalvo J, Suel GM. Ion channels enable electrical communication in bacterial communities. Nature
Nov 5 2015;527(7576):59–63. https://doi.org/10.1038/nature15709.
[168] Stratford JP, et al. Electrically induced bacterial membrane-potential dynamics correspond to cellular proliferation capacity. Proc Natl Acad
Sci USA May 7 2019;116(19):9552–7. https://doi.org/10.1073/pnas.1901788116.
[169] Lee DD, Prindle A, Liu J, Suel GM. SnapShot: electrochemical communication in biofilms. Cell Jun 29 2017;170(1):214–2141.e. https://
doi.org/10.1016/j.cell.2017.06.026.
[170] Bot CT, Prodan C. Quantifying the membrane potential during E. coli growth stages. Biophys Chem Feb 2010;146(2–3):133–7. https://
doi.org/10.1016/j.bpc.2009.11.005.
[171] Yun O, Zeng X-A, Brennan CS, Zhi-wei L. Temperature alters the structure of membrane lipids and pulsed electric field (PEF) resistance of
Salmonella Typhimurium. Int J Food Sci Technol 2017/02/01;52(2):424–30. https://doi.org/10.1111/ijfs.13297.
[172] Peters MJ, Stinstra G, Hendriks M. Estimation of the electrical conductivity of human tissue. Electromagnetics 2001;21(7–8):545–57. https://
doi.org/10.1080/027263401752246199.
[173] Poignard C, Silve A, Wegner L. Different approaches used in modeling of cell membrane electroporation. In: Miklavcic D, editor. Handbook
[174] Jensen SD, Khorokhorina VA, Muratori C, Pakhomov AG, Pakhomova ON. Delayed hypersensitivity to nanosecond pulsed electric field in
electroporated cells. Sci Rep Sep 8 2017;7(1):10992. https://doi.org/10.1038/s41598-017-10825-w.
[175] Muratori C, Casciola M, Pakhomova O. Electric pulse repetition rate: sensitization and desensitization. In: Miklavcic D, editor. Handbook
[176] Valdez CM, Barnes Jr RA, Roth CC, Moen EK, Throckmorton GA, Ibey BL. Asymmetrical bipolar nanosecond electric pulse widths modify
bipolar cancellation. Sci Rep Nov 27 2017;7(1):16372. https://doi.org/10.1038/s41598-017-16142-6.
[177] Pakhomov AG, Grigoryev S, Semenov I, Casciola M, Jiang C, Xiao S. The second phase of bipolar, nanosecond-range electric pulses
determines the electroporation efficiency. Bioelectrochemistry Aug 2018;122:123–33. https://doi.org/10.1016/j.bioelechem.2018.03.014.
[178] Gianulis EC, Casciola M, Xiao S, Pakhomova ON, Pakhomov AG. Electropermeabilization by uni- or bipolar nanosecond electric pulses:
the impact of extracellular conductivity. Bioelectrochemistry Feb 2018;119:10–9. https://doi.org/10.1016/j.bioelechem.2017.08.005.
[179] Valdez CM, Barnes Jr R, Roth CC, Moen E, Ibey B. The interphase interval within a bipolar nanosecond electric pulse modulates bipolar
cancellation. Bioelectromagnetics Sep 2018;39(6):441–50. https://doi.org/10.1002/bem.22134.
[180] Pakhomov AG, et al. Cancellation of cellular responses to nanoelectroporation by reversing the stimulus polarity. Cell Mol Life Sci Nov
2014;71(22):4431–41. https://doi.org/10.1007/s00018-014-1626-z.
[181] Sweeney DC, Rebersek M, Dermol J, Rems L, Miklavcic D, Davalos RV. Quantification of cell membrane permeability induced by monopo-
lar and high-frequency bipolar bursts of electrical pulses. Biochim Biophys Acta Nov 2016;1858(11):2689–98. https://doi.org/10.1016/j.
bbamem.2016.06.024.
245
[182] Orlacchio R, Carr L, Palego C, Arnaud-Cormos D, Leveque P. High-voltage 10 ns delayed paired or bipolar pulses for in vitro bioelectric
experiments. Bioelectrochemistry Feb 2021;137:107648. https://doi.org/10.1016/j.bioelechem.2020.107648.
[183] Gowrishankar TR, Stern JV, Smith KC, Weaver JC. Nanopore occlusion: a biophysical mechanism for bipolar cancellation in cell membranes.
Biochem Biophys Res Commun Sep 10 2018;503(3):1194–9. https://doi.org/10.1016/j.bbrc.2018.07.024.
[184] Schoenbach KH, Pakhomov AG, Semenov I, Xiao S, Pakhomova ON, Ibey BL. Ion transport into cells exposed to monopolar and bipolar
nanosecond pulses. Bioelectrochemistry Jun 2015;103:44–51. https://doi.org/10.1016/j.bioelechem.2014.08.015.
[185] Pillet F, Formosa-Dague C, Baaziz H, Dague E, Rols MP. Cell wall as a target for bacteria inactivation by pulsed electric fields. Sci Rep Feb
1 2016;6(1):19778. https://doi.org/10.1038/srep19778.
[186] Xiao S, Yamada R, Zhou C. Quadrupoles for remote electrostimulation incorporating bipolar cancellation. Bioelectricity Dec 1
2020;2(4):382–90. https://doi.org/10.1089/bioe.2020.0024.
[187] Heller LC, Jaroszeski MJ, Coppola D, McCray AN, Hickey J, Heller R. Optimization of cutaneous electrically mediated plasmid DNA
delivery using novel electrode. Gene Ther Feb 2007;14(3):275–80. https://doi.org/10.1038/sj.gt.3302867.
[188] Rebersek M, et al. Electroporator with automatic change of electric field direction improves gene electrotransfer in-vitro. Biomed Eng Online
Jul 2 2007;6(1):25. https://doi.org/10.1186/1475-925X-6-25.
[189] Polajzer T, Dermol-Cerne J, Rebersek M, O’Connor R, Miklavcic D. Cancellation effect is present in high-frequency reversible and irre-
versible electroporation. Bioelectrochemistry Apr 2020;132:107442. https://doi.org/10.1016/j.bioelechem.2019.107442.
[190] Gianulis EC, Casciola M, Zhou C, Yang E, Xiao S, Pakhomov AG. Selective distant electrostimulation by synchronized bipolar nanosecond
pulses. Sci Rep Sep 11 2019;9(1):13116. https://doi.org/10.1038/s41598-019-49664-2.
[191] Johnson JE, Cornell RB. Amphitropic proteins: regulation by reversible membrane interactions (review). Mol Membr Biol Jul-Sep
1999;16(3):217–35. https://doi.org/10.1080/096876899294544.
[192] Teissie J. Membrane permeabilization lifetime in experiments. In: Miklavcic D, editor. Handbook of electroporation. Cham: Springer Inter-
national Publishing; 2017. p. 1–15, Chapter 79-1.
[193] Smith KC, Weaver JC. Transmembrane molecular transport during versus after extremely large, nanosecond electric pulses. Biochem Biophys
Res Commun Aug 19 2011;412(1):8–12. https://doi.org/10.1016/j.bbrc.2011.06.171.
[194] Sözer EB, Vernier PT. Measurement of molecular transport after electropermeabilization. In: Miklavcic D, editor. Handbook of electropora-
[195] Bennett WF, Sapay N, Tieleman DP. Atomistic simulations of pore formation and closure in lipid bilayers. Biophys J Jan 7
2014;106(1):210–9. https://doi.org/10.1016/j.bpj.2013.11.4486.
[196] Wong-Ekkabut J, Xu Z, Triampo W, Tang IM, Tieleman DP, Monticelli L. Effect of lipid peroxidation on the properties of lipid bilayers: a
molecular dynamics study. Biophys J Dec 15 2007;93(12):4225–36. https://doi.org/10.1529/biophysj.107.112565.
[197] Zhu N, et al. Design of a treatment chamber for low-voltage pulsed electric field sterilization. Innov Food Sci Emerg Technol 2017;42:180–9.
https://doi.org/10.1016/j.ifset.2017.07.016.
[198] Sweeney DC, Douglas TA, Davalos RV. Characterization of cell membrane permeability in vitro part II: computational model of
electroporation-mediated membrane transport. Technol Cancer Res Treat Jan 1 2018;17:1533033818792490. https://doi.org/10.1177/
1533033818792490.
[199] Pakhomov AG, Gianulis E, Vernier PT, Semenov I, Xiao S, Pakhomova ON. Multiple nanosecond electric pulses increase the number but not
the size of long-lived nanopores in the cell membrane. Biochim Biophys Acta Apr 2015;1848(4):958–66. https://doi.org/10.1016/j.bbamem.
2014.12.026.
[200] Gowrishankar TR, Esser AT, Vasilkoski Z, Smith KC, Weaver JC. Microdosimetry for conventional and supra-electroporation in cells with
organelles. Biochem Biophys Res Commun Mar 24 2006;341(4):1266–76. https://doi.org/10.1016/j.bbrc.2006.01.094.
[201] Hu Q, Joshi RP, Schoenbach KH. Simulations of nanopore formation and phosphatidylserine externalization in lipid membranes subjected to
a high-intensity, ultrashort electric pulse. Phys Rev E, Stat Nonlinear Soft Matter Phys Sep 2005;72(3 Pt 1):031902. https://doi.org/10.1103/
PhysRevE.72.031902.
[202] Vernier PT, Sun Y, Gundersen MA. Nanoelectropulse-driven membrane perturbation and small molecule permeabilization. BMC Cell Biol
Oct 19 2006;7(1):37. https://doi.org/10.1186/1471-2121-7-37.
[203] Vernier PT, Ziegler MJ, Sun Y, Chang WV, Gundersen MA, Tieleman DP. Nanopore formation and phosphatidylserine externalization in a
phospholipid bilayer at high transmembrane potential. J Am Chem Soc May 17 2006;128(19):6288–9. https://doi.org/10.1021/ja0588306.
[204] Escoffre JM, et al. Membrane disorder and phospholipid scrambling in electropermeabilized and viable cells. Biochim Biophys Acta Jul
2014;1838(7):1701–9. https://doi.org/10.1016/j.bbamem.2014.02.013.
[205] Vernier PT, Sun Y, Marcu L, Craft CM, Gundersen MA. Nanosecond pulsed electric fields perturb membrane phospholipids in T lym-
phoblasts. FEBS Lett Aug 13 2004;572(1–3):103–8. https://doi.org/10.1016/j.febslet.2004.07.021.
[206] Hu Q, Viswanadham S, Joshi RP, Schoenbach KH, Beebe SJ, Blackmore PF. Simulations of transient membrane behavior in cells subjected
to a high-intensity ultrashort electric pulse. Phys Rev E, Stat Nonlinear Soft Matter Phys Mar 2005;71(3 Pt 1):031914. https://doi.org/10.
1103/PhysRevE.71.031914.
[207] Semenov I, Xiao S, Kang D, Schoenbach KH, Pakhomov AG. Cell stimulation and calcium mobilization by picosecond electric pulses.
Bioelectrochemistry Oct 2015;105:65–71. https://doi.org/10.1016/j.bioelechem.2015.05.013.
[208] Pakhomov AG, Xiao S, Pakhomova ON, Semenov I, Kuipers MA, Ibey BL. Disassembly of actin structures by nanosecond pulsed electric
field is a downstream effect of cell swelling. Bioelectrochemistry Dec 2014;100:88–95. https://doi.org/10.1016/j.bioelechem.2014.01.004.
[209] de Menorval MA, et al. Electric pulses: a flexible tool to manipulate cytosolic calcium concentrations and generate spontaneous-like calcium
oscillations in mesenchymal stem cells. Sci Rep Aug 26 2016;6(1):32331. https://doi.org/10.1038/srep32331.
[210] Urabe G, Sato T, Nakamura G, Kobashigawa Y, Morioka H, Katsuki S. 1.2 MV/cm pulsed electric fields promote transthyretin aggregate
degradation. Sci Rep Jul 20 2020;10(1):12003. https://doi.org/10.1038/s41598-020-68681-0.
246
[211] Hamada Y, et al. Nanosecond pulsed electric fields induce the integrated stress response via reactive oxygen species-mediated heme-regulated
inhibitor (HRI) activation. PLoS ONE 2020;15(3):e0229948. https://doi.org/10.1371/journal.pone.0229948.
[212] Jia J, Xiong ZA, Qin Q, Yao CG, Zhao XZ. Picosecond pulsed electric fields induce apoptosis in a cervical cancer xenograft. Mol Med Rep
Mar 2015;11(3):1623–8. https://doi.org/10.3892/mmr.2014.2953.
[213] Camp JT, et al. Cell death induced by subnanosecond pulsed electric fields at elevated temperatures. IEEE Trans Plasma Sci
2012;40(10):2334–47. https://doi.org/10.1109/tps.2012.2208202.
[214] Xiao S, Guo S, Nesin VV, Heller R, Schoenbach KH. Cell membrane permeabilization and death caused by subnanosecond electric pulses.
Presented at the 2010 IEEE International Power Modulator and High Voltage Conference. https://doi.org/10.1109/ipmhvc.2010.5958327,
2010.
[215] Schoenbach KH, et al. The effect of intense subnanosecond electrical pulses on biological cells. IEEE Trans Plasma Sci 2008;36(2):414–22.
https://doi.org/10.1109/tps.2008.918786.
[216] Garon EB, et al. In vitro and in vivo evaluation and a case report of intense nanosecond pulsed electric field as a local therapy for human
malignancies. Int J Cancer Aug 1 2007;121(3):675–82. https://doi.org/10.1002/ijc.22723.
[217] Ford WE, Ren W, Blackmore PF, Schoenbach KH, Beebe SJ. Nanosecond pulsed electric fields stimulate apoptosis without release of pro-
apoptotic factors from mitochondria in B16f10 melanoma. Arch Biochem Biophys May 2010;497(1–2):82–9. https://doi.org/10.1016/j.abb.
2010.03.008.
[218] Beebe SJ. Regulated and apoptotic cell death after nanosecond electroporation. In: Miklavcic D, editor. Handbook of electroporation. Cham:
[219] Esser AT, Smith KC, Gowrishankar TR, Vasilkoski Z, Weaver JC. Mechanisms for the intracellular manipulation of organelles by conven-
tional electroporation. Biophys J Jun 2 2010;98(11):2506–14. https://doi.org/10.1016/j.bpj.2010.02.035.
[220] Grimnes S, Martinsen ØG. Dielectrics. In: Grimnes S, Martinsen ØG, editors. Bioimpedance and bioelectricity basics. Oxford: Academic
Press; 2015. p. 37–75.
[221] Ivey JW, Latouche EL, Sano MB, Rossmeisl JH, Davalos RV, Verbridge SS. Targeted cellular ablation based on the morphology of malignant
cells. Sci Rep Nov 24 2015;5(1):17157. https://doi.org/10.1038/srep17157.
[222] Lado BH. Characteristics of Listeria monocytogenes important for pulsed electric field process optimization. ProQuest Dissertations Pub-
lishing. 2003. https://www.proquest.com/dissertations-theses/characteristics-i-listeria-monocytogenes/docview/305320230/se-2?accountid=
14723.
[223] Pletnev P, Osterman I, Sergiev P, Bogdanov A, Dontsova O. Survival guide: Escherichia coli in the stationary phase. Acta Nat Oct-Dec
2015;7(4):22–33. https://doi.org/10.32607/20758251-2015-7-4-22-33.
[224] Nair S, Finkel SE. Dps protects cells against multiple stresses during stationary phase. J Bacteriol Jul 2004;186(13):4192–8. https://doi.org/
10.1128/JB.186.13.4192-4198.2004.
[225] Karas VO, Westerlaken I, Meyer AS. The DNA-binding protein from starved cells (Dps) utilizes dual functions to defend cells against
multiple stresses. J Bacteriol Oct 2015;197(19):3206–15. https://doi.org/10.1128/JB.00475-15.
[226] Hanna ES, et al. Evidence for glycosylation on a DNA-binding protein of Salmonella enterica. Microb Cell Fact Apr 2 2007;6(1):11. https://
doi.org/10.1186/1475-2859-6-11.
[227] Calhoun LN, Kwon YM. Structure, function and regulation of the DNA-binding protein Dps and its role in acid and oxidative stress resistance
in Escherichia coli: a review. J Appl Microbiol Feb 2011;110(2):375–86. https://doi.org/10.1111/j.1365-2672.2010.04890.x.
[228] Janissen R, et al. Global DNA compaction in stationary-phase bacteria does not affect transcription. Cell Aug 23 2018;174(5):1188–119914.e.
https://doi.org/10.1016/j.cell.2018.06.049.
[229] Gusbeth C, Frey W, Volkmann H, Schwartz T, Bluhm H. Pulsed electric field treatment for bacteria reduction and its impact on hospital
wastewater. Chemosphere Apr 2009;75(2):228–33. https://doi.org/10.1016/j.chemosphere.2008.11.066.
[230] Polajzer T, Miklavcic D. Development of adaptive resistance to electric pulsed field treatment in CHO cell line in vitro. Sci Rep Jun 19
2020;10(1):9988. https://doi.org/10.1038/s41598-020-66879-w.
[231] Sale AJH, Hamilton WA. Effects of high electric fields on micro-organisms. Biochim Biophys Acta, Biomembr 1968;163(1):37–43. https://
doi.org/10.1016/0005-2736(68)90030-8.
[232] Breton M, Mir LM. Investigation of the chemical mechanisms involved in the electropulsation of membranes at the molecular level. Bioelec-
trochemistry Feb 2018;119:76–83. https://doi.org/10.1016/j.bioelechem.2017.09.005.
[233] Pakhomova ON, et al. Oxidative effects of nanosecond pulsed electric field exposure in cells and cell-free media. Arch Biochem Biophys
Nov 1 2012;527(1):55–64. https://doi.org/10.1016/j.abb.2012.08.004.
[234] Gabriel B, Teissie J. Generation of reactive-oxygen species induced by electropermeabilization of Chinese hamster ovary cells and their
consequence on cell viability. Eur J Biochem Jul 1 1994;223(1):25–33. https://doi.org/10.1111/j.1432-1033.1994.tb18962.x.
[235] González-Arenzana L, et al. Impact of pulsed electric field treatment on must and wine quality. In: Miklavcic D, editor. Handbook of
[236] Teissie J. Involvement of reactive oxygen species in membrane electropermeabilization. In: Miklavcic D, editor. Handbook of electroporation.
Cham: Springer International Publishing; 2017. p. 1–15, Chapter 40-1.
[237] Vernier PT, et al. Electroporating fields target oxidatively damaged areas in the cell membrane. PLoS ONE Nov 23 2009;4(11):e7966. https://
doi.org/10.1371/journal.pone.0007966.
[238] Michel O, Pakhomov AG, Casciola M, Saczko J, Kulbacka J, Pakhomova ON. Electropermeabilization does not correlate with plasma
membrane lipid oxidation. Bioelectrochemistry Apr 2020;132:107433. https://doi.org/10.1016/j.bioelechem.2019.107433.
[239] Bo W, et al. Probing nanoelectroporation and resealing of the cell membrane by the entry of Ca(2+) and Ba(2+) ions. Int J Mol Sci May 11
2020;21(9):3386. https://doi.org/10.3390/ijms21093386.
[240] Rems L. Lipid pores: molecular and continuum models. In: Miklavcic D, editor. Handbook of electroporation. Cham: Springer International
Publishing; 2016. p. 1–21, Chapter 76-1.
247
[241] Weaver JC, Vernier PT. Pore lifetimes in cell electroporation: complex dark pores? https://doi.org/10.48550/arXiv.1708.07478, 2017.
[242] Ribble D, Goldstein NB, Norris DA, Shellman YG. A simple technique for quantifying apoptosis in 96-well plates. BMC Biotechnol May
10 2005;5(1):12. https://doi.org/10.1186/1472-6750-5-12.
[243] García-Gonzalo D, Pagán R. Detection of electroporation in microbial cells: techniques and procedures. In: Miklavcic D, editor. Handbook
[244] Gianulis EC, Pakhomov AG. Experimental determination of lipid electropore size. In: Miklavcic D, editor. Handbook of electroporation.
[245] Napotnik TB. Fluorescent indicators of membrane permeabilization due to electroporation. In: Miklavcic D, editor. Handbook of electropo-
[246] Shi L, Gunther S, Hubschmann T, Wick LY, Harms H, Muller S. Limits of propidium iodide as a cell viability indicator for environmental
bacteria. Cytometry, Part A Aug 2007;71(8):592–8. https://doi.org/10.1002/cyto.a.20402.
[247] Novickij V, et al. Bioluminescence as a sensitive electroporation indicator in sub-microsecond and microsecond range of electrical pulses. J
Photochem Photobiol B Dec 2020;213:112066. https://doi.org/10.1016/j.jphotobiol.2020.112066.
[248] Mutoh H, Akemann W, Knopfel T. Genetically engineered fluorescent voltage reporters. ACS Chem Neurosci Aug 15 2012;3(8):585–92.
https://doi.org/10.1021/cn300041b.
[249] Pakhomov AG, Semenov I, Casciola M, Xiao S. Neuronal excitation and permeabilization by 200-ns pulsed electric field: an optical mem-
brane potential study with FluoVolt dye. Biochim Biophys Acta, Biomembr Jul 2017;1859(7):1273–81. https://doi.org/10.1016/j.bbamem.
2017.04.016.
[250] Silve A, Leray I, Leguebe M, Poignard C, Mir LM. Cell membrane permeabilization by 12-ns electric pulses: not a purely dielectric, but a
charge-dependent phenomenon. Bioelectrochemistry, vol. 106. Pt B; Dec 2015. p. 369–78.
[251] Deal PE, Grenier V, Kulkarni RU, Liu P, Walker AS, Miller EW. Making life visible: fluorescent indicators to probe membrane potential. In:
Toyama Y, Miyawaki A, Nakamura M, Jinzaki M, editors. Make life visible. Singapore: Springer Singapore; 2020. p. 89–104.
[252] Kulkarni RU, Miller EW. Voltage imaging: pitfalls and potential. Biochemistry Oct 3 2017;56(39):5171–7. https://doi.org/10.1021/acs.
biochem.7b00490.
[253] Panzera LC, Hoppa MB. Genetically encoded voltage indicators are illuminating subcellular physiology of the axon. Front Cell Neurosci
2019;13:52. https://doi.org/10.3389/fncel.2019.00052.
[254] Rhee JK, et al. Biophysical parameters of GEVIs: considerations for imaging voltage. Biophys J Jul 7 2020;119(1):1–8. https://doi.org/10.
1016/j.bpj.2020.05.019.
[255] Xu Y, Zou P, Cohen AE. Voltage imaging with genetically encoded indicators. Curr Opin Chem Biol Aug 2017;39:1–10. https://doi.org/10.
1016/j.cbpa.2017.04.005.
[256] Benarroch JM, Asally M. The microbiologist’s guide to membrane potential dynamics. Trends Microbiol Apr 2020;28(4):304–14. https://
doi.org/10.1016/j.tim.2019.12.008.
[257] Lv Y, et al. Analysis of the electric field-dependent current during electroporation pulses. IEEE Access 2020;8:93850–6. https://doi.org/10.
1109/access.2020.2995151.
[258] Cox CD, et al. Removal of the mechanoprotective influence of the cytoskeleton reveals PIEZO1 is gated by bilayer tension. Nat Commun
Jan 20 2016;7(1):10366. https://doi.org/10.1038/ncomms10366.
[259] Ridone P, Vassalli M, Martinac B. Piezo1 mechanosensitive channels: what are they and why are they important. Biophys Rev Oct
2019;11(5):795–805. https://doi.org/10.1007/s12551-019-00584-5.
[260] Zhao Q, et al. Structure and mechanogating mechanism of the Piezo1 channel. Nature Feb 22 2018;554(7693):487–92. https://doi.org/10.
1038/nature25743.
[261] Coste B, et al. Piezo1 and Piezo2 are essential components of distinct mechanically activated cation channels. Science Oct 1
2010;330(6000):55–60. https://doi.org/10.1126/science.1193270.
[262] Chang G, Spencer RH, Lee AT, Barclay MT, Rees DC. Structure of the MscL homolog from Mycobacterium tuberculosis: a gated
mechanosensitive ion channel. Science Dec 18 1998;282(5397):2220–6. https://doi.org/10.1126/science.282.5397.2220.
[263] Zerbib D. Bacterial cell envelopes: composition, architecture, and origin. In: Miklavcic D, editor. Handbook of electroporation. Cham:
