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Experimental macroscopic dosimetry for extremely-low-frequency electric and magnetic fields.

Environmental and laboratory exposure to electric and magnetic fields (EMF) in the extremely-low-frequency range (ELF) produces electrical quantities that interact directly with the exposed biological system on a scale small compared to the size of the human body but large with respect to cellular dimensions. The purpose of this paper is to describe these macroscopic electrical quantities and their characterization through measurements on living systems and experimental models. Electric field exposure results in a total induced current, surface electric fields, internal electric fields, and internal currents. Magnetic field exposure results in internal magnetic field, internal electric fields, and internal currents. Basic properties of fields and matter determine the methods by which these quantities can be measured. Quantification or dosimetry for these parameters on a macroscopic basis can be directed to the whole body, a cross section across the body, a local surface area, or a local volume. Models of varying degrees of sophistication have been used to establish spatial distributions of external fields and internal fields and currents.

Animals↗

Electrical polarization of bioactive glass and assessment of their in vitro apatite deposition.

We evaluated the electrical polarizability of a bioactive glass of the 45S5 type (BG) by thermally stimulated depolarization current (TSDC) measurement based on the effects of the surface charges generated by an electrical polarization treatment on the growth of calcium phosphate crystals in simulated body fluid (SBF). From the results of the TSDC measurements, the polarized BG showed broad peaks in the depolarization current density spectra. The stored electrical charges calculated from the TSDC measurements of the BG polarized with an electrical dc field of 10 V x cm(-1) at 500 degrees C for 1 h was 8.5 mC x cm(-2) and depended on the electrical dc field intensity. The depolarization of the BG was assumed to be due to a migration of sodium ions and influenced by transitions of the BG structure. The immersion tests with the SBF showed that the effects of the electrical polarization produced morphologic changes in the calcium phosphate deposit grown on the BG after the short period of 2 h. We expected the effects to be caused by a change in the ion concentrations in the surrounding SBF due to the surface charges of the polarized BG. The change in the ion concentrations induced the difference in the calcium phosphate deposition. Consequently, we confirmed that BG had an electrical polarizability, a possibility of storing large charges, and differences in the growth of the calcium phosphate deposit due to the effects of the surface charges.

Apatites↗

Ionic Diffusivity, Electrical Conductivity, Membrane and Thermoelectric Potentials in Colloids and Granular Porous Media: A Unified Model.

Ionic diffusivity, electrical conductivity, membrane and thermoelectric potentials in isotropic and homogeneous colloidal suspensions, and granular porous media saturated by a binary symmetric 1:1 electrolyte are four interrelated phenomena. The microstructure and the surface properties of the solid grains-water interface influence directly these properties. The ionic diffusivities (and the electrical conductivity, respectively) in colloids and porous media have contributions from diffusion (and electromigration, respectively) through the bulk solution occupying the pores, together with electromigration occurring at the grains-water interface in the electrical double layer. Surface diffusion in porous materials has no contribution from concentration gradients along the grains-water interface. Instead, surface diffusion is envisioned as a purely electromigration process due to the membrane potential. The tortuosities of the transport of anions and cations are equal to the bulk tortuosity of the pore space only at high ionic strength. As the ionic strength decreases, the dominant paths for transport of the ion corresponding to the counterion of the electrical double layer shift from the pore space to the solid grains-water interface. Because anions and cations do not move independently, the membrane potential created by the charge polarization alters the velocity of the anions and influences the mutual diffusivity coefficient of the salt in the porous material. An electric potential of thermal origin is also produced in nonisothermal conditions. The ionic contributions to the electrical conductivity are based on a differential effective medium approach. These ionic contributions to the electrical conductivity are used to derive the ionic diffusivities and the membrane and thermoelectric potentials. The influence of the temperature and the presence, in the pore space, of a second immiscible and nonwetting phase is also considered in this model. Porosity is shown to affect the membrane potential. Several predictions of the model are checked with success by comparing the model to a set of experimental data previously published. Copyright 1999 Academic Press.

Journal Article↗

The resting membrane potential of cells are measures of electrical work, not of ionic currents.

