Running head: JAMMING AVOIDANCE RESPONSE IN WEAKLY ELECTRIC FISH 1

Jamming Avoidance Response in Weakly Electric Fish

Mackenzie M. Andrews
University of Washington - Neurobiology
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Abstract
The ability to take in information about the environment is crucial to an organism’s survival.
While humans rely on sight, smell, sound, taste, and touch, some species of fish have evolved the
ability to survey their surroundings with electric fields. These weakly electric fish use an electric
organ to produce sinusoidal signals that can be used for navigation, communication, defense, and
prey detection. However, if a nearby fish has an electric organ discharge (EOD) rate close to that
of the fish’s own frequency, signal interference can hinder the ability to take in environmental
information. To avoid interference, weakly electric fish have evolved a jamming avoidance
response (JAR) in which they shift their own frequency away from the neighboring fish’s
frequency. To explore this behavior, we simulated a neighboring fish EOD and recorded the
response of the test fish. Our results confirm that a fish will shift its frequency up when the
neighboring frequency is lower and vice versa. We also show that the magnitude of change is
dependent on the magnitude of difference between the neighbor’s and fish’s own frequency.
Keywords: jamming avoidance response, electric organ discharge, Eigenmannia sp.
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Jamming Avoidance Response in Weakly Electric Fish

Introduction
While many animals rely on the familiar five senses of sight, smell, sound, taste, and
touch to take in environmental information, some species of fish have developed a sixth sense:
the ability to produce and sense electric fields. These species of fish have evolved an electric
organ which they use to produce an electric signal. The electric organ discharge (EOD) can be
used for navigation, communication, defense, and prey detection (Castello, M. E. et al., 2009).
Typically, the organ discharges at a constant rate (Bullock, T., Hamstra Jr., R., & Scheich, H.,
1972) and the fish detects changes in the amplitude and phase of their own signal to gather
information about their surroundings. However, if another nearby electric fish has an EOD of
similar frequency, the signals can interfere or ‘jam.’ In order to reduce the interference, the fish
exhibit a jamming avoidance response (JAR) in which each fish shifts their EOD frequency away
from their neighbor’s.
To investigate the JAR, we recorded the EOD of a weakly electric fish in the genus
Eigenmannia. Eigenmania produce sinusoidal electric signals between 200 and 600 Hz
(Hopkins, C., 1974). To test the JAR, we simulated a neighboring fish by producing a sinusoidal
signal of similar frequency to the test fish and recorded the EOD response of the test fish. We
recorded the response to neighboring signals of higher and lower frequency and analyzed the
time course of the JAR. The difference between the simulated neighboring signal and the test
fish signal was defined as ΔF and fish’s the change in EOD frequency in response to the
neighboring signal was defined as ΔR. We hypothesized that a negative ΔF would correlate to a
negative ΔR because Eigenmannia shift their frequency up when they have a higher EOD rate
than their neighbor and down when they have a lower EOD rate (Watanabe, A. & Takeda, K.,
1963). We also hypothesized that as the magnitude of ΔF increased, the magnitude of ΔR would
decrease because a smaller change in EOD rate would lead to unjamming when the signals have
greater separation.
Methods
Animal Preparation
A fish was placed into a mesh tube. Two sponges were then inserted into either end of the
tube to keep the fish from escaping. The tube was placed in a water tub with the temperature held
at a constant 24.4 degrees Celsius throughout all of the experiments.
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Wiring and Recording Setup