[264] Cantu JC, Tarango M, Beier HT, Ibey BL. The biological response of cells to nanosecond pulsed electric fields is dependent on plasma
membrane cholesterol. Biochim Biophys Acta Nov 2016;1858(11):2636–46. https://doi.org/10.1016/j.bbamem.2016.07.006.
[265] Forrest M. Can the thermodynamic Hodgkin-Huxley model of voltage-dependent conductance extrapolate for temperature? Computation
2014;2(2):47–60. https://doi.org/10.3390/computation2020047.
[266] Eisenberg RS. Maxwell equations without a polarization field, using a paradigm from biophysics. Entropy Jan 30 2021;23(2):172. https://
doi.org/10.3390/e23020172.
[267] Français O, Le Pioufle B. Single cell electrical characterization techniques. In: Miklavcic D, editor. Handbook of electroporation. Cham:
[268] Liu JL, Eisenberg B. Molecular mean-field theory of ionic solutions: a Poisson-Nernst-Planck-Bikerman model. Entropy May 14
2020;22(5):550. https://doi.org/10.3390/e22050550.
[269] Garner AL, Deminsky M, Bogdan Neculaes V, Chashihin V, Knizhnik A, Potapkin B. Cell membrane thermal gradients induced by electro-
magnetic fields. J Appl Phys 2013;113(21):214701. https://doi.org/10.1063/1.4809642.
[270] Montanari C, et al. Heat-assisted pulsed electric field treatment for the inactivation of Saccharomyces cerevisiae: effects of the presence of
citral. Front Microbiol 2019;10:1737. https://doi.org/10.3389/fmicb.2019.01737.
[271] Das S, Jaeger M, Leonetti M, Thaokar RM, Chen PG. Effect of pulse width on the dynamics of a deflated vesicle in unipolar and bipolar
pulsed electric fields. Phys Fluids 2021;33(8):081905. https://doi.org/10.1063/5.0057168.
248
[272] Dastani K, et al. Effect of input voltage frequency on the distribution of electrical stresses on the cell surface based on single-cell dielec-
trophoresis analysis. Sci Rep Jan 9 2020;10(1):68. https://doi.org/10.1038/s41598-019-56952-4.
[273] Nesin OM, Pakhomova ON, Xiao S, Pakhomov AG. Manipulation of cell volume and membrane pore comparison following single cell
permeabilization with 60- and 600-ns electric pulses. Biochim Biophys Acta Mar 2011;1808(3):792–801. https://doi.org/10.1016/j.bbamem.
2010.12.012.
[274] Romeo S, Wu YH, Levine ZA, Gundersen MA, Vernier PT. Water influx and cell swelling after nanosecond electropermeabilization. Biochim
Biophys Acta Aug 2013;1828(8):1715–22. https://doi.org/10.1016/j.bbamem.2013.03.007.
[275] Pakhomov AG, Ibey BL, Bowman AM, Andre FM, Pakhomova ON. World congress on medical physics and biomedical engineering,
September 7 - 12, 2009, Munich, Germany, vol. 25. Berlin, Heidelberg: Springer Berlin Heidelberg; 2009. p. 17–20.
[276] Sun Y, Xiao S, White J, Kolb J, Stacey M, Schoenbach K. Compact, nanosecond, high repetition rate, pulse generator for bioelectric studies.
IEEE Trans Dielectr Electr Insul 2007;14(4):863–70. https://doi.org/10.1109/tdei.2007.4286517.
[277] Sanders JM, Kuthi A, Vernier PT, Wu Y, Jiang C, Gundersen MA. Scalable, compact, nanosecond pulse generator with a high repetition rate
for biomedical applications requiring intense electric fields. In: 2009 IEEE pulsed power conference; 2009. p. 1418–21.
[278] Vernier PT, Sun Y, Marcu L, Craft CM, Gundersen MA. Nanoelectropulse-induced phosphatidylserine translocation. Biophys J Jun
2004;86(6):4040–8. https://doi.org/10.1529/biophysj.103.037945.
[279] Ibey BL, et al. Selective cytotoxicity of intense nanosecond-duration electric pulses in mammalian cells. Biochim Biophys Acta Nov
2010;1800(11):1210–9. https://doi.org/10.1016/j.bbagen.2010.07.008.
[280] Pakhomova ON, Gregory BW, Khorokhorina VA, Bowman AM, Xiao S, Pakhomov AG. Electroporation-induced electrosensitization. PLoS
ONE Feb 9 2011;6(2):e17100. https://doi.org/10.1371/journal.pone.0017100.
[281] Mitsutake K, Satoh A, Mine S, Abe K, Katsuki S, Akiyama H. In: 2010 IEEE international power modulator and high voltage conference.
IEEE; 2010. p. 204–7.
[282] Mitsutake K, Satoh A, Mine S, Abe K, Katsuki S, Akiyama H. Effect of pulsing sequence of nanosecond pulsed electric fields on viability
of HeLa S3 cells. IEEE Trans Dielectr Electr Insul 2012;19(1):337–42. https://doi.org/10.1109/tdei.2012.6148536.
[283] Wijayanti HB, Bansal N, Deeth HC. Stability of whey proteins during thermal processing: a review. Compr Rev Food Sci Food Saf
2014;13(6):1235–51. https://doi.org/10.1111/1541-4337.12105.
[284] Brick T, et al. Effect of processing intensity on immunologically active bovine milk serum proteins. Nutrients Aug 31 2017;9(9):1–14. https://
doi.org/10.3390/nu9090963.
[285] Zhao W, Yang R, Liang Q, Zhang W, Hua X, Tang Y. Electrochemical reaction and oxidation of lecithin under pulsed electric fields (PEF)
processing. J Agric Food Chem Dec 12 2012;60(49):12204–9. https://doi.org/10.1021/jf304236h.
[286] Dan G, et al. Synergetic effects of pulsed electric field and ozone treatments on the degradation of high molecular weight chitosan. Int J Food
Eng 2014;10(4):775–84. https://doi.org/10.1515/ijfe-2014-0100.
[287] Dutreux N, Notermans S, Wijtzes T, Góngora-Nieto MM, Barbosa-Cánovas GV, Swanson BG. Pulsed electric fields inactivation of attached
and free-living Escherichia coli and Listeria innocua under several conditions. Int J Food Microbiol 2000;54(1–2):91–8. https://doi.org/10.
1016/s0168-1605(99)00175-0.
[288] Pol IE, Mastwujk HC, Slump RA, Popa ME, Smid EJ. Influence of food matrix on inactivation of Bacillus cereus by combinations of nisin,
pulsed electric field treatment, and carvacrol. J Food Prot Jul 2001;64(7):1012–8. https://doi.org/10.4315/0362-028x-64.7.1012.
[289] Terebiznik M, Jagus R, Cerrutti P, de Huergo MS, Pilosof AM. Inactivation of Escherichia coli by a combination of nisin, pulsed electric
fields, and water activity reduction by sodium chloride. J Food Prot Aug 2002;65(8):1253–8. https://doi.org/10.4315/0362-028x-65.8.1253.
[290] Ravishankar S. The inactivation of Escherichia coli O157:H7 during pulsed electric field (PEF) treatment in a static chamber. Food Microbiol
2002;19(4):351–61. https://doi.org/10.1006/fmic.2002.0484.
[291] Khadre MA, Yousef AE. Susceptibility of human rotavirus to ozone, high pressure, and pulsed electric field. J Food Prot Sep
2002;65(9):1441–6. https://doi.org/10.4315/0362-028x-65.9.1441.
[292] Rodrigo D, Ruiz P, Barbosa-Canovas GV, Martinez A, Rodrigo M. Kinetic model for the inactivation of Lactobacillus plantarum by pulsed
electric fields. Int J Food Microbiol Mar 25 2003;81(3):223–9. https://doi.org/10.1016/s0168-1605(02)00247-7.
[293] Fleischman GJ, Ravishankar S, Balasubramaniam VM. The inactivation of Listeria monocytogenes by pulsed electric field (PEF) treatment
in a static chamber. Food Microbiol 2004;21(1):91–5. https://doi.org/10.1016/s0740-0020(03)00015-7.
[294] Aronsson K, Borch E, Stenlof B, Ronner U. Growth of pulsed electric field exposed Escherichia coli in relation to inactivation and environ-
mental factors. Int J Food Microbiol May 15 2004;93(1):1–10. https://doi.org/10.1016/S0168-1605(03)00071-0.
[295] Reyns KM, Diels AM, Michiels CW. Generation of bactericidal and mutagenic components by pulsed electric field treatment. Int J Food
Microbiol Jun 1 2004;93(2):165–73. https://doi.org/10.1016/j.ijfoodmicro.2003.10.014.
[296] Rojas MC, Martin SE, Wicklund RA, Paulson DD, Desantos FA, Brewer MS. Effect of high-intensity pulsed electric fields on survival
of Escherichia coli K-12 suspended in meat injection solutions. J Food Saf 2007;27(4):411–25. https://doi.org/10.1111/j.1745-4565.2007.
00086.x.
[297] Pérez MCP, Aliaga DR, Bernat CF, Enguidanos MR, López AM. Inactivation of Enterobacter sakazakii by pulsed electric field in buffered
peptone water and infant formula milk. Int Dairy J 2007;17(12):1441–9. https://doi.org/10.1016/j.idairyj.2007.04.007.
[298] Somolinos M, Garcia D, Manas P, Condon S, Pagan R. Effect of environmental factors and cell physiological state on pulsed electric fields
resistance and repair capacity of various strains of Escherichia coli. Int J Food Microbiol Jun 10 2008;124(3):260–7. https://doi.org/10.1016/
j.ijfoodmicro.2008.03.021.
[299] Somolinos M, Manas P, Condon S, Pagan R, Garcia D. Recovery of Saccharomyces cerevisiae sublethally injured cells after pulsed electric
fields. Int J Food Microbiol Jul 31 2008;125(3):352–6. https://doi.org/10.1016/j.ijfoodmicro.2008.04.023.
[300] Arroyo C, Cebrián G, Pagán R, Condón S. Resistance of Enterobacter sakazakii to pulsed electric fields. Innov Food Sci Emerg Technol
2010;11(2):314–21. https://doi.org/10.1016/j.ifset.2009.11.002.
249
[301] Shin D, Martin BC, Sánchez-Plata MX. Pulsed electric field effects to reduce the level of Campylobacter spp. in scalder and chiller water
during broiler chicken processing. Asian-Australas J Anim Sci 2011;24(9):1314–7. https://doi.org/10.5713/ajas.2011.11075.
[302] Haughton PN, Lyng JG, Cronin DA, Morgan DJ, Fanning S, Whyte P. Efficacy of pulsed electric fields for the inactivation of indicator
microorganisms and foodborne pathogens in liquids and raw chicken. Food Control 2012;25(1):131–5. https://doi.org/10.1016/j.foodcont.
2011.10.030.
[303] Novickij V, Grainys A, Svediene J, Paskevicius A, Novickij J. Controlled inactivation of Trichophyton rubrum using shaped electrical pulse
bursts: parametric analysis. Biotechnol Prog Jul 8 2016;32(4):1056–60. https://doi.org/10.1002/btpr.2276.
[304] Cregenzán-Alberti O, Arroyo C, Dorozko A, Whyte P, Lyng JG. Thermal characterization of Bacillus subtilis endospores and a comparative
study of their resistance to high temperature pulsed electric fields (HTPEF) and thermal-only treatments. Food Control 2017;73:1490–8.
https://doi.org/10.1016/j.foodcont.2016.11.012.
[305] Masood H, Razaeimotlagh A, Cullen PJ, Trujillo FJ. Numerical and experimental studies on a novel Steinmetz treatment chamber for
inactivation of Escherichia coli by radio frequency electric fields. Innov Food Sci Emerg Technol 2017;41:337–47. https://doi.org/10.1016/j.
ifset.2017.04.009.
[306] Krishnaveni S, Subhashini R, Rajini V. Inactivation of bacteria suspended in water by using high frequency unipolar pulse voltage. J Food
Process Eng 2017;40(6):e12574. https://doi.org/10.1111/jfpe.12574.
[307] Pyatkovskyy TI, Shynkaryk MV, Mohamed HM, Yousef AE, Sastry SK. Effects of combined high pressure (HPP), pulsed electric field (PEF)
and sonication treatments on inactivation of Listeria innocua. J Food Eng 2018;233:49–56. https://doi.org/10.1016/j.jfoodeng.2018.04.002.
[308] Wang L-H, Pyatkovskyy T, Yousef A, Zeng X-A, Sastry SK. Mechanism of Bacillus subtilis spore inactivation induced by moderate electric
fields. Innov Food Sci Emerg Technol 2020;62:102349. https://doi.org/10.1016/j.ifset.2020.102349.
[309] Saldaña G, Puértolas E, Álvarez I, Meneses N, Knorr D, Raso J. Evaluation of a static treatment chamber to investigate kinetics of microbial
inactivation by pulsed electric fields at different temperatures at quasi-isothermal conditions. J Food Eng 2010;100(2):349–56. https://doi.
org/10.1016/j.jfoodeng.2010.04.021.
[310] Gomez-Gomez A, Brito-de la Fuente E, Gallegos C, Garcia-Perez JV, Benedito J. Combined pulsed electric field and high-power ultrasound
treatments for microbial inactivation in oil-in-water emulsions. Food Control 2021;130:108348. https://doi.org/10.1016/j.foodcont.2021.
108348.
[311] Furukawa T, Ueno T, Matsumura M, Amarasiri M, Sei K. Inactivation of antibiotic resistant bacteria and their resistance genes in sewage by
applying pulsed electric fields. J Hazard Mater Feb 15 2022;424(Pt A):127382. https://doi.org/10.1016/j.jhazmat.2021.127382.
[312] Zhang Y, et al. Kinetic analysis of the degradation and its color change of cyanidin-3-glucoside exposed to pulsed electric field. Eur Food
Res Technol 2006;224(5):597–603. https://doi.org/10.1007/s00217-006-0343-8.
[313] Zhang Y, Hu XS, Chen F, Wu JH, Liao XJ, Wang ZF. Stability and colour characteristics of PEF-treated cyanidin-3-glucoside during storage.
Food Chem 2008;106(2):669–76. https://doi.org/10.1016/j.foodchem.2007.06.030.
[314] Garde-Cerdán T, Marsellés-Fontanet AR, Arias-Gil M, Ancín-Azpilicueta C, Martín-Belloso O. Influence of SO2 on the evolution of volatile
compounds through alcoholic fermentation of must stabilized by pulsed electric fields. Eur Food Res Technol 2007;227(2):401–8. https://
doi.org/10.1007/s00217-007-0734-5.
[315] Garde-Cerdán T, Marsellés-Fontanet AR, Arias-Gil M, Ancín-Azpilicueta C, Martín-Belloso O. Effect of storage conditions on the volatile
composition of wines obtained from must stabilized by PEF during ageing without SO2. Innov Food Sci Emerg Technol 2008;9(4):469–76.
[316] Sun J, Bai W, Zhang Y, Liao X, Hu X. Effects of electrode materials on the degradation, spectral characteristics, visual colour, and antioxidant
capacity of cyanidin-3-glucoside and cyanidin-3-sophoroside during pulsed electric field (PEF) treatment. Food Chem 2011;128(3):742–7.
https://doi.org/10.1016/j.foodchem.2011.03.099.
[317] Zhao D, Zeng X-A, Sun D-W, Liu D. Effects of pulsed electric field treatment on (+)-catechin–acetaldehyde condensation. Innov Food Sci
Emerg Technol 2013;20:100–5. https://doi.org/10.1016/j.ifset.2013.07.007.
[318] Wang X-Q, Su H-N, Zhang Q-H, Yang P-P. The effects of pulsed electric fields applied to red and white wines during bottle ageing on
organic acid contents. J Food Sci Technol 2013;52(1):171–80. https://doi.org/10.1007/s13197-013-0984-0.
[319] Lyu C, Huang K, Yang N, Wang H, Wang J. Combination of thermosonication and pulsed electric fields treatments for controlling Saccha-
romyces cerevisiae in Chinese rice wine. Food Bioprocess Technol 2016;9(11):1854–64. https://doi.org/10.1007/s11947-016-1769-z.
[320] El Darra N, et al. Effect of pulsed electric field treatment during cold maceration and alcoholic fermentation on major red wine qualitative
and quantitative parameters. Food Chem Dec 15 2016;213:352–60. https://doi.org/10.1016/j.foodchem.2016.06.073.
[321] Martinez JM, Delso C, Aguilar D, Cebrian G, Alvarez I, Raso J. Factors influencing autolysis of Saccharomyces cerevisiae cells induced by
pulsed electric fields. Food Microbiol Aug 2018;73:67–72. https://doi.org/10.1016/j.fm.2017.12.008.
[322] Comuzzo P, Marconi M, Zanella G, Querze M. Pulsed electric field processing of white grapes (cv. Garganega): effects on wine composition
and volatile compounds. Food Chem Oct 30 2018;264:16–23. https://doi.org/10.1016/j.foodchem.2018.04.116.
[323] Martinez JM, Delso C, Maza MA, Alvarez I, Raso J. Pulsed electric fields accelerate release of mannoproteins from Saccharomyces cerevisiae
during aging on the lees of Chardonnay wine. Food Res Int Feb 2019;116:795–801. https://doi.org/10.1016/j.foodres.2018.09.013.
[324] Maza MA, Delso C, Álvarez I, Raso J, Martínez JM. Effect of pulsed electric fields on mannoproteins release from Saccharomyces cerevisiae
during the aging on lees of Caladoc red wine. Lebensm-Wiss Technol 2020;118:108788. https://doi.org/10.1016/j.lwt.2019.108788.
[325] Puertolas E, Lopez N, Condon S, Raso J, Alvarez I. Pulsed electric fields inactivation of wine spoilage yeast and bacteria. Int J Food Microbiol
Mar 15 2009;130(1):49–55. https://doi.org/10.1016/j.ijfoodmicro.2008.12.035.
[326] Marselles-Fontanet AR, Puig A, Olmos P, Minguez-Sanz S, Martin-Belloso O. Optimising the inactivation of grape juice spoilage organisms
by pulse electric fields. Int J Food Microbiol Apr 15 2009;130(3):159–65. https://doi.org/10.1016/j.ijfoodmicro.2008.12.034.
[327] Huang K, Yu L, Liu D, Gai L, Wang J. Modeling of yeast inactivation of PEF-treated Chinese rice wine: effects of electric field intensity,
treatment time and initial temperature. Food Res Int 2013;54(1):456–67. https://doi.org/10.1016/j.foodres.2013.07.046.
250
[328] González-Arenzana L, et al. Inactivation of wine-associated microbiota by continuous pulsed electric field treatments. Innov Food Sci Emerg
Technol 2015;29:187–92. https://doi.org/10.1016/j.ifset.2015.03.009.
[329] Delsart C, et al. Comparison of the effect of pulsed electric field or high voltage electrical discharge for the control of sweet white must fer-
mentation process with the conventional addition of sulfur dioxide. Food Res Int 2015;77(Part 4):718–24. https://doi.org/10.1016/j.foodres.
2015.04.017.
[330] Abca EE, Akdemir Evrendilek G. Processing of red wine by pulsed electric fields with respect to quality parameters. J Food Process Preserv
2015;39(6):758–67. https://doi.org/10.1111/jfpp.12285.
[331] González-Arenzana L, López-Alfaro I, Gutiérrez AR, López N, Santamaría P, López R. Continuous pulsed electric field treatments’ impact
on the microbiota of red Tempranillo wines aged in oak barrels. Food Biosci 2019;27:54–9. https://doi.org/10.1016/j.fbio.2018.10.012.
[332] van Wyk S, Silva FVM, Farid MM. Pulsed electric field treatment of red wine: inactivation of Brettanomyces and potential hazard caused by
metal ion dissolution. Innov Food Sci Emerg Technol 2019;52:57–65. https://doi.org/10.1016/j.ifset.2018.11.001.
[333] Niu D, et al. Effect of ethanol adaption on the inactivation of Acetobacter sp. by pulsed electric fields. Innov Food Sci Emerg Technol
2019;52:25–33. https://doi.org/10.1016/j.ifset.2018.11.009.
[334] Puértolas E, López N, Condón S, Álvarez I, Raso J. Potential applications of PEF to improve red wine quality. Trends Food Sci Technol
2010;21(5):247–55. https://doi.org/10.1016/j.tifs.2010.02.002.
[335] Shynkaryk MV, Lebovka NI, Lanoisellé JL, Nonus M, Bedel-Clotour C, Vorobiev E. Electrically-assisted extraction of bio-products using
high pressure disruption of yeast cells (Saccharomyces cerevisiae). J Food Eng 2009;92(2):189–95. https://doi.org/10.1016/j.jfoodeng.2008.
10.041.
[336] Liu D, Lebovka NI, Vorobiev E. Impact of electric pulse treatment on selective extraction of intracellular compounds from Saccharomyces
cerevisiae yeasts. Food Bioprocess Technol 2011;6(2):576–84. https://doi.org/10.1007/s11947-011-0703-7.
[337] Marsellés-Fontanet ÁR, Puig-Pujol A, Olmos P, Mínguez-Sanz S, Martín-Belloso O. A comparison of the effects of pulsed electric field and
thermal treatments on grape juice. Food Bioprocess Technol 2011;6(4):978–87. https://doi.org/10.1007/s11947-011-0731-3.
[338] Gonzalez-Arenzana L, Lopez-Alfaro I, Garde-Cerdan T, Portu J, Lopez R, Santamaria P. Microbial inactivation and MLF performances of
Tempranillo Rioja wines treated with PEF after alcoholic fermentation. Int J Food Microbiol Mar 23 2018;269:19–26. https://doi.org/10.
1016/j.ijfoodmicro.2018.01.008.
[339] van Wyk S, Farid MM, Silva FVM. SO2, high pressure processing and pulsed electric field treatments of red wine: effect on sensory,
Brettanomyces inactivation and other quality parameters during one year storage. Innov Food Sci Emerg Technol 2018;48:204–11. https://
doi.org/10.1016/j.ifset.2018.06.016.
[340] González-Arenzana L, et al. Pulsed electric field treatment after malolactic fermentation of Tempranillo Rioja wines: influence on microbial,
physicochemical and sensorial quality. Innov Food Sci Emerg Technol 2019;51:57–63. https://doi.org/10.1016/j.ifset.2018.05.019.
[341] van Wyk S, Hong L, Silva FVM. Non-thermal high pressure processing, pulsed electric fields and ultrasound preservation of five different
table wines. Beverages 2021;7(4):69. https://doi.org/10.3390/beverages7040069.
[342] Shen C, et al. The sensory and flavor characteristics of Shaoxing Huangjiu (Chinese rice wine) were significantly influenced by micro-oxygen
and electric field. Food Sci Nutr Nov 2021;9(11):6006–19. https://doi.org/10.1002/fsn3.2531.
[343] Xiang BY, Ngadi MO, Ochoa-Martinez LA, Simpson MV. Pulsed electric field-induced structural modification of whey protein isolate. Food
Bioprocess Technol 2009;4(8):1341–8. https://doi.org/10.1007/s11947-009-0266-z.
[344] Martín O, Qin BL, Chang FJ, Barbosa-Cánovas GV, Swanson BG. Inactivation of Escherichia coli in skim milk by high intensity pulsed
electric fields. J Food Process Eng 1997;20(4):317–36. https://doi.org/10.1111/j.1745-4530.1997.tb00425.x.
[345] Sensoy I, Zhang QH, Sastry SK. Inactivation kinetics of Salmonella dublin by pulsed electric field. J Food Process Eng 1997;20(5):367–81.
[346] Reina LD, Jin ZT, Zhang QH, Yousef AE. Inactivation of Listeria monocytogenes in milk by pulsed electric field. J Food Prot Sep
1998;61(9):1203–6. https://doi.org/10.4315/0362-028x-61.9.1203.
[347] Calderon-Miranda ML, Barbosa-Canovas GV, Swanson BG. Transmission electron microscopy of Listeria innocua treated by pulsed electric
fields and nisin in skimmed milk. Int J Food Microbiol Oct 1 1999;51(1):31–8. https://doi.org/10.1016/s0168-1605(99)00071-9.
[348] Calderon-Miranda ML, Barbosa-Canovas GV, Swanson BG. Inactivation of Listeria innocua in skim milk by pulsed electric fields and nisin.
Int J Food Microbiol Oct 1 1999;51(1):19–30. https://doi.org/10.1016/s0168-1605(99)00069-0.
