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PubMed1996

Transient electric birefringence of colloidal particles immersed in shear flow. Part II. The initial response under the action of a rectangular electric pulse and the behavior at a low alternating electric field.

The time-dependent rotational diffusion equation for rigid macromolecules in solution has been approximately solved for two cases in order to extend the electric birefringence technique to streaming-electric birefringence. One is for the initial period through the application of a rectangular electric pulse to the solution immersed in a low shear flow. The purpose of this is expansion of the distribution function into a function series made by the product of the powers of reduced time (= Thetat) and hydrodynamic field alpha (= G Theta , G: velocity gradient, Theta: rotary diffusion constant) and a surface harmonic P(i)(j)cos jphi. The solution for the build-up process at arbitrary electric field strength is found, but is limited to low hydrodynamic fields. The other is for the response when an alternating electric field is applied to the solution in a shear flow. Here, instead of reduced time, the maximum electric field E(0) is chosen as a parameter for the expansion. The expressions for the intensity of the transmitted light through crossed Nicols are derived in two optical systems where the polarizer is set at an angle of 45 degrees and 0 degrees to the direction of the electric field. The results in the former case show that we can determine four parameters, the ratio of velocity gradient to rotary diffusion constant, the axial ratio of a particle, the anisotropy of electric polarizability, and the optical anisotropy factor, from four values observed in two optical systems, namely, two light intensities before applying an electric field and two initial slopes of the build-up after applying an electric field. On the other hand, when a low alternating electric field with extremely high frequency is applied, the build-up of the light intensity in the former case is the same as that of electric birefringence for pure induced dipole orientation. The build-up for the latter optical system is the same as the expression for pure induced dipole orientation of Eq. (38) shown in a previous work.

Journal Article↗
PubMedDate not supplied

Electric shock, Part III: Deliberately applied electric shocks and the treatment of electric injuries.

Earlier parts of this series have discussed the physics, pathophysiology, and nature of electric injury. This part will discuss deliberately applied electric shocks and the treatment of electric injuries. Electric shocks are deliberately applied to persons during electroshock therapy and with stun guns, shock batons, and electric cattle prods. Electric injuries, whether a complication of deliberate electric shock or due to accidental injury, should be treated to preserve cardiac and respiratory function and to prevent further tissue damage. Safe extrication at the scene, rapid triage, and emergency medical treatment are discussed.

Electric Injuries↗
PubMed1986

Acetylcholine, ATP, and proteoglycan are common to synaptic vesicles isolated from the electric organs of electric eel and electric catfish as well as from rat diaphragm.

Cholinergic synaptic vesicles were isolated from the electric organs of the electric eel (Electrophorus electricus) and the electric catfish (Malapterurus electricus) as well as from the diaphragm of the rat by density gradient centrifugation followed by column chromatography on Sephacryl-1000. This was verified by both biochemical and electron microscopic criteria. Differences in size between synaptic vesicles from the various tissue sources were reflected by their elution pattern from the Sephacryl column. Specific activities of acetylcholine (ACh; in nmol/mg of protein) of chromatography-purified vesicle fractions were 36 (electric eel), 2 (electric catfish), and 1 (rat diaphragm). Synaptic vesicles from all three sources contained ATP in addition to ACh (molar ratios of ACh/ATP, 9-12) as well as binding activity for an antibody raised against Torpedo cholinergic synaptic vesicle proteoglycan. Synaptic vesicles from rat diaphragm contained binding activity for the monoclonal antibody asv 48 raised against a rat brain 65-kilodalton synaptic vesicle protein. Antibody asv 48 binding was absent from electric eel and electric catfish synaptic vesicles. These antibody binding results, which were obtained by a dot blot assay on isolated vesicles, directly correspond to the immunocytochemical results demonstrating fluorescein isothiocyanate staining in the respective nerve terminals. Our results imply that ACh, ATP, and proteoglycan are common molecular constituents of motor nerve terminal-derived synaptic vesicles from Torpedo to rat. In addition to ACh, both ATP and proteoglycan may play a specific role in the process of cholinergic signal transmission.

