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INFLUENCE OF ALUMINUM IN BIOLOGIC EFFECTS OF ELF MAGNETIC FIELD STIMULATION
Romana Ruži a; Dominik Vodnik b; Igor Jerman a a BION—Institute for Bioelectromagnetics and New Biology, Ljubljana, Slovenia b Department of Biology, Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia Online Publication Date: 21 March 2000

To cite this Article Ruži, Romana, Vodnik, Dominik and Jerman, Igor(2000)'INFLUENCE OF ALUMINUM IN BIOLOGIC EFFECTS OF

ELF MAGNETIC FIELD STIMULATION',Electromagnetic Biology and Medicine,19:1,57 — 68
To link to this Article: DOI: 10.1081/JBC-100100296 URL: http://dx.doi.org/10.1081/JBC-100100296

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ELECTRO- AND MAGNETOBIOLOGY, 19(1), 57–68 (2000)

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INFLUENCE OF ALUMINUM IN BIOLOGIC EFFECTS OF ELF MAGNETIC FIELD STIMULATION
1

ˇˇ Romana Ruzic,1* Dominik Vodnik,2 and Igor Jerman1 BION—Institute for Bioelectromagnetics and New Biology 1000 Ljubljana, Slovenia 2 Department of Biology Biotechnical Faculty University of Ljubljana 1000 Ljubljana, Slovenia

ABSTRACT
We studied the influence of a weak, extremely low-frequency magnetic field (MF) with a frequency of 50 Hz and a peak amplitude of 103 µT and aluminum solution (in the form of AlCl3) at different concentrations (0, 40, 70, 100, 130, 160, 400, 800, 2000, and 5000 µM) on the growth of spruce seedlings (Picea abies). The results showed that stimulatory and statistically significant MF effects on the growth of seedlings were observed only with a 100-µM aluminum solution. Slight stimulative effects were also observed within the range of concentrations between 40 and 160 µM Al3 (all the stimulated groups taken together). Germination and fresh weight were not significantly influenced. At these concentrations the aluminum solution alone (without MF) or the MF alone (without Al3 ) did not influence the growth parameters. These results suggest the importance of synergistic action of the MF with environmental factors as well as the existence of ‘‘physiologic windows’’ in addition to the frequency and power ones.

INTRODUCTION
In the effort to understand the variety of magnetic field (MF) effects on plant development, our attention should not be directed simply at the physical characteristics of the applied MFs, but also at the environmental and biologic characteristics of the experiment.
* To whom correspondence should be addressed, at BION—Institute for Bioelectromagnetics ˇ and New Biology, Celovska 264, 1000 Ljubljana, Slovenia. E-mail:romana.ruzic@guest.arnes. sibion@guest.arnes.si

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Copyright © 2000 by Marcel Dekker, Inc. www.dekker.com

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According to many studies, including our own (1–3), it can be expected that the effects of MF stimulation will vary with the same MF, given the varying environments to which the experimental organisms are exposed and also the varying physiologic states of the organisms. In support of this hypothesis, there are some reports on the synergistic effects of MF and stress conditions (4–7) as well as possible physiologic window effects. In this manner, Tsong and Gross (8) reported on the existence of ‘‘concentration windows,’’ i.e., the applied MF yielded effects only at specific ligand concentrations. McLeod et al. (6) described the dependence of the organisms’ physiologic state in response to an MF stimulus. In our previous experiments, MFs with a frequency 50 Hz and amplitude of 105 µT (peak) exhibited highly heterogeneous effects, depending on the environmental conditions; the effects were either inhibitory when drought or pH stress were applied, or stimulatory when an excess of water or Ca2 was used (1–3). For experiments with calcium, the MF stimulation demonstrated a biologic effect only at specific Ca2 concentrations. In this case, however, a possible mechanism could be explained by means of the theory of ion cyclotron resonance (9); so far the physical background for such results has been mostly unknown. The toxic effect of the Al3 ion is primarily expressed as an inhibition of root growth as well as a decreased uptake of divalent cations. These two reactions are the primary events that cause all other physiologic changes in response to aluminum (10). The inhibition of root growth is due to the inhibition of cell division and/or cell elongation. A decrease in the content of Mg2 and Ca2 in roots and needles in response to aluminum exposure was shown in a number of tree species, including spruce, Picea abies (11). Research on seedlings and young spruce trees showed that the inhibitory effect of aluminum arises mostly at concentrations over 100 µM (10). Although aluminum is not considered an essential element for plant life, its growth effects at low concentrations (below 1 ppm) can be stimulative (12,13). The availability and phytotoxicity of aluminum depend on a number of abiotic and biotic factors (14). In the present work we examined whether MFs have any synergistic effects in association with toxic levels of the metal aluminum on the early development of spruce seedlings.

