Comparison of 60-Hz electric fields and incandescent light as aversive stimuli controlling the behavior of rats

Rats were exposed to two procedures which enabled them to press a lever to turn off a 90 or 100 kV/m 60-Hz electric field or, later in the study, illumination from an incandescent lamp. Under one procedure, a response turned off the stimulus for a fixed duration, after which the stimulus was turned on again. A response during the off-period restarted the fixed duration. None of the rats turned the field off reliably. Next, under an alternative procedure, pressing one lever turned the field off; pressing the other lever turned it back on; responding under those conditions differed little from that seen at 0 kV/m. Under both procedures, when illumination from an incandescent lamp served as the stimulus, each rat did turn the stimulus off, and performances varied with stimulus intensity. The results show that a 100 kV/m 60-Hz electric field is not sufficient to function as an aversive stimulus under two procedures where illumination from a lamp does function as an aversive stimulus.     

Growth and metabolism of rodents exposed to 60-Hz electric fields

There have been a number of reports in the literature concerning growth-related changes in various animal species exposed to high-strength electric fields. Many of the laboratories reporting such effects have not documented and controlled for the secondary factors that are associated with generating high-strength electric fields (ie, corona, ozone, harmonic distortion, cage vibration, spark discharge). We have designed an exposure system in which we eliminated or minimized these secondary factors, therefore enabling us to examine only the effects of electric fields per se. Sprague-Dawley rats and Swiss-Webster mice were exposed to 60-Hz electric fields at kV/m for up to four months. In 17 individual experiments, we found a greater number of experiments in which the exposed rats had lower body weights than controls. This trend was not evident in data obtained from 14 individual mouse experiments. In more exhaustive growth studies, we found no significant differences in body weights, organ weights, or O₂ consumption between exposed and sham-exposed controls. Our failure to detect any major changes in growth was probably the result of eliminating or minimizing the secondary factors associated with electric field exposure.     

Studies on prenatal and postnatal development in rats exposed to 60-Hz electric fields

A series of three experiments was performed to determine the effects of 30-day exposures to uniform 60-Hz electric fields (100 kV/m) on reproduction and on growth and development in the fetuses and offspring of rats. In the first experiment, exposure of females for 6 days prior to and during the mating period did not affect their reproductive performance, and continued exposure through 20 days of gestation (dg) did not affect the viability, size, or morphology of their fetuses. In the second experiment, exposure of the pregnant rat was begun on 0 dg and continued until the resulting offspring reached 8 days of age. In the third experiment, exposure began at 17 dg and continued through 25 days of postnatal life. In the second and third experiments, no statistically significant differences suggesting impairment of the growth or survival of exposed offspring were detected. In the second experiment, a significantly greater percentage of the exposed offspring showed movement, standing, and grooming at 14 days of age than among-sham-exposed offspring. There was a significant decrease at 14 days in the percentage of exposed offspring displaying the righting reflex in the second experiment and negative geotropism in the third experiment. These differences were all transient and were not found when the animals were tested again at 21 days of age. Evaluation of the reproductive integrity of the offspring of the second experiment did not disclose any deficits.     

Behavioral studies with mice exposed to DC and 60-Hz magnetic fields

Behavioral measures were evaluated in adult CD-1 and LAF-1 mice continuously exposed for 72 h to a 1.5-Tesla (1 T = 10(4) Gauss) homogeneous DC magnetic field, and in LAF-1 mice continuously exposed for 72 h to a sinusoidal 60-Hz, 1.65-mT (rms) homogeneous AC field. Three types of behavioral tests were employed: (1) Memory of an electroshock-motivated passive avoidance task was assessed in animals that had been trained immediately prior to the field exposure. The strength of memory was varied either by altering the strength of the electric footshock during training, or by administering a cerebral protein synthesis inhibitor, anisomycin, at the time of training. (2) General locomotor activity was measured using a quadrant-crossing test immediately after termination of the magnetic field exposure. (3) Sensitivity of the experimental subjects to the seizure-inducing neuropharmacological agent, pentylenetrazole , was assessed immediately after the field exposure on the basis of three criteria: (a) the percentage of subjects exhibiting a generalized seizure, (b) the mean time to seizure, and (c) the mean seizure level. The results of these studies revealed no behavioral alterations in exposed mice relative to controls in any of the experimental tests with the 1.5-T DC field or the 60-Hz, 1.65-mT (rms) AC field.     

