ISSN 1061933X, Colloid Journal, 2010, Vol. 72, No. 6, pp. 830836. Pleiades Publishing, Ltd., 2010.

Original Russian Text V.I. Podolska, L.N. Yakubenko, Z.R. Ulberg, V.N. Ermakov, N.I. Grischenko, 2010, published in Kolloidnyi Zhurnal, 2010, Vol. 72, No. 6, pp. 822829.
830
INTRODUCTION
The experimental study of the effect of weak elec
tromagnetic (electric) fields on cell suspensions runs
into the complexity of the choice of unbiased criteria
of their action. Colloidal and biochemical parameters
of cells are sensitive indicators of external actions on
the microbial cell that allow us to evaluate their
response to stress factors and, at the same time, to
clarify the mechanism of cell adaptation to these fac
tors. It is known that biocolloids differ from inorganic
colloidal systems by the nonequilibrium state associ
ated with the permanent exchange of substance and
energy with external system [1]. The electrical charac
teristic of living cells is the transmembrane potential
(TMP). The main contribution to the degree of power
capacity of membrane is introduced by its respiratory
centers (RCs). The transfer of electrons from substrate
to oxygen and the release of energy needed for ATP
synthesis occur in these nanosized subunits. In turn,
TMP is a binding link between electrosurface proper
ties and energydriven membrane processes [2].
Experimental data that support the stimulating
effect of weak electric and electromagnetic fields on
microorganisms are available. Data on the accelera
tion of growth, the adaptation to stress conditions, the
release of excess metabolites, and the activation of
enzymatic processes are reported [3, 4]. In the major
ity of cases, the mechanisms of these actions are not
entirely known and electromagnetic fields are
imposed empirically.
We demonstrated earlier that the treatment of
P. fluorescens suspension by d.c. electric field with
strength of 25 V/cm (the duration is 1530 s, the
pause is 3060 s) in solutions of copper and silver cya
nide complexes improved the kinetics and enhanced
the efficiency of cyanide destruction [5, 6]. Similar
stimulating effect is also caused by the halfhour treat
ment of P. fluorescens inoculate by the electromagnetic
field with the frequency of 54 GHz and the power of
10 mW/cm
2
before the seeding into cyanidecontain
ing solution [7]. In subsequent experiments, we dis
covered the respiratory response of culturedestructor
to the treatment by weak pulse electric fields [8]. The
data obtained confirmed the hypothesis according to
which the conditions of electron transfer along the
respiratory chain in RCs are changed upon the exposi
tion of bacterial cell to external pulse electric field
(PEF). This transfer is adequately described by the
mechanism of nonlinear resonance tunneling (NRT)
[9, 10]. In this case, depending on the number of func
tioning RCs, both the blocking and unblocking of
electron transfer chains that determine the magnitude
and direction of respiratory response can take place.
The goal of this work is to study the effect of weak
nonuniform external pulse electric field on the
destruction of silver cyanide complex by Pseudomonas
resistant bacteria. In addition to destruction, we mea
sured the parameters of surface and electrosurface
properties of bacteria. Prior to seeding into cyanide
containing solution, bacterial inoculate was subjected
to electrotreatment. Experiments were performed so
Effect of Weak Pulse Electric Fields on Surface Properties
and Destructive Activity of Pseudomonas Bacteria
V. I. Podolska
a
, L. N. Yakubenko
a
, Z. R. Ulberg
a
, V. N. Ermakov
b
, and N. I. Grischenko
a
a
Ovcharenko Institute of Biocolloid Chemistry, National Academy of Sciences of Ukraine,
bulv. Akademika Vernadskogo 42, Kiev142, 03142 Ukraine
b
Bogoliubov Institute of Theoretical Physics, National Academy of Sciences of Ukraine,
ul. Metrologicheskaya 14b, Kiev143, 03143 Ukraine
Received September 21, 2009
AbstractThe effect of weak pulse electric fields on the destructive activity and surface properties of
Pseudomonas bacteria is studied. It is revealed that the physiological response of bacteria on their treatment
by external pulse fields depends on field parameters and is accompanied by the changes in the values of elec
trokinetic potential and hydrophobicity. The treatment of bacteria by the electric field with pulse duration of
110 ms and frequency of 100500 Hz tends to increase by 2030% the destruction of NaAg(CN)
2
. It is sug
gested that changes in surface properties and the enhancement of biochemical activity of bacteria are based
on the metabolic reaction of the cell related to the stimulation or suppression of the bacteria respiration upon
the imposition of external field.
DOI: 10.1134/S1061933X10060153
COLLOID JOURNAL Vol. 72 No. 6 2010
EFFECT OF WEAK PULSE ELECTRIC FIELDS 831
as to exclude changes in the concentration of electron
active components due to electrode reactions.
EXPERIMENTAL
Microorganisms and Growth Conditions
We used Pseudomonas fluorescens RCIM B5040 and
Pseudomonas fluorescens NCIMB 11764 bacteria.
Destructive properties of these cultures are described
in [11, 12]. The cultivation was performed in medium
with the following composition (g/l): glucose, 2.0;
peptone, 0.5; KH
2
PO
4
, 2.0; K
2
HPO
4
, 1.0; MgSO
4