[349] Terebiznik MR, Jagus RJ, Cerrutti P, de Huergo MS, Pilosof AM. Combined effect of nisin and pulsed electric fields on the inactivation of
Escherichia coli. J Food Prot Jun 2000;63(6):741–6. https://doi.org/10.4315/0362-028x-63.6.741.
[350] Wouters PC, Alvarez I, Raso J. Critical factors determining inactivation kinetics by pulsed electric field food processing. Trends Food Sci
Technol 2001;12(3–4):112–21. https://doi.org/10.1016/s0924-2244(01)00067-x.
[351] Smith K, Mittal GS, Griffiths MW. Pasteurization of milk using pulsed electrical field and antimicrobials. J Food Sci 2002;67(6):2304–8.
[352] Alvarez I, Pagan R, Condon S, Raso J. The influence of process parameters for the inactivation of Listeria monocytogenes by pulsed electric
fields. Int J Food Microbiol Oct 15 2003;87(1–2):87–95. https://doi.org/10.1016/s0168-1605(03)00056-4.
[353] Evrendilek GA, Zhang QH, Richter ER. Application of pulsed electric fields to skim milk inoculated with Staphylococcus aureus. Biosyst
Eng 2004;87(2):137–44. https://doi.org/10.1016/j.biosystemseng.2003.11.005.
[354] Li SQ, Zhang QH. Inactivation of E. coli 8739 in enriched soymilk using pulsed electric fields. J Food Sci 2006;69(7):169–74. https://
doi.org/10.1111/j.1365-2621.2004.tb13616.x.
[355] Sepulveda DR, Góngora-Nieto MM, San-Martin MF, Barbosa-Cánovas GV. Influence of treatment temperature on the inactivation of Listeria
innocua by pulsed electric fields. Lebensm-Wiss Technol 2005;38(2):167–72. https://doi.org/10.1016/j.lwt.2004.05.011.
[356] Sepulveda DR, Góngora-Nieto MM, Guerrero JA, Barbosa-Cánovas GV. Production of extended-shelf life milk by processing pasteurized
milk with pulsed electric fields. J Food Eng 2005;67(1–2):81–6. https://doi.org/10.1016/j.jfoodeng.2004.05.056.
251
[357] Evrendilek GA, Zhang QH. Effects of pulse polarity and pulse delaying time on pulsed electric fields-induced pasteurization of E. coli
O157:H7. J Food Eng 2005;68(2):271–6. https://doi.org/10.1016/j.jfoodeng.2004.06.001.
[358] Fernandez-Molina JJ, Altunakar B, Bermudez-Aguirre D, Swanson BG, Barbosa-Canovas GV. Inactivation of Pseudomonas fluorescens in
skim milk by combinations of pulsed electric fields and organic acids. J Food Prot Jun 2005;68(6):1232–5. https://doi.org/10.4315/0362-
028x-68.6.1232.
[359] Fernandez-Molina JJ, Fernandez-Gutierrez SA, Altunakar B, Bermudez-Aguirre D, Swanson BG, Barbosa-Canovas GV. The combined
effect of pulsed electric fields and conventional heating on the microbial quality and shelf life of skim milk. J Food Process Preserv
2005;29(5–6):390–406. https://doi.org/10.1111/j.1745-4549.2005.00036.x.
[360] Fernandez-Molina JJ, Barbosa-Canovas GV, Swanson BG. Skim milk processing by combining pulsed electric fields and thermal treatments.
J Food Process Preserv 2005;29(5–6):291–306. https://doi.org/10.1111/j.1745-4549.2005.00029.x.
[361] Floury J, Grosset N, Lesne E, Jeantet R. Continuous processing of skim milk by a combination of pulsed electric fields and conventional heat
treatments: does a synergetic effect on microbial inactivation exist? Le Lait 2006;86(3):203–11. https://doi.org/10.1051/lait:2006002.
[362] Sampedro F, Rivas A, Rodrigo D, Martínez A, Rodrigo M. Effect of temperature and substrate on Pef inactivation of Lactobacillus plantarum
in an orange juice–milk beverage. Eur Food Res Technol 2006;223(1):30–4. https://doi.org/10.1007/s00217-005-0096-9.
[363] Rivas A, Sampedro F, Rodrigo D, Martínez A, Rodrigo M. Nature of the inactivation of Escherichia coli suspended in an orange juice and
milk beverage. Eur Food Res Technol 2006;223(4):541–5. https://doi.org/10.1007/s00217-005-0234-4.
[364] Sampedro F, Rivas A, Rodrigo D, Martínez A, Rodrigo M. Pulsed electric fields inactivation of Lactobacillus plantarum in an orange juice–
milk based beverage: effect of process parameters. J Food Eng 2007;80(3):931–8. https://doi.org/10.1016/j.jfoodeng.2006.08.013.
[365] Alkhafaji SR, Farid M. An investigation on pulsed electric fields technology using new treatment chamber design. Innov Food Sci Emerg
Technol 2007;8(2):205–12. https://doi.org/10.1016/j.ifset.2006.11.001.
[366] Shin J-K, Jung K-J, Pyun Y-R, Chun M-S. Application of pulsed electric fields with square wave pulse to milk inoculated with E. coli,
P. fluorescens, and B. stearothermophilus. Food Sci Biotechnol 2007;16(6):1082–4 [Online]. Available: https://www.earticle.net/Article/
A79297.
[367] Otunola A, El-Hag A, Jayaram S, Anderson WA. Effectiveness of pulsed electric fields in controlling microbial growth in milk. Int J Food
Eng 2008;4(7):1. https://doi.org/10.2202/1556-3758.1494.
[368] Craven HM, et al. Evaluation of pulsed electric field and minimal heat treatments for inactivation of pseudomonads and enhancement of milk
shelf-life. Innov Food Sci Emerg Technol 2008;9(2):211–6. https://doi.org/10.1016/j.ifset.2007.11.002.
[369] Walkling-Ribeiro M, Noci F, Cronin DA, Lyng JG, Morgan DJ. Inactivation of Escherichia coli in a tropical fruit smoothie by a combination
of heat and pulsed electric fields. J Food Sci Oct 2008;73(8):M395–9. https://doi.org/10.1111/j.1750-3841.2008.00927.x.
[370] Alkhafaji S, Farid M. Modelling the inactivation of Escherichia coli ATCC 25922 using pulsed electric field. Innov Food Sci Emerg Technol
2008;9(4):448–54. https://doi.org/10.1016/j.ifset.2008.02.003.
[371] Noci F, Walkling-Ribeiro M, Cronin DA, Morgan DJ, Lyng JG. Effect of thermosonication, pulsed electric field and their combination on
inactivation of Listeria innocua in milk. Int Dairy J 2009;19(1):30–5. https://doi.org/10.1016/j.idairyj.2008.07.002.
[372] Sampedro F, Geveke DJ, Fan X, Rodrigo D, Zhang QH. Shelf-life study of an orange juice-milk based beverage after PEF and thermal
processing. J Food Sci Mar 2009;74(2):S107–12. https://doi.org/10.1111/j.1750-3841.2009.01068.x.
[373] Sepulveda DR, Góngora-Nieto MM, Guerrero JA, Barbosa-Cánovas GV. Shelf life of whole milk processed by pulsed electric fields in
combination with PEF-generated heat. Lebensm-Wiss Technol 2009;42(3):735–9. https://doi.org/10.1016/j.lwt.2008.10.005.
[374] Pina-Perez MC, Silva-Angulo AB, Rodrigo D, Martinez-Lopez A. Synergistic effect of pulsed electric fields and CocoanOX 12% on the inac-
tivation kinetics of Bacillus cereus in a mixed beverage of liquid whole egg and skim milk. Int J Food Microbiol Apr 15 2009;130(3):196–204.
https://doi.org/10.1016/j.ijfoodmicro.2009.01.021.
[375] Pina-Pérez MC, Rodrigo D, López AM. Sub-lethal damage in Cronobacter sakazakii subsp. sakazakii cells after different pulsed electric
field treatments in infant formula milk. Food Control 2009;20(12):1145–50. https://doi.org/10.1016/j.foodcont.2009.03.006.
[376] Guerrero-Beltrán JÁ, Sepulveda DR, Góngora-Nieto MM, Swanson B, Barbosa-Cánovas GV. Milk thermization by pulsed electric fields
(PEF) and electrically induced heat. J Food Eng 2010;100(1):56–60. https://doi.org/10.1016/j.jfoodeng.2010.03.027.
[377] Walkling-Ribeiro M, Rodriguez-Gonzalez O, Jayaram S, Griffiths MW. Microbial inactivation and shelf life comparison of ‘cold’ hurdle
processing with pulsed electric fields and microfiltration, and conventional thermal pasteurisation in skim milk. Int J Food Microbiol Jan 5
2011;144(3):379–86. https://doi.org/10.1016/j.ijfoodmicro.2010.10.023.
[378] Sampedro F, Rodrigo D, Martínez A. Modelling the effect of pH and pectin concentration on the PEF inactivation of Salmonella enterica
serovar Typhimurium by using the Monte Carlo simulation. Food Control 2011;22(3–4):420–5. https://doi.org/10.1016/j.foodcont.2010.09.
013.
[379] Rodriguez-Gonzalez O, Walkling-Ribeiro M, Jayaram S, Griffiths MW. Factors affecting the inactivation of the natural microbiota of
milk processed by pulsed electric fields and cross-flow microfiltration. J Dairy Res Aug 2011;78(3):270–8. https://doi.org/10.1017/
S0022029911000367.
[380] Rodríguez-González O, Walkling-Ribeiro M, Jayaram S, Griffiths MW. Cross-protective effects of temperature, pH, and osmotic and
starvation stresses in Escherichia coli O157:H7 subjected to pulsed electric fields in milk. Int Dairy J 2011;21(12):953–62. https://
doi.org/10.1016/j.idairyj.2011.07.003.
[381] Bermúdez-Aguirre D, Dunne CP, Barbosa-Cánovas GV. Effect of processing parameters on inactivation of Bacillus cereus spores in milk
using pulsed electric fields. Int Dairy J 2012;24(1):13–21. https://doi.org/10.1016/j.idairyj.2011.11.003.
[382] Palgan I, et al. Effectiveness of combined pulsed electric field (PEF) and manothermosonication (MTS) for the control of Listeria innocua in
a smoothie type beverage. Food Control 2012;25(2):621–5. https://doi.org/10.1016/j.foodcont.2011.11.009.
[383] Pina-Pérez MC, Martínez-López A, Rodrigo D. Cinnamon antimicrobial effect against Salmonella typhimurium cells treated by pulsed
electric fields (PEF) in pasteurized skim milk beverage. Food Res Int 2012;48(2):777–83. https://doi.org/10.1016/j.foodres.2012.06.027.
252
[384] Seratlić S, et al. Behavior of the surviving population of Lactobacillus plantarum 564 upon the application of pulsed electric fields. Innov
Food Sci Emerg Technol 2013;17:93–8. https://doi.org/10.1016/j.ifset.2012.11.011.
[385] Zhao W, Yang R, Shen X, Zhang S, Chen X. Lethal and sublethal injury and kinetics of Escherichia coli, Listeria monocytogenes and
Staphylococcus aureus in milk by pulsed electric fields. Food Control 2013;32(1):6–12. https://doi.org/10.1016/j.foodcont.2012.11.029.
[386] Pina-Pérez MC, Martínez-López A, Rodrigo D. Cocoa powder as a natural ingredient revealing an enhancing effect to inactivate Cronobacter
sakazakii cells treated by pulsed electric fields in infant milk formula. Food Control 2013;32(1):87–92. https://doi.org/10.1016/j.foodcont.
2012.11.014.
[387] Halpin RM, Cregenzán-Alberti O, Whyte P, Lyng JG, Noci F. Combined treatment with mild heat, manothermosonication and pulsed electric
fields reduces microbial growth in milk. Food Control 2013;34(2):364–71. https://doi.org/10.1016/j.foodcont.2013.05.008.
[388] Pina-Pérez MC, Guillier L, Rodrigo D, Martínez A. Modelling the exposure to Cronobacter sakazakii by consumption of a cocoa-milk-based
beverage processed by pulsed electric fields. Ir J Agric Food Res 2014;53(1):75–89.
[389] Sharma P, Bremer P, Oey I, Everett DW. Bacterial inactivation in whole milk using pulsed electric field processing. Int Dairy J
2014;35(1):49–56. https://doi.org/10.1016/j.idairyj.2013.10.005.
[390] Halpin RM, Duffy L, Cregenzán-Alberti O, Lyng JG, Noci F. The effect of non-thermal processing technologies on microbial inactivation:
an investigation into sub-lethal injury of Escherichia coli and Pseudomonas fluorescens. Food Control 2014;41(1):106–15. https://doi.org/
10.1016/j.foodcont.2014.01.011.
[391] Walter L, Knight G, Ng SY, Buckow R. Kinetic models for pulsed electric field and thermal inactivation of Escherichia coli and Pseudomonas
fluorescens in whole milk. Int Dairy J 2016;57:7–14. https://doi.org/10.1016/j.idairyj.2016.01.027.
[392] Lee GJ, Han BK, Choi HJ, Kang SH, Baick SC, Lee DU. Inactivation of Escherichia coli, Saccharomyces cerevisiae, and Lactobacillus
brevis in low-fat milk by pulsed electric field treatment: a pilot-scale study. Korean J Food Sci Anim Resour 2015;35(6):800–6. https://
doi.org/10.5851/kosfa.2015.35.6.800.
[393] Simonis P, et al. Pulsed electric field effects on inactivation of microorganisms in acid whey. Int J Food Microbiol Feb 16 2019;291:128–34.
https://doi.org/10.1016/j.ijfoodmicro.2018.11.024.
[394] Schottroff F, Johnson K, Johnson NB, Bédard MF, Jaeger H. Challenges and limitations for the decontamination of high solids protein
solutions at neutral pH using pulsed electric fields. J Food Eng 2020;268:109737. https://doi.org/10.1016/j.jfoodeng.2019.109737.
[395] Khoori E, Hakimzadeh V, Mohammadi Sani A, Rashidi H. Effect of ozonation, UV light radiation, and pulsed electric field processes on the
reduction of total aflatoxin and aflatoxin M1 in acidophilus milk. J Food Process Preserv 2020;44(10). https://doi.org/10.1111/jfpp.14729.
[396] Li SQ, Zhang QH, Lee YZ, Pham TV. Effects of pulsed electric fields and thermal processing on the stability of bovine immunoglobulin G
(IgG) in enriched soymilk. J Food Sci 2003;68(4):1201–7. https://doi.org/10.1111/j.1365-2621.2003.tb09625.x.
[397] Yang R, Li SQ, Zhang QH. Effects of pulsed electric fields on the activity and structure of pepsin. J Agric Food Chem Dec 1
2004;52(24):7400–6. https://doi.org/10.1021/jf049183p.
[398] Li SQ, Bomser JA, Zhang QH. Effects of pulsed electric fields and heat treatment on stability and secondary structure of bovine immunoglob-
ulin G. J Agric Food Chem Feb 9 2005;53(3):663–70. https://doi.org/10.1021/jf048923r.
[399] Wan J, et al. Emerging processing technologies for functional foods. Aust J Dairy Technol 2005;60(2):167–9 [Online]. Available: https://
www.proquest.com/scholarly-journals/emerging-processing-technologies-functional-foods/docview/199505200/se-2?accountid=14723.
[400] Rivas A, Rodrigo D, Company B, Sampedro F, Rodrigo M. Effects of pulsed electric fields on water-soluble vitamins and ACE inhibitory
peptides added to a mixed orange juice and milk beverage. Food Chem 2007;104(4):1550–9. https://doi.org/10.1016/j.foodchem.2007.02.
034.
[401] Li YQ, Chen Q, Liu XH, Chen ZX. Inactivation of soybean lipoxygenase in soymilk by pulsed electric fields. Food Chem Jul 15
2008;109(2):408–14. https://doi.org/10.1016/j.foodchem.2008.01.001.
[402] Riener J, Noci F, Cronin DA, Morgan DJ, Lyng JG. Combined effect of temperature and pulsed electric fields on soya milk lipoxygenase
inactivation. Eur Food Res Technol 2008;227(5):1461–5. https://doi.org/10.1007/s00217-008-0868-0.
[403] Luis RD, et al. Effect of high-intensity pulse electric fields on denaturation of bovine whey proteins. Milchwissenschaft 2009;64:422–6.
[404] Riener J, Noci F, Cronin DA, Morgan DJ, Lyng JG. Effect of high intensity pulsed electric fields on enzymes and vitamins in bovine raw
milk. Int J Dairy Technol 2009;62(1):1–6. https://doi.org/10.1111/j.1471-0307.2008.00435.x.
[405] Garcia-Amezquita LE, Primo-Mora AR, Barbosa-Cánovas GV, Sepulveda DR. Effect of nonthermal technologies on the native size distri-
bution of fat globules in bovine cheese-making milk. Innov Food Sci Emerg Technol 2009;10(4):491–4. https://doi.org/10.1016/j.ifset.2009.
03.002.
[406] Sampedro F, Geveke DJ, Fan X, Zhang HQ. Effect of PEF, HHP and thermal treatment on PME inactivation and volatile compounds
concentration of an orange juice–milk based beverage. Innov Food Sci Emerg Technol 2009;10(4):463–9. https://doi.org/10.1016/j.ifset.
2009.05.006.
[407] Yu LJ, Ngadi M, Raghavan GSV. Effect of temperature and pulsed electric field treatment on rennet coagulation properties of milk. J Food
Eng 2009;95(1):115–8. https://doi.org/10.1016/j.jfoodeng.2009.04.013.
[408] Zulueta A, Esteve MJ, Frígola A. Ascorbic acid in orange juice–milk beverage treated by high intensity pulsed electric fields and its stability
during storage. Innov Food Sci Emerg Technol 2010;11(1):84–90. https://doi.org/10.1016/j.ifset.2009.07.007.
[409] Zulueta A, Barba FJ, Esteve MJ, Frígola A. Effects on the carotenoid pattern and vitamin A of a pulsed electric field-treated orange juice–milk
beverage and behavior during storage. Eur Food Res Technol 2010;231(4):525–34. https://doi.org/10.1007/s00217-010-1304-9.
[410] Sui Q, Roginski H, Williams RPW, Versteeg C, Wan J. Effect of pulsed electric field and thermal treatment on the physicochemical properties
of lactoferrin with different iron saturation levels. Int Dairy J 2010;20(10):707–14. https://doi.org/10.1016/j.idairyj.2010.03.013.
[411] Bermúdez-Aguirre D, Yáñez JA, Dunne CP, Davies NM, Barbosa-Cánovas GV. Study of strawberry flavored milk under pulsed electric field
processing. Food Res Int 2010;43(8):2201–7. https://doi.org/10.1016/j.foodres.2010.07.021.
[412] Buckow R, Schroeder S, Berres P, Baumann P, Knoerzer K. Simulation and evaluation of pilot-scale pulsed electric field (PEF) processing.
J Food Eng 2010;101(1):67–77. https://doi.org/10.1016/j.jfoodeng.2010.06.010.
253
[413] Sui Q, Roginski H, Williams RP, Wooster TJ, Versteeg C, Wan J. Effect of the ionic strength of pulsed electric field treatment medium on
the physicochemical and structural characteristics of lactoferrin. J Agric Food Chem Nov 24 2010;58(22):11725–31. https://doi.org/10.1021/
jf102171u.
[414] Zhang S, Yang R, Zhao W, Hua X, Zhang W, Zhang Z. Influence of pulsed electric field treatments on the volatile compounds of milk in
comparison with pasteurized processing. J Food Sci Jan-Feb 2011;76(1):C127–32. https://doi.org/10.1111/j.1750-3841.2010.01916.x.
[415] Hemar Y, Augustin MA, Cheng L, Sanguansri P, Swiergon P, Wan J. The effect of pulsed electric field processing on particle size and
viscosity of milk and milk concentrates. Milchwissenschaft 01/01 2011;66:126–8.
[416] Sui Q, Roginski H, Williams RPW, Versteeg C, Wan J. Effect of pulsed electric field and thermal treatment on the physicochemical and
functional properties of whey protein isolate. Int Dairy J 2011;21(4):206–13. https://doi.org/10.1016/j.idairyj.2010.11.001.
[417] Xiang BY, Simpson MV, Ngadi MO, Simpson BK. Flow behaviour and viscosity of reconstituted skimmed milk treated with pulsed electric
field. Biosyst Eng 2011;109(3):228–34. https://doi.org/10.1016/j.biosystemseng.2011.04.004.
[418] Salvia-Trujillo L, Morales-de la Pena M, Rojas-Grau A, Martin-Belloso O. Changes in water-soluble vitamins and antioxidant capacity of
fruit juice-milk beverages as affected by high-intensity pulsed electric fields (HIPEF) or heat during chilled storage. J Agric Food Chem Sep
28 2011;59(18):10034–43. https://doi.org/10.1021/jf2011497.
[419] Xiang BY, Ngadi MO, Simpson BK, Simpson MV. Pulsed electric field induced structural modification of soy protein isolate as studied by
fluorescence spectroscopy. J Food Process Preserv 2011;35(5):563–70. https://doi.org/10.1111/j.1745-4549.2010.00501.x.
[420] Xiang BY, Simpson MV, Ngadi MO, Simpson BK. Effect of pulsed electric field on the rheological and colour properties of soy milk. Int J
Food Sci Nutr Dec 2011;62(8):787–93. https://doi.org/10.3109/09637486.2011.584860.
[421] Yu LJ, Ngadi M, Raghavan V. Proteolysis of cheese slurry made from pulsed electric field-treated milk. Food Bioprocess Technol
2010;5(1):47–54. https://doi.org/10.1007/s11947-010-0341-5.
[422] Buckow R, Semrau J, Sui Q, Wan J, Knoerzer K. Numerical evaluation of lactoperoxidase inactivation during continuous pulsed electric field
processing. Biotechnol Prog Sep-Oct 2012;28(5):1363–75. https://doi.org/10.1002/btpr.1582.
[423] Chugh A, Khanal D, Walkling-Ribeiro M, Corredig M, Duizer L, Griffiths MW. Change in color and volatile composition of skim milk
processed with pulsed electric field and microfiltration treatments or heat pasteurization. Foods Apr 23 2014;3(2):250–68. https://doi.org/10.
3390/foods3020250.
[424] Liu Z, et al. Pulsed electric field treatment of reconstituted skim milks at alkaline pH or with added EDTA. J Food Eng 2015;144:112–8.
[425] Sharma P, Oey I, Everett DW. Interfacial properties and transmission electron microscopy revealing damage to the milk fat globule system
after pulsed electric field treatment. Food Hydrocoll 2015;47:99–107. https://doi.org/10.1016/j.foodhyd.2015.01.023.
[426] Xu S, Walkling-Ribeiro M, Griffiths MW, Corredig M. Pulsed electric field processing preserves the antiproliferative activity of the milk fat
globule membrane on colon carcinoma cells. J Dairy Sci May 2015;98(5):2867–74. https://doi.org/10.3168/jds.2014-8839.
[427] Zhang Z-H, Yu Q, Zeng X-A, Han Z, Sun D-W, Muhammad-Aadil R. Effects of pulsed electric field on selected properties of L-tryptophan.
Int J Food Sci Technol 2015;50(5):1130–6. https://doi.org/10.1111/ijfs.12758.
[428] Wu L, Zhao W, Yang R, Yan W. Pulsed electric field (PEF)-induced aggregation between lysozyme, ovalbumin and ovotransferrin in multi-
protein system. Food Chem May 15 2015;175:115–20. https://doi.org/10.1016/j.foodchem.2014.11.136.
[429] Sharma P, Oey I, Everett DW. Thermal properties of milk fat, xanthine oxidase, caseins and whey proteins in pulsed electric field-treated
bovine whole milk. Food Chem Sep 15 2016;207:34–42. https://doi.org/10.1016/j.foodchem.2016.03.076.
[430] Subramanian V, Shanmugam N, Ranganathan K, Kumar S, Reddy R. Effect of combination processing on aflatoxin reduction: process
optimization by response surface methodology. J Food Process Preserv 2017;41(6):e13230. https://doi.org/10.1111/jfpp.13230.
[431] Zhang J, Tang T, Jiang Z, Liu Y, Jiang A. The modification of ovalbumin surface properties treated by pulsed electric field combined with
divalent metal ions. Food Chem Sep 30 2019;293:455–62. https://doi.org/10.1016/j.foodchem.2019.05.016.