Living cells create electric potential force, E, between their various phases by at least three distinct mechanisms. Charge separation, F = [equation: see text] (Eqn 1) creates the potential, E = [equation: see text] of -120 to -145 mV between cytoplasmic and mitochondrial phases by unbalanced proton expulsion powered by the redox energy of the respiratory chain. Electrically unbalanced flow of Na+ through voltage gated Na+ channels raises the potential of nerve from -85 to +30 mV. The so-called resting potential of cells, which varies from -85 mV in heart to -4.5 mV in red cell, does not appear to result from the unbalanced flow of ions between phases, but rather to be a measure of the work required to move ions between phases. Movement of an ion between phases entails three types of energy. Concentration work is that required to move an ion between phases containing different concentrations of ions: [equation: see text] Electrical work is that work required to move an ion from phases with differing electric potentials: [equation: see text] The Nernst potential of an ion existing at different concentrations in two phases is: [equation: see text] The osmotic work term is small and can generally be ignored. In heart the measured resting potential between extra- and intracellular phases, EN is approximately -85 mV. The calculated Nernst potential of K+, E [K+]out/in, is -85 mV (Eqn 4). This means that in heart, K+ distributes itself between the two phases as if it moved through an open ion channel. Its concentration work (Eqn 2) is equal in magnitude but opposite in sign to its electrical work (Eqn 3). This makes net K+ current flow, I, equal 0, indicating that this potential cannot be a diffusion potential. In liver the resting potential ranges from -28 to -40 mV, and is equivalent to the E[Cl-]out/in, while in red cell the resting potential is about -4.5 mV, which is equivalent to the potential of all nine major inorganic ion species except Na+, K+ and Ca2+. Therefore the resting potential between extra- and intracellular phases of cells should be thought of, not as a diffusion potential but rather as a measure of the electrical work: [equation: see text] required to transport the most permeant ions in a Gibbs-Donnan near-equilibrium system, either K+ or Cl- or both, between the phases of an aqueous system during the flow of current required to measure potentials with intracellular KCl electrodes or during ion movements brought about during normal cellular activity. The resting electrical potential results from the existence of a mono-ionic Gibbs-Donnan near-equilibrium system between the extra- and intracellular phases of cell wherein the activity of free H2O within all phases of the system is equal and the energy of the gradients of the nine major inorganic ions, delta G[ionz]out/in, are in near-equilibrium with one another, with the potential between the phases, EN, and with the energy of ATP hydrolysis. delta GATP Hydrolysis. ranges from a low of -55 to slightly over -60 kJ/mole in all cell types.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Measurement of cardiac output in pregnancy by thoracic electrical bioimpedance and thermodilution. A preliminary report.

Thoracic electrical bioimpedance is a noninvasive, continuous method of obtaining cardiac output that requires no operator skill. However, the most recent thoracic electrical bioimpedance technology has not been validated in pregnancy. We therefore compared two methods of measuring cardiac output in pregnancy, thoracic electrical bioimpedance and thermodilution. We studied 11 patients who required pulmonary artery catheterization for peripartum management and measured cardiac output simultaneously by thoracic electrical bioimpedance and thermodilution. Among eight of nine patients, there was agreement (within +/- 20%) between the two methods. Bivariate linear regression with these nine cases showed excellent correlation (r = 0.91, p less than 0.001) with a slope of 1.04, which indicated a one-to-one relationship between thoracic electrical bioimpedance and thermodilution. The remaining two cases were removed from analysis because of septic shock in one case (which invalidates thoracic electrical bioimpedance) and 4+ tricuspid regurgitation in another case (which invalidates thermodilution). These data support that thoracic electrical bioimpedance measurement of cardiac output may be valid in most peripartum patients.

Cardiac Output↗

Horizontal cell electrical coupling in the giant danio: synaptic modulation by dopamine and synaptic maintenance by calcium.

Electrical synapses, and their structural manifestation, gap junctions, are critical elements of retinal circuitry. These synapses are subject to both rapid modulation and slower structural changes by physiological signals which mediate changes in the adaptational state of the retina. The electrical synapses of fish retinal horizontal cells are an excellent preparation for in vitro studies of electrical synapses. We have examined the rapid modulation of electrical coupling by dopamine and effects on the expression and maintenance of electrical synapses by cell calcium in pairs of horizontal cells isolated from retinas of the giant danio (Danio aquipinnatus). We report that rapid modulation by dopamine reduces junctional conductance by modifying gap junction channel gating, while maintaining cells in reduced calcium medium, and lowering intracellular calcium concentration, results in the loss of electrical coupling. The effects of calcium on synaptic maintenance may be related to structural changes observed in horizontal cell electrical synapses during light adaptation.