Acetylcholine↗
PubMed1997

Ontogeny of the electric organ discharge and the electric organ in the weakly electric pulse fish Brachyhypopomus pinnicaudatus (Hypopomidae, Gymnotiformes).

I recorded the electric organ discharges (EODs) of 331 immature Brachyhypopomus pinnicaudatus 6-88 mm long. Larvae produced head-positive pulses 1.3 ms long at 7 mm (6 days) and added a second, small head-negative phase at 12 mm. Both phases shortened duration and increased amplitude during growth. Relative to the whole EOD, the negative phase increased duration until 22 mm and amplitude until 37 mm. Fish above 37 mm produced a "symmetric" EOD like that of adult females. I stained cleared fish with Sudan black, or fluorescently labeled serial sections with anti-desmin (electric organ) or anti-myosin (muscle). From day 6 onward, a single electric organ was found at the ventral margin of the hypaxial muscle. Electrocytes were initially cylindrical, overlapping, and stalk-less, but later shortened along the rostrocaudal axis, separated into rows, and formed caudal stalks. This differentiation started in the posterior electric organ in 12-mm fish and was complete in the anterior region of fish with "symmetric" EODs. The lack of a distinct "larval" electric organ in this pulse-type species weakens the hypothesis that all gymnotiforms develop both a temporary (larval) and a permanent (adult) electric organ.

Animals↗
PubMed2006

Modeling the electric field of weakly electric fish.

Weakly electric fish characterize the environment in which they live by sensing distortions in their self-generated electric field. These distortions result in electric images forming across their skin. In order to better understand electric field generation and image formation in one particular species of electric fish, Apteronotus leptorhynchus, we have developed three different numerical models of a two-dimensional cross-section of the fish's body and its surroundings. One of these models mimics the real contour of the fish; two other geometrically simple models allow for an independent study of the effects of the fish's body geometry and conductivity on electric field and image formation. Using these models, we show that the fish's tapered body shape is mainly responsible for the smooth, uniform field in the rostral region, where most electroreceptors are located. The fish's narrowing body geometry is also responsible for the relatively large electric potential in the caudal region. Numerical tests also confirm the previous hypothesis that the electric fish body acts approximately like an ideal voltage divider; this is true especially for the tail region. Next, we calculate electric images produced by simple objects and find they vary according to the current density profile assigned to the fish's electric organ. This explains some of the qualitative differences previously reported for different modeling approaches. The variation of the electric image's shape as a function of different object locations is explained in terms of the fish's geometrical and electrical parameters. Lastly, we discuss novel cues for determining an object's rostro-caudal location and lateral distance using these electric images.

Animals↗
PubMed1991

Coupled in vivo activity of creatine phosphokinase and the membrane-bound (Na+,K+)-ATPase in the resting and stimulated electric organ of the electric fish Narcine brasiliensis.

Physiological control of the plasma membrane sodium pump, (Na+,K+)-ATPase, is essential for proper function of eukaryotic cells. In the electric organ of the elasmobranch Narcine brasiliensis, the normal demands placed upon the pump during the process of generation of electrical currents call for large and rapid changes in activity of this enzyme, making this a good model for the study of its cellular regulation. 31P NMR spectroscopic techniques were used to study metabolic regulation of membrane pump function in resting and stimulated electric organ and in skeletal muscle of the live, intact N. brasiliensis. Because the ATP synthetic abilities of the electric organ by glycolysis or oxidative phosphorylation are extremely limited, depletion of phosphocreatinine (PCr) could be used to determine the activity of the (Na+,K+)-ATPase after the electric organ was stimulated to discharge, and to measure the net flux from PCr to ATP through the creatine phosphokinase (CPK) reaction in the electric organ. Saturation transfer, an NMR technique which measures exchange rates, was applied to determine the unidirectional flux in the forward direction through the same reaction in the electric organ and in skeletal muscle as a control. The pseudo first-order rate constant kf for the CPK reaction at 24 degrees C in resting electric organ was 0.000 +/- 0.002 s-1 (n = 10) and in skeletal muscle was 0.08 +/- 0.03 s-1 (n = 3). The results demonstrate that in resting electric organ, which is well supplied with CPK, there was no measurable flux through this reaction, although CPK when extracted is highly active. Measured and calculated levels of all substrates for the creatine kinase reaction in the electric organ are similar to those in unstimulated skeletal muscle, where the creatine phosphokinase reaction rates are high in vivo. In contrast to the resting electric organ, during stimulation of the electric organ the measured net rate constant was greater than 0.08 s-1. In addition, as shown by lack of PCr depletion, there was virtually no net turnover of ATP in the resting organ compared to the stimulated organ. The marked difference in the (Na+,K+)-ATPase activity in the resting and activated electric organ confirmed earlier results (Blum, H., Nioka, S., and Johnson, R. G., Jr. (1990) Proc. Natl. Acad. Sci. U. S. A. 87, 1247-1251). Together, these results suggest that there is a novel method of coordinate regulation of cellular enzymes of great sensitivity and rapidity.