Relationship between field strength and arousal response in mice exposed to 60-Hz electric fields

White-footed mice, Peromyscus leucopus, were exposed to 60-Hz electric fields to study the relationship between field strength and three measures of the transient arousal response previously reported to occur with exposures at 100 kV/m. Five groups of 12 mice each were given a series of four 1-h exposures, separated by an hour, with each group exposed at one of the following field strengths: 75, 50, 35, 25, and 10 kV/m; 8 additional mice were sham-exposed with no voltage applied to the field generator. All mice were experimentally naive before the start of the experiment, and all exposures occurred during the inactive (lights-on) phase of the circadian cycle. The first exposure produced immediate increases in arousal measures, but subsequent exposures had no significant effect on any measure. These arousal responses were defined by significant increases of gross motor activity, carbon dioxide production, and oxygen consumption, and were frequently recorded with field strengths of 50 kV/m or higher. Significant arousal responses rarely occurred with exposures at lower field strengths. Responses of mice exposed at 75 and 50 kV/m were similar to previously described transient arousal responses in mice exposed to 100-kV/m electric fields. Less than half of the mice in each of the field strength groups below 50 kV/m showed arousal responses based on Z (standard) scores, but the arousals of the mice that did respond were similar to those of mice exposed at higher field strengths. Polynomial regression was used to calculate the field strength producing the greatest increases for each of the arousal measures. The results show that the amplitude of the transient arousal response is related to the strength of the electric field, but different measures of arousal may have different relationships to field strength.     

Neuroendocrine parameters in the rat exposed to 60-Hz electric fields

This study was designed to assess the neuroendocrine response of male Long-Evans rats to sustained or intermittent 60-Hz electric fields when exposed for 1 or 3 h at 100 kV/m. No significant differences were noted in corticosterone, prolactin, or thyrotropin levels between exposed and sham-exposed rats. A statistically significant increase (P less than .01) in growth hormone was noted in rats exposed to intermittent electric fields for 3 h. Emphasis was placed on good experimental design and the need to avoid standard laboratory stressors (excessive handling, temperature extremes, transportation, noise, etc.) known to be present in many biomedical studies. The importance of avoiding reactions due to extraneous factors in experiments predicated on investigating physiological function in relation to electric field exposure is discussed.     

Cardiovascular response of rats exposed to 60-Hz electric fields

Recently, it has been reported that exposure to high-strength electric fields can influence electrocardiogram (ECG) patterns, heart rates, and blood pressures in various species of animals. Our studies were designed to evaluate these reported effects and to help clarify some of the disagreement present in the literature. Various cardiovascular variables were measured in Sprague-Dawley rats exposed or sham-exposed to 60-Hz electric fields at 80 or 100 kV/m for periods up to four months. No significant differences in heart rates, ECG patterns, blood pressures, or vascular reactivity were observed between exposed and sham-exposed rats after 8 hours, 40 hours, 1 month, or 4 months of exposure. Blood pressure and heart rate measurements, made during exposure to a 100-kV/m electric field for one hour, revealed no significant differences between exposed and sham-exposed groups. In addition, physiologic reserve capacity, measured in rats subjected to low temperature after exposure to 100 kV/m for one month, showed that electric-field exposure had no significant effect on physiological response to cold stress. Our studies cannot be directly compared to the work of other investigators because of differences in animal species and electric-field characteristics. However, our failure to detect any cardiovascular changes may have been the result of 1) eliminating secondary field effects such as shocks, audible noise, corona, and ozone; 2) minimizing steady-state microcurrents between the mouth of the animal and watering devices; and 3) minimizing electric-field-induced vibration of the electrodes and animal cages.     

60-Hz electric fields: detection by female rats

Female rats were trained to detect a vertical, 60-Hz electric field using the same apparatus and procedure we used previously to study behavioral detection of the field by male rats. Each rat was trained individually to press a lever in the presence of the field and not to press in its absence. Correct detections occasionally produced a food pellet. The probability of detecting the field increased as field strength increased. The threshold of detection--ie, the field strength required for detections at a probability of 0.5 after correction for errors--varied among rats between 3 and 10 kV/m. Behavioral detection by female rats was indistinguishable from that by male rats.     