MATERIALS AND METHODS
Experimental Conditions and Exposure to MF The P. abies seeds were directly exposed to MF and simultaneously watered, first with distilled and then with deionized water (pure or with various concentrations of AlCl3 ). There were two sets of experiments. In the first set we tested the effects of various MF Al3 solutions (40–5000 µM). In the second we examined concentration levels of Al3 around 100 µM, since at the latter concentration we obtained significant stimulatory results (see below). The MF used was a horizontal, continuous, pure sine wave ( f 50 Hz, 103 4 µT peak) for 12 h/day (11 a.m. to 11 p.m.). MF was applied through a slightly modified system of a pair of Helmholtz-like coils (radius 20 cm, separation 40 cm, copper wire 1.4 mm, number of turns 140), connected to an amplifier (40 W) and computer program to generate the chosen field parameters and the regime of daily exposure (Fig. 1). The coil configuration was a compromise between ideal MF conditions and space requirements, but the active volume had nearly homogeneous fields. Orientation in respect to the dishes was vertical (Fig. 1). The composite static geomagnetic field was 46 4 µT measured

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FIGURE 1. The experimental system and the position of the Petri dishes in the coil. The coils were aligned in the N-S axis.

by Proton Magnetometer GMS-8. The axis of the coils was in the N-S direction, in which the horizontal geomagnetic field was 22 3 µT measured by a Hall probe (Gaussmeter RFL 912, RFL, Boonton, NY). The background sinusoidal MF did not exceed 0.2 µT, as measured by a Holaday HI 3604 ELF Survey Meter designed to evaluate electric and magnetic fields. It measures MF from 10 5 to 2 mT. The experimental room was inside a larger room with approximately the same temperature, so it was like a box. In addition, the coils (without other equipment) were inside a pasteboard (70 70 60-cm) box isolated with styrofoam and grounded aluminum foil protecting the seeds from external electric fields. No one except the experimenter was allowed to enter the experimental room and uncover the pasteboard box during the experiment. Electrical growth chambers or incubators were avoided because of complex electromagnetic noise appearing in them. The experiments were performed in the dark. In the present work, an improvement was made: the Petri dishes were additionally enclosed in a plastic box to prevent greater exhaustion of humidity, and the plastic boxes were aerated every day (i.e., opened for a few minutes). The average room temperature during the performance of all the experiments was 24.1 0.8°C. However, during any single experiment the temperature difference between the exposed groups and the control ones did not exceed 0.2°C and was checked twice daily.

Growth Conditions The experimental group consisted of 20 plastic Petri dishes (diameter 7 cm; 10 exposed to MF 10 not exposed to MF, both for every aluminum concentration) covered with filter paper. Each Petri dish contained 30 spruce seeds [P. abies (L.) Karsten, 1992, ˇ provenance Loski potok, Slovenia] aligned in rows and watered with pure (distilled deionized) water (i.e., the appropriate amount of water was added over filter paper) or with a specific composition of aluminum solution: 2.5 ml on day 1 and 1.5 ml on day 4. Prior to the experiments the seeds were stored at 5°C.