Adenosine Triphosphate↗
PubMed2003

Cyclic AMP modulates electrical signaling in a weakly electric fish.

Many species of electric fish show diurnal or socially elicited variation in electric organ discharge amplitude. In Sternopygus macrurus, activation of protein kinase A by 8-bromo-cAMP increases electrocyte sodium current magnitude. To determine whether the behavioral plasticity in electric organ discharge amplitude is controlled by electrocyte biophysical properties, we examined whether the effects of phosphorylation on ion currents in the electric organ translate directly into electric organ discharge changes. We injected the electric organ of restrained fish with 8-bromo-cAMP and monitored the electric organ discharge. The effect of protein kinase A activation on electrocyte action potentials was examined in isolated electric organ using two-electrode current clamp. Electric organ discharge and action potential amplitude and pulse duration increased in response to 8-bromo-cAMP. Pulse and action potential duration both increased by about 25%. However, the increase in electric organ discharge amplitude (approximately 400%) was several-fold greater than the action potential amplitude increase (approximately 40%). Resting membrane resistance decreased in electrocytes exposed to 8-bromo-cAMP. We propose that in the Thevenin equivalent circuit of the electric organ a moderate increase in action potential amplitude combined with a decrease in internal resistance produces a greater voltage drop across the external resistance (the water around the fish), accounting for the large increase in the externally recorded electric organ discharge.

8-Bromo Cyclic Adenosine Monophosphate↗
PubMed1994

Correlations among indices of electric and magnetic field exposure in electric utility workers.

Power-frequency electric and magnetic fields are known to exhibit marked temporal variation, yet in the absence of clear biological indications, the most appropriate summary indices for use in epidemiologic studies are unknown. In order to assess the statistical patterns among candidate indices, data on 4383 worker-days for magnetic fields and 2082 worker-days for electric fields collected for the Electric and Magnetic Field Project for Electric Utilities using the EMDEX meter [Bracken (1990): Palo Alto, CA: Electric Power Research Institute] were analyzed. We examined correlations at the individual and job title group levels among indices of exposure to both electric and magnetic fields, including the arithmetic mean, geometric mean, median, 20th and 90th percentiles, time above lower cutoffs of 20 V/m and 0.2 microT, and time above higher cutoffs of 100 V/m and 2.0 microT. For both electric and magnetic fields, the arithmetic mean was highly correlated with the 90th percentile; moderately correlated with the geometric mean, median, and lower and higher cutoff scores; and weakly correlated with the 20th percentile. Electric and magnetic field indices were generally weakly correlated with one another. Rank-order correlation coefficients were consistently greater than product-moment correlation coefficients. Job title group summary scores showed higher correlations among electric field indices and magnetic field indices and between electric and magnetic field indices than was found for individual worker-days, with only the 20th percentile clearly independent of the others. These results suggest that individuals' exposures are adequately characterized by a measure of central tendency for electric and magnetic fields, such as the arithmetic or geometric mean, and an indicator of a lower threshold or cutoff for each field type, such as the 20th percentile or proportion of time above 20 V/m or 0.2 microT. A single measure of central tendency for each type of field appears to be adequate when exposures are assessed at the job title level.