[432] Manzoor MF, et al. Impact of pulsed electric field on rheological, structural, and physicochemical properties of almond milk. J Food Process
Eng 2019;42(8):e13299. https://doi.org/10.1111/jfpe.13299.
[433] Mohamad A, Shah NNAK, Sulaiman A, Mohd Adzahan N, Aadil RM. Impact of the pulsed electric field on physicochemical properties,
fatty acid profiling, and metal migration of goat milk. J Food Process Preserv 2020;44(12):e14940. https://doi.org/10.1111/jfpp.14940.
[434] Yang S, Suwal S, Andersen U, Otte J, Ahrné L. Effects of pulsed electric field on fat globule structure, lipase activity, and fatty acid
composition in raw milk and milk with different fat globule sizes. Innov Food Sci Emerg Technol 2021;67:102548. https://doi.org/10.1016/
j.ifset.2020.102548.
[435] Evrendilek GA, Dantzer WR, Streaker CB, Ratanatriwong P, Zhang QH. Shelf-life evaluations of liquid foods treated by pilot plant pulsed
electric field system. J Food Process Preserv 2001;25(4):283–97. https://doi.org/10.1111/j.1745-4549.2001.tb00461.x.
[436] Bendicho Sl, Barbosa-Cánovas GV, Martín O. Milk processing by high intensity pulsed electric fields. Trends Food Sci Technol
2002;13(6–7):195–204. https://doi.org/10.1016/s0924-2244(02)00132-2.
[437] Michalac S, Alvarez V, Ji T, Zhang QH. Inactivation of selected microorganisms and properties of pulsed electric field processed milk. J
Food Process Preserv 2003;27(2):137–51. https://doi.org/10.1111/j.1745-4549.2003.tb00507.x.
[438] Floury J, Grosset N, Leconte N, Pasco M, Madec M-N, Jeantet R. Continuous raw skim milk processing by pulsed electric field at non-lethal
temperature: effect on microbial inactivation and functional properties. Le Lait 2005;86(1):43–57. https://doi.org/10.1051/lait:2005039.
[439] Shamsi K, Versteeg C, Sherkat F, Wan J. Alkaline phosphatase and microbial inactivation by pulsed electric field in bovine milk. Innov Food
Sci Emerg Technol 2008;9(2):217–23. https://doi.org/10.1016/j.ifset.2007.06.012.
[440] Jaeger H, Schulz A, Karapetkov N, Knorr D. Protective effect of milk constituents and sublethal injuries limiting process effectiveness during
PEF inactivation of Lb. rhamnosus. Int J Food Microbiol Aug 31 2009;134(1–2):154–61. https://doi.org/10.1016/j.ijfoodmicro.2009.06.007.
[441] Jaeger H, Meneses N, Knorr D. Impact of PEF treatment inhomogeneity such as electric field distribution, flow characteristics and tem-
perature effects on the inactivation of E. coli and milk alkaline phosphatase. Innov Food Sci Emerg Technol 2009;10(4):470–80. https://
doi.org/10.1016/j.ifset.2009.03.001.
254
[442] Jaeger H, Meneses N, Moritz J, Knorr D. Model for the differentiation of temperature and electric field effects during thermal assisted PEF
processing. J Food Eng 2010;100(1):109–18. https://doi.org/10.1016/j.jfoodeng.2010.03.034.
[443] Bermudez-Aguirre D, Fernandez S, Esquivel H, Dunne PC, Barbosa-Canovas GV. Milk processed by pulsed electric fields: evalua-
tion of microbial quality, physicochemical characteristics, and selected nutrients at different storage conditions. J Food Sci Jun-Jul
2011;76(5):S289–99. https://doi.org/10.1111/j.1750-3841.2011.02171.x.
[444] Salvia-Trujillo L, Morales-de la Peña M, Rojas-Graü MA, Martín-Belloso O. Microbial and enzymatic stability of fruit juice-milk beverages
treated by high intensity pulsed electric fields or heat during refrigerated storage. Food Control 2011;22(10):1639–46. https://doi.org/10.
1016/j.foodcont.2011.03.022.
[445] Zulueta A, Barba FJ, Esteve MJ, Frígola A. Changes in quality and nutritional parameters during refrigerated storage of an orange juice–
milk beverage treated by equivalent thermal and non-thermal processes for mild pasteurization. Food Bioprocess Technol 2012;6(8):2018–30.
https://doi.org/10.1007/s11947-012-0858-x.
[446] Sharma P, Oey I, Bremer P, Everett DW. Reduction of bacterial counts and inactivation of enzymes in bovine whole milk using pulsed electric
fields. Int Dairy J 2014;39(1):146–56. https://doi.org/10.1016/j.idairyj.2014.06.003.
[447] McAuley CM, Singh TK, Haro-Maza JF, Williams R, Buckow R. Microbiological and physicochemical stability of raw, pasteurised or pulsed
electric field-treated milk. Innov Food Sci Emerg Technol 2016;38:365–73. https://doi.org/10.1016/j.ifset.2016.09.030.
[448] Sharma P, Oey I, Bremer P, Everett DW. Microbiological and enzymatic activity of bovine whole milk treated by pulsed electric fields. Int J
Dairy Technol 2018;71(1):10–9. https://doi.org/10.1111/1471-0307.12379.
[449] Schottroff F, Gratz M, Krottenthaler A, Johnson NB, Bédard MF, Jaeger H. Pulsed electric field preservation of liquid whey protein for-
mulations – influence of process parameters, pH, and protein content on the inactivation of Listeria innocua and the retention of bioactive
ingredients. J Food Eng 2019;243:142–52. https://doi.org/10.1016/j.jfoodeng.2018.09.003.
[450] Manzoor MF, et al. Effect of pulsed electric field and thermal treatments on the bioactive compounds, enzymes, microbial, and physical
stability of almond milk during storage. J Food Process Preserv 2020;44(7):e14541. https://doi.org/10.1111/jfpp.14541.
[451] Mohamad A, Abdul Karim Shah NN, Sulaiman A, Mohd Adzahan N, Aadil RM. Pulsed electric field of goat milk: impact on Escherichia
coli ATCC 8739 and vitamin constituents. J Food Process Eng 2021;44(9):e13779. https://doi.org/10.1111/jfpe.13779.
[452] Shinde GP, Kumar R, Reddy KR, Nadanasabhapathi S, Dutt Semwal A. Effect of pulsed electric field processing on reduction of sulfamet-
hazine residue content in milk. J Food Sci Technol May 2022;59(5):1931–8. https://doi.org/10.1007/s13197-021-05207-0.
[453] Akarca G, Ozkan M, Ozcan T. The impact of combination of solution plasma processing and pulsed electric field on the viability of probiotic
bacteria, microbial growth, and structure of yoghurt drink. J Food Process Preserv 2022. https://doi.org/10.1111/jfpp.16616.
[454] Zhao W, Yang R. The effect of pulsed electric fields on the inactivation and structure of lysozyme. Food Chem Sep 15 2008;110(2):334–43.
https://doi.org/10.1016/j.foodchem.2008.02.008.
[455] Zhao W, Yang R. Comparative study of inactivation and conformational change of lysozyme induced by pulsed electric fields and heat. Eur
Food Res Technol 2008;228(1):47–54. https://doi.org/10.1007/s00217-008-0905-z.
[456] Marco-Molés R, Pérez-Munuera I, Quiles A, Hernando I. Effect of pulsed electric fields on the main chemical components of liquid egg and
stability at 4 ◦ C. Czech J Food Sci 2009;27(Special Issue 1):S109–12. https://doi.org/10.17221/622-cjfs.
[457] Zhao W, Yang R, Tang Y, Zhang W, Hua X. Investigation of the protein-protein aggregation of egg white proteins under pulsed electric fields.
J Agric Food Chem May 13 2009;57(9):3571–7. https://doi.org/10.1021/jf803900f.
[458] Marco-Moles R, Rojas-Grau MA, Hernando I, Perez-Munuera I, Soliva-Fortuny R, Martin-Belloso O. Physical and structural changes in
liquid whole egg treated with high-intensity pulsed electric fields. J Food Sci Mar 2011;76(2):C257–64. https://doi.org/10.1111/j.1750-3841.
2010.02016.x.
[459] Zhao W, et al. Oxidation of oleic acid under pulsed electric field processing. Food Res Int 2011;44(5):1463–7. https://doi.org/10.1016/j.
foodres.2011.03.022.
[460] Lin S, Guo Y, You Q, Yin Y, Liu J. Preparation of antioxidant peptide from egg white protein and improvement of its activities assisted by
high-intensity pulsed electric field. J Sci Food Agric May 2012;92(7):1554–61. https://doi.org/10.1002/jsfa.4742.
[461] Monfort S, Mañas P, Condón S, Raso J, Álvarez I. Physicochemical and functional properties of liquid whole egg treated by the application of
pulsed electric fields followed by heat in the presence of triethyl citrate. Food Res Int 2012;48(2):484–90. https://doi.org/10.1016/j.foodres.
2012.04.015.
[462] Lin S, Guo Y, You QI, Yin Y, Liu J, Luo PG. Effects of high-intensity pulsed electric field on antioxidant attributes of hydrolysates derived
from egg white protein. J Food Biochem 2013;37(1):45–52. https://doi.org/10.1111/j.1745-4514.2011.00598.x.
[463] Wang K, et al. Effects on DPPH inhibition of egg-white protein polypeptides treated by pulsed electric field technology. J Sci Food Agric
May 2013;93(7):1641–8. https://doi.org/10.1002/jsfa.5941.
[464] Lin S, et al. Research on the preparation of antioxidant peptides derived from egg white with assisting of high-intensity pulsed electric field.
Food Chem Aug 15 2013;139(1–4):300–6. https://doi.org/10.1016/j.foodchem.2013.01.048.
[465] Wu L, Zhao W, Yang R, Chen X. Effects of pulsed electric fields processing on stability of egg white proteins. J Food Eng 2014;139:13–8.
[466] Wu L, Zhao W, Yang R, Yan W, Sun Q. Aggregation of egg white proteins with pulsed electric fields and thermal processes. J Sci Food Agric
Aug 2016;96(10):3334–41. https://doi.org/10.1002/jsfa.7512.
[467] Qian J-Y, Ma L-J, Wang L-J, Jiang W. Effect of pulsed electric field on structural properties of protein in solid state. Lebensm-Wiss Technol
2016;74:331–7. https://doi.org/10.1016/j.lwt.2016.07.068.
[468] Liu YF, Oey I, Bremer P, Carne A, Silcock P. Effects of pH, temperature and pulsed electric fields on the turbidity and protein aggregation
of ovomucin-depleted egg white. Food Res Int Jan 2017;91:161–70. https://doi.org/10.1016/j.foodres.2016.12.005.
[469] Liu YF, Oey I, Bremer P, Silcock P, Carne A. In vitro peptic digestion of ovomucin-depleted egg white affected by pH, temperature and
pulsed electric fields. Food Chem Sep 15 2017;231:165–74. https://doi.org/10.1016/j.foodchem.2017.03.136.
255
[470] Yang W, et al. Immunogenic and structural properties of ovalbumin treated by pulsed electric fields. Int J Food Prop 2018;20(sup3):S3164–76.
https://doi.org/10.1080/10942912.2017.1396479.
[471] Liu YF, Oey I, Bremer P, Silcock P, Carne A. Proteolytic pattern, protein breakdown and peptide production of ovomucin-depleted egg white
processed with heat or pulsed electric fields at different pH. Food Res Int Jun 2018;108:465–74. https://doi.org/10.1016/j.foodres.2018.03.
075.
[472] Liu YF, Oey I, Bremer P, Silcock P, Carne A, McConnell M. Pulsed electric fields treatment at different pH enhances the antioxidant and
anti-inflammatory activity of ovomucin-depleted egg white. Food Chem Mar 15 2019;276:164–73. https://doi.org/10.1016/j.foodchem.2018.
10.009.
[473] Li Y, Zhang S, Jiang P, Zhong L, Lin S, Sun N. Exploration of structure-activity relationship between IgG1 and IgE binding ability and
spatial conformation in ovomucoid with pulsed electric field treatment. Lebensm-Wiss Technol 2021;141:110891. https://doi.org/10.1016/j.
lwt.2021.110891.
[474] Calderon-Miranda ML, Barbosa-Canovas GV, Swanson BG. Inactivation of Listeria innocua in liquid whole egg by pulsed electric fields and
nisin. Int J Food Microbiol Oct 1 1999;51(1):7–17. https://doi.org/10.1016/s0168-1605(99)00070-7.
[475] Góngora-Nieto MM, Pedrow PD, Swanson BG, Barbosa-Cánovas GV. Energy analysis of liquid whole egg pasteurized by pulsed electric
fields. J Food Eng 2003;57(3):209–16. https://doi.org/10.1016/s0260-8774(02)00299-6.
[476] Hermawan N, Evrendilek GA, Dantzer WR, Zhang QH, Richter ER. Pulsed electric field treatment of liquid whole egg inoculated with
Salmonella Enteritidis. J Food Saf 2004;24(1):71–85. https://doi.org/10.1111/j.1745-4565.2004.tb00376.x.
[477] Amiali M, Ngadi MO, Raghavan VGS, Smith JP. Inactivation of Escherichia coli O157:H7 in liquid dialyzed egg using pulsed electric fields.
Food Bioprod Process 2004;82(2):151–6. https://doi.org/10.1205/0960308041614936.
[478] Amiali M, Ngadi M, Smith JP, Raghavan V. Inactivation of Escherichia coli 0157:H7 and Salmonella Enteritidis in liquid egg using continu-
ous pulsed electric field system. Int J Food Eng 2006;1(5). https://doi.org/10.2202/1556-3758.1058.
[479] Bazhal MI, Ngadi MO, Raghavan GSV, Smith JP. Inactivation of Escherichia coli O157:H7 in liquid whole egg using combined pulsed
electric field and thermal treatments. Lebensm-Wiss Technol 2006;39(4):420–6. https://doi.org/10.1016/j.lwt.2005.02.013.
[480] Amiali M, Ngadi MO, Raghavan VGS, Nguyen DH. Electrical conductivities of liquid egg products and fruit juices exposed to high pulsed
electric fields. Int J Food Prop 2006;9(3):533–40. https://doi.org/10.1080/10942910600596456.
[481] Amiali M, Ngadi MO, Smith JP, Raghavan VGS. Inactivation of Escherichia coli O157:H7 and Salmonella enteritidis in liquid egg white
using pulsed electric field. J Food Sci 2006;71(3):M88–94. https://doi.org/10.1111/j.1365-2621.2006.tb15637.x.
[482] Amiali M, Ngadi MO, Smith JP, Raghavan GSV. Synergistic effect of temperature and pulsed electric field on inactivation of Escherichia
coli O157:H7 and Salmonella enteritidis in liquid egg yolk. J Food Eng 2007;79(2):689–94. https://doi.org/10.1016/j.jfoodeng.2006.02.029.
[483] Jin T, Zhang H, Hermawan N, Dantzer W. Effects of pH and temperature on inactivation of Salmonella typhimurium DT104 in liquid whole
egg by pulsed electric fields. Int J Food Sci Technol 2009;44(2):367–72. https://doi.org/10.1111/j.1365-2621.2008.01759.x.
[484] Monfort S, et al. Inactivation of Salmonella Typhimurium and Staphylococcus aureus by pulsed electric fields in liquid whole egg. Innov
Food Sci Emerg Technol 2010;11(2):306–13. https://doi.org/10.1016/j.ifset.2009.11.007.
[485] Monfort S, Gayan E, Raso J, Condon S, Alvarez I. Evaluation of pulsed electric fields technology for liquid whole egg pasteurization. Food
Microbiol Oct 2010;27(7):845–52. https://doi.org/10.1016/j.fm.2010.05.011.
[486] Monfort S, Gayan E, Condon S, Raso J, Alvarez I. Design of a combined process for the inactivation of Salmonella Enteritidis in liquid
whole egg at 55 degrees C. Int J Food Microbiol Feb 28 2011;145(2–3):476–82. https://doi.org/10.1016/j.ijfoodmicro.2011.01.027.
[487] Monfort S, Saldana G, Condon S, Raso J, Alvarez I. Inactivation of Salmonella spp. in liquid whole egg using pulsed electric fields, heat,
and additives. Food Microbiol Jun 2012;30(2):393–9. https://doi.org/10.1016/j.fm.2012.01.004.
[488] Monfort S, Sagarzazu N, Condón S, Raso J, Álvarez I. Liquid whole egg ultrapasteurization by combination of PEF, heat, and additives.
Food Bioprocess Technol 2012;6(8):2070–80. https://doi.org/10.1007/s11947-012-0918-2.
[489] Espina L, Monfort S, Alvarez I, Garcia-Gonzalo D, Pagan R. Combination of pulsed electric fields, mild heat and essential oils as an alter-
native to the ultrapasteurization of liquid whole egg. Int J Food Microbiol Oct 17 2014;189:119–25. https://doi.org/10.1016/j.ijfoodmicro.
2014.08.002.
[490] Sanz-Puig M, Santos-Carvalho L, Cunha LM, Pina-Pérez MC, Martínez A, Rodrigo D. Effect of pulsed electric fields (PEF) combined with
natural antimicrobial by-products against S. Typhimurium. Innov Food Sci Emerg Technol 2016;37:322–8. https://doi.org/10.1016/j.ifset.
2016.09.004.
[491] Sanz-Puig M, et al. Resistance changes in Salmonella enterica serovar Typhimurium treated by high hydrostatic pressure and pulsed electric
fields and assessment of virulence changes by using Caenorhabditis elegans as a test organism. Innov Food Sci Emerg Technol 2019;51:51–6.
[492] Guillen S, Marcen M, Alvarez I, Manas P, Cebrian G. Stress resistance of emerging poultry-associated Salmonella serovars. Int J Food
Microbiol Dec 16 2020;335:108884. https://doi.org/10.1016/j.ijfoodmicro.2020.108884.
[493] Jeantet R, Baron F, Nau F, Roignant M, Brule G. High intensity pulsed electric fields applied to egg white: effect on Salmonella Enteritidis
inactivation and protein denaturation. J Food Prot Dec 1999;62(12):1381–6. https://doi.org/10.4315/0362-028x-62.12.1381.
[494] Zhao W, Yang R, Tang Y, Lu R. Combined effects of heat and PEF on microbial inactivation and quality of liquid egg whites. Int J Food Eng
2007;3(4):12. https://doi.org/10.2202/1556-3758.1256.
[495] W˛esierska E, Trziszka T. Evaluation of the use of pulsed electrical field as a factor with antimicrobial activity. J Food Eng 2007;78(4):1320–5.
[496] Jia M, Howard Zhang Q, Min DB. Pulsed electric field processing effects on flavor compounds and microorganisms of orange juice. Food
Chem 1999;65(4):445–51. https://doi.org/10.1016/s0308-8146(98)00186-1.
[497] Jin ZT, Zhang QH. Pulsed electric field inactivation of microorganisms and preservation of quality of cranberry juice. J Food Process Preserv
1999;23(6):481–97. https://doi.org/10.1111/j.1745-4549.1999.tb00399.x.
256
[498] Yeom HW, Streaker CB, Zhang QH, Min DB. Effects of pulsed electric fields on the activities of microorganisms and pectin methyl esterase
in orange juice. J Food Sci 2000;65(8):1359–63. https://doi.org/10.1111/j.1365-2621.2000.tb10612.x.
[499] Walkling-Ribeiro M, Noci F, Cronin DA, Riener J, Lyng JG, Morgan DJ. Reduction of Staphylococcus aureus and quality changes in apple
juice processed by ultraviolet irradiation, pre-heating and pulsed electric fields. J Food Eng 2008;89(3):267–73. https://doi.org/10.1016/j.
jfoodeng.2008.05.001.
[500] Akın E, Evrendilek GA. Effect of pulsed electric fields on physical, chemical, and microbiological properties of formulated carrot juice.
Food Sci Technol Int 2009;15(3):275–82. https://doi.org/10.1177/1082013209341414.
[501] Walkling-Ribeiro M, Noci F, Cronin DA, Lyng JG, Morgan DJ. Shelf life and sensory evaluation of orange juice after exposure to thermoson-
ication and pulsed electric fields. Food Bioprod Process 2009;87(2):102–7. https://doi.org/10.1016/j.fbp.2008.08.001.
[502] Walkling-Ribeiro M, Noci F, Riener J, Cronin DA, Lyng JG, Morgan DJ. The impact of thermosonication and pulsed electric fields on
Staphylococcus aureus inactivation and selected quality parameters in orange juice. Food Bioprocess Technol 2008;2(4):422–30. https://
doi.org/10.1007/s11947-007-0045-7.
[503] Altuntas J, Evrendilek GA, Sangun MK, Zhang HQ. Effects of pulsed electric field processing on the quality and microbial inactivation of
sour cherry juice. Int J Food Sci Technol 2010;45(5):899–905. https://doi.org/10.1111/j.1365-2621.2010.02213.x.
[504] Walkling-Ribeiro M, Noci F, Cronin DA, Lyng JG, Morgan DJ. Shelf life and sensory attributes of a fruit smoothie-type beverage processed
with moderate heat and pulsed electric fields. Lebensm-Wiss Technol 2010;43(7):1067–73. https://doi.org/10.1016/j.lwt.2010.02.010.
[505] Azhuvalappil Z, Fan X, Geveke DJ, Zhang HQ. Thermal and nonthermal processing of apple cider: storage quality under equivalent process
conditions. J Food Qual 2010;33(5):612–31. https://doi.org/10.1111/j.1745-4557.2010.00342.x.
[506] Zhang YI, Gao BEI, Zhang M, Shi J, Xu Y. Pulsed electric field processing effects on physicochemical properties, flavor compounds and
microorganisms of longan juice. J Food Process Preserv 2010;34(6):1121–38. https://doi.org/10.1111/j.1745-4549.2009.00441.x.
[507] Caminiti IM, et al. The effect of pulsed electric fields (PEF) in combination with high intensity light pulses (HILP) on Escherichia coli
inactivation and quality attributes in apple juice. Innov Food Sci Emerg Technol 2011;12(2):118–23. https://doi.org/10.1016/j.ifset.2011.01.
003.
[508] Ait-Ouazzou A, Cherrat L, Espina L, Lorán S, Rota C, Pagán R. The antimicrobial activity of hydrophobic essential oil constituents acting
alone or in combined processes of food preservation. Innov Food Sci Emerg Technol 2011;12(3):320–9. https://doi.org/10.1016/j.ifset.2011.
04.004.
[509] Timmermans RAH, et al. Comparing equivalent thermal, high pressure and pulsed electric field processes for mild pasteurization of orange
juice. Part I: impact on overall quality attributes. Innov Food Sci Emerg Technol 2011;12(3):235–43. https://doi.org/10.1016/j.ifset.2011.05.
001.
[510] Altuntaş J, Akdemir Evrendilek G, Sangun MK, Zhang HQ. Processing of peach nectar by pulsed electric fields with respect to physical and
chemical properties and microbial inactivation. J Food Process Eng 2011;34(5):1506–22. https://doi.org/10.1111/j.1745-4530.2009.00549.x.
[511] Ertugay MF, Başlar M, Ortakci F. Effect of pulsed electric field treatment on polyphenol oxidase, total phenolic compounds, and microbial
growth of apple juice. Turk J Agric For 2013;37(6):772–80. https://doi.org/10.3906/tar-1211-17.
[512] Guo M, Jin TZ, Geveke DJ, Fan X, Sites JE, Wang L. Evaluation of microbial stability, bioactive compounds, physicochemical properties,
and consumer acceptance of pomegranate juice processed in a commercial scale pulsed electric field system. Food Bioprocess Technol
2013;7(7):2112–20. https://doi.org/10.1007/s11947-013-1185-6.
[513] Xiang B, Sundararajan S, Mis Solval K, Espinoza-Rodezno L, Aryana K, Sathivel S. Effects of pulsed electric fields on physicochemical
properties and microbial inactivation of carrot juice. J Food Process Preserv 2014;38(4):1556–64. https://doi.org/10.1111/jfpp.12115.