Growth rate and mitotic index analysis of Vicia faba L. roots exposed to 60-Hz electric fields

Growth, mitotic index, and growth rate recovery were determined for Vicia faba L. roots exposed to 60-Hz electric fields of 200, 290, and 360 V/m in an aqueous inorganic nutrient medium (conductivity 0.07-0.09 S/m). Root growth rate decreased in proportion to the increasing strength; the electric field threshold for a growth rate effect was about 230 V/m. The induced transmembrane potential at the threshold exposure was about 4-7 mV. The mitotic index was not affected by an electric field exposure sufficient to reduce root growth rate to about 35% of control. Root growth rate recovery from 31-96% of control occurred in 4 days after cessation of the 360 V/m exposure. The results support the postulate that the site of action of the applied electric fields is the cell membrane.     

Effects of 60-Hz electric fields on specific humoral and cellular components of the immune system

We evaluated humoral and cellular functions of the immune system of Swiss-Webster mice exposed to 60-Hz electric fields at 100 kV/m. No significant differences were observed in primary antibody response to keyhole limpet hemocyanin (precipitating antibody levels) between exposed (30 or 60 days) and control mice, nor were there significant changes in mitogen-stimulation response of spleen cells from mice similarly exposed for 90 or 150 days when compared to sham-exposed animals.     

Detection of 60-hertz vertical electric fields by rats

Rats were trained to press levers to indicate the presence or absence of 60-Hz vertical electric fields at intensities from 0 to 27 kV/m (rms). The probability of detecting the field increased as the strength of the field increased. The shape of the detection curve (psychometric function) for most subjects (Ss) was similar whether the discriminative stimulus was the electric field or a tone. Two protocols were used to estimate the minimum field intensity necessary to detect the field (Reiz Limen, RL). The RL was estimated to be 13.3 kV/m (rms) when using one protocol (the staircase method) and 7.9 kV/m (rms) when using another protocol (the method of constant stimuli).     

A system for simultaneous exposure of small animals to 60-Hz electric and magnetic fields

Equipment designed for simultaneous exposure of rodents to 60-Hz electric and magnetic fields is described. Three identical systems were constructed, each capable of continuous exposure of 256 rats or 640 mice to a nominal electric field at less than 50 kV/m, and to horizontal and vertical magnetic fields at less than 1 mT. Design features, construction details, and results of various tests of the systems are described. Tests were made: of phase relations between electric and magnetic fields; of uniformity of electric and magnetic fields; of changes across time in electric-field intensity as a result of animals' soiling of cages and various washing routines; of resistance of bedding material during humid and dry conditions; and of acoustic noise due to background, to field-generation equipment, and to air conditioning equipment. The results demonstrated that fields were effectively generated but that significant and troublesome changes in electric-field intensity occurred because of cage-soiling. However, when cages were frequently cleaned, field intensities were consistent from one exposure to another.     

Endocrinological effects of strong 60-Hz electric fields on rats

Adult male rats were exposed or sham-exposed to 60-Hz electric fields without spark discharges, ozone, or significant levels of other secondary variables. No effects were observed on body weights or plasma hormone levels after 30 days of exposure at an effective field strength of 68 kV/m. After 120 days of exposure (effective field strength = 64 kV/m), effects were inconsistent, with significant reductions in body weight and plasma levels of follicle-stimulating hormone and corticosterone occurring in one replicate experiment but not in the other. Plasma testosterone levels were significantly reduced after 120 days of exposure in one experiment, with a similar but not statistically significant reduction in a replicate experiment. Weanling rats, exposed or sham-exposed in electric fields with an effective field strength of 80 kV/m from 20 to 56 days of age, exhibited identical or closely similar growth trends in body and organ weights. Hormone levels in exposed and sham-exposed groups were also similar. However, there was an apparent phase shift between the two groups in the cyclic variations of concentrations of hormones at different stages of development, particularly with respect to follicle-stimulating hormone and corticosterone. We concluded that 60-Hz electric fields may bring about subtle changes in the endocrine system of rats, and that these changes may be related to alterations in episodic rhythms.     

Effects of 60-Hz electric fields on avoidance behavior and activity of rats

In repeated short-term tests (four sessions, each of 45-minute duration), and one longer test (a 23.5-hour session), behavior of rats was evaluated in a long, narrow shuttlebox. One side of the box was exposed to an electric field at various strengths, while a visually identical opposite side was shielded from exposure. In the short-term tests, rats generally remained shielded from electric fields of 90 kV/m and greater during the first session, and maintained this response in subsequent sessions. In the longer test, this same preference response was demonstrated at field strengths of 75 kV/m and greater; however, at 25 and 50 kV/m, rats exhibited a statistically significant preference for the exposed region of the shuttlebox, but only during the light portion of a 12-hour light: 12-hour dark cycle. Exposed animals made more traverses than sham-exposed controls between the two ends of the shuttlebox during the first hour of the test. The experimental data support the hypothesis that the observed behavioral effects are the result of direct interaction of the electric field with the animal, and not the result of secondary factors such as electric shock, corona discharge, audible noise, ozone, or vibration of the experimental apparatus.     