Two recording electrodes were placed into the mesh tube anterior and posterior to the
fish. The electrodes were connected to the extracellular amplifier with a gain of 100X. The signal
was then processed through the Humbug to remove 60 Hz noise. The output was then sent to
both the oscilloscope and audio monitor to monitor the signal in real time and to the PowerLab
box to be collected through LabChart and saved on the computer. A ground wire was placed in
the tank to ground the tank water.
To set the stimulus, a function generator was used. The output of the function generator
was sent to the PowerLab, stimulating electrodes, and to the oscilloscope to be displayed and
adjusted in real time. The stimulating electrodes were taped to the opposite side of the tank as the
fish and placed approximately a fish-length apart. Tape was placed on the ends of the electrodes
to dampen the signal.
Data Collection and Analysis
Sample data of the fish’s resting EOD was taken to get a baseline frequency value. All
frequency analyses were done in LabChart’s Fast Fourier Transform (FFT) spectrum analyzer.
We then found the dynamic range of the fish’s EOD rate. We did this by ‘chasing’ the fish’s EOD
by setting the stimulator frequency just below that of the fish and increased gradually until the
fish no longer increased its discharge rate. We repeated this starting just above the fish’s
frequency and chasing down. We then set the function generator to produce a frequency 5 Hz
below that of the fish and recorded the JAR. We repeated this for a stimulating frequency 5 Hz
greater than the fish as well. Time course data was taken by performing a frequency analysis for
every 2 seconds during and after stimulation. Since LabChart’s spectrum analyzer produces
discrete frequency bins, duplicate frequency points were deleted to produce a representative time
course curve without discrete step transitions. To produce a ΔF vs. ΔR plot, stimulator
frequencies of 0, 2, 5, 10, and 15 Hz above and below the resting EOD rate were delivered such
that ΔF = Fstimulator - Ffish and the change in the fish’s EOD was calculated such that ΔR = Fbefore –
Fafter.
For the previously explained experiments, constant stimulus amplitudes were used to
observe the fish’s JAR to a known ΔF. The fish was given time to recover to it’s natural EOD
rate between each test. The amplitude of recordings was dependent on the fish’s location relative
to the recording electrodes and does not reflect the strength of the fish’s signal.
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Results
Dynamic Range
In order to find the dynamic range of the fish’s EOD rate, we ‘chased’ the fish’s EOD
with our stimulator as described in the Methods. We found that the fish was able to modulate its
discharge rate between _____ Hz and _____ Hz, but did not modulate above or below this range
regardless of the stimulator frequency.
Resting EOD
To find the resting EOD frequency of the fish, we took a recording of the fish without
stimulation (Figure 1.a) and performed an FFT on the signal (Figure 1.c). The fish had a resting
discharge rate of 308 Hz.
JAR
Fstimulus Above Ffish
To observe the JAR of the fish to a stimulating frequency greater than that of the fish’s
resting EOD rate, the stimulator frequency was set to 313 Hz (5 Hz above the fish’s EOD).
When the stimulator was turned on, a 5 Hz beat frequency was created by the summation of the
fish and stimulator signals (Figure 1.b). As the fish began its JAR, the beat frequency increased.
The fish got to a steady adapted EOD rate of 303 Hz (Figure 1.d) after 35 seconds of stimulation.
Fstimulus Below Ffish
Conversely, to observe the JAR of the fish to a stimulating frequency lower than that of
the fish’s resting EOD rate, the stimulator frequency was set to 303 Hz (5 Hz below the fish’s
EOD). When the stimulator was turned on, a similar 5 Hz beat frequency was created. The fish
got to a steady adapted EOD rate of 313 Hz after 57 seconds of stimulation (Figure 2).
Time Course of Response
As shown in Figure 2, the JAR of the fish had a seemingly logarithmic rise in frequency
before reaching the maximum frequency of 313 Hz at 57 seconds of stimulation. This frequency
was maintained until the stimulator was disconnected at 92 seconds. The frequency then fell in
an inversely logarithmic fashion for the duration of the recording. The time course of recovery
was longer than that of initial adaptation. We recorded the recovery for 62 seconds after the
stimulus was tuned off and the EOD rate only got to 310 Hz during the recording period.
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ΔF vs. ΔR
To determine the relationship between the difference in stimulator frequency (ΔF) and the
change in EOD frequency (ΔR), we stimulated at ΔF’s of -15, 10, 5, 2, 0, 2, 5, 10, and 15 Hz.
The fish responded with ΔR’s of 0, -2, -4, -5, -7, 7, 4, 1, and 0 Hz, respectively (Figure 3). This
demonstrates that the threshold to elicit a JAR is a 10 Hz difference between the neighboring
frequency and the fish’s own frequency. This also confirms our first hypothesis that a negative
ΔF evokes a negative ΔR. In addition, the change in frequency during the JAR increases when
the neighboring frequency is closer to the resting EOD rate of the fish, confirming our second
hypothesis. When the stimulator was set to a ΔF of 0, the fish increased its EOD rate. Likely, our
stimulator frequency was slightly below that of the fish rather than exactly equal. It is unclear
how the fish would respond to an exact match in frequency.
Discussion
Our results show a number of characteristics of the JAR of Eigenmannia. As
demonstrated in Figure 1, Eigenmannia is able to adjust its EOD frequency in response to a
neighboring signal that is similar to its resting discharge rate. If the neighboring signal frequency
is lower than the fish’s EOD rate, the fish will increase its EOD rate in response. Likewise, if the
neighboring signal is of higher frequency, the fish will shift its EOD down (Figure 3). All of our
experiments show that the fish will adjust its EOD frequency until there is approximately a 10
Hz difference between the fish’s EOD and the neighboring signal. This can be explained by the
fact that the fish use low frequency phase and amplitude changes to survey their surroundings.
The low frequency beat created by a neighboring signal jams the ability of the fish to recognize
the amplitude and phase modulations of the environment. By increasing the difference between
the two signals, the beat frequency also increases and thus can be separated from the lower
frequency environmental modulations.
We also showed that the time course of recovery from an adapted signal is slower than
the initial adaptation (Figure 2). This could be explained by the fact that there is a survival need
for the fish to rapidly adjust its EOD rate when presented with an interfering signal. However,
when that signal is no longer present, the fish is capable of surveying its environment at the
adjusted frequency with equal efficacy to its preferred resting frequency, thus the need to return
the EOD back to rest is not as urgent. In addition, it is possible that the slower recovery allows
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for the fish to adapt faster if the same neighboring signal comes back into interference range
which would be likely in the case of two fish occupying a similar territory.
All of our experiments were performed with a constant stimulating frequency, which is
not representative of a natural JAR. In normal conditions, both fish would adjust their EOD rates
in opposite directions and thus decrease the time course and magnitude of the response. To
understand the naturalistic time course and magnitude of the JAR, experiments in which two fish
were present instead of an artificial signal would be required.
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References
Bosma, M., Perkel, D., Kennedy, M., Canfield, J., Hass, C., Sisneros, J. (2017). Introduction to
Systems and Behavioral Neurobiology.
Bullock, T., Hamstra Jr., R., Scheich, H. (1972) The Jamming Avoidance Response of High
Frequency Electric Fish. J. Comp. Physiol. 77:1-22.
Castello, M. E., Rodriguez-Cattaneo, A., Aguilera, L. Iribarne, P. A., Pereira, A. C., Caputi, A. A.
(2009). Waveform generation in the weakly electric fish Gymnotus coropinae
(Hoedeman): the electric organ and the electric organ discharge. Journal of Experimental
Biology, 212 (9): 1351–1364.
Hopkins, C. (1974) Electric communication: functions in the social behavior of Eigenmannia
virescens. Behaviour 50: 270-305.
Watanabe, A., Takeda, K. (1963) The change of discharge frequency by A.C. stimulus in a weak
electric fish. J. Exp. Biol. 40: 57-66.
JAMMING AVOIDANCE RESPONSE IN WEAKLY ELECTRIC FISH 9