Electricity↗
PubMed1994

Transient electric birefringence of agarose gels. I. Unidirectional electric fields.

The orientation of agarose gels in pulsed electric fields has been studied by the technique of transient electric birefringence. The unidirectional electric fields ranged from 2 to 20 V/cm in amplitude and 1 to 100 s in duration, values within the range typically used for pulsed field gel electrophoresis (PFGE). Agarose gels varying in concentration from 0.3 to 2.0% agarose were studied. The sign of the birefringence varied randomly from one gel to another, as described previously [J. Stellwagen & N.C. Stellwagen (1989), Nucleic Acids Research, Vol. 17, 1537-1548]. The sign and amplitude of the birefringence also varied randomly at different locations within each gel, indicating that agarose gels contain multiple subdomains that orient independently in the electric field. Three or four relaxation times of alternating sign were observed during the decay of the birefringence. The various relaxation times, which range from 1 to approximately 120 s, can be attributed to hierarchies of aggregates that orient in different directions in the applied electric field. The orienting domains range up to approximately 22 microns in size, depending on the pulsing conditions. The absolute amplitude of the birefringence of the agarose gels increased approximately as the square of the electric field strength. The measured Kerr constants are approximately 5 orders of magnitude larger than those observed when short, high-voltage pulses are applied to agarose gels. The increase in the Kerr constants in the low-voltage regime parallels the increase in the relaxation times in low-voltage electric fields. Birefringence saturation curves in both the low- and high-voltage regimes can be fitted by theoretical curves for permanent dipole orientation. The apparent permanent dipole moment increases approximately as the 1.6 power of fiber length, consistent with the presence of overlapping agarose helices in the large fiber bundles orienting in low-voltage electric fields. The optical factor is approximately independent of fiber length. Therefore, the marked increase in the Kerr constants observed in the low-voltage regime is due to the large increase in the electrical orientation factor, which is due in turn to the increased length of the fiber bundles and domains orienting in low-voltage electric fields. Since the size of the fiber bundles and domains approximates the size of the DNA molecules being separated by PFGE, the orientation of the agarose matrix in the applied electric field may facilitate the migration of large DNA molecules during PFGE.

Birefringence↗
PubMed1998

Plasticity of the electric organ discharge waveform of the electric fish Brachyhypopomus pinnicaudatus. I. Quantification of day-night changes.

The electric organ discharge of the gymnotiform fish Brachyhypopomus pinnicaudatus is a biphasic waveform. The female's electric organ discharge is nearly symmetric but males produce a longer second phase than first phase. In this study, infrared-sensitive video cameras monitored the position of unrestrained fish, facilitating precise measurement of electric organ discharge duration and amplitude every 2 h for 24 h. Males (n = 27) increased electric organ discharge duration by 37 +/- 12% and amplitude by 24 +/- 9% at night and decreased it during the day. In contrast, females (n = 8) exhibited only minor electric organ discharge variation over time. Most of a male's increase occurred rapidly within the first 2-3 h of darkness. Electric organ discharge values gradually diminished during the second half of the dark period and into the next morning. Modulation of the second phase of the biphasic electric organ discharge produced most of the duration change in males, but both phases changed amplitude by similar amounts. Turning the lights off at mid-day triggered an immediate increase in electric organ discharge, suggesting modification of existing ion channels in the electric organ, rather than altered genomic expression. Exaggeration of electric organ discharge sex differences implies a social function. Daily reduction of duration and amplitude may reduce predation risk or energy expenditure.

Action Potentials↗
PubMed2006

Regulation and modulation of electric waveforms in gymnotiform electric fish.

