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Animals across species compete for limited resources. Whereas in some species competition behavior is solely based on the individual's own abilities, other species assess their opponents to facilitate these interactions. Using cues and communication signals, contestants gather information about their opponent, adjust their behavior accordingly, and can thereby avoid high costs of escalating fights. We tracked electrocommunication signals known as ‘rises’ and agonistic behaviors of the gymnotiform electric fish Apteronotus leptorhynchus in staged competition experiments. A larger body size relative to the opponent was the sole significant predictor for winners. Sex and the frequency of the continuously emitted electric field only mildly influenced competition outcome. In males, correlations of body size and winning were stronger than in females and, especially when losing against females, communication and agonistic interactions were enhanced, suggesting that males are more motivated to compete. Fish that lost competitions emitted the majority of rises, but their quantity depended on the competitors’ relative size and sex. The emission of a rise could be costly since it provoked ritualized biting or chase behaviors by the other fish. Despite winners being accurately predictable based on the number of rises after the initial 25 min, losers continued to emit rises. The number of rises emitted by losers and the duration of chase behaviors depended in similar ways on physical attributes of contestants. Detailed evaluation of these correlations suggests that A. leptorhynchus adjusts its competition behavior according to mutual assessment, where rises could signal a loser's motivation to continue assessment through ritualized fighting., Summary: Electric fish adjust their competition behavior according to mutual assessment, where electrocommunication with so-called ‘rises’ could signal a loser's motivation to continue assessment through ritualized fighting.

Understanding how individuals detect and recognize signals emitted by conspecifics is fundamental to discussions of animal communication. The species pair Gymnotus omarorum and Brachyhypopomus gauderio, found in syntopy in Uruguay, emit species-specific electric organ discharges that can be sensed by both species. The aim of this study was to unveil whether either of these species are able to identify a conspecific electric organ discharge, and to investigate distinctive recognition signal features. We designed a forced-choice experiment using a natural behavior (i.e. tracking electric field lines towards their source) in which each fish had to choose between a conspecific and a heterospecific electric field. We found a clear pattern of preference for a conspecific waveform even when pulses were played within 1 Hz of the same rate. By manipulating the time course of the explored signals, we found that the signal features for preference between conspecific and heterospecific waveforms were embedded in the time course of the signals. This study provides evidence that pulse Gymnotiformes can recognize a conspecific exclusively through species-specific electrosensory signals. It also suggests that the key signal features for species differentiation are probably encoded by burst coder electroreceptors. Given these results, and because receptors are sharply tuned to amplitude spectra and also tuned to phase spectra, we extend the electric color hypothesis used in evaluation of objects to apply to communication signals.

Mormyrid weakly electric fish have a special electrosensory modality that allows them to actively sense their environment and to communicate with conspecifics by emitting sequences of electric signals. Electroreception is mediated by different types of dermal electroreceptor organs for active electrolocation and electrocommunication, respectively. During electrocommunication, mormyrids exhibit stereotyped discharge sequences and locomotor patterns, which can be induced by playback of electric signals. This raises the question, what sensory information is required to initiate and sustain social interactions, and which electrosensory pathway mediates such interactions. By experimentally excluding stimuli from vision and the lateral line system, we show that Mormyrus rume proboscirostris can rely exclusively on its electrosensory system to track a mobile source of electric communication signals. Detection of electric playback signals induced discharge cessations, followed by double-pulse patterns. The animals tried to interact with the moving signal source and synchronized their discharge activity to the playback. These behaviors were absent in control trials without playback. Silencing the electric organ in some fish did not impair their ability to track the signal source. Silenced fish followed on trajectories similar to those obtained from intact animals, indicating that active electrolocation is no precondition for close-range interactions based on electrocommunication. However, some silenced animals changed their strategy when searching for the stationary playback source, which indicates passive sensing. Social interactions among mormyrids can therefore be induced and mediated by passive reception of electric communication signals without the need for perception of the location of the signal source through other senses.

In this study we examined electrocommunication behavior in Sternarchogiton nattereri (Apteronotidae), a weakly electric fish from South America. We focused on variation between females and males lacking external dentition and used playbacks of simulated conspecifics to elicit chirps (modulations of their electric organ discharge, EOD). Chirp responses were not affected by the frequency of the playback stimulus. EOD frequency, chirp rate, and chirp duration were not sexually dimorphic; however, the amount of chirp frequency modulation was significantly greater in toothless males than in females. These results reinforce that sex differences in chirp structure are highly diverse and widespread in the Apteronotidae.

Summary The ghost knifefishes (family Apteronotidae) are one of the most successful and diverse families of electric fish. Like other weakly electric fish, apteronotids produce electric organ discharges (EODs) that function in electrolocation and communication. This review highlights the diversity in the structure, function and sexual dimorphism of electrocommunication signals within and across apteronotid species. EOD frequency (EODf) and waveform vary as a function of species, sex and/or social rank. Sex differences in EODf are evolutionarily labile; apteronotid species express every pattern of sexual dimorphism in EODf (males>females; males

Electrocommunication in weakly electric fish has various functions, such as species recognition and information transfer during social encounters. For communication, weakly electric fish rely on their electric organ discharges (EOD), which are generated by an electric organ located in the fish’s tail. While in mormyrids the species-specific EOD-waveform remains rather constant, the highly variable inter-discharge intervals (IDIs) enable the fish to modify their signalling behaviour on a short time scale. Different IDI-patterns were found to correlate with certain behavioural situations. We investigated electrocommunication in a social group of five freely swimming Mormyrus rume during different group behaviours. We found several electrical discharge patterns, which occurred during group foraging, aggressive encounters, fleeing and resting. We were able to describe several so far unknown signal patterns, such as synchronous discharging of two fish, and fixed-order-signalling in groups of up to five fish. Our results show that electrocommunication in social groups consists of complex and rapidly changing interactions between different individuals.

SUMMARY Environmental complexity and season both influence brain cell proliferation in adult vertebrates, but their relative importance and interaction have not been directly assessed. We examined brain cell proliferation during both the breeding and non-breeding seasons in adult male electric fish, Brachyhypopomus gauderio, exposed to three environments that differed in complexity: (1) a complex natural habitat in northern Uruguay, (2) an enriched captive environment where fish were housed socially and (3) a simple laboratory setting where fish were isolated. We injected fish with BrdU 2.5 h before sacrifice to label newborn cells. We examined the hindbrain and midbrain and quantified the density of BrdU+ cells in whole transverse sections, proliferative zones and two brain nuclei in the electrocommunication circuitry (the pacemaker nucleus and the electrosensory lateral line lobe). Season had the largest effect on cell proliferation, with fish during the breeding season having three to seven times more BrdU+ cells than those during the non-breeding season. Although the effect was smaller, fish from a natural environment had greater rates of cell proliferation than fish in social or isolated captive environments. For most brain regions, fish in social and isolated captive environments had equivalent levels of cell proliferation. However, for brain regions in the electrocommunication circuitry, group-housed fish had more cell proliferation than isolated fish, but only during the breeding season (season × environment interaction). The regionally and seasonally specific effect of social environment on cell proliferation suggests that addition of new cells to these nuclei may contribute to seasonal changes in electrocommunication behavior.