Figure 1: Resting Discharge and Jamming Avoidance Response - a) Time domain recording
of the resting EOD showing a consistent sinusoidal frequency over a 55 millisecond time course.
b) Time domain recording of the beat frequency produced by interference between the 313 Hz
stimulus and fish’s EOD. The beat starts at 5 Hz and increases as the fish adjusts its EOD rate to
be farther away from the stimulus frequency. c) Frequency spectrum depiction of the resting
EOD rate from trace a of 308 Hz. Additional small peaks at 616 Hz and 824 Hz represent the
first and second harmonic frequencies. d) Frequency spectrum depiction of the adapted EOD
frequency during the stimulation in trace b. The fish adapted to a frequency of 303 Hz (tall peak)
to separate its self from the stimulator frequency of 313 Hz (smaller peak).
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Figure 2: Time Course of JAR and Recovery – The fish’s EOD frequency rose at a decreasing
rate from its resting frequency (308 Hz) before reaching the maximum frequency of 313 Hz at 57
seconds of stimulation. This frequency was maintained until the stimulator was disconnected at
92 seconds (blue dots). The frequency then fell at a decreasing rate for the remaining of the
recording, reaching 310 Hz 62 seconds after the stimulus was shut off (orange dots).
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Figure 3: ΔF vs ΔR - Using the real-time readout of the fish’s EOD on the oscilloscope, the
function generator was set to provide stimuli with ΔF’s of -15, 10, 5, 2, 0, 2, 5, 10, and 15 Hz.
After being given ample time to adapt, the fish responded with ΔR’s of 0, -2, -4, -5, -7, 7, 4, 1,
and 0 Hz, respectively.