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ˇ ˇ RUZIC, VODNIK, AND JERMAN Aluminum Treatment

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To see the effects at markedly different Al3 (composed of AlCl3) concentrations, the first set of experiments included the following Al3 concentrations: 100, 200, 400, 800, 2000, and 5000 µM. Each concentration represented one experimental group. To see whether any stimulative MF effect could also be demonstrated in Al3 concentrations close to 100 µM Al3 (at which concentration the effects were statistically significant), a second set of experiments was performed at 40, 70, 130, and 160 µM Al3 . Each concentration represented one experimental group. The solutions were calibrated to pH 4 (optimal growth conditions for spruce seeds) with 0.1 M HCl and stored at room temperature. Measurements and Calculations Three parameters were observed: the length of all the radicles, the number of germinated seeds, and the fresh weight of all the germinated seedlings. The seeds started to germinate on day 4 of the experiment. They were considered germinated when their radicles exceeded their lengths by 2 mm. The number of germinated seeds was evaluated daily and at same hour (8 a.m.). The average number of germinated seeds per Petri dish was calculated, and then the data of repeated experiments for every Al3 concentration were pooled. The results are presented (Fig. 2) as the time course of germination, i.e., all

FIGURE 2. Synergistic effects of MF [50 Hz, 103 µT (peak)] and Al3 solution on spruce seeds (Picea abies): time course of germination. The seeds were watered with distilled deionized water (day 0), with added Al3 ions as AlCl3 (40–5000 µM) on days 1 and 4. For each repetition of the experiment, 300 seeds were exposed to the MF (30 seeds 10 Petri dishes), and 300 seeds constituted the control—both for each concentration of Al3 . The data are calculated as the averages of germinated seeds per Petri dish presented as the percentages of control (MF treated seeds divided by control and multiplied by 100 for each Al3 concentration). All the repetitions of the experiments (with the same Al3 solution) were pooled. 100% means there was no difference between the exposed and the control seeds. No single result was statistically significant.

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the germinated seeds in a measured day (3) and as percentages of the control, where 100% means no difference between the MF treated seeds and the control ones. At the end of every experiment (day 7), the growth parameters were evaluated. The length of the radicle of every seedling and the fresh weight of all the seedlings in one Petri dish (subsequently normalized to 100 seedlings) were measured only at the end of the experiment. Measurements of weight included germinated seedlings together with seed coats. Experiments of the same group (i.e., at the same concentration of Al3 ) were repeated three to six times depending on the group. Since the distribution of the radicle lengths was only seldom normal, we calculated their geometric mean, standard deviation, and confidence interval for the geometric mean. The statistical significance ( p) of the results concerning every individual experiment was measured by using Student’s t-test for comparison between the two groups (two-tailed). The significances are marked as follows: *, p 0.05; **, p 0.01; and ***, p 0.001. The data were also analyzed by a two-way analysis of variance (ANOVA) with two factors: Al treatment and MF treatment. The significance between the means was tested by using the LSD test. In addition, we also calculated the average geometric mean of the lengths of the radicles at 100 µM Al3 by pooling the data of all the repeated experiments, and we also calculated the significance by means of Student’s t-test (see Table 5). Summary results (all the data together) for 40, 70, 130, and 160 µM Al3 (taken in percentages of the control) for the growth of seedlings were also tested by using the Wilcoxon nonparametric test. To see the effects of various aluminum concentrations (without MF) on the length of spruce seedlings after 7 days, the data comparing the controls with the MF experiments were used. The average lengths at various Al3 solutions were compared with the lengths of seedlings that had received the distilled deionized water. Statistical significance was determined by Student’s t-test.

RESULTS
First Set of Experiments Aluminum Effects The results of different Al solutions on the growth of seedlings are presented in Table 1 as percentages of the ‘‘control’’ (i.e., the control was distilled deionized water in this case). Al3 concentrations of 100 and 800 µM had no consistent or statistically significant effect on growth. At 400 µM the effect was stimulative. At 2000 µM the Al3 solution produced a slight (non-significant) inhibition of growth (by 10%), but at 5000 µM the inhibition of growth amounted to more than 40% and was highly statistically significant. Magnetic Field Effects The results of experiments I–III (Table 2) showed an MF-aluminum interaction at 100 µM Al3 . Here the MF demonstrated a stimulative and statistically significant effect on the growth of seedlings. This effect was also confirmed in experiment VI (Table 3). In experiment V no bioeffects of the MF were observed; however, the pooled data of all the repetitions of 100-µM experiments strongly indicate stimulative (by 13%) and statistically significant MF effects (Table 4). There were no effects of the MF at 400 and 800 µM Al3 (Table 2), nor were there any at 2000 and 5000 µM Al3 , except in one experiment
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Table 1. Effects of Different Aluminum Concentrations on the Length of Seedlings (mm; without MF)a Al (MM)c Exp.b I II III V VI VII
a