7H
2
O, 0.3; Na
2
CO
3
, 0.5; NaCl, 0.1; and NaCN,
0.094; pH 7.8. In experiments, we used 2,5h culture
(the end of the exponential growth phase).
Procedures of PEF Treatment
Bacteria of the same harvest were separated from
the nutrient medium on a centrifuge at 3700 g, washed
in distilled water, and resuspended in the abovemen
tioned medium that contians NaAg(CN)
2
. Suspension
(3.0 ml) was stirred on a vortex for 1 min, transferred
to the cell, and treated with PEF.
Field treatment was performed in experimental
setup which is schematically shown in [8]. The scheme
composed of dismountable cell made of isolator mate
rial and equipped with stirring unit and system of air
supply via microcompressor. Seven needlelike steel
electrodes were mounted on the cell lid. Flat electrode
of opposite sign was placed beyond the cell and
arranged under its bottom. The distance between elec
trodes is equal to 1 cm. The pulse field was supplied by
a G554 pulse generator (Russia) and controlled with
an oscillograph. Needlelike electrodes with low
capacitance ensure the abrupt switching of the exter
nal electric field.
A nonuniform field with strength modulus E,
which is typical for the chosen point of the cell, is gen
erated upon switching the d.c. voltage. According to
estimations, at the applied voltage of 30 V, the maxi
mum strength of field at the needle tip in the solution
does not exceed 10
3
V/cm. This field gives rise to the
displacement of current carriers, which in turn leads to
the formation of induced electric field
p
oppositely
directed toward the external field. In the absence of
current, these fields are equal by strength and oppo
sitely directed in each point in electrolyte medium.
The balance of field is violated with variations in
applied voltage and the additional displacement of
current carriers (displacement current) is needed to
restore this balance. In general, the balance equation
for an induced field has the form
(1) ( )
p
p
r
1
,
dE
E E
dt
=

where E and
p
are timedependent values and
r
is the
relaxation time (the time of electroneutrality restora
tion). As follows from Eq. (1), at constant E values, we
arrive at
p
= , i.e., the action of external field is com
pletely compensated for by the response of environ
ment. This compensation is violated in the a.c. field
that is related to the delay in the response of medium.
According to Eq. (1), thus emerging total electric field
has the form shown in Fig. 1 (solid line). As can be
seen from this figure, the field produced by voltage
pulse with duration (dashed line) generates compen
sating field

with a certain delay (dotted line). After

switching off the voltage pulse, compensating field

also disappears with a certain delay.

The delay time is determined by relaxation time
r
.
Upon imposing the potential difference, types of
polarization with shorter relaxation times are gener
ated. In 1 mM electrolyte, the relaxation time of ion
atmospheres is equal to ~10
7
s. The relaxation time of
macrostructural (dielectric) polarization related to the
cell heterogeneity varies from 10
8
to 10
3
s. The relax
ation time of surface polarization related to the pres
ence of electrical double layer ranges from 10
3
to 1 s.
At the pulse duration = 10
3
s, polarization effects
with relaxation time
r
< 10
3
s, i.e., the first two of the
aforementioned effects, become important. The total
field between switching a voltage pulse on and off is
equal to zero. Due to the multiplicity of the conditions
of field formation in a microbial suspension associated
0.6
0.4
0
0.2
0.4
0.6
0.8
1.0
1.2
25 20 15 10 5 0 5
0.8
1.0
1.2
0.2
30 35 40 45 50
/
r
1/(f
r
)
t/
r
E(t), E
p
(t), E(t)/E
0
Fig. 1. Time dependence of total field (solid line) gen
erated upon imposition of unipolar rectangular pulse elec
tric field with amplitude
0
and frequency f (dashed line)
on a system. The dotted line shows the induced field E
p
,
is the puls duration, and
r
is the relaxation time.
832
COLLOID JOURNAL Vol. 72 No. 6 2010
PODOLSKA et al.
with spatial nonuniformity, some inductance of the
system, and polarization of a different origin, it rather
difficult to evaluate the maximal strength of the filed;
however, in all cases, this value is proportional to the
applied voltage. The total field acting on the cells was
identical in all events of switching and was indepen
dent of the pulse frequency.
Microbial Destruction and Analysis
for Cyanide Content
We used stable cyanide complex NaAg(CN)
2
with