The South American weakly-electric knifefish (Apteronotidae) produce highly diverse and readily quantifiable electrocommunication signals. The electric organ discharge frequency (EODf), and EOD modulations (chirps and gradual frequency rises (GFRs)), vary dramatically across sexes and species, presenting an ideal opportunity to examine the proximate and ultimate bases of sexually dimorphic behavior. We complemented previous studies on the sexual dimorphism of apteronotid communication signals by investigating electric signal features and their hormonal correlates in Apteronotus bonapartii, a species which exhibits strong sexual dimorphism in snout morphology. Electrocommunication signals were evoked and recorded using a playback paradigm, and were analyzed for signal features including EOD frequency and the structure of EOD modulations. To investigate the androgenic correlates of sexually dimorphic EOD signals, we measured plasma concentrations of testosterone and 11-ketotestosterone. A. bonapartii responded robustly to stimulus playbacks. EODf was sexually monomorphic, and males and females produced chirps with similar durations and amounts of frequency modulation. However, males were more likely than females to produce chirps with multiple frequency peaks. Sexual dimorphism in apteronotid electrocommunication signals appears to be highly evolutionarily labile. Extensive interspecific variation in the magnitude and direction of sex differences in EODf and in different aspects of chirp structure suggest that chirp signals may be an important locus of evolutionary change within the clade. The weakly-electric fish represent a rich source of data for understanding the selective pressures that shape, and the neuroendocrine mechanisms that underlie, diversity in the sexual dimorphism of behavior.

The great variety of species-typical electric signals (electric organ discharges, EOD) emitted by weakly electric mormyrid fish might be the result of evolutionary pressures stemming from the two main functions of the electro-sensory-motor system: electrocommunication and electrolocation. Employing a conditioned discrimination task we tested four species of mormyrids, emitting EODs differing in waveform, for their ability to detect capacitive properties of objects during electrolocation. Each fish could discriminate capacitive objects within a certain range of capacitive values, which was species specific. The upper and lower limits (upper and lower thresholds) of this detectable range were determined for each fish. In fish species emitting long duration EODs composed of mainly low spectral frequencies both the lower and the upper thresholds were shifted to larger capacitive values compared to fish species emitting shorter EODs. The upper limit of the detectable range was much more variable between species than the lower limit, which was relatively low in all fish. We interpret this as an adaptation of mormyrids to detect small capacitive objects, for example food items. All mormyrids could discriminate between a resistive object and a capacitive object even if the complex impedances of the two objects were identical. This implies that the fish are highly sensitive to small waveform distortions of their self produced EODs.

Weakly electric fishes emit electric organ discharges (EODs) from their tail electric organs and sense feedback signals from their EODs by electroreceptors in the skin. The electric sense is utilized for various behaviors, including electrolocation, electrocommunication, and the Jamming avoidance response (JAR). For each behavior, various types of sensory Information are embedded in the transient electrical signals produced by the fish. These temporal signals are sampled, encoded, and further processed by peripheral and central neurons specialized for time coding. There are time codes for the sex or species Identities of other fish or the resistance and capacitance of objects. In the central nervous system, specialized neural elements exist for decoding time codes for different behavioral functions. Comparative studies allow phylogenetic comparison of time-coding neural systems among weakly electric fishes.

Signals emitted preceding and during combat can aid in sequential assessment of opponent quality and motivation. Signal reliability can be maintained by physical constraints, by costs of production or receiver retaliation, or can be cost-free, when contestants have a common interest. In staged dyadic contests over a shelter, male brown ghost knifefish, Apteronotus leptorhynchus, modulate the frequency of their electric organ discharge (EODF) and perform increasingly costly behaviours as fights escalate. Relative body length was the best predictor of fight duration and victory, and although there were initially no differences in electrical signalling rates between contestants, through the course of the interaction ultimate winners made increasingly more abrupt EODF increases (‘chirps’) and fewer gradual frequency rises (‘GFRs’) than losers. This is consistent with previous hypotheses that chirps signal aggression and dominance, whereas GFRs indicate submission. However, fine temporal analysis revealed that both signal types are good predictors of impending attack: paradoxically, just prior to bouts of combat, combat initiators gradually increase EODF significantly more than receivers and continue to do so throughout the bout. We discuss the putative functions of these signals.

SUMMARY During social interactions, apteronotid electric fish modulate their electric organ discharges (EODs) to produce transient communication signals known as chirps. Chirps vary widely across species and sex in both number and structure. In Apteronotus leptorhynchus, males chirp far more than females and their chirps have greater frequency modulation than those of females. High-frequency chirps are produced by males most often in response to female-like electric signals. As such, they have been hypothesized to function in courtship. The more common low-frequency chirps, produced by both males and females in response to same-sex signals, are hypothesized to function as aggressive signals. To determine whether the two chirp types in the closely related Apteronotus albifrons have similar functions, we stimulated chirping in male and female A. leptorhynchus and A. albifrons with playbacks simulating the EODs of same-sex versusopposite-sex conspecifics. As in A. leptorhynchus, male and female A. albifrons produced low-frequency chirps most often to same-sex signals. Unlike A. leptorhynchus, however, A. albifrons also produced more high-frequency chirps to same-sex stimuli than to opposite-sex stimuli. This suggests that high-frequency chirps in A. albifrons,unlike those in A. leptorhynchus, may not function as courtship signals and that the function of similar chirp types has diversified in Apteronotus. Examples such as this, in which the function of a communication signal has changed in closely related species, are rare. The electrocommunication signals of apteronotids may thus provide a remarkable opportunity to investigate the evolutionary interactions of signal structure and function.

SUMMARY Mormyrid electric fish rely on the waveform of their electric organ discharges (EODs) for communicating species, sex, and social status, while they use the sequences of pulse intervals (SPIs) for communicating rapidly changing behavioral states and motivation. Little is known of electric signaling during courtship behavior because of two major difficulties: (1) the fish are not easily bred in captivity and (2) there is no reliable means of separating electric signals from several individuals in natural communication settings. Through simulating artificial rain conditions, we have successfully induced courtship and succeeded in breeding a mormyrid electric fish(Brienomyrus brachyistius) in the laboratory. We have also developed a system of video recording and editing combined with cross correlation analysis to precisely record and view behavior and separate EODs from two individuals in non-breeding and breeding contexts. Knowing the electrical and motor patterns during courtship allows for further exploration of topics such as mate choice and neural basis of pattern generation in these fish. Here we describe nine common motor displays and 11 SPIs. Analysis of frequency of occurrences suggests that some SPI patterns are sex and season specific. We also observed electrical duetting called `rasp matching' during courtship signaling among pairs; males and females exchange `rasps' and`bursts', respectively, in alternation. Our study employs new techniques to separate and document SPIs in the context of courtship. We show that some SPIs correlate with specific behavioral acts around the time of spawning.