100 100 118* 90 106 88 —

400 119** 152*** 109 — — —

800 108 132*** 105 — — —

2000 — — — 91 90 —

5000 — — — 69*** 62*** 62***

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In % of the control (in this case the control is di de water with which the data were compared (data of the lengths of seedlings grown in conditions without MF, i.e., as control for MF experiments)—, data not available. b Exp., number of the experiment (see length data in Tables 2 and 3). c Concentration of Al compared with control (i.e., di de water). Level of significance (t-test): * p 0.05. **p 00.1. ***p 0.001.

(Table 3). Only in the first experiment (I) did the ANOVA test reveal a significant MF– Al interaction. The results for germination were generally inconsistent; however, in most of the experiments the combination 100 µM Al MF enhanced germination. The pooled data (Fig. 2) demonstrate a stimulative effect of 35% on day 4, 9% on day 5, 10% on day 6, and 7% on day 7. In the remaining aluminum solutions the pooled data demonstrate a stimulative MF effect higher than 10% (on day 4) at 800 and 2000 µM Al3 (Fig. 2); however, when considering the individual experiments, these effects were statistically insignificant (not shown). There were no observed MF-Al effects on the fresh weight of seedlings. Second Set of Experiments The data (Table 5) show only a weak, statistically insignificant MF-Al interaction for the length of seedlings if a single experimental group is considered. A statistically significant stimulative MF effect at 130 µM Al3 was present in only one experiment. However, the effect of MF and Al3 in concentrations around 100 µM (taking all the data for 40, 70, 130 and 160 µM Al3 together) was stimulatory in 12 groups out of 16 ( p 0.02 by means of Wilcoxon test); therefore there is good evidence of a stimulatory tendency within this concentration range. The results for germination (pooled data, Fig. 2) showed a stimulative effect of more than 10% (on day 5) at 40 and 130 µM Al3 . However, if individual experiments are considered, these results were not statistically significant.

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Table 2. Effects of Aluminum (0, 100, 200, 400, and 800 µM) and the Magnetic Field (50-Hz, 103-µT peak) on the Length of Seedlings (mm) After 7 Days of Germination—First Type of Experimentsa Magnetic field treatment (Al, µm) 0 9.9 9.6–10.2 1.8 9.9 9.6–10.2 1.9 11.8 11.5–12.1 1.9 100 400 800 0 100 400 Control (no MF; Al, µm) 800 10.7 10.4–11.0 1.9 T 22.9

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10.1 9.8–10.4 1.9 102 ns 9.4 8.8–9.7 2.0 109 ns 10.3 10.0–10.6 1.9 120 0.01 9.8 9.5–10.1 2.1 94 ns 11.1 10.8–11.4 1.8 111 ns 9.6 9.3–9.9 1.9 11.8 11.5–12.1 1.9 117 0.03 13.2 12.9–13.5 1.9 102 ns 11.0 10.7–11.3 1.9 97 ns 8.6 8.3–8.9 1.9 10.1 9.8–10.4 1.9

11.6 11.3–11.9 1.9 118 0.01

10.7 10.4–11.0 2.0 91 ns

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I/Sept.’97 AV CI SD % p II/Oct.’97 AV CI SD % p III/Oct.’97 AV CI SD % p 8.6 8.3–8.9 1.9

9.7 9.4–10.0 1.9 101 ns

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Exp/month, number of experiments/time of experiment; AV, average length (geometric mean) of seedlings (mm) (radicle without seed); CI, confidence interval (AV 2 SE); SD, standard deviation of the mean; %, percentage of control (control 100%); T, average temperature of the boxes during the experiment; p, level of significance (t-test); ns, not significant; —, data are not available.