= 8 10
22
[13]. The CN

concentration was deter

mined by photometry at = 590 nm using pyridine
barbiturate reagent. The sensitivity of the method was
2 10
3
mM. After the PEF treatment, bacterial inoc
ulate was transferred into 250ml flasks filled with cya
nidecontaining solution (70 ml). Flasks were placed
on a shaker (120 rpm) at constant temperature (26C).
Experiments were carried out with a series of bacteria
from the same harvest. The concentration of bacteria
upon seeding was 0.340.38 mg/l that corresponded to
a suspension optical density D
540
= 0.40 0.03.
The degree of total destruction, W
T
, was calculated
from kinetic dependences by the formula
(2)
where
0
is the initial CN

concentration and
T
is the
concentration in solution after destruction. The
degree of biological destruction, W
B
, was calculated by
the equation
(3)
where
B
is the CN

concentration after destruction

with PEFuntreated bacteria.
The degree of biological electrodestruction, W
EB
,
was calculated as the difference between the biological
destruction and fieldactivated destruction as follows:
(4)
where C
EB
is the CN

concentration after destruction

with PEFtreated bacteria.
Surface hydrophobicity H was determined via the
adhesion of cells to noctane [14] by measuring the
distribution coefficient between organic and aqueous
phases as follows: H = 100(1 D
x
/D
0
), where D
0
and D
x
are optical densities of aqueous bacterial suspension
before and after its contact with noctane, respectively.
Cells for these experiments were grown as described
above. Washing, PEFtreatment, and measurements
of hydrophobicity were performed in 5 mM trisHCl
solution (pH 7.5). The electrophoretic mobility of
bacteria was measured in a closed cell by the micro
electrophoresis technique in distilled water [15]. The
electrokinetic potential was calculated by the
Smoluchowski formula.
All experiments were performed at least three
times.
T
T
0
0
100%,
C C
W
C

=
B
B
0
0
100%,
C C
W
C

=
B EB
EB
0
100%,
C C
W
C

=
0.7
0.6
0.5
0.4
0.3
60 40 20 0
0.42
0.40
0.38
0.36
0.34
1
2
3
4
5
6
7
8
9
10
40
30
20
10
0
70 40 20 10 0
(W
T
)
(W
B
)
(W
EB
)
CN

, mM D
540
t, h
W, %
U, V
()
(b)
Fig. 2. (a) Kinetic curves of microbial destruction of (37)
NaAg(CN)
2
and optical density D
540
of (810) P. fluores
cence B5040 cells as a function of PEF strength: (1) the
spontaneous destruction of the complex in solution;
(2) the destruction after treatment at a voltage of 40 V;
(3) microbial destruction, (47) destruction by bacteria
treated at a voltage of 10, 20, 40, and 70 V, respectively; and
(810) after treatment at a voltage of 20, 40, and 70 V,
respectively. (b) Contribution of electrobiodestruction,
W
EB
and biological destruction, W
B
, to the total destruction
W
T
.
CN
= 0.10 mM, f = 100 Hz, = 1 ms, and t = 15 min.
COLLOID JOURNAL Vol. 72 No. 6 2010
EFFECT OF WEAK PULSE ELECTRIC FIELDS 833
RESULTS AND DISCUSSION
Effect of Value of PEF Voltage
Kinetic curves of NaAg(CN)
2
destruction as a func
tion of applied PEF voltage are shown in Fig. 2a. In
this series of experiments, the pulse duration and fre
quency were 1 ms and 100 Hz, respectively. The spon
taneous destruction of complex did not exceed 1.0%
(Fig. 2a, curve 1). The destruction by P. fluorescence
B5040 culture proceeded with the delay of about 20 h;
after 46 h, the CN