The fixed part of the electrocommunication signal, the electric organ discharge (EOD) waveform, is well differentiated in the two vicariant dwarf stonebasher species, Pollimyrus castelnaui and P. marianne. However, differentiation regarding the variable, situation-dependent part, i.e., inter-discharge interval (IDI) patterns, has never been studied in a pair of sibling species of mormyrid fish. We here compare the electrical signalling that accompanies different motor behaviours (such as resting and swimming, territorial agonistic interactions, courtship and spawning) in the two species. Double pulse patterns of regularly alternating short IDIs of 8-11 ms and long ones of 16-100 ms accompanied threat displays in both species. In three pairs of P. marianne and five pairs of P. castelnaui, courtship was characterised by nest building, territory patrolling and acoustic displays (advertisment calls) that were accompanied by long discharge breaks in the male and highly regular IDIs around 50 ms in the female of both species. Nest-tending males showed IDI sequences consisting of regularly alternating double pulse patterns, similar to threat displays. During spawning both sexes generated stereotyped IDI sequences of a low discharge rate. All IDI patterns occurring in one species were also found in the other, and no species-specifity was identified at that level. Playback experiments contrasting conspecific and heterospecific IDI sequences (that had been recorded from nocturnally swimming fish) revealed preferences in none of the six experimental subjects. Double pulse patterns, high discharge rate displays (HD) and regularisations of the IDI sequence accompanying specific behaviours occurred in similar form in both dwarf stonebasher species of the present study. Therefore, we conclude that in the speciation of P. castelnaui and P. marianne the fixed part of the EOD, its waveform, was under more differential selection pressure than its variable part, the patterns of IDI.

SUMMARYElectrocommunication signals of electric fish vary across species, sexes and individuals. The diversity of these signals and the relative simplicity of the neural circuits controlling them make them a model well-suited for studying the mechanisms, evolution and sexual differentiation of behavior. In most wave-type gymnotiform knifefishes, electric organ discharge (EOD)frequency and EOD modulations known as chirps are sexually dimorphic. In the most speciose gymnotiform family, the Apteronotidae, EOD frequency is higher in males than females in some species, but lower in males than females in others. Sex differences in EOD frequency and chirping, however, have been examined in only three apteronotid species in a single genus, Apteronotus. To understand the diversity of electrocommunication signals, we characterized these behaviors in another genus, Adontosternarchus. Electrocommunication signals of Adontosternarchus devenanzii differed from those of Apteronotus in several ways. Unlike in Apteronotus, EOD frequency was not sexually dimorphic in A. devenanzii. Furthermore,although A. devenanzii chirped in response to playbacks simulating conspecific EODs, the number of chirps did not vary with different stimulus frequencies. A. devenanzii chirps also differed in structure from Apteronotus chirps. Whereas Apteronotus species produce functionally distinct chirp types differing in frequency modulation (FM), A. devenanzii produced only high-frequency chirps that had either single or multiple frequency peaks. Males produced more multi-peaked chirps than females. Thus, the temporal structure of chirps, rather than the amount of FM, delineated chirp types in A. devenanzii. Our results demonstrate that the structure, function and sexual dimorphism of electrocommunication signals are evolutionary labile in apteronotids and may be useful for understanding the diversity of sexually dimorphic behavior.

The electric organ discharge (EOD) in Brachyhypopomus pinnicaudatus is modified by temperature and reproductive state. We studied the influence of these variables upon a complex behavior, the jamming avoidance response (JAR). Experiments were performed in non-reproductive fish and in two groups of fish after the induction of reproductive state (by nature or by acclimation at 28 degrees C). JARs were elicited at 20 and 30 degrees C by free-run electric stimuli with different deltaLs (interval difference between stimulus and EOD). In non-reproductive fish, JARs induced by stimuli with +deltaLs showed temperature sensitivity, with smaller responses at 30 degrees C. Conversely, similar JARs were obtained at both temperatures in reproductive fish. These observations were replicated in curarized preparations. Stimuli with -deltaLs were almost ineffective in non-reproductive fish at 30 degrees C, whereas adequate JARs were shown by reproductive fish. Phase-locked stimuli were used to evaluate the duration of the low-threshold electrosensory periods preceding and following the EOD. In non-reproductive fish, the temperature step induced a shortening of these periods. The opposite effect was observed in reproductive fish, probably explaining the differences in JAR capability. A prolongation of the low-threshold periods would favor the perception of electrocommunication signals during courtship. JAR changes would be a consequence of this adaptation.

Brown ghost knife fish, Apteronotus leptorhynchus, produce a continuous electric organ discharge (EOD) that they use for communication. While interacting aggressively, males also emit brief EOD modulations termed chirps. The simplicity of this behaior and its underlying neural circuitry has made it an important model system in neuroethology. Chirping is typically assayed by confining a fish in a tube (‘chirp chamber’) and presenting it with sine wave electrical stimuli that partially mimic EODs of other fish. We presented male fish with progressively more realistic social stimuli to examine whether some of the stimulus complexities during dyadic interaction influence the production of chirps. In a chirp chamber, fish chirped less to a recording of an EOD containing chirps than to a recording of an EOD alone and to sine wave stimuli. Free-swimming fish chirped more to stimulus fish than to sine wave stimuli presented through electrodes. Fish chirped more when interacting directly than when interacting across a perforated barrier. Together, these studies demonstrate that the presence of chirps, electric field complexity, and/or non-electric social stimuli are important in eliciting chirp production in brown ghosts.

SUMMARYWeakly electric fish communicate and electrolocate objects in the dark by discharging their electric organs (EOs) and monitoring the spatiotemporal pattern of current flow through their skin. In the South-American pulse-type gymnotid fish these organs often are intriguingly complex, comprising several hundreds of electrogenic cells (electrocytes) of various morphologies,innervation patterns and abilities to generate a spike, distributed over nearly the full length of the fish. An attractive idea is that different parts of the organ may serve distinct functions in electrocommunication and electrolocation. Recent studies support this notion and suggest that the currents produced during the final phase of the electric organ discharge (EOD)are used for communication. Here, we explore a method to directly assess the relevance of the various currents for electrolocation. In this new method, the pattern of current flow during a gymnotid EOD is changed selectively at distinct phases of the EOD so that currents generated by known electrocyte groups are affected. We have studied the roles played by the various currents for the detection of novel feedback at the trunk/tail region of the gymnotid fish Gymnotus carapo. An experimental animal rested in a cage and two electrodes were placed at a close distance to its trunk and tail. An electronic switch briefly connected these electrodes during a selected phase within an EOD and the shunting of EOD current that resulted from switch closure was directly monitored. G. carapo responded with an acceleration of its discharge rate to novelties in the EOD feedback that occurred only for a fraction of a single EOD. Controls in which the switch was closed during the silent intervals between successive EODs showed that the fish responded to the changes in EOD feedback and not to unrelated artefacts of the brief switch closure. Fish responded to shunting of current during all phases; the sensitivity was highest during the final headnegative phase but the magnitude of shunted current was largest in the preceeding phase. The current produced during the final part of the EOD is thus not reserved for communication as previously suggested but plays a predominant role in electrolocation at the trunk and tail region of G. carapo.

We measured and mapped the electric fields produced by three species of neotropical electric fish of the genus Brachyhypopomus (Gymnotiformes, Rham phichthyoidea, Hypopomidae), formerly Hypopomus. These species produce biphasic pulsed discharges from myogenic electric organs. Spatio-temporal false-color maps of the electric organ discharges measured on the skin show that the electric field is not a simple dipole in Brachyhypopomus. Instead, the dipole center moves rostro-caudally during the 1st phase (P1) of the electric organ discharge, and is stationary during the 2nd phase (P2). Except at the head and tip of tail, electric field lines rotate in the lateral and dorso-ventral planes. Rostrocaudal differences in field amplitude, field lines, and spatial stability suggest that different parts of the electric organ have undergone selection for different functions; the rostral portions seem specialized for electrosensory processing, whereas the caudal portions show adaptations for d.c. signal balancing and mate attraction as well. Computer animations of the electric field images described in this paper are available on web sites http:/(/) www.bbb.caltech.edu/ElectricFish or http:/(/)www.fiu.edu/-stoddard/electric fish.html.