Animals↗

Integrated electrical activity and number of zero crossings during a gradual increase in muscle force in patients with neuromuscular diseases.

In the brachial biceps muscle of 17 patients with myopathy and of 14 patients with neurogenic disorders the integrated electrical activity and number of zero crossings per unit time were analysed in 3 sites of each muscle every 100 msec during a gradual increase in force from zero to maximum within 10 sec. The analysis of integrated electrical activity could not discriminate between patients with myopathy and patients with neurogenic disorders. The slope of the linear relation between the square root of integrated electrical activity and force expressed in kilograms was increased in 44% of the patients with myopathy and in 64% of the patients with neurogenic disorders. The increase in slope may be due to increase in the ratio of electrical activity and force of individual motor units. The integrated electrical activity related to a force of 40% of maximum was decreased in about two-thirds of patients with myopathy and in about half of the patients with neurogenic disorders. This decrease in integrated electrical activity may be due to random loss of muscle fibres or to loss of whole motor units respectively. The integrated electrical activity was linearly related to mean amplitude between potential reversals per unit time (turns). The number of zero crossings was increased in 29% of the patients with myopathy and decreased in 70% of the patients with neurogenic disorders at a force of 30% of maximum. The number of zero crossings was linearly related to turns.(ABSTRACT TRUNCATED AT 250 WORDS)

Action Potentials↗

Effects of transcutaneous thermal and electrical stimulation of the teat on pituitary luteinizing hormone, prolactin and oxytocin secretion in ovariectomized, estradiol-treated beef cows following acute weaning.

The objective of the study was to determine if chronic electrical or thermal stimulation of sensory neurons on the surface of the teat is able to activate pathways that suppress the weaning-induced increase in luteinizing hormone (LH) secretion in beef cows. Treatment groups (n = 5 per group) consisted of: 1) control suckled (CS); 2) weaned plus electrical stimulation of the teat (ESTT); 3) weaned plus electrical stimulation of the tail (ESTL); 4) weaned plus thermal stimulation of the teat (TTT); 5) weaned plus thermal stimulation of the tail (TTL) and 6) weaned (WN). Cows were ovariectomized on Day 5 post partum (PP) and were treated with estradiol-17beta to maintain a constant tonic baseline. Beginning on Days 17 to 21 post partum, cows were suckled by their own calf (control), weaned or weaned and electrically or thermally stimulated for 10 minutes every 6 hours for 4 days. Chronic transcutaneous electrical and thermal stimulation of the teat or tail failed to impede the unambiguous rise (P < 0.001) in LH pulse frequency and amplitude following weaning. Positive and negative feedback of estradiol on LH secretion was not affected by treatments. Relatively consistent episodes of oxytocin and prolactin release were observed following control-suckling, but responses to electrical and thermal stimulation were inconsistent. Chronic electrical or thermal stimulation of teat-specific or nonspecific loci did not attenuate heightened secretion of LH after weaning. The results are further evidence against a role for mammary somatosensory neurons in the suckling-mediated inhibition of LH secretion.

Journal Article↗

Relation of growth process to spatial patterns of electric potential and enzyme activity in bean roots.

The electric spatial pattern and invertase activity distribution in growing roots of azuki bean (Phaseolus chrysanthos) have been studied. The electric potential near the surface along the root showed a banding pattern with a spatial period of about 2 cm. It was found that the enzyme activity has a peak around 3-7 mm from the root tip, in good agreement with the position of the first peak of the electric potential, which is located a little behind the elongation zone. An inhomogeneous distribution of ATP content was also detected along the root. Experiments on the electric isolation of the elongation zone from the mature zone and acidification treatment showed that H+ is transported from the mature-side to elongation-side regions, causing tip elongation through an acid-growth mechanism. Both acidification and electric disturbance on growing roots affected growth significantly. Simultaneous measurements of electric potential and enzyme activity clearly showed a good correlation between these two quantities and growth speed. From an analogy with the Characean banding, the spatio-temporal organization via the cell membrane in electric potential and enzyme activity can be regarded as a dissipative structure arising far from equilibrium. These experimental results can be interpreted with a new mechanism that the dissipative structure is formed spontaneously along the whole root, accompanied by energy metabolism, to make H+ flow into the root tip.