concentration decreased by 18.9%

from 0.74 to 0.6 mM (Fig. 2a, curve 3). The field
treatment of bacterial inoculate at a voltage of 10 and
20 V slightly affects the degree and kinetics of
NaAg(CN)
2
destruction (curves 4 and 5). The destruc
tion is intensified after treating at a voltage of 40 and 70
V; in this case, the duration of lagphase noticeably
decreases (Fig. 2a, curves 6 and 7). Cells retain their
viability and a small increase in biomass was observed
(Fig. 2a, curves 810).
The degree of cyanide destruction as a function of
field parameters was calculated by Eqs. (1)(3) using
kinetic dependences. As can be seen from Fig. 1b, the
fraction of fieldactivated electrobiodestruction in the
overall destruction of silver cyanide complex with
P. fluorescence B5040 culture rises from 5% to more
than 50% with an increase in voltage from 10 to 70 V at a
fixed frequency of 100 Hz and pulse duration of 1 ms.
The electrokinetic potential of bacteria, which was
measured immediately after the PEF switchingoff,
changes nonmonotonically depending on the applied
voltage (Fig. 3a). The potential of bacteria differs
from the reference value for untreated cells by no more
than 3.0 mV within the 1030 V range. At potential
values beyond this voltage range, the potential
markedly decreases compared to the reference value.
Previously, we demonstrated [16] that the electroki
netic potential changes simultaneously with a trans
membrane potential that reflects the processes that
involve P. fluorescence B5040 culture upon its adapta
tion to Ag, Au, and Cu cyanide complexes, as well as
to simple cyanide.
Changes in the charge values correlate with
changes in hydrophobichydrophilic properties of
bacteria; the potential decreases with an increase in
hydrophobicity. After PEF treatment, the affinity of
bacteria to organic phase varies depending on the field
strength and solution composition (see Fig. 3b). For
single episodes of the record of field treatment, we
used cells of different harvests; hence, experimental
values of hydrophobicity, H
EB
, were normalized to the
hydrophobicity of untreated cells, H
C
. Relative
H
EB
/H
C
values are shown in Fig. 3. As can be seen, the
response of bacteria to the PEF treatment depends on
the concentration of cyanide in solution. The degree
of hydrophilization of bacteria surface increases with
the voltage on the cell in the solution containing no
cyanide and at its content lower than 1.2 mM (Fig. 3b,
curves 1 and 2). At the cyanide concentration of
1.2 mM, the degree of hydrophobicity remains con
stant throughout the studied voltage range (Fig. 3b,
curve 3). At concentrations of 1.9 and 2.6 mM, the
reverse run of dependences is observed, i.e., the hydro
phobicity increases with the voltage; moreover, the
2.5
2.0
1.5
1.0
0.5
0
80 60 40 20
25
20
15
10
40 30 20 10 0
, mV
Control (22 mV)
U, V
5
4
3
2
1
U, V
H
EB
/H
C
()
(b)
6
Fig. 3. Dependences of electrokinetic potential and rel
ative hydrophobicity of bacteria, H
EB
/H
C
, on the applied
voltage: (a) f = 100 Hz, = 1 ms, and t = 15 min; (b) f =
100 Hz, = 1 ms, and t = 75 min; C
CN
: (1) 0, (2) 0.4,
(3) 1.2, (4) 1.9, and (5) 2,7 mM; dotted line 6 indicates the
theoretical approximation of curve 1 using formula (5).
834
COLLOID JOURNAL Vol. 72 No. 6 2010
PODOLSKA et al.
faster the higher the content of cyanide (Fig. 3b,
curves 4 and 5).
The dependences of relative hydrophobicity on
voltage shown in Fig. 3 are distinguished by some
interesting features. In particular, they are nonmono
tonic and contain parts, which are well approximated
by exponential dependences. These features can be
explained if it is assumed that the degree of bacteria
hydrophobicity is determined by the biochemical
reaction of cells, whereas the rate of reaction is
described by the Arrhenius equation. Variations in the
relative hydrophobicity of bacteria as a function of
external voltage can be described by the following
equation:
(5)
where A is dimensionless preexponential factor, E
a
is
the activation energy that, in our case, depends on
external field, k
B
is Boltzmanns constant, and is the
absolute temperature. According to the microscopic
kinetic interpretation of the Arrhenius equation, the
activation energy is characterized by a threshold pat
tern and due to the existence of a barrier, U
b
, for reac
tion components that vary hydrophobic properties of
the cell surface. In the electric field, the height of the
potential barrier can be changed, which leads to varia
tions in the activation energy. This change is propor
tional to the strength of external electric field multi
plied by the width of barrier. In turn, the strength of
field is proportional to applied voltage. With allowance
for reasons mentioned above, the activation energy in
the field can be represented as where
is the proportionality coefficient accounting for the
efficiency of field action. At the potential
barrier is transformed into the potential well with two
possible variants. If the rate of the relaxation processes
is low compared to the time of reaction, the rate of
reaction does not depend on the field. If the rate of
relaxation processes is high, the depth of the potential
well becomes new activation energy. In this case, the
dependence of the activation energy on the field can
be expressed as Then, factor A in Eq.
(5) is defined by relation and the
equation describes nonmonotonic dependence on the
field. A comparison of Eq. (5) and experimental
dependence in the absence of cyanide (Fig. 3, curves 6
and 1) permits us to calculate the values of
and =(1.3 0.08) 10
3
,
where is the electron charge. The maximum on
curve 6 at voltage U = U
b
/ = 8.3 V corresponds to the
transformation of the potential barrier into the well.
The addition of cyanides to solution (Fig. 3, curves 2
4) leads to the appearance of new factors that give rise
to changes in the cell hydrophobicity. Equation (5) is
a EB
C B
exp ,
E H
A
H k T