Weakly electric gymnotiform fish specialize in the regulation and modulation of the action potentials that make up their multi-purpose electric signals. To produce communication signals, gymnotiform fish modulate the waveforms of their electric organ discharges (EODs) over timescales spanning ten orders of magnitude within the animal's life cycle: developmental, reproductive, circadian, and behavioral. Rapid changes lasting milliseconds to seconds are the result of direct neural control of action potential firing in the electric organ. Intermediate-term changes taking minutes to hours result from the action of melanocortin peptides, the pituitary hormones that induce skin darkening and cortisol release in many vertebrates. Long-term changes in the EOD waveform taking days to weeks result from the action of sex steroids on the electrocytes in the electric organ as well as changes in the neural control structures in the brain. These long-term changes in the electric organ seem to be associated with changes in the expression of voltage-gated ion channels in two gene families. Electric organs express multiple voltage-gated sodium channel genes, at least one of which seems to be regulated by androgens. Electric organs also express multiple subunits of the shaker (Kv1) family of voltage-gated potassium channels. Expression of the Kv1 subtype has been found to vary with the duration of the waveform in the electric signal. Our increasing understanding of the mechanisms underlying precise control of electric communication signals may yield significant insights into the diversity of natural mechanisms available for modifying the performance of ion channels in excitable membranes. These mechanisms may lead to better understanding of normal function in a wide range of physiological systems and future application in treatment of disease states involving pathology of excitable membranes.

Action Potentials↗
PubMed2001

Electrical stimulation alters protein adsorption and nerve cell interactions with electrically conducting biomaterials.

Electrical charges have been shown to enhance nerve regeneration; however, the mechanisms for this effect are unclear. One hypothesis is that an electrical stimulus alters the local electrical fields of extracellular matrix molecules, changing protein adsorption. We have investigated this hypothesis--that electrical stimulation increases the adsorption of serum proteins, specifically fibronectin (FN), to the electrically conducting polymer polypyrrole (PP), thereby, increasing neurite extension. PP was used because electrical stimulation of PP has been shown to significantly enhance neurite outgrowth, and more importantly, PP can be formed into conduits to guide nerve regeneration in vivo. Here, we looked at the effects of electrical stimulation on protein adsorption when an electrical current was applied to PP (1) during protein adsorption (immediate stimulation) and (2) several hours after protein adsorption (delayed stimulation). We found that immediate stimulation of PP increases FN adsorption from purified FN and serum-containing solutions. Correspondingly, PC-12 cells grown on PP films that had been previously adsorbed with FN during immediate stimulation expressed longer neurites. However, for delayed stimulation, no significant differences in adsorption or neurite outgrowth were observed. These studies suggest that increased FN adsorption with immediate electrical stimulation may explain enhanced neurite extension on electrically stimulated PP.

Adsorption↗
PubMed2000

The electric image in weakly electric fish: perception of objects of complex impedance.

Weakly electric fish explore the environment using electrolocation. They produce an electric field that is detected by cutaneous electroreceptors; external objects distort the field, thus generating an electric image. The electric image of objects of complex impedance was investigated using a realistic model, which was able to reproduce previous experimental data. The transcutaneous voltage in the presence of an elementary object is modulated in amplitude and waveform on the skin. Amplitude modulation (measured as the relative change in the local peak-to-peak amplitude) consists of a 'Mexican hat' profile whose maximum relative slope depends on the distance of the fish from the object. Waveform modulation depends on both the distance and the electrical characteristics of the object. Changes in waveform are indicated by the amplitude ratio of the larger positive and negative phases of the local electric organ discharge on the skin. Using the peak-to-peak amplitude and the positive-to-negative amplitude ratio of this discharge, a perceptual space can be defined and correlated with the capacitance and resistance of the object. When the object is moved away, the perceptual space is reduced but keeps the same proportions (homothetically): for a given object, the positive-to-negative amplitude ratio is a linear function of the peak-to-peak amplitude. This linear function depends on the electrical characteristics of the object. However, there are 'families' of objects with different electrical characteristics that produce changes in the parameters of the local electric organ discharge that are related by the same linear function. We propose that these functions code the perceptual properties of an object related to its impedance.

Animals↗
PubMed1994

Transient electric birefringence of agarose gels. II. Reversing electric fields and comparison with other polymer gels.