Weakly electric fish produce electric signals (electric organ discharges, EODs) with a specialised electric organ creating an electric field around their body. Objects within this field alter the EOD-induced current at epidermal electroreceptor organs, which are distributed over almost the entire body surface. The detection, localisation and analysis of objects performed by monitoring self-produced electric signals is called active electrolocation. Electric fish employ active electrolocation to detect objects that are less than 12 cm away and have electric properties that are different from those of the surrounding water. Within this range, the mormyrid Gnathonemus petersii can also perceive the distance of objects. Depth perception is independent of object parameters such as size, shape and material. The mechanism for distance determination through electrolocation involves calculating the ratio between two parameters of the electric image that the object projects onto the fish’s skin. Electric fish can not only locate objects but can also analyse their electrical properties. Fish are informed about object impedance by measuring local amplitude changes at their receptor organs evoked by an object. In addition, all electric fish studied so far can independently determine the capacitative and resistive components of objects that possess complex impedances. This ability allows the fish to discriminate between living and non-living matter, because capacitance is a property of living organisms. African mormyrids and South American gymnotiforms use different mechanisms for capacitance detection. Mormyrids detect capacitance-evoked EOD waveform distortions, whereas gymnotiforms perform time measurements. Gymnotiforms measure the temporal phase shift of their EODs induced at body parts close to the object relative to unaffected body parts further away.

How do the communication discharges produced by electric fish evolve to accommodate the unique design features for the modality? Two design features are considered: first, the limited range of signaling imposed on the electric modality by the physics of signal transmission from dipole sources; and second, the absence of signal echoes and reverberations for electric discharges, which are non-propagating electrostatic fields. Electrostatic theory predicts that electric dicharges from fish will have a short range because of the inverse cube law of geometric spreading around an electrostatic dipole. From this, one predicts that the costs of signaling will be high when fish attempt to signal over a large distance. Electric fish may economize in signal production whenever possible. For example, some gymnotiform fish appear to be impedance-matched to the resistivity of the water; others modulate the amplitude of their discharge seasonally and diurnally. The fact that electric signals do not propagate, but exist as electrostatic fields, means that, unlike sound signals, electric organ discharges produce no echoes or reverberations. Because temporal information is preserved during signal transmission, receivers may pay close attention to the temporal details of electric signals. As a consequence, electric organs have evolved with mechanisms for controlling the fine structure of electric discharge waveforms.

Gymnotiform electric fish emit an electric organ discharge that, in several species, is sexually dimorphic and functions in gender recognition. In addition, some species produce frequency modulations of the electric organ discharge, known as chirps, that are displayed during aggression and courtship. We report that two congeneric species (Apteronotus leptorhynchus and A. albifrons) differ in the expression of sexual dimorphism in these signals. In A. leptorhynchus, males chirp more than females, but in A. albifrons chirping is monomorphic. The gonadosomatic index and plasma levels of 11-ketotestosterone were equivalent in both species, suggesting that they were in similar reproductive condition. Corresponding to this difference in dimorphism, A. leptorhynchus increases chirping in response to androgens, but chirping in A. albifrons is insensitive to implants of testosterone, dihydrotestosterone or 11-ketotestosterone. Species also differ in the sexual dimorphism and androgen sensitivity of electric organ discharge frequency. In A. leptorhynchus, male discharge at higher frequencies than females, and androgens increase electric organ discharge frequency. In A. albifrons, males discharge at lower frequencies than females, and androgens decrease electronic organ discharge frequency. Thus, in both chirping and electric organ discharge frequency, evolutionary changes in the presence or direction of sexual dimorphism have been accompanied and perhaps caused by changes in the androgen regulation of the electric organ discharge.

Summary The balance between the costs and benefits of conspicuous animal communication signals ensures that signal expression relates to the quality of the bearer. Signal plasticity enables males to enhance conspicuous signals to impress mates and competitors and to reduce signal expression to lower energetic and predation-related signaling costs when competition is low. While signal plasticity may benefit the signaler, it can compromise the reliability of the information conveyed by the signals. In this paper we review the effect of signal plasticity on the reliability of the electrocommunication signal of the gymnotiform fish Brachyhypopomus gauderio. We (1) summarize the endocrine regulation of signal plasticity, (2) explore the regulation of signal plasticity in females, (3) examine the information conveyed by the signal, (4) show how that information changes when the signal changes, and (5) consider the energetic strategies used to sustain expensive signaling. The electric organ discharge (EOD) of B. gauderio changes in response to social environment on two time scales. Two hormone classes, melanocortins and androgens, underlie the short-term and long-term modulation of signal amplitude and duration observed during social interaction. Population density drives signal amplitude enhancement, unexpectedly improving the reliability with which the signal predicts the signaler's size. The signal's second phase elongation predicts androgen levels and male reproductive condition. Males sustain signal enhancement with dietary intake, but when food is limited, they ‘go for broke’ and put extra energy into electric signals. Cortisol diminishes EOD parameters, but energy-limited males offset cortisol effects by boosting androgen levels. While physiological constraints are sufficient to maintain signal amplitude reliability, phenotypic integration and signaling costs maintain reliability of signal duration, consistent with theory of honest signaling.

Social interactions dramatically affect the brain and behavior of animals. Studies in birds and mammals indicate that socially induced changes in adult neurogenesis participate in the regulation of social behavior, but little is known about this relationship in fish. Here, we review studies in electric fish ( Apteronotus leptorhychus ) that link social stimulation, changes in electrocommunication behavior and adult neurogenesis in brain regions associated with electrocommunication. Compared with isolated fish, fish living in pairs have greater production of chirps, an electrocommunication signal, during dyadic interactions and in response to standardized artificial social stimuli. Social interaction also promotes neurogenesis in the periventricular zone, which contributes born cells to the prepacemaker nucleus, the brain region that regulates chirping. Both long-term chirp rate and periventricular cell addition depend on the signal dynamics (amplitude and waveform variation), modulations (chirps) and novelty of the stimuli from the partner fish. Socially elevated cortisol levels and cortisol binding to glucocorticoid receptors mediate, at least in part, the effect of social interaction on chirping behavior and brain cell addition. In a closely related electric fish ( Brachyhypopomus gauderio ), social interaction enhances cell proliferation specifically in brain regions for electrocommunication and only during the breeding season, when social signaling is most elaborate. Together, these studies demonstrate a consistent correlation between brain cell addition and environmentally regulated chirping behavior across many social and steroidal treatments and suggest a causal relationship. * 11-KT : 11-ketotestosterone BrdU : bromodeoxyuridine CP : central posterior thalamic nucleus dF : difference frequency EOD : electric organ discharge PPn : prepacemaker nucleus PVZ : periventricular zone

During the day, weakly electric snoutfish, such as Marcusenius altisambesi from the Okavango delta, avoid visually oriented predators by hiding in sheltered, dark places where they discharge their electric organs at a low and variable rate, interspersed with occasional short bursts (mean discharge rate, 4–12 Hz). Hence, histograms of inter-discharge intervals (IDI) are broad and bimodal (IDI range, about 15–500 ms; “variable IDI pattern”). We report here that with a female neighbor in electrical communication reach, captive males of M. altisambesi (N = 4) each showed a novel type of IDI resting pattern that was characterized by a higher and more constant discharge rate (16–28 Hz). These IDI histograms were unimodal and narrow (IDI range, about 11–100 ms; “regularized IDI pattern”). In each of these males, the regularized pattern vanished when the female neighbors were replaced by males, and the common variable IDI pattern of low rate was observed instead. In an unforced choice paradigm, six M. altisambesi experimental females were allowed to choose between two electric fish decoys, one playing back the novel regularized IDI pattern and one playing back the variable IDI pattern. Five experimental females significantly preferred staying close to the decoy playing back the regularized IDI pattern, whereas one female showed the opposite preference. It appears that males advertise to females during their diurnal period of overt inactivity, with an inconspicuous signal that neither threatens conspecifics nor alerts predators by overt behavior. A secondary function of the regularized male IDI pattern could be to advance the reproductive cycle of females.