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Table 3. Effects of Aluminum (0, 100, 2000, and 5000 µM) and the Magnetic Field (50-Hz, 103-µT peak) on the Length of Seedlings (mm) After 7 Days of Germination—First Type of Experimentsa Magnetic field treatment (Al, µM) 0 — — — — — — — — 7.9 7.6–8.2 2.1 8.4 8.1–8.7 1.9 9.8 9.5–10.1 1.9 110 ns 7.5 7.3–7.7 1.7 100 ns — — — — 8.9 8.6–9.2 1.9 7.5 7.5–7.7 1.7 100 2000 5000 0 100 2000 Control (no MF; Al, µM) 5000 — — — T 23.4

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8.3 8.0–8.6 2.0 105 ns 8.5 8.2–8.8 2.0 96 ns 9.7 9.4–10.0 2.0 107 ns — — — — — — — — — — 5.5 5.3–5.7 1.7 98 ns 9.0 8.7–9.3 1.9 116 0.05 7.2 7.0–7.4 1.8 90 ns 6.4 6.2–6.6 1.5 117 0.01 8.9 2.6–9.2 2.0

8.4 8.1–8.7 2.0 100 ns

7.6 7.4–7.8 1.7 104 ns

5.3 5.2–5.5 1.7 96 ns

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IV/March ’98 AV CI SD % p V/April ’98 AV CI SD % p VI/April ’98 AV CI SD % p VII/May ’98 AV CI SD % p 9.0 8.7–9.3 2.0 — — — — — —

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For abbreviations, see Table 2 footnote.

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Table 4. Length of Seedings Exposed to MF 100 µM Ala AV SD SE Z p N
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1.13 1.9 0.1 2.273 0.03 1085

Pooled results of the repeated experiments. Expected value is 1. SE, standard error; N, number of all the germinated seedlings; Z, z value calculated as (AV 1)/SE (expected value 1). For other abbreviations, see Table 2 footnote.

DISCUSSION
According to our previous work, a weak sinusoidal MF with an amplitude of 105 µT (50 Hz) manifested biologic effects only under specific environmental conditions (1– 3). This finding was also evident in our present experiments, although a slightly different field density (103-µT peak) was applied. It triggered growth effects only in the presence of a 100 µM Al3 solution (Tables 2–4). At this time neither the aluminum solution (at 100 µM) nor the MF alone produced any growth response; it became evident only when both MF and aluminum were combined. The same kind of effect was found in our previous work, when the MF produced a statistically significant effect only at 0.5 µM calcium (3). However, the biologic response was completely different. In the presence of calcium, the MF inhibited germination of seedlings, while in the presence of the appropriate concentrations of aluminum it stimulated the growth of seedlings. Additional experiments at concentrations close to 100 µM Al3 (Table 5) did not generate MF effects of any significance. Nevertheless, taking into account that all the experimental groups around 100 µM produced a similar, statistically significant, although less pronounced, influence, it therefore appears as if the 103-µT and 50-Hz MF bioeffects in connection with Al3 had a relatively narrow window of their own (at 100 µM Al3 ) and a larger, more moderate one as well in the range of concentrations around 100 µM Al3 . It should be noted, however, that these windows are not the classical power or frequency ones introduced by Adey (15), but instead are physiologic, i.e., they manifest under certain physiologic conditions of the organisms treated. Similar findings have already been reported (3,8); however, the physiologic basis is still to be explained. It is difficult to explain why only at 100 µM Al3 or near this value the MF consistently influenced plant growth, while higher concentrations that are inhibitory by themselves (without MF; Table 1) did not demonstrate any consistent evidence of similar MF effects. According to the previous research by our team and by others, studying the association of MF bioeffects and stress conditions (1,2,4,5), effects at stronger concentrations would be expected. One possible reason could be that the inhibitory growth effect of aluminum alone was so strong that it obscured any additional MF effect. Something similar was encountered in our previous investigations with drought stress, when stronger drought conditions (a higher concentration of polyethylene glycol applied together with the MF) usually did not result in an increase in the MF effect in comparison with a lower concentration (1). This by itself, however, cannot explain the stimulatory effect at 100 µM Al3 .