Journal Article↗

Temperature-, electric field- and solute-induced percolation in water-in-oil microemulsions.

We report investigations on the percolation of the aqueous phase in water-in-oil microemulsions, comparing systems stabilized by ionic AOT and non-ionic Igepal amphiphiles. First, we briefly review the opposite effect of temperature on the two systems and compare electric conductivity with viscosity data. In the second part, we show that percolation can be induced by high electric fields resulting in a shift of the percolation curve. The electric field measurements allow to investigate the dynamics of clustering of the water droplets to form a network of percolating channels. We examine the slow build-up and the fast decay of the percolating structure, monitoring simultaneously electric conductivity and electric birefringence. In the third part we discuss the effect of some solutes on the percolation curve, especially of small molecules which act as protein denaturants and of native and denatured proteins like methemoglobin, chymotrypsin and gelatin. The spectroscopic determination of the dimerization of hemin, released from denatured hemoglobin, reflects the incorporation of the hemin monomers in the surfactant monolayer. In the gelatin system time resolved electric birefringence shows that even at low concentrations it is the macromolecule which determines the structure of the aqueous domain. In the appendix, a simple estimate of the intrinsic Kerr-constant is given for microemulsion droplets deformed in an electric field.

Journal Article↗

Embryonic bone matrix formation is increased after exposure to a low-amplitude capacitively coupled electric field, in vitro.

In order to investigate the mechanism(s) of electric field-stimulated osteogenesis, we have developed an in vitro model in which embryonic chick tibiae have consistently demonstrated increased bone matrix formation in response to a low amplitude (estimated 10(-5) V/m in the serum-free culture medium), capacitively coupled, 10 Hz sinusoidal electric field. Initial applications of this model revealed that 72 h of continuous exposure to the electric field increased tibial collagen production by 29% compared to untreated controls, P less than 0.01. Additional studies further revealed: (a) that when electric field exposure was limited to 30 min/day during the 72 h in vitro incubation, embryonic bone matrix formation was increased by 83%, compared to non-treated controls (P less than 0.001), suggesting an inductive mechanism; (b) that the osteogenic response to electric field exposure in vitro was not unique to embryonic chick tibiae, since a similar response was also seen with newborn mouse calvaria (+133%, P less than 0.02); (c) that electric field-exposure-stimulated chick bone matrix formation was associated with increased bone cell proliferation; and (d) that this mitogenic response to in vitro electric field exposure could also be observed with embryonic chick calvarial cells in monolayer, serum-free cultures.

Animals↗

Prevalence and predictors of electrical storm in patients with implantable cardioverter-defibrillator.

Identifying predictors of electrical storm in patients with implantable cardioverter-defibrillators (ICDs) could help identify those at risk and reduce the incidence of this emergency situation, which has a detrimental effect on mortality and morbidity in patients with ICDs. This retrospective study sought to determine the prevalence and predictors of electrical storm in patients with ICDs. One hundred sixty-two patients (126 men; mean age 58 +/- 13 years) who received ICDs from January 2001 to January 2005 were included in the study. Clinical, electrocardiographic, and ICD stored data and electrograms were collected and analyzed. Twenty-two patients (14%) developed electrical storm during a mean follow-up of 14.3 +/- 10 months. Using Cox multiple regression analysis, it was found that an ejection fraction <25% (p = 0.007), QRS width > or =120 ms (p = 0.002), and a lack of adjunctive angiotensin-converting enzyme inhibitor and beta-blocker therapy (both p < 0.001) were correlated with a greater probability of electrical storm. Adjunctive amiodarone and digoxin therapy, indication of ICD implantation, and age were not correlated with the occurrence of electrical storm during follow-up (all p = NS). In conclusion, electrical storm is not uncommon in patients with ICDs. Optimum medical therapy with beta blockers and angiotensin-converting enzyme inhibitors could reduce the occurrence of electrical storm, and this especially should be considered in those at greater risk for this complication (i.e., those with left ventricular ejection fractions <25% and QRS widths > or =120 ms).