Apteronotus leptorhynchus, a gymnotiform fish, produces highly regular electric organ discharges of 600–1000 Hz. Short-term modulations of the electric organ discharge ("chirps") were elicited by imitating the discharges of neighboring fish. Chirps displayed an increase in frequency of approximately 100 Hz, a duration of about 15 ms, and an absolute amplitude of 0.5–2 mV. Since, similar to natural conditions, chirps summated with the beat caused by interference of the fish's own electric organ discharge and the imitating discharge, the size and shape of the chirp's amplitude envelope varied greatly according to its phase relative to the beat cycle; however, the frequency of the chirp amplitude modulation was always 50–100 Hz. All 21 males examined chirped, but their rate of chirping varied considerably (range 2–59 chirps/30 s; mean 22 chirps/30 s). In contrast, only one out of nine females chirped (mean 0.25 chirps/30 s). The latency between stimulus onset and first chirp was variable and often long (range 1.0–25.0 s; median 3.3 s). We propose that chirps are not a sensory reflex but a communicatory behavior regulated by hypothalamic peptidergic input.

SUMMARY Brown ghost knife fish (Apteronotus leptorhynchus) can briefly increase their electric organ discharge (EOD) frequency to produce electrocommunication signals termed chirps. The chirp rate increases when fish are presented with conspecific fish or high-frequency (700–1100 Hz) electric signals that mimic conspecific fish. We examined whether A. leptorhynchus also chirps in response to artificial low-frequency electric signals and to heterospecific electric fish whose EOD contains low-frequency components. Fish chirped at rates above background when presented with low-frequency (10–300 Hz) sine-wave stimuli; at 30 and 150 Hz, the threshold amplitude for response was 1 mV cm–1. Low-frequency (30 Hz) stimuli also potentiated the chirp response to high-frequency (∼900 Hz) stimuli. Fish increased their chirp rate when presented with two heterospecific electric fish, Sternopygus macrurus and Brachyhypopomus gauderio, but did not respond to the presence of the non-electric fish Carassius auratus. Fish chirped to low-frequency (150 Hz) signals that mimic those of S. macrurus and to EOD playbacks of B. gauderio. The response to the B. gauderio playback was reduced when the low-frequency component (

[Marcusenius pongolensis (South Africa) and M. altisambesi (Upper Zambezi/Okavango) differ in the sex difference of their electric organ discharges (EODs). It is permanent and graded in the former and seasonally dimorphic in the latter. Four out of six experimental female M. altisambesi gave significantly stronger responses to the long playback EODs of breeding males rather than short EODs, whereas four of seven experimental males responded in the opposite way (with one exception in association time). When re-tested after a year, the female preference for long EODs had vanished. Both sexes discriminated between playback EODs as either long or short (unlike M. pongolensis), with the cutoff between 1420 and 2470 μs (female EOD average, 350 μs). Exposure to social stimuli from tank neighbours of both sexes did not induce growth of EOD duration in males (unlike M. pongolensis). We suggest that in M. altisambesi the sexually dimorphic male EOD has evolved under female (inter-sexual) selection (in M. pongolensis, both inter- and intra-sexual selection); that there is no evidence for male–male competition (intra-sexual selection, present in M. pongolensis); that ecological differences between a floodplain (M. altisambesi) and a more riverine (M. pongolensis) species may be at the origin of the differentiation., Marcusenius pongolensis (South Africa) and M. altisambesi (Upper Zambezi/Okavango) differ in the sex difference of their electric organ discharges (EODs). It is permanent and graded in the former and seasonally dimorphic in the latter. Four out of six experimental female M. altisambesi gave significantly stronger responses to the long playback EODs of breeding males rather than short EODs, whereas four of seven experimental males responded in the opposite way (with one exception in association time). When re-tested after a year, the female preference for long EODs had vanished. Both sexes discriminated between playback EODs as either long or short (unlike M. pongolensis), with the cutoff between 1420 and 2470 μs (female EOD average, 350 μs). Exposure to social stimuli from tank neighbours of both sexes did not induce growth of EOD duration in males (unlike M. pongolensis). We suggest that in M. altisambesi the sexually dimorphic male EOD has evolved under female (inter-sexual) selection (in M. pongolensis, both inter- and intra-sexual selection); that there is no evidence for male–male competition (intra-sexual selection, present in M. pongolensis); that ecological differences between a floodplain (M. altisambesi) and a more riverine (M. pongolensis) species may be at the origin of the differentiation.]

Waveform analysis of the electric organ discharges (EODs) of the weakly electric fish Gymnotus carapo found significant variation between individuals but little variation in EODs from the same individual. That is, EOD waveforms are individually distinctive. A playback experiment obtained results that were comparable to those used to infer neighbour recognition in other territorial species. These two results show that the electric signals of this species of weakly electric fish have individually distinctive variation and that individual variation can be perceived by the fish, independently of possible cues contained within the inter-pulse interval distribution. This is the first time that individual discrimination has been demonstrated in any species of electric fish.

Weakly electric fi shes have been an important model system in behavioral neuroscience for more than 40 years. These fi shes use a specialized electric organ to produce an electric fi eld that is typically below 1 volt/cm and serves in many behaviors including social communication and prey detection. Electrical behaviors are easy to study because inexpensive and widely available tools enable continuous monitoring of the electric fi eld of individual or groups of interacting fi sh. Weakly electric fi sh have been routinely used in tightly controlled neurophysiological experiments in which the animal is immobilized using neuromuscular blockers (e.g., curare). Although experiments that involve immobilization are generally discouraged because it eliminates movement-based behavioral signs of pain and distress, many observable electrosensory behaviors in fipersist when the animal is immobilized. Weakly electric fi sh thus offer a unique opportunity to assess the effects of immobilization on behaviors including those that may refl ect pain and distress. We investigated the effects of both immobilization and restraint on a variety of electrosensory behaviors in four species of weakly electric fi shes and observed minor effects that were not consistent between the species tested or between particular behaviors. In general, we observed small increases and decreases in response magnitude to particular electrosensory stimuli. Stressful events such as asphyxiation and handling, however, resulted in signifi cant changes in the fi shes’ electrosensory behaviors. Signs of pain and distress include marked reductions in responses to electrosensory stimuli, inconsistent responses, and reductions in or complete cessation of the autogenous electric fi eld.