[514] Aganovic K, et al. Impact of different large scale pasteurisation technologies and refrigerated storage on the headspace fingerprint of tomato
juice. Innov Food Sci Emerg Technol 2014;26:431–44. https://doi.org/10.1016/j.ifset.2014.10.007.
[515] Kayalvizhi V, Pushpa AJS, Sangeetha G, Antony U. Effect of pulsed electric field (PEF) treatment on sugarcane juice. J Food Sci Technol
Mar 2016;53(3):1371–9. https://doi.org/10.1007/s13197-016-2172-5.
[516] Aganovic K, et al. Headspace fingerprinting and sensory evaluation to discriminate between traditional and alternative pasteurization of
watermelon juice. Eur Food Res Technol 2015;242(5):787–803. https://doi.org/10.1007/s00217-015-2586-8.
[517] Agcam E, Akyildiz A, Akdemir Evrendilek G. A comparative assessment of long-term storage stability and quality attributes of orange juice
in response to pulsed electric fields and heat treatments. Food Bioprod Process 2016;99:90–8. https://doi.org/10.1016/j.fbp.2016.04.006.
[518] Jin TZ, Yu Y, Gurtler JB. Effects of pulsed electric field processing on microbial survival, quality change and nutritional characteristics of
blueberries. Lebensm-Wiss Technol 2017;77:517–24. https://doi.org/10.1016/j.lwt.2016.12.009.
[519] Katiyo W, Yang R, Zhao W. Effects of combined pulsed electric fields and mild temperature pasteurization on microbial inactivation and
physicochemical properties of cloudy red apple juice (Malus pumilaNiedzwetzkyana (Dieck)). J Food Saf 2017;37(4):e12369. https://doi.
org/10.1111/jfs.12369.
[520] Evrendilek GA. Impacts of pulsed electric field and heat treatment on quality and sensory properties and microbial inactivation of
pomegranate juice. Food Sci Technol Int Dec 2017;23(8):668–80. https://doi.org/10.1177/1082013217715369.
[521] Aadil RM, et al. Combined effects of pulsed electric field and ultrasound on bioactive compounds and microbial quality of grapefruit juice.
J Food Process Preserv 2017;42(2):e13507. https://doi.org/10.1111/jfpp.13507.
[522] Sotelo KAG, et al. Red cherries (Prunus avium var. Stella) processed by pulsed electric field - physical, chemical and microbiological
analyses. Food Chem Feb 1 2018;240:926–34. https://doi.org/10.1016/j.foodchem.2017.08.017.
[523] Lee SJ, Bang IH, Choi HJ, Min SC. Pasteurization of mixed mandarin and Hallabong tangor juice using pulsed electric field processing
combined with heat. Food Sci Biotechnol Jun 2018;27(3):669–75. https://doi.org/10.1007/s10068-018-0311-7.
[524] Dziadek K, et al. Effect of pulsed electric field treatment on shelf life and nutritional value of apple juice. J Food Sci Technol Mar
2019;56(3):1184–91. https://doi.org/10.1007/s13197-019-03581-4.
257
[525] Akdemir Evrendilek G, Keskin E, Golge O. Interaction and multi-objective effects of multiple non-thermal treatments of sour cherry juice:
pesticide removal, microbial inactivation, and quality preservation. J Sci Food Agric Mar 15 2020;100(4):1653–61. https://doi.org/10.1002/
jsfa.10178.
[526] Wu S, Yang N, Jin Y, Xu X, Jin Z, Xie Z. Effects of induced electric field (IEF) on the reduction of Saccharomyces cerevisiae and quality of
fresh apple juice. Food Chem Apr 30 2020;325:126943. https://doi.org/10.1016/j.foodchem.2020.126943.
[527] Sharma SK, Zhang QH, Chism GW. Development of a protein fortified fruit beverage and its quality when processed with pulsed electric
field treatment. J Food Qual 1998;21(6):459–73. https://doi.org/10.1111/j.1745-4557.1998.tb00536.x.
[528] Yeom HW, Streaker CB, Zhang QH, Min DB. Effects of pulsed electric fields on the quality of orange juice and comparison with heat
pasteurization. J Agric Food Chem Oct 2000;48(10):4597–605. https://doi.org/10.1021/jf000306p.
[529] Ayhan Z, Yeom HW, Zhang QH, Min DB. Flavor, color, and vitamin C retention of pulsed electric field processed orange juice in different
packaging materials. J Agric Food Chem Feb 2001;49(2):669–74. https://doi.org/10.1021/jf000984b.
[530] Yeom HW, Zhang QH, Chism GW. Inactivation of pectin methyl esterase in orange juice by pulsed electric fields. J Food Sci
[531] Ayhan Z, Zhang QH, Min DB. Effects of pulsed electric field processing and storage on the quality and stability of single-strength orange
juice. J Food Prot Oct 2002;65(10):1623–7. https://doi.org/10.4315/0362-028x-65.10.1623.
[532] Min S, Jin ZT, Min SK, Yeom H, Zhang QH. Commercial-scale pulsed electric field processing of orange juice. J Food Sci
[533] Min S, Jin ZT, Zhang QH. Commercial scale pulsed electric field processing of tomato juice. J Agric Food Chem May 21
2003;51(11):3338–44. https://doi.org/10.1021/jf0260444.
[534] Min S, Zhang QH. Effects of commercial-scale pulsed electric field processing on flavor and color of tomato juice. J Food Sci
[535] Min S, Min SK, Zhang QH. Inactivation kinetics of tomato juice lipoxygenase by pulsed electric fields. J Food Sci 2003;68(6):1995–2001.
[536] Rodrigo D, Barbosa-Canovas GV, Martinez A, Rodrigo M. Pectin methyl esterase and natural microflora of fresh mixed orange and carrot
juice treated with pulsed electric fields. J Food Prot Dec 2003;66(12):2336–42. https://doi.org/10.4315/0362-028x-66.12.2336.
[537] Sanchez-Moreno C, Plaza L, Elez-Martinez P, De Ancos B, Martin-Belloso O, Cano MP. Impact of high pressure and pulsed electric fields
on bioactive compounds and antioxidant activity of orange juice in comparison with traditional thermal processing. J Agric Food Chem Jun
1 2005;53(11):4403–9. https://doi.org/10.1021/jf048839b.
[538] Giner J, Bailo E, Gimeno V, Martín-Belloso O. Models in a Bayesian framework for inactivation of pectinesterase in a commercial enzyme
formulation by pulsed electric fields. Eur Food Res Technol 2005;221(3–4):255–64. https://doi.org/10.1007/s00217-005-1143-2.
[539] Roodenburg B, Morren J, Berg HE, de Haan SWH. Metal release in a stainless steel pulsed electric field (PEF) system. Innov Food Sci
Emerg Technol 2005;6(3):337–45. https://doi.org/10.1016/j.ifset.2005.04.004.
[540] Rivas A, Rodrigo D, Martínez A, Barbosa-Cánovas GV, Rodrigo M. Effect of PEF and heat pasteurization on the physical–chemical charac-
teristics of blended orange and carrot juice. Lebensm-Wiss Technol 2006;39(10):1163–70. https://doi.org/10.1016/j.lwt.2005.07.002.
[541] Torregrosa F, Esteve MJ, Frígola A, Cortés C. Ascorbic acid stability during refrigerated storage of orange–carrot juice treated by high pulsed
electric field and comparison with pasteurized juice. J Food Eng 2006;73(4):339–45. https://doi.org/10.1016/j.jfoodeng.2005.01.034.
[542] Cserhalmi Z, Sass-Kiss Á, Tóth-Markus M, Lechner N. Study of pulsed electric field treated citrus juices. Innov Food Sci Emerg Technol
2006;7(1–2):49–54. https://doi.org/10.1016/j.ifset.2005.07.001.
[543] Giner-Seguí J, Bailo-Ballarín E, Gorinstein S, Martín-Belloso O. New kinetic approach to the evolution of polygalacturonase (EC 3.2.1.15)
activity in a commercial enzyme preparation under pulsed electric fields. J Food Sci 2006;71(6):E262–9. https://doi.org/10.1111/j.1750-
3841.2006.00054.x.
[544] Plaza L, Sánchez-Moreno C, Elez-Martínez P, de Ancos B, Martín-Belloso O, Cano MP. Effect of refrigerated storage on vitamin C and
antioxidant activity of orange juice processed by high-pressure or pulsed electric fields with regard to low pasteurization. Eur Food Res
Technol 2006;223(4):487–93. https://doi.org/10.1007/s00217-005-0228-2.
[545] Garde-Cerdán T, Arias-Gil M, Marsellés-Fontanet AR, Ancín-Azpilicueta C, Martín-Belloso O. Effects of thermal and non-thermal process-
ing treatments on fatty acids and free amino acids of grape juice. Food Control 2007;18(5):473–9.
[546] Aguilar-Rosas SF, Ballinas-Casarrubias ML, Nevarez-Moorillon GV, Martin-Belloso O, Ortega-Rivas E. Thermal and pulsed electric fields
pasteurization of apple juice: effects on physicochemical properties and flavour compounds. J Food Eng 2007;83(1):41–6. https://doi.org/10.
1016/j.jfoodeng.2006.12.011.
[547] Marsellés-Fontanet ÁR, Martín-Belloso O. Optimization and validation of PEF processing conditions to inactivate oxidative enzymes of
grape juice. J Food Eng 2007;83(3):452–62. https://doi.org/10.1016/j.jfoodeng.2007.04.001.
[548] Noci F, Riener J, Walkling-Ribeiro M, Cronin DA, Morgan DJ, Lyng JG. Ultraviolet irradiation and pulsed electric fields (PEF) in a hurdle
strategy for the preservation of fresh apple juice. J Food Eng 2008;85(1):141–6. https://doi.org/10.1016/j.jfoodeng.2007.07.011.
[549] Cortés C, Esteve MJ, Frígola A. Effect of refrigerated storage on ascorbic acid content of orange juice treated by pulsed electric fields and
thermal pasteurization. Eur Food Res Technol 2007;227(2):629–35. https://doi.org/10.1007/s00217-007-0766-x.
[550] Aguiló-Aguayo I, Soliva-Fortuny R, Martín-Belloso O. Comparative study on color, viscosity and related enzymes of tomato juice treated
by high-intensity pulsed electric fields or heat. Eur Food Res Technol 2007;227(2):599–606. https://doi.org/10.1007/s00217-007-0761-2.
[551] Schilling S, et al. Comparative study of pulsed electric field and thermal processing of apple juice with particular consideration of juice
quality and enzyme deactivation. J Agric Food Chem Jun 25 2008;56(12):4545–54. https://doi.org/10.1021/jf0732713.
[552] Riener J, Noci F, Cronin DA, Morgan DJ, Lyng JG. Combined effect of temperature and pulsed electric fields on apple juice peroxidase and
polyphenoloxidase inactivation. Food Chem Jul 15 2008;109(2):402–7. https://doi.org/10.1016/j.foodchem.2007.12.059.
[553] Riener J, Noci F, Cronin DA, Morgan DJ, Lyng JG. Combined effect of temperature and pulsed electric fields on pectin methyl esterase
inactivation in red grapefruit juice (Citrus paradisi). Eur Food Res Technol 2008;228(3):373–9. https://doi.org/10.1007/s00217-008-0943-6.
258
[554] Esteve MJ, Barba FJ, Palop S, Frígola A. The effects of non-thermal processing on carotenoids in orange juice. Czech J Food Sci
2009;27(Special Issue 1):S304–6. https://doi.org/10.17221/1094-cjfs.
[555] Cortés C, Esteve MJ, Frígola A. Anteroxanthin concentration during refrigerated storage in orange juice treated by PEF. Czech J Food Sci
2009;27(Special Issue 1):S307–9. https://doi.org/10.17221/1083-cjfs.
[556] Chen F, et al. Degradation behaviour of methamidophos and chlorpyrifos in apple juice treated with pulsed electric fields. Food Chem
2009;112(4):956–61. https://doi.org/10.1016/j.foodchem.2008.07.016.
[557] Jin ZT, et al. Quality of applesauces processed by pulsed electric fields and HTST pasteurisation. Int J Food Sci Technol 2009;44(4):829–39.
https://doi.org/10.1111/j.1365-2621.2009.01912.x.
[558] Sanchez-Vega R, Mujica-Paz H, Marquez-Melendez R, Ngadi MO, Ortega-Rivas E. Enzyme inactivation on apple juice treated by ultra-
pasteurization and pulsed electric fields technology. J Food Process Preserv 2009;33(4):486–99. https://doi.org/10.1111/j.1745-4549.2008.
00270.x.
[559] Zhao W, Yang R, Hua X, Zhang W, Tang Y, Chen T. Inactivation of polyphenoloxidase of pear by pulsed electric fields. Int J Food Eng
2010;6(3). https://doi.org/10.2202/1556-3758.1853.
[560] Caminiti IM, et al. Impact of selected combinations of non-thermal processing technologies on the quality of an apple and cranberry juice
blend. Food Chem 2011;124(4):1387–92. https://doi.org/10.1016/j.foodchem.2010.07.096.
[561] Hartyáni P, Dalmadi I, Cserhalmi Z, Kántor D-B, Tóth-Markus M, Sass-Kiss Á. Physical–chemical and sensory properties of pulsed electric
field and high hydrostatic pressure treated citrus juices. Innov Food Sci Emerg Technol 2011;12(3):255–60. https://doi.org/10.1016/j.ifset.
2011.04.008.
[562] Vervoort L, et al. Comparing equivalent thermal, high pressure and pulsed electric field processes for mild pasteurization of orange juice.
Innov Food Sci Emerg Technol 2011;12(4):466–77. https://doi.org/10.1016/j.ifset.2011.06.003.
[563] Zhang Y, et al. Reduction of diazinon and dimethoate in apple juice by pulsed electric field treatment. J Sci Food Agric Mar 15
2012;92(4):743–50. https://doi.org/10.1002/jsfa.4636.
[564] Barba FJ, Jäger H, Meneses N, Esteve MJ, Frígola A, Knorr D. Evaluation of quality changes of blueberry juice during refrigerated storage
after high-pressure and pulsed electric fields processing. Innov Food Sci Emerg Technol 2012;14:18–24. https://doi.org/10.1016/j.ifset.2011.
12.004.
[565] Caminiti IM, Noci F, Morgan DJ, Cronin DA, Lyng JG. The effect of pulsed electric fields, ultraviolet light or high intensity light pulses
in combination with manothermosonication on selected physico-chemical and sensory attributes of an orange and carrot juice blend. Food
Bioprod Process 2012;90(3):442–8. https://doi.org/10.1016/j.fbp.2011.11.006.
[566] Chen C, Zhao W, Yang R, Zhang S. Effects of pulsed electric field on colloidal properties and storage stability of carrot juice. Int J Food Sci
Technol 2012;47(10):2079–85. https://doi.org/10.1111/j.1365-2621.2012.03072.x.
[567] Vervoort L, et al. Headspace fingerprinting as an untargeted approach to compare novel and traditional processing technologies: a case-study
on orange juice pasteurisation. Food Chem Oct 15 2012;134(4):2303–12. https://doi.org/10.1016/j.foodchem.2012.03.096.
[568] Quintao-Teixeira LJ, Soliva-Fortuny R, Mota Ramos A, Martin-Belloso O. Kinetics of peroxidase inactivation in carrot juice treated with
pulsed electric fields. J Food Sci Feb 2013;78(2):E222–8. https://doi.org/10.1111/1750-3841.12019.
[569] Aguilar-Rosas S, Ballinas-Casarrubias M, Elias-Ogaz L, Martin-Belloso O, Ortega-Rivas E. Enzyme activity and colour changes in apple
juice pasteurised thermally and by pulsed electric fields. Acta Aliment 03/01 2013;42(1):45–54. https://doi.org/10.1556/AAlim.42.2013.1.5.
[570] Meneses N, et al. Modelling of polyphenoloxidase inactivation by pulsed electric fields considering coupled effects of temperature and
electric field. Innov Food Sci Emerg Technol 2013;20:126–32. https://doi.org/10.1016/j.ifset.2012.12.009.
[571] Bansal V, Sharma A, Ghanshyam C, Singla ML. Coupling of chromatographic analyses with pretreatment for the determination of bioactive
compounds in Emblica officinalis juice. Anal Methods 2014;6(2):410–8. https://doi.org/10.1039/c3ay41375f.
[572] Agcam E, Akyildiz A, Akdemir Evrendilek G. Comparison of phenolic compounds of orange juice processed by pulsed electric fields (PEF)
and conventional thermal pasteurisation. Food Chem Jan 15 2014;143:354–61. https://doi.org/10.1016/j.foodchem.2013.07.115.
[573] Chen J, et al. Influence of pulsed electric field and thermal treatments on the quality of blueberry juice. Int J Food Prop 2014;17(7):1419–27.
https://doi.org/10.1080/10942912.2012.713429.
[574] Jin TZ, Guo M, Zhang HQ. Upscaling from benchtop processing to industrial scale production: more factors to be considered for pulsed
electric field food processing. J Food Eng 2015;146:72–80. https://doi.org/10.1016/j.jfoodeng.2014.08.020.
[575] Leong SY, Oey I, Burritt DJ. A novel strategy using pulsed electric fields to modify the thermostability of ascorbic acid oxidase in different
carrot cultivars. Food Bioprocess Technol 2014;8(4):811–23. https://doi.org/10.1007/s11947-014-1448-x.
[576] Evrendilek GA, Avsar YK, Evrendilek F. Modelling stochastic variability and uncertainty in aroma active compounds of PEF-treated peach
nectar as a function of physical and sensory properties, and treatment time. Food Chem Jan 1 2016;190:634–42. https://doi.org/10.1016/j.
foodchem.2015.06.010.
[577] Evrendilek GA. Change regime of aroma active compounds in response to pulsed electric field treatment time, sour cherry juice apricot and
peach nectars, and physical and sensory properties. Innov Food Sci Emerg Technol 2016;33:195–205. https://doi.org/10.1016/j.ifset.2015.
11.020.
[578] Medina-Meza IG, Boioli P, Barbosa-Cánovas GV. Assessment of the effects of ultrasonics and pulsed electric fields on nutritional and
rheological properties of raspberry and blueberry purees. Food Bioprocess Technol 2015;9(3):520–31. https://doi.org/10.1007/s11947-015-
1642-5.
[579] Carbonell-Capella JM, et al. Changes of antioxidant compounds in a fruit juice-stevia rebaudiana blend processed by pulsed electric tech-
nologies and ultrasound. Food Bioprocess Technol 2016;9(7):1159–68. https://doi.org/10.1007/s11947-016-1706-1.
[580] Aschoff JK, Knoblauch K, Hüttner C, Vásquez-Caicedo AL, Carle R, Schweiggert RM. Non-thermal pasteurization of orange (Citrus sinensis
(L.) Osbeck) juices using continuous pressure change technology (PCT): a proof-of-concept. Food Bioprocess Technol 2016;9(10):1681–91.
https://doi.org/10.1007/s11947-016-1754-6.
259
[581] Andreou V, Dimopoulos G, Katsaros G, Taoukis P. Comparison of the application of high pressure and pulsed electric fields technologies
on the selective inactivation of endogenous enzymes in tomato products. Innov Food Sci Emerg Technol 2016;38:349–55. https://doi.org/10.
1016/j.ifset.2016.07.026.
[582] Carbonell-Capella JM, Buniowska M, Cortes C, Zulueta A, Frigola A, Esteve MJ. Influence of pulsed electric field processing on the quality
of fruit juice beverages sweetened with Stevia rebaudiana. Food Bioprod Process 2017;101:214–22. https://doi.org/10.1016/j.fbp.2016.11.
012.
[583] Sulaiman A, Farid M, Silva FV. Quality stability and sensory attributes of apple juice processed by thermosonication, pulsed electric field
and thermal processing. Food Sci Technol Int Apr 2017;23(3):265–76. https://doi.org/10.1177/1082013216685484.
[584] Buniowska M, Carbonell-Capella JM, Frigola A, Esteve MJ. Bioaccessibility of bioactive compounds after non-thermal processing of an
exotic fruit juice blend sweetened with Stevia rebaudiana. Food Chem Apr 15 2017;221:1834–42. https://doi.org/10.1016/j.foodchem.2016.
10.093.
[585] Moussa-Ayoub TE, Jäger H, Knorr D, El-Samahy SK, Kroh LW, Rohn S. Impact of pulsed electric fields, high hydrostatic pressure, and
thermal pasteurization on selected characteristics of Opuntia dillenii cactus juice. Lebensm-Wiss Technol 2017;79:534–42. https://doi.org/
10.1016/j.lwt.2016.10.061.
[586] Lasekan O, Ng S, Azeez S, Shittu R, Teoh L, Gholivand S. Effect of pulsed electric field processing on flavor and color of liquid foods. J
Food Process Preserv 2017;41(3):e12940. https://doi.org/10.1111/jfpp.12940.
[587] Akdemir Evrendilek G, Celik P, Agcam E, Akyildiz A. Assessing impacts of pulsed electric fields on quality attributes and furfural and
hydroxymethylfurfural formations in apple juice. J Food Process Eng 2017;40(5):e12524. https://doi.org/10.1111/jfpe.12524.
[588] Tian Y, Wang S, Yan W, Tang Y, Yang R, Zhao W. Inactivation of apple (Malus domestica Borkh) polyphenol oxidases by radio frequency
combined with pulsed electric field treatment. Int J Food Sci Technol 2018;53(9):2054–63. https://doi.org/10.1111/ijfs.13781.
[589] Katiyo W, Yang R, Zhao W. Phenolic composition and antioxidant activity of Chinese red-fleshed apples (Malus pumila Niedzwetzkyana
(Dieck) and effect of different pasteurization treatments on the cloudy juice. Int Food Res J 2018;25(5):2185–94.
[590] Kebede B, et al. A chemometrics approach comparing volatile changes during the shelf life of apple juice processed by pulsed electric fields,
high pressure and thermal pasteurization. Foods Oct 17 2018;7(10):169. https://doi.org/10.3390/foods7100169.
[591] Kumar R, Bawa AS, Rajeswara Reddy K, Kathiravan T, Subramanian V, Nadanasabapathi S. Pulsed electric field and combination processing
of mango nectar: effect on volatile compounds and HMF formation. Croat J Food Sci Technol 2015;7(2):58–67. https://doi.org/10.17508/
cjfst.2015.7.2.02.
[592] Wibowo S, et al. Comparing the impact of high pressure, pulsed electric field and thermal pasteurization on quality attributes of cloudy apple
juice using targeted and untargeted analyses. Innov Food Sci Emerg Technol 2019;54:64–77. https://doi.org/10.1016/j.ifset.2019.03.004.
[593] Gagneten M, Leiva G, Salvatori D, Schebor C, Olaiz N. Optimization of pulsed electric field treatment for the extraction of bioactive
compounds from blackcurrant. Food Bioprocess Technol 2019;12(7):1102–9. https://doi.org/10.1007/s11947-019-02283-1.
[594] Mannozzi C, et al. Influence of pulsed electric field and ohmic heating pretreatments on enzyme and antioxidant activity of fruit and vegetable
juices. Foods Jul 8 2019;8(7):247. https://doi.org/10.3390/foods8070247.
[595] Zhong S, et al. Novel processing technologies as compared to thermal treatment on the bioaccessibility and Caco-2 cell uptake of carotenoids
from tomato and kale-based juices. J Agric Food Chem Sep 11 2019;67(36):10185–94. https://doi.org/10.1021/acs.jafc.9b03666.
[596] Stübler A-S, Lesmes U, Heinz V, Rauh C, Shpigelman A, Aganovic K. Digestibility, antioxidative activity and stability of plant protein-