Adult↗

Transient electroosmotic flow induced by DC or AC electric fields in a curved microtube.

This study investigates transient electroosmotic flow in a rectangular curved microtube in which the fluid is driven by the application of an external DC or AC electric field. The resultant flow-field evolutions within the microtube are simulated using the backwards-Euler time-stepping numerical method to clarify the relationship between the changes in the axial-flow velocity and the intensity of the applied electric field. When the electric field is initially applied or varies, the fluid within the double layer responds virtually immediately, and the axial velocity within the double layer tends to follow the varying intensity of the applied electric field. The greatest net charge density exists at the corners of the microtube as a result of the overlapping electrical double layers of the two walls. It results in local maximum or minimum axial velocities in the corners during increasing or decreasing applied electric field intensity in either the positive or negative direction. As the fluid within the double layer starts to move, the bulk fluid is gradually dragged into motion through the diffusion of momentum from the double layer. A finite time is required for the full momentum of the double layer to diffuse to the bulk fluid; hence, a certain phase shift between the applied electric field and the flow response is inevitable. The patterns of the axial velocity contours during the transient evolution are investigated in this study. It is found that these patterns are determined by the efficiency of momentum diffusion from the double layer to the central region of the microtube.

Journal Article↗

Electrical-to-mechanical coupling in purple membranes: membrane as electrostrictive medium.

In this paper, we present acousto-electrical measurements performed on dry films of purple membranes (PM) of Halobacterium salinarium. The purpose of these measurements is to determine the relation between mechanical and electrical phenomena in bacteriorhodopsin and to define the role of the protein in the proton transfer process. Electrical-to-mechanical coupling in PMs manifests itself as direct and inverse piezoelectric effects. Measurements performed on the samples with different degrees of PM orientation and at various values of the externally applied cross-membrane electric field indicate that piezoelectric phenomena in PMs arise from the electric asymmetry of the membranes, i.e., they originate from electrostriction. Experiments with samples made of oriented PMs allow estimation of the value of the intrinsic cross-membrane electric field, which is approximately 10(8) V/m. A hypothetical model of PM is presented where the electrical-to-mechanical coupling is suggested to be the main driving force for the proton translocation against the Coulomb forces acting in the membrane.

Acoustics↗

Recognition and processing of randomly fluctuating electric signals by Na,K-ATPase.

Previous work has shown that Na,K-ATPase of human erythrocytes can extract free energy from sinusoidal electric fields to pump cations up their respective concentration gradients. Because regularly oscillating waveform is not a feature of the transmembrane electric potential of cells, questions have been raised whether these observed effects are biologically relevant. Here we show that a random-telegraph fluctuating electric field (RTF) consisting of alternating square electric pulses with random lifetimes can also stimulate the Rb(+)-pumping mode of the Na,K-ATPase. The net RTF-stimulated, ouabain-sensitive Rb+ pumping was monitored with 86Rb+. The tracer-measured, Rb+ influx exhibited frequency and amplitude dependencies that peaked at the mean frequency of 1.0 kHz and amplitude of 20 V/cm. At 4 degrees C, the maximal pumping activity under these optimal conditions was 28 Rb+/RBC-hr, which is approximately 50% higher than that obtained with the sinusoidal electric field. These findings indicate that Na,K-ATPase can recognize an electric signal, either regularly oscillatory or randomly fluctuating, for energy coupling, with high fidelity. The use of RTF for activation also allowed a quantitative theoretical analysis of kinetics of a membrane transport model of any complexity according to the theory of electroconformational coupling (ECC) by the diagram methods. A four-state ECC model was shown to produce the amplitude and the frequency windows of the Rb(+)-pumping if the free energy of interaction of the transporter with the membrane potential was to include a nonlinear quadratic term. Kinetic constants for the ECC model have been derived. These results indicate that the ECC is a plausible mechanism for the recognition and processing of electric signals by proteins of the cell membrane.

Biophysical Phenomena↗

On the efficiency and reversibility of active ligand transport induced by alternating rectangular electric pulses.