SUMMARY Brown ghost knifefish, Apteronotus leptorhynchus, are a species of weakly electric fish that produce a continuous electric organ discharge (EOD)that is used in navigation, prey capture and communication. Stereotyped modulations of EOD frequency and amplitude are common in social situations and are thought to serve as communication signals. Of these modulations, the most commonly studied is the chirp. This study presents a quantitative analysis of chirp production in pairs of free-swimming, physically interacting male and female A. leptorhynchus. Under these conditions, we found that in addition to chirps, the fish commonly produce a second signal type, a type of frequency rise called abrupt frequency rises, AFRs. By quantifying the behaviours associated with signal production, we find that Type 2 chirps tend to be produced when the fish are apart, following periods of low aggression,whereas AFRs tend to be produced when the fish are aggressively attacking one another in close proximity. This study is the first to our knowledge that quantitatively describes both electrocommunication signalling and behavioural correlates on a subsecond time-scale in a wave-type weakly electric fish.

SUMMARYTo understand the evolution of sexually dimorphic communication signals, we must quantify their costs, including their energetic costs, the regulation of these costs, and the difference between the costs for the sexes. Here, we provide the first direct measurements of the relative energy expended on electric signals and show for the focal species Brachyhypopomus pinnicaudatus that males spend a significantly greater proportion of their total energy budget on signal generation (11–22%) compared with females (3%). Both sexes significantly reduce the energy spent on electric signals during daylight hours through circadian modulation of the amplitude,duration and repetition rate of the electric signal, but this effect is more marked in males. Male body condition predicted the energy spent on electric signals (R2=0.75). The oxygen consumed by males for signal production closely paralleled the product of the electric signal's waveform area (R2=0.99) and the discharge rate(R2=0.59), two signal parameters that can be assessed directly by conspecifics. Thus the electric communication signal of males carries the information to reveal their body condition to prospective mates and competing males. Because the electric signal constitutes a significant fraction of the energy budget, energy savings, along with predation avoidance,provides an adaptive basis for the production of circadian rhythms in electric signals.

SUMMARYElectrocommunication signals in electric fish are diverse, easily recorded and have well-characterized neural control. Two signal features, the frequency and waveform of the electric organ discharge (EOD), vary widely across species. Modulations of the EOD (i.e. chirps and gradual frequency rises) also function as active communication signals during social interactions, but they have been studied in relatively few species. We compared the electrocommunication signals of 13 species in the largest gymnotiform family,Apteronotidae. Playback stimuli were used to elicit chirps and rises. We analyzed EOD frequency and waveform and the production and structure of chirps and rises. Species diversity in these signals was characterized with discriminant function analyses, and correlations between signal parameters were tested with phylogenetic comparative methods. Signals varied markedly across species and even between congeners and populations of the same species. Chirps and EODs were particularly evolutionarily labile, whereas rises differed little across species. Although all chirp parameters contributed to species differences in these signals, chirp amplitude modulation, frequency modulation (FM) and duration were particularly diverse. Within this diversity,however, interspecific correlations between chirp parameters suggest that mechanistic trade-offs may shape some aspects of signal evolution. In particular, a consistent trade-off between FM and EOD amplitude during chirps is likely to have influenced the evolution of chirp structure. These patterns suggest that functional or mechanistic linkages between signal parameters(e.g. the inability of electromotor neurons increase their firing rates without a loss of synchrony or amplitude of action potentials) constrain the evolution of signal structure.

The weakly electric fish Apteronotus leptorhynchus produces wave-like electric organ discharges distinguished by a high degree of regularity. Transient amplitude and frequency modulations ("chirps") can be evoked in males by stimulation with the electric field of a conspecific. During these interactions, the males examined in this study produced six types of chirps, including two novel ones. Stimulation of a test fish with a conspecific at various distances showed that two electrically interacting fish must be within 10 cm of each other to evoke chirping behavior in the neighboring fish. The chirp rate of all but one chirp type elicited by the neighboring fish was found to be negatively correlated with the absolute value of the frequency difference between the two interacting fish, but independent of the sign of this difference. Correlation analysis of the instantaneous rates of chirp occurrence revealed two modes of interactions characterized by reciprocal stimulation and reciprocal inhibition. Further analysis of the temporal relationship between the chirps generated by the two fish during electric interactions showed that the chirps generated by one individual follow the chirps of the other with a short latency of approximately 500-1,000 ms. We hypothesize that this "echo response" serves a communicatory function.

Bulldog fish (Marcusenius macrolepidotus) generate short (

Mormyrid fish generate electric organ discharges (EODs) continuously for electrocommunication and electrolocation. EOD pulse waveforms are, within species limits, individually specific and usually stable over long periods of time. We conducted playback experiments with the African weakly electric snoutfish,Marcusenius macrolepidotus (South African form), or bulldog fish, originating from the Incomati River System in South Africa. Previous field observations had shown an increase of male, but not female, EOD pulse duration with body size. Therefore, we tested the hypothesis that EOD pulse duration might be of relevance in territorial male-male contests. We determined whether or not aggression scores recorded for males kept in adjacent aquarium compartments depended on playback EOD duration. Playback EODs had been pre-recorded either from familiar or from unfamiliar conspecific males, and varied naturally in duration. As independently confirmed in all five experimental subjects, the attack rate on a dipole model playing back strangers' EODs from the territorial boundary increased significantly with stimulus EOD pulse duration (that varied from 329 μs to 975 μs in our field sample of playback EODs). This result was confirmed when one specific EOD, the duration of which was artificially decreased or increased, was used for playback, showing that in male M. macrolepidotus EODs of so long duration signal a greater threat than EODs of shorter duration. Therefore, EOD duration may be relevant for males in the assessment of their rivals' size and fighting potential. Moreover, experimental subjects confronted with familiar playback EODs showed a significantly lower rate of attack than expected for unfamiliar EODs of the same duration. An individual familiar neighbour's EOD evoked a significantly weaker attack rate when played back from the regular rather than from the opposite territorial boundary. These observations are compatible with the 'dear enemy' hypothesis.

The present paper highlights the relationship between social status and production of gradual frequency rises in interacting Apteronotus albifrons. The gradual frequency rise production was mathematically inferred and a discrete classification deliberately avoided. The results showed little gradual frequency rise production before the hierarchy settlement. Afterwards, only the dominant fish kept this gradual frequency rise production at low levels, while the subdominant fish drastically increased it in all following interaction contexts. The hypothesis of gradual frequency rises being involved in communication as submissive signals was thus strengthened.

Female communication behaviors are often overlooked by researchers in favor of male behaviors, which are usually more overt and easier to elicit. Very little is known about female electrocommunication behaviors in brown ghost knifefish, a weakly electric wavetype Gymnotiform fish. Most behavioral studies have focused on males, and fish are usually restrained and played a stimulus near their own electric organ discharge frequency to evoke chirps (abrupt short-term frequency rises) or the jamming avoidance response. Our study focuses on categorizing and describing spontaneous and evoked electric organ discharge modulations in free-swimming female fish that were either electrically coupled to tanks containing a conspecific (male or female), or left isolated. Cluster analysis of signals produced under isolated and social conditions revealed three categories of rises: short rise, medium rise and long rise; and one category of frequency decrease (dip). Females produce significantly more short rises when electrically coupled to tanks containing lower-frequency females, and produce more long rises when electrically coupled to tanks containing males. Short rises may have an intrasexual aggressive function, while long rises may serve as an advertisement of status or reproductive condition in intersexual interactions.