=


a b
, E U U =
b
, U U >
a b
. E U U =
( )
b B
exp , A U k T =
( )
b B
0.463 0.016 U k T =
modified in these cases, which requires additional
study.
As can be seen from data obtained, there is a
threshold value of cyanide concentration
p
at which
the inversion of the effect of field on bacteria takes
place. The existence of threshold value C
th
was also
demonstrated using the hypothesis of nonlinear reso
nance tunneling (NRT) in [8] devoted to the effect of
pulse filed on respiratory activity of P. fluorescence
B5040 culture. The concept of using the NRT phe
nomenon is based on the assumption that the transfer
of electrons between molecules in a respiratory chain
proceeds by a tunneling mechanism; in this case, con
ditions arise that are characteristic for resonance tun
neling. Because electrons are transferred along the
chain in pairs, the resonance tunneling becomes non
linear and very sensitive to the action of external fields.
These fields can change electron transport and, thus,
significantly affect the respiratory activity of the chain.
Being added to the respiratory chain, cyanide blocks
the electrons transfer [17], thus creating competitive
conditions between different types of actions. At a cya
nide concentration C
CN
<
th
, electric field blocks the
cell respiration, while, at C
CN
>
th
, the field stimulates
the cell respiration. If the threshold concentration is
lower than critical value conforming to the termina
tion of aerobic respiration, the electric filed causes
stimulating effect on the respiratory activity of the
cells.
Taking into account the conclusions drawn in [7]
and our data, we can propose the hypothesis that the
observed changes in the surface charge and hydropho
bichydrophilic properties under the imposition of
electric field are based on the metabolic reaction of the
cell related to the stimulation or suppression of bacte
ria respiration and the relevant ejection of protons
outside the cell. The composition of extracellular
metabolites including more hydrophobic proteins and
amine acids, or more hydrophilic polysaccharides,
glycolipids, and lipopolysaccharides can also directly
affect the surface properties of bacteria.
Effect of PEF Frequency
The study of partial dependences demonstrated
that, in the 2550 Hz range, PEF (at a voltage of 40 V
and treatment time of 15 min) does not markedly
affect the rate of microbial destruction (Fig. 4a). As in
the untreated reference sample, the destruction by
fieldtreated bacteria begins after a 20h delay. The
contribution of electrobiodestruction is evidenced at
frequencies higher then 100 Hz. The rate of destruc
tion also increases with increasing frequency. At a con
stant pulse duration of 1 ms, this dependence reaches
a plateau in the 250500 Hz range.
COLLOID JOURNAL Vol. 72 No. 6 2010
EFFECT OF WEAK PULSE ELECTRIC FIELDS 835
Three regions can conventially be specified on the
dependence of hydrophobic surface properties of
studied bacteria on the PEF frequency. At frequencies
that range from 25 to 700 Hz, after 75 min of treat
ment, the culture becomes more hydrophilic than the
reference sample. At frequencies varying from 1000 to
3000 Hz, the dependence changes in an extreme man
ner. The hydrophobicity of the culture increases
almost sixfold. Finally, the third frequency range (up
to 10 000 Hz) is characterized by the tendency of cell
surfaces to hydrophilize.
The sensitivity of the studied system to the fre
quency can be attributed to the fact that the real dura
tion of field action on bacteria is connected with the
time of its relaxation in a suspension; in fact, it can be
shorter by several orders of magnitude than the preset
pulse duration of 1 ms (Fig. 1). Intervals between
pulses become considerably shorter with an increase in
frequency when the total field is equal to zero. In this
case, the total duration of cell exposure to the field
increases.
Effects of Treatment Duration
and Pulse Energy
Results obtained when varying the duration of
pulse and frequency support the above assumption.
Based on these data, it was revealed that, at a constant
field pulse energy f, where f is the field frequency and
is the duration of pulse, the rate of destruction
depends on the duration of pulse. Figure 5a shows data
on the rate of 24h destruction at constant f = 0.1 (the
frequency increases from 100 to 10 000 Hz and the
duration of pulse decreases from 1000 to 10 s). The
tendency of the rate of destruction to increase with a
decrease in pulse duration was observed with a simul
taneous increase in field frequency.
At fixed field frequency (100 Hz) and pulse dura
tion (1 ms), bacteria respond to PEF treatment prima
rily via a decrease in the hydrophobicity. As can be
seen from Fig. 5b, the surface hydrophilicity of bacte
rial culture rises with an increase in the time of expo
sure. Comparing the dependences of hydrophobic and
electrosurface properties of bacteria on treatment time
(Figs. 5b and 5c), we can note the correlation between
the electrokinetic potential and surface hydrophobic
ity at short times of exposure to the field. At a duration
of treatment to 30 min, the surface becomes hydro
philic. At longer treatment (6090 min), the hydro
phobicity increases with a simultaneous decrease in
the potential.
In conclusion, note that the use of weak PEF pre
sents new possibilities for studying the functions of cell
metabolism because surface characteristics carry
information, not only about the chemical composition
of cell wall, but also about physiological processes.
The possibility of the electrochemical destruction of
cyanide on electrodes and direct physical action of the
field on the cell (the heating of environment, the dis
tortion of membrane permeability, the action of oxi
dizers, etc.) was excluded in experiments. Changes in
surface properties and destructive activity of bacteria
are conserved after the switchingoff the field. There
fore, we can assume that the pulse electric field acts as
a modulator of respiratory activity, which changes the
0.15
0.10
0.05
500 400 300 200 100 0
6
4
2
10000 8000 6000 4000 2000 0
V
d
, mM [CN