rich products after processing and formulation with polyphenol-rich juices: kale and kale–strawberry as a model. Eur Food Res Technol
2019;245(11):2499–514. https://doi.org/10.1007/s00217-019-03362-5.
[597] Manzoor MF, et al. Novel processing techniques and spinach juice: quality and safety improvements. J Food Sci Apr 2020;85(4):1018–26.
https://doi.org/10.1111/1750-3841.15107.
[598] Rahaman A, et al. Effect of pulsed electric fields processing on physiochemical properties and bioactive compounds of apricot juice. J Food
Process Eng 2020;43(8):e13449. https://doi.org/10.1111/jfpe.13449.
[599] Stübler A-S, et al. Impact of pilot-scale processing (thermal, PEF, HPP) on the stability and bioaccessibility of polyphenols and proteins
in mixed protein- and polyphenol-rich juice systems. Innov Food Sci Emerg Technol 2020;64:102426. https://doi.org/10.1016/j.ifset.2020.
102426.
[600] Yildiz S, Pokhrel PR, Unluturk S, Barbosa-Cánovas GV. Changes in quality characteristics of strawberry juice after equivalent high pressure,
ultrasound, and pulsed electric fields processes. Food Eng Rev 2020;13(3):601–12. https://doi.org/10.1007/s12393-020-09250-z.
[601] Akdemir Evrendilek G, Agcam E, Akyildiz A. Effects of pulsed electric fields on sour cherry juice properties and formations of furfural and
hydroxymethylfurfural. Int J Food Eng 2021;17(3):217–26. https://doi.org/10.1515/ijfe-2020-0189.
[602] Faisal Manzoor M, et al. Probing the combined impact of pulsed electric field and ultra-sonication on the quality of spinach juice. J Food
Process Preserv 2021;45(5):e15475. https://doi.org/10.1111/jfpp.15475.
[603] Pallarés N, Berrada H, Tolosa J, Ferrer E. Effect of high hydrostatic pressure (HPP) and pulsed electric field (PEF) technologies on reduction
of aflatoxins in fruit juices. Lebensm-Wiss Technol 2021;142:111000. https://doi.org/10.1016/j.lwt.2021.111000.
[604] Qin B-L, Chang F-J, Barbosa-Cánovas GV, Swanson BG. Nonthermal inactivation of Saccharomyces cerevisiae in apple juice using pulsed
electric fields. Lebensm-Wiss Technol 1995;28(6):564–8. https://doi.org/10.1016/0023-6438(95)90002-0.
[605] Harrison SL, Barbosa-Cánovas GV, Swanson BG. Structural changes induced by pulsed electric field treatment. Lebensm-Wiss Technol
1997;30(3):236–40. https://doi.org/10.1006/fstl.1996.0171.
[606] Raso J, Calderón MaL, Góngora M, Barbosa-Cánovas G, Swanson BG. Inactivation of mold ascospores and conidiospores suspended in fruit
juices by pulsed electric fields. Lebensm-Wiss Technol 1998;31(7–8):668–72. https://doi.org/10.1006/fstl.1998.0426.
[607] Raso J, Calderón ML, Góngora M, Barbosa-Cánovas GV, Swanson BG. Inactivation of Zygosaccharomyces bailii in fruit juices by heat,
high hydrostatic pressure and pulsed electric fields. J Food Sci 2006;63(6):1042–4. https://doi.org/10.1111/j.1365-2621.1998.tb15850.x.
[608] Evrendilek GA, Zhang QH, Richter ER. Inactivation of Escherichia coli O157:H7 and Escherichia coli 8739 in apple juice by pulsed electric
fields. J Food Prot Jul 1999;62(7):793–6. https://doi.org/10.4315/0362-028x-62.7.793.
260
[609] McDonald CJ, Lloyd SW, Vitale MA, Petersson K, Innings F. Effects of pulsed electric fields on microorganisms in orange juice using
electric field strengths of 30 and 50 kV/cm. J Food Sci 2000;65(6):984–9. https://doi.org/10.1111/j.1365-2621.2000.tb09404.x.
[610] Liang Z, Mittal GS, Griffiths MW. Inactivation of Salmonella Typhimurium in orange juice containing antimicrobial agents by pulsed electric
field. J Food Prot Jul 2002;65(7):1081–7. https://doi.org/10.4315/0362-028x-65.7.1081.
[611] Hodgins AM, Mittal GS, Griffiths MW. Pasteurization of fresh orange juice using low-energy pulsed electrical field. J Food Sci
[612] Liang Z, Cheng Z, Mittal GS. Inactivation of spoilage microorganisms in apple cider using a continuous flow pulsed electric field system.
Lebensm-Wiss Technol 2006;39(4):351–7. https://doi.org/10.1016/j.lwt.2005.02.019.
[613] Alvarez I, Manas P, Condon S, Raso J. Resistance variation of Salmonella enterica serovars to pulsed electric fields treatments. J Food Sci
[614] Sen Gupta B, Masterson F, Magee TRA. Inactivation of E. coli K12 in apple juice by high voltage pulsed electric field. Eur Food Res Technol
2003;217(5):434–7. https://doi.org/10.1007/s00217-003-0756-6.
[615] Molinari P, Pilosof AMR, Jagus RJ. Effect of growth phase and inoculum size on the inactivation of Saccharomyces cerevisiae in fruit juices,
by pulsed electric fields. Food Res Int 2004;37(8):793–8. https://doi.org/10.1016/j.foodres.2004.03.010.
[616] Selma MV, Salmerón MC, Valero M, Fernández PS. Control of Lactobacillus plantarum and Escherichia coli by pulsed electric fields in
MRS Broth, Nutrient Broth and orange–carrot juice. Food Microbiol 2004;21(5):519–25. https://doi.org/10.1016/j.fm.2003.12.004.
[617] Java S, Varadharaju N, Kennedy ZJ. Inactivation of microorganisms in the fruit juice using pulsed electric field. J Food Sci Technol, Mysore
2004;41:652–5.
[618] Lebovka NI, Praporscic I, Vorobiev E. Combined treatment of apples by pulsed electric fields and by heating at moderate temperature. J
Food Eng 2004;65(2):211–7. https://doi.org/10.1016/j.jfoodeng.2004.01.017.
[619] Zhang Y, Mittal G. Inactivation of spoilage microorganisms in mango juice using low energy pulsed electric field in combination with
antimicrobials. Ital J Food Sci 2005;17:167–76.
[620] Wu Y, Mittal GS, Griffiths MW. Effect of pulsed electric field on the inactivation of microorganisms in grape juices with and without
antimicrobials. Biosyst Eng 2005;90(1):1–7. https://doi.org/10.1016/j.biosystemseng.2004.07.012.
[621] Garcia D, Hassani M, Manas P, Condon S, Pagan R. Inactivation of Escherichia coli O157:H7 during the storage under refrigeration of apple
juice treated by pulsed electric fields. J Food Saf 2005;25(1):30–42. https://doi.org/10.1111/j.0149-6085.2005.25552.x.
[622] Gómez N, García D, Álvarez I, Raso J, Condón S. A model describing the kinetics of inactivation of Lactobacillus plantarum in a buffer
system of different pH and in orange and apple juice. J Food Eng 2005;70(1):7–14. https://doi.org/10.1016/j.jfoodeng.2004.09.007.
[623] Zhong K, Chen F, Wang Z, Wu J, Liao X, Hu X. Inactivation and kinetic model for the Escherichia coli treated by a co-axial pulsed electric
field. Eur Food Res Technol 2005;221(6):752–8. https://doi.org/10.1007/s00217-005-0015-0.
[624] Zhong KUI, et al. Kinetics of inactivation of Escherichia coli in carrot juice by pulsed electric field. J Food Process Eng 2005;28(6):595–609.
https://doi.org/10.1111/j.1745-4530.2005.00041.x.
[625] Selma MV, Salmeron MC, Valero M, Fernandez PS. Efficacy of pulsed electric fields for Listeria monocytogenes inactivation and control in
horchata. J Food Saf 2006;26(2):137–49. https://doi.org/10.1111/j.1745-4565.2006.00038.x.
[626] Zhang H, Wang Z, Yang RJ, Xu SY. Inactivation of microorganisms in cloudy ginkgo (Ginkgo biloba Linn.) juice by pulsed electric fields.
Food Sci Technol Int 2016;13(2):83–90. https://doi.org/10.1177/1082013207078522.
[627] Charles-Rodríguez AV, Nevárez-Moorillón GV, Zhang QH, Ortega-Rivas E. Comparison of thermal processing and pulsed electric fields
treatment in pasteurization of apple juice. Food Bioprod Process 2007;85(2):93–7. https://doi.org/10.1205/fbp06045.
[628] Mosqueda-Melgar J, Raybaudi-Massilia RM, Martin-Belloso O. Influence of treatment time and pulse frequency on Salmonella Enteritidis,
Escherichia coli and Listeria monocytogenes populations inoculated in melon and watermelon juices treated by pulsed electric fields. Int J
Food Microbiol Jun 30 2007;117(2):192–200. https://doi.org/10.1016/j.ijfoodmicro.2007.04.009.
[629] Mosqueda-Melgar J, Raybaudi-Massilia RM, Martin-Belloso O. Combination of high-intensity pulsed electric fields with natural an-
timicrobials to inactivate pathogenic microorganisms and extend the shelf-life of melon and watermelon juices. Food Microbiol May
2008;25(3):479–91. https://doi.org/10.1016/j.fm.2008.01.002.
[630] Evrendilek GA, Tok FM, Soylu EM, Soylu S. Inactivation of Penicillum expansum in sour cherry juice, peach and apricot nectars by pulsed
electric fields. Food Microbiol Aug 2008;25(5):662–7. https://doi.org/10.1016/j.fm.2008.03.009.
[631] Gachovska TK, Kumar S, Thippareddi H, Subbiah J, Williams F. Ultraviolet and pulsed electric field treatments have additive effect on
inactivation of E. coli in apple juice. J Food Sci Nov 2008;73(9):M412–7. https://doi.org/10.1111/j.1750-3841.2008.00956.x.
[632] Evrendilek G, Tok F, Soylu EM, Soylu S. Effect of pulsed electric fields on germination tube elongation and spore germination of Botrytis
cinerea inoculated into sour cherry juice, apricot and peach nectars. Ital J Food Sci 2009;21:171–82.
[633] García D, Somolinos M, Hassani M, Álvarez I, Pagán R. Modeling the inactivation kinetics of Escherichia coli O157:H7 during the storage
under refrigeration of apple juice treated by pulsed electric fields. J Food Saf 2009;29(4):546–63. https://doi.org/10.1111/j.1745-4565.2009.
00176.x.
[634] McNamee C, Noci F, Cronin DA, Lyng JG, Morgan DJ, Scannell AG. PEF based hurdle strategy to control Pichia fermentans, Listeria
innocua and Escherichia coli k12 in orange juice. Int J Food Microbiol Mar 31 2010;138(1–2):13–8. https://doi.org/10.1016/j.ijfoodmicro.
2009.12.001.
[635] Gurtler JB, Rivera RB, Zhang HQ, Geveke DJ. Selection of surrogate bacteria in place of E. coli O157:H7 and Salmonella Typhimurium for
pulsed electric field treatment of orange juice. Int J Food Microbiol Apr 30 2010;139(1–2):1–8. https://doi.org/10.1016/j.ijfoodmicro.2010.
02.023.
[636] Gurtler JB, Bailey RB, Geveke DJ, Zhang HQ. Pulsed electric field inactivation of E. coli O157:H7 and non-pathogenic surrogate E. coli in
strawberry juice as influenced by sodium benzoate, potassium sorbate, and citric acid. Food Control 2011;22(10):1689–94. https://doi.org/
10.1016/j.foodcont.2011.03.029.
261
[637] Saldana G, Puertolas E, Monfort S, Raso J, Alvarez I. Defining treatment conditions for pulsed electric field pasteurization of apple juice. Int
J Food Microbiol Nov 15 2011;151(1):29–35. https://doi.org/10.1016/j.ijfoodmicro.2011.07.033.
[638] Palgan I, et al. Combined effect of selected non-thermal technologies on Escherichia coli and Pichia fermentans inactivation in an apple and
cranberry juice blend and on product shelf life. Int J Food Microbiol Nov 15 2011;151(1):1–6. https://doi.org/10.1016/j.ijfoodmicro.2011.
07.019.
[639] Marx G, Moody A, Bermudez-Aguirre D. A comparative study on the structure of Saccharomyces cerevisiae under nonthermal technologies:
high hydrostatic pressure, pulsed electric fields and thermo-sonication. Int J Food Microbiol Dec 15 2011;151(3):327–37. https://doi.org/10.
1016/j.ijfoodmicro.2011.09.027.
[640] Ait-Ouazzou A, Espina L, García-Gonzalo D, Pagán R. Synergistic combination of physical treatments and carvacrol for Escherichia coli
O157:H7 inactivation in apple, mango, orange, and tomato juices. Food Control 2013;32(1):159–67. https://doi.org/10.1016/j.foodcont.2012.
11.036.
[641] Huang K, Yu L, Wang W, Gai L, Wang J. Comparing the pulsed electric field resistance of the microorganisms in grape juice: application of
the Weibull model. Food Control 2014;35(1):241–51. https://doi.org/10.1016/j.foodcont.2013.07.011.
[642] Moody A, Marx G, Swanson BG, Bermúdez-Aguirre D. A comprehensive study on the inactivation of Escherichia coli under nonthermal
technologies: high hydrostatic pressure, pulsed electric fields and ultrasound. Food Control 2014;37(1):305–14. https://doi.org/10.1016/j.
foodcont.2013.09.052.
[643] Timmermans RA, Nierop Groot MN, Nederhoff AL, van Boekel MA, Matser AM, Mastwijk HC. Pulsed electric field processing of dif-
ferent fruit juices: impact of pH and temperature on inactivation of spoilage and pathogenic micro-organisms. Int J Food Microbiol Mar 3
2014;173:105–11. https://doi.org/10.1016/j.ijfoodmicro.2013.12.022.
[644] Huang K, Jiang T, Wang W, Gai L, Wang J. A comparison of pulsed electric field resistance for three microorganisms with different biological
factors in grape juice via numerical simulation. Food Bioprocess Technol 2014;7(7):1981–95. https://doi.org/10.1007/s11947-014-1272-3.
[645] Jin TZ, Guo M, Yang R. Combination of pulsed electric field processing and antimicrobial bottle for extending microbiological shelf-life of
pomegranate juice. Innov Food Sci Emerg Technol 2014;26:153–8. https://doi.org/10.1016/j.ifset.2014.07.011.
[646] Agcam E, Akyildiz A, Evrendilek GA. Effects of PEF and heat pasteurization on PME activity in orange juice with regard to a new inactiva-
tion kinetic model. Food Chem Dec 15 2014;165:70–6. https://doi.org/10.1016/j.foodchem.2014.05.097.
[647] Bansal V, Sharma A, Ghanshyam C, Singla ML, Kim K-H. Influence of pulsed electric field and heat treatment on Emblica officinalis juice
inoculated with Zygosaccharomyces bailii. Food Bioprod Process 2015;95:146–54. https://doi.org/10.1016/j.fbp.2015.05.005.
[648] Tao X, Chen J, Li L, Zhao L, Zhang M, Sun A. Influence of pulsed electric field on Escherichia coli and Saccharomyces cerevisiae. Int J
Food Prop 2014;18(7):1416–27. https://doi.org/10.1080/10942912.2014.917098.
[649] Geveke DJ, Aubuchon I, Zhang HQ, Boyd G, Sites JE, Bigley ABW. Validation of a pulsed electric field process to pasteurize strawberry
purée. J Food Eng 2015;166:384–9. https://doi.org/10.1016/j.jfoodeng.2015.05.008.
[650] Timmermans RA, Nederhoff AL, Nierop Groot MN, van Boekel MA, Mastwijk HC. Effect of electrical field strength applied by PEF
processing and storage temperature on the outgrowth of yeasts and moulds naturally present in a fresh fruit smoothie. Int J Food Microbiol
Aug 2 2016;230:21–30. https://doi.org/10.1016/j.ijfoodmicro.2016.04.014.
[651] de Carvalho RJ, de Souza GT, Pagán E, García-Gonzalo D, Magnani M, Pagán R. Nanoemulsions of Mentha piperita L. essential oil in
combination with mild heat, pulsed electric fields (PEF) and high hydrostatic pressure (HHP) as an alternative to inactivate Escherichia coli
O157: H7 in fruit juices. Innov Food Sci Emerg Technol 2018;48:219–27. https://doi.org/10.1016/j.ifset.2018.07.004.
[652] Timmermans RAH, et al. Moderate intensity pulsed electric fields (PEF) as alternative mild preservation technology for fruit juice. Int J Food
Microbiol Jun 2 2019;298:63–73. https://doi.org/10.1016/j.ijfoodmicro.2019.02.015.
[653] Nierop Groot M, Abee T, van Bokhorst-van de Veen H. Inactivation of conidia from three Penicillium spp. isolated from fruit juices by
conventional and alternative mild preservation technologies and disinfection treatments. Food Microbiol Aug 2019;81:108–14. https://doi.
org/10.1016/j.fm.2018.06.004.
[654] Yildiz S, Pokhrel PR, Unluturk S, Barbosa-Cánovas GV. Identification of equivalent processing conditions for pasteurization of strawberry
juice by high pressure, ultrasound, and pulsed electric fields processing. Innov Food Sci Emerg Technol 2019;57:102195. https://doi.org/10.
1016/j.ifset.2019.102195.
[655] Zhu N, Zhang S-l, Li J-p, Qu C, Sun A-d, Qiao X-l. Design and optimization of a microchip operating at low-voltage pulsed electric field for
juice sterilization. Food Bioprocess Technol 2019;12(10):1696–707. https://doi.org/10.1007/s11947-019-02333-8.
[656] Mendes-Oliveira G, Jin TZ, Campanella OH. Modeling the inactivation of Escherichia coli O157:H7 and Salmonella Typhimurium in juices
by pulsed electric fields: the role of the energy density. J Food Eng 2020;282:110001. https://doi.org/10.1016/j.jfoodeng.2020.110001.
[657] Yildiz S, Pokhrel PR, Unluturk S, Barbosa-Canovas GV. Shelf life extension of strawberry juice by equivalent ultrasound, high pressure, and
pulsed electric fields processes. Food Res Int Feb 2021;140:110040. https://doi.org/10.1016/j.foodres.2020.110040.
[658] Li L, Yang R, Zhao W. The effect of pulsed electric fields (PEF) combined with temperature and natural preservatives on the quality and
microbiological shelf-life of cantaloupe juice. Foods Oct 28 2021;10(11):2606. https://doi.org/10.3390/foods10112606.
[659] Mendes-Oliveira G, Jin TZ, Campanella OH. Microbial safety and shelf-life of pulsed electric field processed nutritious juices and their
potential for commercial production. J Food Process Preserv 2021. https://doi.org/10.1111/jfpp.16249.
[660] Kantala C, Supasin S, Intra P, Rattanadecho P. Evaluation of pulsed electric field and conventional thermal processing for microbial inacti-
vation in Thai orange juice. Foods Apr 12 2022;11(8):1102. https://doi.org/10.3390/foods11081102.
[661] Jin TZ, Aboelhaggag RM, Guo M. Apple juice preservation using combined nonthermal processing and antimicrobial packaging. J Food
Prot Sep 1 2021;84(9):1528–38. https://doi.org/10.4315/JFP-21-035.
[662] Wu S, Xu X, Yang N, Jin Y, Jin Z, Xie Z. Inactivation of Escherichia coli O157:H7 in apple juice via induced electric field (IEF) and its
bactericidal mechanism. Food Microbiol Apr 2022;102:103928. https://doi.org/10.1016/j.fm.2021.103928.
[663] Mitta GS. Inactivation of microorganisms in apple cider using spice powders, extracts and oils as antimicrobials with and without low-energy
pulsed electric field; 2003.
262
[664] Azhu Valappil Z, Fan X, Zhang HQ, Rouseff RL. Impact of thermal and nonthermal processing technologies on unfermented apple cider
aroma volatiles. J Agric Food Chem Feb 11 2009;57(3):924–9. https://doi.org/10.1021/jf803142d.
[665] Ulmer HM, Heinz V, Ganzle MG, Knorr D, Vogel RF. Effects of pulsed electric fields on inactivation and metabolic activity of Lactobacillus
plantarum in model beer. J Appl Microbiol 2002;93(2):326–35. https://doi.org/10.1046/j.1365-2672.2002.01699.x.
[666] Gad A, Jayaram SH. Processing of carbonated beer by pulsed electric fields. IEEE Trans Ind Appl 2015;51(6):4759–65. https://doi.org/10.
1109/tia.2015.2448523.
[667] Mihindukulasuriya SDF, Jayaram SH. Release of electrode materials and changes in organoleptic profiles during the processing of liquid
foods using pulse electric field treatment. IEEE Trans Ind Appl 2020;56(1):711–7. https://doi.org/10.1109/tia.2019.2955659.
[668] Poojary MM, et al. Impact of pulsed electric fields on enzymes. In: Miklavcic D, editor. Handbook of electroporation. Cham: Springer
International Publishing; 2016. p. 1–21, Chapter 173-1.
[669] Zhao W, Yang R. Pulsed electric fields for inactivation of endogenous enzymes in foods. In: Miklavcic D, editor. Handbook of electroporation.
[670] Silva ES, et al. Effect of pulsed electric fields on food constituents. In: Miklavcic D, editor. Handbook of electroporation. Cham: Springer
International Publishing; 2016. p. 1–19, Chapter 31-2.
[671] Zhao W, Yang R, Wang M, Lu R. Effects of pulsed electric fields on bioactive components, colour and flavour of green tea infusions. Int J
Food Sci Technol 2009;44(2):312–21. https://doi.org/10.1111/j.1365-2621.2008.01714.x.
[672] Zhang ZH, Zeng XA, Brennan CS, Brennan M, Han Z, Xiong XY. Effects of pulsed electric fields (PEF) on vitamin C and its antioxidant
properties. Int J Mol Sci Oct 13 2015;16(10):24159–73. https://doi.org/10.3390/ijms161024159.
[673] Zhang S, Sun L, Ju H, Bao Z, Zeng XA, Lin S. Research advances and application of pulsed electric field on proteins and peptides in food.
Food Res Int Jan 2021;139:109914. https://doi.org/10.1016/j.foodres.2020.109914.
[674] Odriozola-Serrano I, Soliva-Fortuny R, Hernández-Jover T, Martín-Belloso O. Carotenoid and phenolic profile of tomato juices processed
by high intensity pulsed electric fields compared with conventional thermal treatments. Food Chem 2009;112(1):258–66. https://doi.org/10.
1016/j.foodchem.2008.05.087.
[675] Liu YY, Zeng XA, Deng Z, Yu SJ, Yamasaki S. Effect of pulsed electric field on the secondary structure and thermal properties of soy protein
isolate. Eur Food Res Technol 2011;233(5):841–50. https://doi.org/10.1007/s00217-011-1580-z.
[676] Reineke K, Schottroff F, Meneses N, Knorr D. Sterilization of liquid foods by pulsed electric fields-an innovative ultra-high temperature
process. Front Microbiol 2015;6(MAY):400. https://doi.org/10.3389/fmicb.2015.00400.
[677] Wang LH, Wang MS, Zeng XA, Liu ZW. Temperature-mediated variations in cellular membrane fatty acid composition of Staphylococcus
aureus in resistance to pulsed electric fields. Biochim Biophys Acta Aug 2016;1858(8):1791–800. https://doi.org/10.1016/j.bbamem.2016.
05.003.
[678] Banaschik R, Burchhardt G, Zocher K, Hammerschmidt S, Kolb JF, Weltmann KD. Comparison of pulsed corona plasma and pulsed electric
fields for the decontamination of water containing Legionella pneumophila as model organism. Bioelectrochemistry Dec 2016;112:83–90.
https://doi.org/10.1016/j.bioelechem.2016.05.006.
