Functions of EODs and Chirps

Because EOD frequency and chirping vary systematically within and between species, these signals can convey information about species identity, sex, reproductive condition, dominance, and/or motivation. How receivers use chirps and EOD frequency to evaluate conspecifics, however, is only partly understood. One of the main projects in our laboratory is to understand how fish respond to different types of electrocommunication signals.

We use several different approaches to assess the functions of EOD frequency and chirping in several different apteronotid species that differ in the degree and direction of sexual dimorphism in these signals:

(1) Playbacks of simulated electrocommunication signals to fish in a “chirp chamber.” One way in which we study the function of EODs and chirping is to record the responses of fish to playbacks of simulated EODs in a chirp chamber. The fish is placed in a shelter tube on the bottom or a tank or in a net “hammock” suspended in a tank with recording electrodes placed at the ends of the tank opposite the head and tail of the fish and playback electrodes placed on the other two sides of the tank. We can vary the frequency of the stimulus to simulate the presence of a fish of the same or opposite sex, a fish with a higher or lower EOD frequency (which in some species is an indicator of dominance or subordinance), or a fish of the same or different species. Most fish respond to playbacks by chirping. Indeed, some fish produce more than a hundred chirps during a two-minute playback. An advantage of using a chirp chamber is that because the fish, recording electrodes, and stimulus electrodes are in a fixed orientation, we can use relatively simple algorithms to isolate the subject fish’s EOD from the playback stimulus. This allows us to use customized software to precisely measure structural parameters of chirps (i.e. the amount of frequency modulation, duration, and amplitude modulation). This method thus allows us to compare how the structure of chirps of different species and sexes are affected by different types of social stimuli (Turner et al. 2007).

Chirp Chamber. A 10-gallon tank positioned in a cooler filled with water and with a heater connected to a temperature controller that maintains the tank temperature within 0.2 degrees C. The fish is placed in a shelter tube at the bottom of the tank or in a net ‘hammock” suspended in the middle of the tank. Carbon recording electrodes opposite the head and tail of the fish record the subject’ EOD. Playback electrodes located on either side of the fish are used to present simulated EODs of other fish.

Audio recording of a typical chirp chamber playback experiment. At the beginning of the recording, you can hear a single tone, which is the EOD of a male brown ghost knifefish in a chirp chamber (converted into a sound). Then, a second tone can be heard, which is a simulated EOD (also converted to a sound) being played by the second set of electrodes in the chirp chamber. The ‘blips’ in the recording are the subject fish producing chirps, in which it increases its EOD frequency and reduces the amplitude (loudness) of its EOD for a fraction of a second.

(2) Playbacks of electrocommunication signals to freely-swimming fish. Chirp chamber experiments allow us to precisely quantify the structure of chirps and other electrocommunication signals. Because fish in the chirp chamber remain in the recording tube or net hammock during the playback, the recording configuration is not very naturalistic, and we are unable to use this technique to study the physical interactions of the fish with different stimuli.  To study the range of behaviors that accompany electrocommunication signals, we use infrared-sensitive video cameras to record the behavior of freely-swimming fish interacting with playback stimuli or with the EODs of fish in an adjacent tank. We use this technique to study how the EODs and chirping of other fish affect aggression and courtship behavior.

EOD Playbacks to freely-swimming fish. Video taken in the dark (with infrared illumination) of a female black ghost knifefish (A. albifrons) during the presentation of a simulated EOD of a female conspecific through playback electrodes on the right side of the tank. Both males and females in this species are aggressive towards conspecifics. The female in this experiment repeatedly bit the electrodes during the playback. The continuous tone during the recording is the EOD of the subject fish being recorded by carbon electrodes on the four corners of the tank combined with a simulated “playback” EOD being presented from a pair of carbon electrodes on the right side of the tank. These electrical signals are converted into sounds so that we can hear them.

(3) Recording EODs and behavior of freely interacting fish. We also use electrode arrays and infrared video recording to analyze electrical signals and other behaviors. Although it is difficult to analyze the fine structure of chirps using this method, it allows us to examine how fish chirp in semi-natuarlistic social contexts. For example, we can determine whether dominant fish chirp more than subordinate fish and quantify behaviors that fish perform before and after they chirp or are chirped at.

Live interactions. Recording of a live dyadic interaction between two elephant nose ghost knifefish (Sternarchorhynchus sp.) recorded under infrared lighting (but otherwise completely dark). The sounds in the recording are the EODs of the two fish converted into sounds so that we can perceive them. The “blips” in the tones are chirps being produced by the fish.

We use these experimental techniques to address the following questions:

What information is conveyed by EOD frequency?

Do males and females (or fish of different species) respond differently to playbacks male vs. female EOD frequencies or to same-sex EODs with frequencies higher versus lower than the fishes’ own EOD? For example, both brown ghost knifefish and black ghost knifefish both produce two different types of chirps (high-frequency chirps and low-frequency chirps). By examining the stimuli that elicit these chirps, we found that the social context in which different types of chirps are produced differs between species. Males and females in both species produce low frequency chirps most often during same-sex interactions. In brown ghosts, however, high frequency chirps are produced mostly by males in response to female EODs, whereas in black ghosts both sexes produce high frequency chirps mostly in response to same-sex EODs (Kolodziejski et al. 2007). These results suggest that the function of analogous signals has changed in closely-related species.

What are the function of chirps and rises?

Do males and females (or fish of different species) respond differently to chirping versus non-chirping EODs? Do they respond differently to different types of chirps? Does the condition of the fish (reproductive condition, dominance status) influence how a fish chirps or responds to chirps? Do fish chirp more or less (or produce different types of chirps) at different times of day/night? Do fish behave more aggressively when they chirp or are chirped at? Do fish chirp back and forth at each other with “echo responses?” By correlating the behaviors fish perform before and after they chirp as well as the social, physiological, and physical contexts in which they chirp during live interactions, we are elucidating the meaning and function of chirps and other EOD modulations.

References

Kolodziejski, J.A., Sanford, S.E., and Smith, G.T. 2007. Stimulus frequency differentially affects chirping in two species of weakly electric fish: implications for the evolution of signal structure and function. J. Exp. Biol. 210:2501-2509.

Turner, C.R., Derylo, M., de Santana, C.D., Alves-Gomes, J.A., and Smith, G.T. 2007. Phylogenetic comparative analysis of electric communication signals in ghost knifefishes (Gymnotiformes: Apteronotidae). J. Exp. Biol. 210:4104-4122.