The transient electric birefringence of low electroendosmosis (LE) agarose gels oriented by pulsed unidirectional electric fields was described in detail in Part I [J. Stellwagen and N. C. Stellwagen (1994), Biopolymers, Vol. 34, p. 187]. Here, the birefringence of LE agarose gels in rapidly reversing electric fields, similar in amplitude and duration to those used for field inversion gel electrophoresis, is reported. Symmetric reversing electric fields cause the sign of the birefringence of LE agarose gels, and hence the direction of orientation of the agarose fibers, to oscillate in phase with the applied electric field. Because of long-lasting memory effects, the alternating sign of the birefringence appears to be due to metastable changes in gel structure induced by the electric field. If the reversing field pulses are equal in amplitude but different in duration, the orientation behavior depends critically on the applied voltage. If E < 7 V/cm, the amplitude of the birefringence gradually decreases with increasing pulse number and becomes unmeasurably small. However, if E > 7 V/cm, the amplitude of the birefringence increases more than 10-fold after approximately 20 pulses have been applied to the gel, suggesting that a cooperative change in gel structure has occurred. Because there is no concomitant change in the relaxation times of the orienting particles, the large increase in the amplitude of the birefringence must be due to an increase in the number of agarose fibers and/or fiber bundles orienting in the electric field, which in turn indicates a cooperative breakdown of the noncovalent "junction zones" that cross-link the fibers into the gel matrix. The sign of the birefringence of LE agarose gels is always positive after extensive junction zone breakdown, indicating that the agarose fibers and fiber bundles preferentially orient parallel to the electric field when they are freed from the constraints of the gel matrix. Three other gel-forming polymers, high electroendosmosis (HEEO) agarose (a more highly charged agarose), beta-carrageenan (a stereoisomer of agarose), and polyacrylamide (a chemically cross-linked polymer) were also studied in unidirectional and rapidly reversing electric fields. The birefringence of HEEO agarose gels in reversing fields is very similar to that of LE agarose gels, suggesting that the orientation anomalies are not due to the occasional charged residues on the agarose backbone chain. The beta-carrageenan gels exhibit variable orientation behavior in reversing electric fields, suggesting that its internal gel structure is not as tightly interconnected as that of agarose gels.(ABSTRACT TRUNCATED AT 400 WORDS)

Birefringence↗
PubMed1995

Electric parameters of Na+/K(+)-ATPase by measurements of the fluorescence-detected electric dichroism.

The electric parameters of Na+/K(+)-ATPase labeled by FITC have been characterized by measurements of the fluorescence-detected electric dichroism. The fluorescence emission was measured with polarizers at the magic angle and the light for excitation was usually polarized parallel to the field vector. The FITC-Na+/K(+)-ATPase preparations exhibit a negative electric dichroism at field strengths up to about 600 V/cm and a positive dichroism at higher field strengths. Pulse reversal experiments reveal a dominant permanent electric moment at low electric field strengths and an increasing contribution from an induced electric moment at higher field strengths. The dichroism rise curves and the transients upon pulse reversal show two relaxation processes with opposite amplitudes, whereas the dichroism decay curves in most cases can be represented by single exponentials at a reasonable accuracy. The amplitude A2 associated with the slower of the rise processes is dominant at low field strengths and also approaches saturation already at low field strengths. The dependence of A2 on the electric field strength is consistent with the orientation function for permanent dipoles and cannot be represented by the orientation function for induced dipoles. The fitted permanent dipole moment is in the range of 3.5 x 10(-24) Cm [1 x 10(6) D] and shows only a relatively small decrease with increasing ionic strength. The stationary values of the electric dichroism up to field strengths E < or = 800 V/cm can be represented with high accuracy by an orientation function for disk-shaped particles with a permanent moment along the particle symmetry axis and an induced moment along the semi-major axis. The permanent electric moment determined according to this function is consistent with the one obtained from the amplitudes A2. In summary, our measurements indicate that Na+/K(+)-ATPase is associated with a large permanent electric moment directed perpendicular to the membrane plane. The dipole moment per ATPase monomer unit is estimated to be 1.4 x 10(-27) Cm [430 +/- 50 D].

Animals↗