[679] Sutcliffe IC. A phylum level perspective on bacterial cell envelope architecture. Trends Microbiol Oct 2010;18(10):464–70. https://doi.org/
10.1016/j.tim.2010.06.005.
[680] Vollmer W, Seligman SJ. Architecture of peptidoglycan: more data and more models. Trends Microbiol Feb 2010;18(2):59–66. https://
doi.org/10.1016/j.tim.2009.12.004.
[681] Mosqueda-Melgar J, Elez-Martinez P, Raybaudi-Massilia RM, Martin-Belloso O. Effects of pulsed electric fields on pathogenic microorgan-
isms of major concern in fluid foods: a review. Crit Rev Food Sci Nutr Sep 2008;48(8):747–59. https://doi.org/10.1080/10408390701691000.
[682] Drees KP, Abbaszadegan M, Maier RM. Comparative electrochemical inactivation of bacteria and bacteriophage. Water Res May
2003;37(10):2291–300. https://doi.org/10.1016/S0043-1354(03)00009-5.
[683] Liu C, et al. Conducting nanosponge electroporation for affordable and high-efficiency disinfection of bacteria and viruses in water. Nano
Lett Sep 11 2013;13(9):4288–93. https://doi.org/10.1021/nl402053z.
[684] Lee C, Kim J, Yoon J. Inactivation of MS2 bacteriophage by streamer corona discharge in water. Chemosphere Feb 2011;82(8):1135–40.
https://doi.org/10.1016/j.chemosphere.2010.11.036.
[685] Jurgelevičiūtė J, Bičkovas N, Sakalauskas A, Novickij V, Smirnovas V, Lastauskienė E. Effects of pulsed electric fields on yeast with prions
and the structure of amyloid fibrils. Appl Sci 2021;11(6):2684. https://doi.org/10.3390/app11062684.
[686] Yuan AH, Hochschild A. A bacterial global regulator forms a prion. Science Jan 13 2017;355(6321):198–201. https://doi.org/10.1126/
science.aai7776.
[687] Fleming E, Yuan AH, Heller DM, Hochschild A. A bacteria-based genetic assay detects prion formation. Proc Natl Acad Sci USA Mar 5
2019;116(10):4605–10. https://doi.org/10.1073/pnas.1817711116.
[688] Arroyo C, Lyng JG. Pulsed electric fields in hurdle approaches for microbial inactivation. In: Miklavcic D, editor. Handbook of electropora-
[689] García D, Gómez N, Raso J, Pagán R. Bacterial resistance after pulsed electric fields depending on the treatment medium pH. Innov Food
Sci Emerg Technol 2005/12/01;6(4):388–95. https://doi.org/10.1016/j.ifset.2005.04.003.
[690] Guerrero-Beltrán JA, Barbosa-Cánovas GV. Advantages and limitations on processing foods by UV light. Food Sci Technol Int
2016;10(3):137–47. https://doi.org/10.1177/1082013204044359.
[691] Tanino T, Hirosawa M, Moteki R, Matsui M, Ohshima T. Engineering of pulsed electric field treatment using carbon materials as electrode
and application to pasteurization of sake. J Electrost 2020;104:103424. https://doi.org/10.1016/j.elstat.2020.103424.
[692] Yogesh K. Pulsed electric field processing of egg products: a review. J Food Sci Technol Feb 2016;53(2):934–45. https://doi.org/10.1007/
s13197-015-2061-3.
[693] Pashley RM, Rzechowicz M, Pashley LR, Francis MJ. De-gassed water is a better cleaning agent. J Phys Chem B Jan 27 2005;109(3):1231–8.
https://doi.org/10.1021/jp045975a.
263
[694] Francis MJ. Effect of degassing on the electrical conductivity of pure water and potassium chloride solutions. J Phys Chem C
2008;112(37):14563–9. https://doi.org/10.1021/jp8037686.
[695] Wetz DA, Mankowski JJ, Dickens JC, Kristiansen M. The impact of field enhancements and charge injection on the pulsed breakdown
strength of water. IEEE Trans Plasma Sci 2006;34(5):1670–9. https://doi.org/10.1109/tps.2006.881891.
[696] Góngora-Nieto MM, Pedrow PD, Swanson BG, Barbosa-Cánovas GV. Impact of air bubbles in a dielectric liquid when subjected to high
field strengths. Innov Food Sci Emerg Technol 2003;4(1):57–67. https://doi.org/10.1016/s1466-8564(02)00067-x.
[697] Sunka P, et al. Generation of chemically active species by electrical discharges in water. Plasma Sources Sci Technol 1999;8(2):258–65.
https://doi.org/10.1088/0963-0252/8/2/006.
[698] Sun B, Xin Y, Zhu X, Gao Z, Yan Z, Ohshima T. Effects of shock waves, ultraviolet light, and electric fields from pulsed discharges in water
on inactivation of Escherichia coli. Bioelectrochemistry Apr 2018;120:112–9. https://doi.org/10.1016/j.bioelechem.2017.11.011.
[699] U.S. Department of Health and Human Services and Food and Drug Administration. Grade A Pasteurized Milk Ordinance (PMO).
[700] Clifford V, et al. What are optimal bacteriological screening test cut-offs for pasteurized donor human milk intended for feeding preterm
infants? J Hum Lact Feb 2021;37(1):43–51. https://doi.org/10.1177/0890334420981013.
[701] Mabrook MF, Petty MC. Effect of composition on the electrical conductance of milk. J Food Eng 2003;60(3):321–5. https://doi.org/10.1016/
s0260-8774(03)00054-2.
[702] Lawton BA, Pethig R. Determining the fat content of milk and cream using AC conductivity measurements. Meas Sci Technol
1993;4(1):38–41. https://doi.org/10.1088/0957-0233/4/1/007.
[703] Bijl E, van Valenberg HJ, Huppertz T, van Hooijdonk AC. Protein, casein, and micellar salts in milk: current content and historical perspec-
tives. J Dairy Sci Sep 2013;96(9):5455–64. https://doi.org/10.3168/jds.2012-6497.
[704] Prentice JH. The conductivity of milk—the effect of the volume and degree of dispersion of the fat. J Dairy Res 2009;29(2):131–9. https://
doi.org/10.1017/s0022029900017738.
[705] Dhungana P, Truong T, Bansal N, Bhandari B. Effect of fat globule size and addition of surfactants on whippability of native and homogenised
dairy creams. Int Dairy J 2020/06/01;105:104671. https://doi.org/10.1016/j.idairyj.2020.104671.
[706] Ho TM, Dhungana P, Bhandari B, Bansal N. Effect of the native fat globule size on foaming properties and foam structure of milk. J Food
Eng 2021/02/01;291:110227. https://doi.org/10.1016/j.jfoodeng.2020.110227.
[707] Sharma P, Oey I, Everett DW. Effect of pulsed electric field processing on the functional properties of bovine milk. Trends Food Sci Technol
2014;35(2):87–101. https://doi.org/10.1016/j.tifs.2013.11.004.
[708] Truong T, Palmer M, Bansal N, Bhandari B. Effect of milk fat globule size on the physical functionality of dairy products. Cham: Springer
International Publishing AG; 2015.
[709] Dallas DC, Murray NM, Gan J. Proteolytic systems in milk: perspectives on the evolutionary function within the mammary gland and the
infant. J Mammary Gland Biol Neoplasia Dec 2015;20(3–4):133–47. https://doi.org/10.1007/s10911-015-9334-3.
[710] Chen PW, Lin YL, Huang MS. Profiles of commensal and opportunistic bacteria in human milk from healthy donors in Taiwan. J Food Drug
Anal Oct 2018;26(4):1235–44. https://doi.org/10.1016/j.jfda.2018.03.004.
[711] Adhikari BM, Truong T, Bansal N, Bhandari B. Use of gases in dairy manufacturing: a review. Crit Rev Food Sci Nutr 2018;58(15):2557–69.
https://doi.org/10.1080/10408398.2017.1333488.
[712] Chantry CJ, et al. Effect of flash-heat treatment on antimicrobial activity of breastmilk. Breastfeed Med Jun 2011;6(3):111–6. https://doi.org/
10.1089/bfm.2010.0078.
[713] Gopal N, Hill C, Ross PR, Beresford TP, Fenelon MA, Cotter PD. The prevalence and control of bacillus and related spore-forming bacteria
in the dairy industry. Front Microbiol 2015;6:1418. https://doi.org/10.3389/fmicb.2015.01418.
[714] Grady EN, MacDonald J, Liu L, Richman A, Yuan ZC. Current knowledge and perspectives of Paenibacillus: a review. Microb Cell Fact
Dec 1 2016;15(1):203. https://doi.org/10.1186/s12934-016-0603-7.
[715] Scheldeman P, Herman L, Foster S, Heyndrickx M. Bacillus sporothermodurans and other highly heat-resistant spore formers in milk. J Appl
Microbiol Sep 2006;101(3):542–55. https://doi.org/10.1111/j.1365-2672.2006.02964.x.
[716] Wiencek KM, Klapes NA, Foegeding PM. Adhesion ofbacillusspores to inanimate materials: effects of substratum and spore hydrophobic-
ityab. Biofouling 2009;3(2):139–49. https://doi.org/10.1080/08927019109378168.
[717] Husmark U, Rönner U. The influence of hydrophobic, electrostatic and morphologic properties on the adhesion of Bacillus spores. Biofouling
1992;5(4):335–44. https://doi.org/10.1080/08927019209378253.
[718] Pol IE, van Arendonk WG, Mastwijk HC, Krommer J, Smid EJ, Moezelaar R. Sensitivities of germinating spores and carvacrol-adapted
vegetative cells and spores of Bacillus cereus to nisin and pulsed-electric-field treatment. Appl Environ Microbiol Apr 2001;67(4):1693–9.
https://doi.org/10.1128/AEM.67.4.1693-1699.2001.
[719] Choi J, et al. Inactivation of spores using pulsed electric field in a pressurized flow system. J Appl Phys 2008;104(9):094701. https://
doi.org/10.1063/1.3006440.
[720] Pothakamury UR, Monsalve-Gonzàlez A, Barbosa-Cánovas GV, Swanson BG. Inactivation of Escherichia coli and Staphylococcus aureus
in model foods by pulsed electric field technology. Food Res Int 1995;28(2):167–71. https://doi.org/10.1016/0963-9969(95)90801-g.
[721] Roos YH. Water and pathogenic viruses inactivation—food engineering perspectives. Food Eng Rev 2020;12(3):251–67. https://doi.org/10.
1007/s12393-020-09234-z.
[722] Walker GJ, et al. SARS-CoV-2 in human milk is inactivated by Holder pasteurisation but not cold storage. J Paediatr Child Health Dec
2020;56(12):1872–4. https://doi.org/10.1111/jpc.15065.
[723] Bidawid S, Farber JM, Sattar SA, Hayward S. Heat inactivation of hepatitis A virus in dairy foods. J Food Prot Apr 2000;63(4):522–8. https://
doi.org/10.4315/0362-028x-63.4.522.
[724] Hewitt J, Rivera-Aban M, Greening GE. Evaluation of murine norovirus as a surrogate for human norovirus and hepatitis A virus in heat
inactivation studies. J Appl Microbiol Jul 2009;107(1):65–71. https://doi.org/10.1111/j.1365-2672.2009.04179.x.
264
[725] Terpstra FG, et al. Antimicrobial and antiviral effect of high-temperature short-time (HTST) pasteurization applied to human milk. Breastfeed
Med Mar 2007;2(1):27–33. https://doi.org/10.1089/bfm.2006.0015.
[726] Sauerbrei A, Wutzler P. Testing thermal resistance of viruses. Arch Virol 2009;154(1):115–9. https://doi.org/10.1007/s00705-008-0264-x.
[727] Hirneisen KA, Black EP, Cascarino JL, Fino VR, Hoover DG, Kniel KE. Viral inactivation in foods: a review of traditional and novel
food-processing technologies. Compr Rev Food Sci Food Saf Jan 2010;9(1):3–20. https://doi.org/10.1111/j.1541-4337.2009.00092.x.
[728] Morren J, Roodenburg B, de Haan SWH. Electrochemical reactions and electrode corrosion in pulsed electric field (PEF) treatment chambers.
Innov Food Sci Emerg Technol 2003;4(3):285–95. https://doi.org/10.1016/s1466-8564(03)00041-9.
[729] L’Anthoën NC, Ingledew WM. Heat resistance of bacteria in alcohol-free beer. J Am Soc Brew Chem 1996/01/01;54(1):32–6. https://
doi.org/10.1094/asbcj-54-0032.
[730] Zufall C, Tyrell T. The influence of heavy metal ions on beer flavour stability. J Inst Brew 2008;114(2):134–42. https://doi.org/10.1002/j.
2050-0416.2008.tb00318.x.
[731] The International Organization of Vine and Wine. Resolution OIV-OENO 634-2020 [Online]. Available: https://www.oiv.int/public/medias/
7594/oiv-oeno-634-2020-en.pdf.
[732] Fauster T, et al. Impact of a combined pulsed electric field (PEF) and enzymatic mash treatment on yield, fermentation behaviour and
composition of white wine. Eur Food Res Technol 2020;246(3):609–20. https://doi.org/10.1007/s00217-020-03427-w.
[733] Saldana G, Cebrian G, Abenoza M, Sanchez-Gimeno C, Alvarez I, Raso J. Assessing the efficacy of PEF treatments for improving polyphenol
extraction during red wine vinifications. Innov Food Sci Emerg Technol Feb 2017;39:179–87. https://doi.org/10.1016/j.ifset.2016.12.008.
[734] Lonvaud-Funel A. Lactic acid bacteria in the quality improvement and depreciation of wine. Antonie Van Leeuwenhoek Jul-Nov
1999;76(1–4):317–31. https://doi.org/10.1023/A:1002088931106.
[735] Food and drug regulations [Online] Available: https://laws-lois.justice.gc.ca/eng/regulations/c.r.c.,_c._870/index.html, 2022.
[736] Heresztyn T. Metabolism of volatile phenolic compounds from hydroxycinnamic acids by Brettanomyces yeast. Arch Microbiol
1986;146(1):96–8. https://doi.org/10.1007/bf00690165.
[737] Chatonnet P, Dubourdieu D, Boidron JN. The influence of Brettanomyces/Dekkera sp. yeasts and lactic acid bacteria on the ethylphenol
content of red wines. Am J Enol Vitic 1995;46(4):463–8 [Online]. Available: https://www.ajevonline.org/content/ajev/46/4/463.full.pdf.
[738] Peter G, Dlauchy D, Tobias A, Fulop L, Podgorsek M, Cadez N. Brettanomyces acidodurans sp. nov., a new acetic acid producing yeast
species from olive oil. Antonie Van Leeuwenhoek May 2017;110(5):657–64. https://doi.org/10.1007/s10482-017-0832-8.
[739] Freer SN. Acetic acid production by Dekkera/Brettanomyces yeasts. World J Microbiol Biotechnol 2002;18(3):271–5. https://doi.org/10.
1023/a:1014927129259.
[740] Couto JA, Neves F, Campos F, Hogg T. Thermal inactivation of the wine spoilage yeasts Dekkera/Brettanomyces. Int J Food Microbiol Oct
25 2005;104(3):337–44. https://doi.org/10.1016/j.ijfoodmicro.2005.03.014.
[741] Cocolin L, Rantsiou K, Iacumin L, Zironi R, Comi G. Molecular detection and identification of Brettanomyces/Dekkera bruxellensis and
Brettanomyces/Dekkera anomalus in spoiled wines. Appl Environ Microbiol Mar 2004;70(3):1347–55. https://doi.org/10.1128/AEM.70.3.
1347-1355.2004.
[742] Usseglio-Tomasset L. Properties and use of sulphur dioxide. Food Addit Contam Sep-Oct 1992;9(5):399–404. https://doi.org/10.1080/
02652039209374090.
[743] Henick-Kling, Edinger, Daniel, Monk. Selective effects of sulfur dioxide and yeast starter culture addition on indigenous yeast populations
and sensory characteristics of wine. J Appl Microbiol 1998;84(5):865–76. https://doi.org/10.1046/j.1365-2672.1998.00423.x.
[744] Picinelli A, Bakker J, Bridle P. Model wine solutions: effect of sulphur dioxide on colour and composition during ageing. Vitis
1994;33(1):31–5.
[745] Bakker J, Bridle P, Bellworthy SJ, Garcia-Viguera C, Reader HP, Watkins SJ. Effect of sulphur dioxide and must extraction on colour,
phenolic composition and sensory quality of red table wine. J Sci Food Agric 1998;78(3):297–307. https://doi.org/10.1002/(SICI)1097-
0010(199811)78:3<297::AID-JSFA117>3.0.CO;2-G.
[746] Saldaña G, Luengo E, Puértolas E, Álvarez I, Raso J. Pulsed electric fields in wineries: potential applications. In: Miklavcic D, editor.
Handbook of electroporation. Cham: Springer International Publishing; 2016. p. 1–18, Chapter 155-1.
[747] Novickij V, et al. Low concentrations of acetic and formic acids enhance the inactivation of Staphylococcus aureus and Pseudomonas
aeruginosa with pulsed electric fields. BMC Microbiol Apr 3 2019;19(1):73. https://doi.org/10.1186/s12866-019-1447-1.
[748] Heinz V, Knorr D. Effect of pH, ethanol addition and high hydrostatic pressure on the inactivation of Bacillus subtilis by pulsed electric
fields. Innov Food Sci Emerg Technol 2000;1(2):151–9. https://doi.org/10.1016/s1466-8564(00)00013-8.
[749] Iu J, Mittal GS, Griffiths MW. Reduction in levels of Escherichia coli O157:H7 in apple cider by pulsed electric fields. J Food Prot
2001;64(7):964–9. https://doi.org/10.4315/0362-028X-64.7.964.
[750] Kato T, Shimoda M, Suzuki J, Kawaraya A, Igura N, Hayakawa I. Changes in the odors of squeezed apple juice during thermal processing.
Food Res Int 2003;36(8):777–85. https://doi.org/10.1016/s0963-9969(03)00072-3.
[751] Chen BH, Peng HY, Chen HE. Changes of carotenoids, color, and vitamin A contents during processing of carrot juice. J Agric Food Chem
2002;43(7):1912–8. https://doi.org/10.1021/jf00055a029.
[752] Buckow R, Ng S, Toepfl S. Pulsed electric field processing of orange juice: a review on microbial, enzymatic, nutritional, and sensory quality
and stability. Compr Rev Food Sci Food Saf Sep 2013;12(5):455–67. https://doi.org/10.1111/1541-4337.12026.
[753] Pallarés N, Barba FJ, Berrada H, Tolosa J, Ferrer E. Pulsed electric fields (PEF) to mitigate emerging mycotoxins in juices and smoothies.
Appl Sci 2020;10(19):6989. https://doi.org/10.3390/app10196989.
[754] Doyle MP. Escherichia coli O157:H7 and its significance in foods. Int J Food Microbiol Apr 1991;12(4):289–301. https://doi.org/10.1016/
0168-1605(91)90143-d.
[755] Conner DE, Kotrola JS. Growth and survival of Escherichia coli O157:H7 under acidic conditions. Appl Environ Microbiol Jan
1995;61(1):382–5. https://doi.org/10.1128/aem.61.1.382-385.1995.
265
[756] Vamos-Vigyazo L. Polyphenol oxidase and peroxidase in fruits and vegetables. Crit Rev Food Sci Nutr 1981;15(1):49–127. https://doi.org/
10.1080/10408398109527312.
[757] Foegeding PM, Leasor SB. Heat resistance and growth of Listeria monocytogenes in liquid whole egg. J Food Prot Jan 1990;53(1):9–14.
https://doi.org/10.4315/0362-028X-53.1.9.
[758] Uysal RS, Boyaci IH, Sumnu G. Determination of pasteurization treatment of liquid whole egg by measuring physical and rheological
properties of cake cream. J Food Process Eng 2019;42(6):e13167. https://doi.org/10.1111/jfpe.13167.
[759] Molina-Hoppner A, Doster W, Vogel RF, Ganzle MG. Protective effect of sucrose and sodium chloride for Lactococcus lactis during sublethal
and lethal high-pressure treatments. Appl Environ Microbiol Apr 2004;70(4):2013–20. https://doi.org/10.1128/AEM.70.4.2013-2020.2004.
[760] Vyrides I, Stuckey DC. Compatible solute addition to biological systems treating waste/wastewater to counteract osmotic and other environ-
mental stresses: a review. Crit Rev Biotechnol Nov 2017;37(7):865–79. https://doi.org/10.1080/07388551.2016.1266460.
[761] Sleator RD, Hill C. Compatible solutes: the key to Listeria’s success as a versatile gastrointestinal pathogen? Gut Pathogens Dec 10
2010;2(1):20. https://doi.org/10.1186/1757-4749-2-20.
[762] da Costa MS, Santos H, Galinski EA. An overview of the role and diversity of compatible solutes in Bacteria and Archaea. Adv Biochem
Eng Biotechnol 1998;61:117–53. https://doi.org/10.1007/BFb0102291.
[763] Empadinhas N, da Costa MS. Diversity, biological roles and biosynthetic pathways for sugar-glycerate containing compatible solutes in
bacteria and archaea. Environ Microbiol Aug 2011;13(8):2056–77. https://doi.org/10.1111/j.1462-2920.2010.02390.x.
[764] Pleitner A, Zhai Y, Winter R, Ruan L, McMullen LM, Ganzle MG. Compatible solutes contribute to heat resistance and ribosome stability
in Escherichia coli AW1.7. Biochim Biophys Acta Dec 2012;1824(12):1351–7. https://doi.org/10.1016/j.bbapap.2012.07.007.
[765] Ng H, Garibaldi JA. Death of Staphylococcus aureus in liquid whole egg near pH 8. Appl Microbiol Jun 1975;29(6):782–6. https://doi.org/
10.1128/am.29.6.782-786.1975.
[766] Darvishi H, Khoshtaghaza MH, Zarein M, Azadbakht M. Ohmic processing of liquid whole egg, white egg and yolk. CIGR E-J
2012;14(4):224–30.
[767] Techer C, Daoud A, Madec MN, Gautier M, Jan S, Baron F. Microbial quality of industrial liquid egg white: assumptions on spoiling issues
in egg-based chilled desserts. J Food Sci Feb 2015;80(2):M389–98. https://doi.org/10.1111/1750-3841.12764.
[768] Stadelman WJ. Contaminants of liquid egg products. In: Board RG, Fuller R, editors. Microbiology of the avian egg. Boston, MA: Springer
US; 1994. p. 139–51, Chapter 7.
[769] Abenoza M, Benito M, Saldaña G, Álvarez I, Raso J, Sánchez-Gimeno AC. Effects of pulsed electric field on yield extraction and quality of
olive oil. Food Bioprocess Technol 2012;6(6):1367–73. https://doi.org/10.1007/s11947-012-0817-6.
[770] Grahl T, Markl H. Killing of microorganisms by pulsed electric fields. Appl Microbiol Biotechnol Mar 1996;45(1–2):148–57. https://doi.
org/10.1007/s002530050663.
[771] Atılgan MR, Unluturk S. Rheological properties of liquid egg products (LEPS). Int J Food Prop 04/01 2008;11(2):296–309. https://doi.org/
10.1080/10942910701329658.
[772] Abbasnezhad B, Hamdami N, Monteau JY, Vatankhah H. Numerical modeling of heat transfer and pasteurizing value during thermal pro-
cessing of intact egg. Food Sci Nutr Jan 2016;4(1):42–9. https://doi.org/10.1002/fsn3.257.
[773] Vilela A. Use of nonconventional yeasts for modulating wine acidity. Fermentation 2019;5(1). https://doi.org/10.3390/fermentation5010027.
[774] Fabjanowicz M, Kosek K, Płotka-Wasylka J, Namieśnik J. Evaluation of the influence of grapevine growing conditions on wine quality.
Monatsh Chem 2019/09/01;150(9):1579–84. https://doi.org/10.1007/s00706-019-02454-y.
[775] Lee JH, et al. Physicochemical characteristics and electric conductivity of various fruit wines. Int Food Res J 2013;20(6):2987–93.
[776] Sáez ÁB, Lopez-Cordon EN, Fernández FC, Sáez SB. Refrigeration in winemaking industry. In: Refrigeration. IntechOpen; 2017, Chapter
4.
[777] Ağçam E, Akyıldız A, Dündar B. Thermal pasteurization and microbial inactivation of fruit juices. In: Rajauria G, Tiwari BK, editors. Fruit
juices. San Diego: Academic Press; 2018. p. 309–39.
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Advances in pulsed electric stimuli as a physical method for treating