The stationary-state kinetic properties of a simplified two-state electro-conformational coupling model (ECC) in the presence of alternating rectangular electric potential pulses are derived analytically. Analytic expressions for the transport flux, the rate of electric energy dissipation, and the efficiency of the transducing system are obtained as a function of the amplitude and frequency of the oscillation. These formulas clarify some fundamental concept of the ECC model and are directly applicable to the interpretation and design of experiments. Based on these formulas, the reversibility and the degree of coupling of the system can be studied quantitatively. It is found that the oscillation-induced free energy transduction is reversible and tight-coupled only when the amplitude of the oscillating electric field is infinitely large. In general, the coupling is not tight when the amplitude of the electric field is finite. Furthermore, depending on the kinetic parameters of the model, there may exist a "critical" electric field amplitude, below which free energy transduction is not reversible. That is, energy may be transduced from the electric to the chemical, but not from the chemical to the electric.

Carrier Proteins↗

Light chain-dependent myosin structural dynamics in solution investigated by transient electrical birefringence.

The technique of transient electrical birefringence was used to compare some of the electric and structural dynamic properties of myosin subfragment 1 (S1(elc, rlc)), which has both the essential and regulatory light chains bound, to S1(elc), which has only an essential light chain. The rates of rotational Brownian motion indicate that S1(elc, rlc) is larger, as expected. The permanent electric dipole moment of S1(elc, rlc) is also larger, indicating that the regulatory light chain portion of S1(elc, rlc) has a dipole moment and that it is aligned head-to-tail with the dipole moment of the S1(elc) portion. The permanent electric dipoles decrease with increasing ionic strength, apparently because of ion binding to surface charges. Both S1(elc, rlc) and S1(elc) have intrinsic segmental flexibility, as detected by the ability to selectively align segments with a brief weak electric field. However, unlike S1(elc), which can be structurally distorted by the action of a brief strong electric field, S1(elc, rlc) is stiffer and cannot be distorted by fields as high as 7800 V/cm applied to its approximately 8000 D permanent electric dipole moment. The S1 . MgADP . Pi analog S1 . MgADP . Vi is smaller than S1 . MgADP, for both S1(elc, rlc) and S1(elc). Interestingly, the smaller, stiffer S1(elc, rlc) . MgADP . Vi complex retains intrinsic segmental flexibility. These results are discussed within a framework of current hypotheses of force-producing mechanisms that involve S1 segmental motion and/or the loss of cross-bridge flexibility during force production.

Adenosine Diphosphate↗

Fluctuation-driven directional flow in biochemical cycle: further study of electric activation of Na,K pumps.

Directional flow of information and energies is characteristic of many types of biochemical reactions, for instance, ion transport, energy coupling during ATP synthesis, and muscle contraction. Can a fluctuating force field, or a noise, induce such a directional flux? Previous work has shown that Na,K-ATPase of human erythrocyte can absorb free energy from an externally applied random-telegraph-noise (RTN) electric field to pump Rb+ up its concentration gradient. However, the RTN field used in these experiments was constant in amplitude and would not mimic fluctuating electric fields of a cell membrane. Here we show that electric fields which fluctuate both in life time and in amplitude, and thus, better mimicking the transmembrane electric fields of a cell, can also induce Rb+ pumping by Na,K-ATPase. A Gaussian-RTN-electric field, or a field with amplitude fluctuating according to the Gaussian distribution, with varied standard deviation (sigma), induced active pumping of Rb+ in human erythrocyte, which was completely inhibited by ouabain. Increased values for sigma led to a nonmonotonic reduction in pumping efficiency. A general formula for calculating the ion transport in a biochemical cycle induced by fluctuating electric field has been derived and applied to a simple four-state electroconformational coupling (ECC) model. It was found that the calculated efficiency in the energy coupling decreased with increasing sigma value, and this effect was relatively small and monotonic, whereas experimental data were more complex: monotonic under certain sets of conditions but nonmonotonic under different sets. The agreement in general features but disagreement in some fine features suggest that there are other properties of the electric activation process for Na,K-ATPase that cannot be adequately described by the simple ECC model, and further refinement of the ECC model is required.

Biophysical Phenomena↗