Electric fields induced in chicken eggs by 60-Hz magnetic fields and the dosimetric importance of biological membranes.

Chicken eggs are convenient models for observing the effects of inhomogeneities and variations, such as those found in biological membranes and in cellular conductivities, on the distribution of internal electric fields as induced by exposure to magnetic fields. The vitelline membrane separates the yolk, which has a conductivity of 0.26 S/m, from the white, which has a conductivity of 0.85 S/m. A miniaturized probe with 2.4-mm resolution was used to measure induced fields in eggs placed in a uniform, 1-mT magnetic field at 60 Hz. The E fields induced in eggs with homogenized contents agreed with expectations based on simple theory. Results were similar to intact eggs unless the probe moved the yolk off-center, which greatly perturbed the induced fields. A more reproducible arrangement, which consisted of saline-agar filled dishes with a hole cut for test samples, was developed to enhance definition of electrical parameters. With this test system, the vitelline membrane was found to be responsible for most of the perturbation of the induced field, because it electrically isolates the yolk from the surrounding white. From a theoretical viewpoint, this dosimetry for the macroscopic egg yolk is analogous to the interaction of fields with microscopic cells. These findings may have important implications for research on biological effects of ELF electromagnetic fields, especially for studies of avian embryonic development.     

Attempts to produce taste-aversion learning in rats exposed to 60-Hz electric fields

A measure of taste-aversion (TA) learning was used in three experiments to 1) determine whether exposure to intense 60-Hz electric fields can produce TA learning in male Sprague-Dawley rats, and 2) establish a dose-response function for the behavior in question. In Experiment 1, four groups of eight rats each were distributed into one of two exposures (69 ± 5 kV/m or 133 ± 10 kV/m) or into one of two sham-exposure groups. Conditioning trials paired 0.1% sodium saccharin in water with 3 h of exposure to a 60-Hz electric field. Following five conditioning trials, a 20-min, two-bottle preference test between water and saccharin-flavored water failed to reveal TA conditioning in exposed groups. In Experiment 2, four groups of eight rats each (34 ± 2 kV/m or 133 ± 10 kV/m and two sham-exposed groups) were treated as before. Electric-field exposure had no effect on TA learning. Experiment 3 tested for a possible synergy between a minimal dose (for TA learning) of cyclophosphamide (6 mg/kg) and 5 h of exposure to 133 ± 10 kV/m electric fields in a dark environment under conditions otherwise similar to those of Experiments 1 and 2. The results indicated no TA learning as reflected in the relative consumption of saccharin.     

A behavioral response of swine to a 60-Hz electric field

It has been shown that rats, given the choice, will spend more time out of a 60-Hz electric field than in it at field strengths ? 75 kV/m. This paper describes research to examine the relevance of these data to a different species, the pig. Miniature pigs that had been exposed to a 60-Hz electric field at 30 kV/m for 20 h/day, 7 days/week for as long as 6 months, were tested for their preference for the presence or absence of the field during a 23.5-h period. Similar to earlier results with rats, miniature pigs spent more time out of the electric field than in it during the sleeping period.     

Effects of intermittent 60-Hz high voltage electric fields on metabolism,activity, and temperature in mice

Transient effects of 100-kV/m extremely low frequency electric fields were studied in the white footed deermouse, Peromyscus leucopus. Gross motor activity, carbon dioxide production, oxygen consumption, and core body temperature were monitored before, during, and after intermittent field exposures (four hour-long exposures, at one-hour intervals). Thirty-four mice were exposed in cages with plastic floors floating above ground potential, and 21 mice were exposed in cages with grounded metal floor plates. The first field exposure produced an immediate, transient increase of activity and gas measures during the inactive phase of the circadian cycle. All measures returned to baseline levels before the second exposure and were not significantly changed throughout the remainder of the exposures. The rapid habituation of field-induced arousal suggests that significant metabolic changes will not be measured in experiments in which the interval between exposure and measurement is greater than two hours.