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Table 5. Effects of Aluminum (40, 60, 130, and 160 µM) and the Magnetic Field (50-Hz, 103-µT peak) on the Length of Seedlings (mm) After 7 Days of Germination—Second Type of Experimenta Magnetic field treatment (µM) 40 9.1 2.8–9.4 1.9 91 ns 6.1 5.8–6.4 2.0 7.5 7.1–7.9 2.3 9.3 9.0–9.6 1.9 106 ns 9.4 9.1–9.7 2.0 95 ns 9.8 9.5–10.1 2.0 109 ns 9.9 9.6–10.2 1.8 8.8 8.5–9.2 1.9 9.9 9.6–10.2 2.0 70 130 160 40 70 130 Control (µM) 160 9.1 8.8–9.4 2.1 T 24.3

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X/June 98 AV CI SD % 7.2 6.9–7.5 2.1 117 ns 11.5 11.2–11.8 1.9 105 ns 8.1 7.8–8.4 2.0 108 ns 7.9 7.6–8.2 2.0 105 ns 9.1 8.8–9.4 2.0 110 ns 11.2 10.9–11.5 2.0 111 ns 11.3 11.0–11.6 1.9 103 ns 9.8 9.5–10.1 2.0 7.4 7–7.8 2.2 99 ns 8.1 7.8–8.4 2.0 121 0.03 7.3 7.0–7.6 2.0 96 ns

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For abbreviations, see Table 2 footnote.

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On the basis of the present experiments and our previous work, it can be concluded that the MF effects provoked by combining MF and environmental stresses is not a simple phenomenon, (meaning that a stronger stress would evoke more pronounced MF effects). This is somewhat different from more conventional (e.g., chemical) environmental agents: a stronger stress normally also produces stronger effects on plants. If it is assumed that MF acts as a sort of a stressor, we would expect that the combined effects with the Al stress would be even stronger. However, our results show this is not the case. It is also known that environmental stress like that induced by heavy metals provokes the formation of different stress proteins in plants (16). The role of stress proteins in MF effects has already been suggested (4,17). The mechanism of specific MF effects in combination with a stress condition should probably be looked for in the specific production or inhibition of stress (or other) proteins susceptible to certain MF conditions. Calculations did not show the involvement of ion cyclotron resonance effects at the applied parameters in our experiments, for either Al3 or Cl . However, at suitable ion cyclotron resonance conditions (for instance if we change the frequency) it might be possible to enhance the bioeffects. This remains to be elucidated. Our research leaves much to be done in the future. Since two different influencing factors were used, many experiments should be performed to determine various possible windows, their sizes and their nature. Future research in this direction could be very lengthy, or a quicker method could be used. One possibility would be to test the synergistic effects of MFs and some stress factor by examining changes in biochemical markers known to be elicited in stress conditions, for example, putrescin.

ACKNOWLEDGMENTS
ˇ ˇ The authors express their gratitude to Prof. Dr. Anton Jeglic and Prof. Dr. Dusan Fefer of the Faculty for Electrical and Computer Engineering for their technical assistance and for kind advice. We also thank Mojca Piskernik, who edited the manuscript for English.

REFERENCES
ˇˇ 1. Ruzic, R., Jerman, I., and Gogala, N.: Water stress eveals effects of ELF magnetic fields on the growth of seedlings, Electro- Magnetobiol. 17, 17–30, 1998. ˇˇ 2. Ruzic, R., Jerman, I., and Gogala, N.: Effects of weak low-frequency magnetic fields on spruce seed germination under acid conditions, Can. J. For. Res. 28, 609–616, 1998. ˇˇ 3. Ruzic, R. and Jerman, I.: Influence of Ca2 in biological effects of direct and indirect ELF magnetic field stimulation, Electro- Magnetobiol. 17, 203–214, 1998. ¨ 4. Mittenzwey, R., Suβmuth, R., and Mei, W.: Effects of extremely low-frequency electromagnetic fields on bacteria—the question of co-stressing factor. Bioelectrochem. Bioenerg. 40, 21–27, 1996. 5. Bolognani, L., Francia, F., Venturelli, T., and Volpi, N.: Fermentative activity of cold-stressed yeast and effect of electromagnetic pulsed field, Electro- Magnetobiol. 11, 11–17, 1992.

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