Received 24 January 2023; accepted 28 January 2023

Keywords: Pulsed electric field; Liquid foods; Safety; Quality

3.4. Weibull distribution model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216

2. Pulse waveform, generation, and delivery

3. Inactivation kinetics of microorganisms by PEF

3.1. Bigelow model

First-order equation which uses treatment time [128,129].

3.2. Peleg model

A sigmoid function which uses the electric field strength [128,130,131].

3.3. Hülsheger and Niemann model

Uses electric field strength to characterise model [120].

3.4. Weibull distribution model

4. Electroporation, electropermeabilization, and associated mechanisms

Epore (r) = 2γ r − πr 2 (17)

Epore energy (r, Vm ) = 2γ r − πr 2 − 0.5CLW Vm2 πr 2 (18)

Vm (t) = −fs .Ef ield (t) .R. cos θ + Vrest (26)

5. Quality and safety of liquid foods treated with PEFs

• Smooth electrodes which avoid sharp points [85].

• Pressurising the sample to prevent bubble formation [85,695].

5.5. Fruit juices and cider

5.6. Liquid egg

Declaration of competing interest

Liquid Egg Whole egg 8.1-8.3 [480] -

Physics of Life Reviews 44 (2023) 207–266

Physics of Life Reviews 44 (2023) 207–266

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Epore (r) = 2γ r − πr 2 (17)

Epore energy (r, Vm ) = 2γ r − πr 2 − 0.5CLW Vm2 πr 2 (18)