] dm
3
day
1
H
EB
/H
C
f, Hz
f, Hz
()
(b)
Fig. 4. Dependences of rate of 24h destruction of cyanide,
V
d
, and relative hydrophobicity of P. fluorescence B5040
bacteria on the PEF frequency: (a) U = 40 V, t = 15 min,
and = 1 ms; (b) U = 20 V, t = 75 min, and = 1 ms.
836
COLLOID JOURNAL Vol. 72 No. 6 2010
PODOLSKA et al.
physiological potential of bacteria. The use of weak
PEF can also have applied prospects in biotechnology
and medicine for controlling the conditions of bacte
rial adhesion, sorption and desorption of metals in
biological films and grounds, as well as for enhancing
the physiological activity of cultures and curing some
diseases.
ACKNOWLEDGMENTS
This work was partially supported by the National
Academy of Sciences of Ukraine under the auspices of
targeted program.
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0.3
0.2
0.1
0
4 3 2 1
1.0
0.5
400 300 200 100 0
30
25
30 20 10
20
15
10
t, min
Control (22 mV)
, mV
t, min
H
EB
/H
C
V
d
, mM [CN

] dm
3
day
1
()
(b)
(c)
Fig. 5. (a) Rate of NaAg(CN)
2
biodestruction at constant
pulse field energy f = 0.1, U = 40 V, t = 15 min: (1) no
imposed field, f: (2) 100, (3) 1000, and (4)10 000 Hz. Clear
dashed regions denote total destruction; dark dashed
regions denote the contribution of electrobiodestruction;
(b) the dependence of bacteria relative hydrophobicity on
the duration of PEF treatment at U = 40 V, f = 100 Hz, =
1 ms, and t = 75 min; (c) the dependence of bacteria
potential on the time of field treatment at U = 40 V, f = 100 Hz,
= 1 ms, and t = 15 min.
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