Some gymnotiform electric fish modulate their electric organ discharge for intraspecific communication. In Apteronotus leptorhynchus, chirps are usually rapid (10-30 ms) modulations that are activated through non- N-methyl- d-aspartate (non-NMDA) glutamate receptors in the hindbrain pacemaker nucleus. Males produce longer chirp types than females and chirp at higher rates. In Apteronotus albifrons, chirp rate is sexually monomorphic, but chirp structure (change in frequency and amplitude during a chirp) was unknown. To better understand the neural regulation and evolution of chirping behavior, we compared chirp structure in these two species under identical stimulus regimes. A. albifrons, like A. leptorhynchus, produced distinct types of chirps that varied, in part, by frequency excursion. However, unlike in A. leptorhynchus, chirp types in A. albifrons varied little in duration, and chirps were all longer (70-200 ms) than those of A. leptorhynchus. Chirp type production was not sexually dimorphic in A. albifrons, but within two chirp types males produced longer chirps than females. We suggest that species differences in chirp duration might be attributable to differences in the relative proportions of fast-acting (non-NMDA) and slow-acting (NMDA) glutamate receptors in the pacemaker. Additionally, we map species difference onto a phylogeny and hypothesize an evolutionary sequence for the diversification of chirp structure.

Aperonotus leptorhynchus (Gymnotiformes) produces wave-like electric organ discharges distinguished by a high degree of constancy. Transient frequency and amplitude modulations of these discharges occur both spontaneously and during social interactions, which can be mimicked by external electrical stimulation. The so-called chirps can be divided into four different types. Independent of the type of chirp produced under spontaneous conditions, the fish generate only significant numbers of type-2 chirps under evoked conditions. The rate of production of chirps of this type is largely determined by the frequency relative to the fish's frequency and signal intensity. Frequencies of + 10 Hz of the fish's own discharge frequency most effectively elicit chirps. Type-2 chirps can also be evoked through stimulation at or near the higher harmonic frequencies of the fish's frequency, but the chirp rate decreases with increasing number of the higher harmonic component. Over a certain range, the rate of production of type-2 chirps increases with increasing stimulus intensity. At very high intensities the generation of type-2 chirps is accompanied by the production of a novel type of electrical signal ("abrupt frequency rise") characterized by a frequency increase of approximately 20 Hz and high repetition rates of roughly 10 s(-1). We hypothesize that the different types of electric modulations subserve different behavioral functions.

South African bulldogs (Marcusenius macrolepidotus, Mormyridae) generate brief (less than 1 ms) electric organ discharges (EODs), separated by much longer and highly variable inter-discharge intervals (IDIs). The diurnal and nocturnal overt behaviour and electrical activity were studied under various conditions: in isolated fish, in pairs of fish, and in a group of four fish that were kept in a “natural” large aquarium. EODs from up to four individuals were recorded simultaneously and identified. While resting during the day, isolated fish showed a broad inter-individual variability of IDI patterns, with distribution histogram modes ranging from 85.7 ms to 325.8 ms. When foraging during the day, IDI modes were shorter and less variable (36.3–48.3 ms). Behaviour patterns displayed during nocturnal agonistic encounters were retreating, parallel swimming, anti-parallel display, attack, and fleeing/chasing. High-discharge-rate (HD) displays were observed at several stages of these encounters, for example, during antiparallel display (a period of low overt motor activity), or following attacks. IDI durations as short as 11 ms occurred during HD displays, which followed low-rate inter-HD activity almost without transition. IDI distribution histogram modes when fish showed anti-parallel display were 15.4 ms and 24.8 ms, and 30.0 ms during nocturnal nonagonistic interactions. No overt fighting was observed once a dominance relationship was established. In a large aquarium, an approaching dominant male evoked a simultaneous discharge arrest in a group of three subdominant males.

SUMMARY South American weakly electric fish produce a variety of electric organ discharge (EOD) amplitude and frequency modulations including chirps or rapid increases in EOD frequency that function as agonistic and courtship and mating displays. In Apteronotus leptorhynchus, chirps are readily evoked by the presence of the EOD of a conspecific or a sinusoidal signal designed to mimic another EOD, and we found that the frequency difference between the discharge of a given animal and that of an EOD mimic is important in determining which of two categories of chirp an animal will produce. Type-I chirps (EOD frequency increases averaging 650Hz and lasting approximately 25ms) are preferentially produced by males in response to EOD mimics with a frequency of 50–200Hz higher or lower than that of their own. The EOD frequency of Apteronotus leptorhynchus is sexually dimorphic: female EODs range from 600 to 800Hz and male EODs range from 800 to 1000Hz. Hence, EOD frequency differences effective in evoking type-I chirps are most likely to occur during male/female interactions. This result supports previous observations that type-I chirps are emitted most often during courtship and mating. Type-II chirps, which consist of shorter-duration frequency increases of approximately 100Hz, occur preferentially in response to EOD mimics that differ from the EOD of the animal by 10–15Hz. Hence these are preferentially evoked when animals of the same sex interact and, as previously suggested, probably represent agonistic displays. Females typically produced only type-II chirps. We also investigated the effects of arginine vasotocin on chirping. This peptide is known to modulate communication and other types of behavior in many species, and we found that arginine vasotocin decreased the production of type-II chirps by males and also increased the production of type-I chirps in a subset of males. The chirping of most females was not significantly affected by arginine vasotocin.

Local electric fields generated by the electric organ discharge of Gymnotus carapo were explored at selected points on the skin of an emitter fish (‘local self-generated fields’) and on the skin of a conspecific (‘local conspecific-generated fields’) using a specially designed probe. Local self-generated fields showed a constant pattern along the body of the fish. At the head, these fields were collimated, much stronger than elsewhere on the fish, and had a time waveform that was site-independent. This waveform consisted of a slow head-negative wave followed by a faster head-positive wave. In contrast, time waveforms in the trunk and tail regions were site-specific, with field vectors that changed direction over time. Local conspecific-generated fields were similar to the head-to-tail field, but their spatio-temporal pattern at the skin depended on the relative orientation between the receiving fish and the emitting fish. Because self-generated fields had a slow early component at the head region, they displayed a low-frequency peak in their power spectral density histograms. In contrast, the conspecific-generated fields had time waveforms with a sharper phase reversal, resulting in a peak at higher frequency than in the self-generated field. Lesions in emitting fish demonstrated that waveform components generated by the trunk and tail regions of the electric organ predominate in conspecific-generated fields, whereas waveform components generated by the abdominal region prevail in self-generated fields. Similar results were obtained from Brachyhypopomus pinnicaudatus. These results suggest that, in pulse-emitting gymnotids, electrolocation and electrocommunication signals may be carried by different field components generated by different regions of the electric organ.

Brown ghosts, Apteronotus leptorhynchus, are weakly electric gymnotiform fish whose wave-like electric organ discharges are distinguished by their enormous degree of regularity. Despite this constancy, two major types of transient electric organ discharge modulations occur: gradual frequency rises, which are characterized by a relatively fast increase in electric organ discharge frequency and a slow return to baseline frequency; and chirps, brief and complex frequency and amplitude modulations. Although in spontaneously generated gradual frequency rises both duration and amount of the frequency increase are highly variable, no distinct subtypes appear to exist. This contrasts with spontaneously generated chirps which could be divided into four "natural" subtypes based on duration, amount of frequency increase and amplitude reduction, and time-course of the frequency change. Under non-evoked conditions, gradual frequency rises and chirps occur rather rarely. External stimulation with an electrical sine wave mimicking the electric field of a neighboring fish leads to a dramatic increase in the rate of chirping not only during the 30 s of stimulation, but also in the period immediately following the stimulation. The rate of occurrence of gradual frequency rises is, however, unaffected by such a stimulation regime.

Beyond continuation of currently active areas, some less studied ones are selected for predictions of marked advance. (1) Most discoveries will be in cellular components and molecular mechanisms for different classes of receptor or central pathways. (2) More major taxa, possibly birds, reptiles or invertebrates, will be found to have electroreceptive species representing independent evolutionary ‘inventions’. (3) Electric organs with weak and episodic electric discharges will be found in new taxa; first, among siluriforms. (4) New examples are to be expected, such as lampreys, where synchronized muscle action potentials sum to voltages in the range of weakly electric fish. Some of these will look like intermediates in the evolution of electric organs. (5) Ethological significance will be found for a variety of known physiological features; e.g. uranoscopids, skates and weakly electric catfish with episodic electric discharges; electroreceptive ability of animals such as lampreys, chimaeras, lungfish, sturgeons, paddlefish and salamanders with Lorenzinian-type ampullae; gymnotiform and mormyrid detection of the capacitative component of impedance. (6) The organization of some higher functions in the cerebellum and forebrain will gradually come to light.

Weakly electric fish generate an electric field surrounding their body by means of an electric organ typically located within the trunk and tail. Electroreceptors scattered over the surface of the body encode the amplitude and timing of the electric organ discharge (EOD), and central components of the electrosensory system analyze the information provided by the electroreceptor afferents. The electrosensory system is used for electrolocation, for the detection and analysis of objects near the fish which distort the EOD and for electrocommunication. Since the electric organ is typically located in the tail, any movement of this structure relative to the rest of the body alters the EOD field, resulting in large changes in receptor afferent activity. The amplitude of these reafferent stimuli can exceed the amplitudes of near-threshold electrolocation signals by several orders of magnitude. This review summarizes recent studies of the South American weakly electric fish Apteronotus leptorhynchus aimed at determining how the animals differentiate self-generated or reafferent electrosensory stimuli from those that are more behaviorally relevant. Cells within the earliest stages of central electrosensory processing utilize an adaptive filtering technique which allows the system preferentially to attenuate reafferent as well as other predictable patterns of sensory input without degrading responses to more novel stimuli. Synaptic plasticity within the system underlies the adaptive component of the filter and enables the system to learn to reject new stimulus patterns if these become predictable. A Ca2+-mediated form of postsynaptic depression contributes to this synaptic plasticity. The filter mechanism seen in A. leptorhynchus is surprisingly similar to adaptive filters described previously in mormyrid weakly electric fish and in elasmobranchs, suggesting that this mechanism may be a common feature of sensory processing systems.

Behavioral and electrophysiological experiments were conducted to examine whether frequency-modulated electrocommunication signals are detected by the ampullary electroreceptor system in Eigenmannia. First, frequency-modulated electric organ discharges were found to contain a low-frequency component that could be detected by the ampullary system. Second, fish were successfully trained to distinguish a frequency-modulated signal, which contained a low-frequency component as in natural signals, from an artificial signal in which the low-frequency component was eliminated but still modulated in frequency. Subsequently, the trained fish responded without reinforcement to a low-frequency sinusoidal signal which mimicked the low-frequency component in the frequency-modulated signal, suggesting that the fish used the ampullary system to detect frequency modulation. Finally, physiological recording from ampullary afferent fibers demonstrated that they respond to frequency-modulated signals as predicted from the signal's low-frequency component. Electrophysiological study also showed that detection of frequency modulation by the ampullary system is immune to the presence of other constant electric organ discharges.

We present detailed measurements of the electric organ discharge of the weakly electric fish, Eigenmannia sp. These maps illuminate, with high resolution in both space and time, the electric organ discharge potential and electric field patterns in the water about the fish and on the skin surface itself. The results demonstrate that the electric organ discharge of Eigenmannia approximates a simple oscillating dipole, which confirms previous descriptions and assumptions, but is in contrast to the electric organ discharges of several other gymnotiform species. Over each cycle of Eigenmannia's electric organ discharge, the electric field amplitude measured at any point near the fish oscillates from positive to negative, but the field vector remains nearly constant in direction. This electric organ discharge pattern is correlated with known anatomical and physiological features of the fish's electric organ, and confirms that the activation of electrocytes comprising the organ is well synchronized. As a result, the relatively simple electric organ discharge leads to a fairly uniform pattern of electrosensory stimuli along the body surface, which may facilitate central processing of electrosensory images. Electric organ discharge maps and animations resulting from this series of studies are available via the Internet (http:@www.bbb.caltech.edu/ElectricFish, or www.fiu.edu/ approximately stoddard/electricfish.html).

The present study describes a measurement-based model of electric image generation in the weakly electric mormyrid fish Gnathonemus petersii. Measurements of skin impedance, internal resistivity and fish body dimensions have been used to generate an electrical-equivalent model of the fish and to calculate electrical images and equivalent dipole sources for elementary resistive objects. These calculations allow us to understand how exafferent and reafferent signals are sensed by electroreceptors. An object's electric image consists of the modulation of the transcutaneous voltage profile generated by the fish's own discharge. The results suggest a set of rules for electrolocation: (1) the side of the fish where modulation is larger indicates the side on which the object is situated; (2) the object's position in the electroreceptive field is indicated by the point of maximum modulation of the transcutaneous voltage; (3) the degree of focus of the image indicates the distance to the object. In addition, center-surround opposition originating at pre-receptor level is proposed. Both experimental measurements and modeling indicate that fish skin impedance is relatively low (400-11 000IMG src="/images/symbols/capomega.gif" WIDTH="13" HEIGHT="13" ALIGN="BOTTOM" NATURALSIZEFLAG="3"cmSUP2/SUP) and mainly resistive. This low skin impedance appears to enhance the local electric organ discharge modulation, the center-surround effect, the signal-to-noise ratio for electrolocation and the active space for electrocommunication.

The electric organ discharges (EODs) of Marcnsenius senegalensis, a West African freshwater fish, are bipolar pulses of short duration (220 ± SE 13µs )I.n males (n= 10; 10.1-13.1 cm standard length-which is around the size of getting mature), the duration of EOD pulses was of significantly greater variance than in females (n = 9; 9.8-12.8 cm standard length). Male EODs also showed a tendency for a longer duration than female EODs. Groups of three as well as of 14 M. senegalensis formed temporary schools in a 'naturally' equipped 720-l tank. While swimming slowly in a loose school during their nocturnal active phase, fish discharged in irregular long-short-long inter-EOD interval patterns. Near neighbours displayed a tendency to discharge in intervals of similar duration (nearest neighbour distance < 1/2 fish length). On removal of a plastic partition that had separated a pair of fish for at least 3 days, mutual threat displays followed by fighting were observed. During threatening, the fish alternated regularly between bursts of a high discharge rate and short discharge breaks; the rate of change was 4/s. The subdominant animal in a group of two was attacked frequently and often ceased discharging when the dominant fish approached. Courtship behaviour involving gonadally mature fish was accompanied by highdischarge- rate displays with intervals of constant duration in both fish, and the reciprocal display of 'preferred' EOD latencies in the 12 ms range. The results demonstrate electric communication by distinct inter-discharge interval patterns in the social behaviour of this mormyrid fish.