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Electrical discharges (EDs) in the African sharptooth catfish Clarias gariepinus were studied. Irregular monopolar EDs, lasting for 8–10 ms and longer, and more complex EDs, lasting for 20–50 ms and characterized by specific indentation of the discharge pattern, were recorded in pairs of individuals exhibiting both aggressive and defensive behavior. The pattern of long discharges was reconstructed by adding short EDs with different latencies and amplitudes. The conditions for the formation of long EDs with asynchronous activity of electromotor neurons and the possibility of coordinated operation of the electric generator and electroreceptor systems in catfish were considered, since an increase in the duration of EDs lowers the threshold of their perception. [ABSTRACT FROM AUTHOR]

The patterns of electric discharges were studied in two species of upside-down catfishes, Synodontis clarias and S. membranaceus. In S. clarias, constant generation of weak electrical pulses with a relatively stable frequency (~30 s–1) and an amplitude reaching 200 μV was found. In representatives of both species, both relatively short two-phase discharges of two types, differing in the total duration and ratio of positive and negative phases, and long-term impulses of various shapes with characteristic frontal irregularities were recorded at aggressive-defensive behavior in pairs of specimens. In S. membranaceus, short biphasic discharges of both the first and second types were recorded. Using a simple computer model, an attempt was made to reconstruct the process of the formation of long-term electric discharges from short elementary pulses. [ABSTRACT FROM AUTHOR]

Environmental signals act as input and are processed across successive stages in the brain to generate a meaningful behavioral output. However, a ubiquitous observation is that descending feedback projections from more central to more peripheral brain areas vastly outnumber ascending feedforward projections. Such projections generally act to modify how sensory neurons respond to afferent signals. Recent studies in the electrosensory system of weakly electric fish have revealed novel functions for feedback pathways in that their transformation of the afferent input generates neural firing rate responses to sensory signals mediating perception and behavior. In this review, we focus on summarizing these novel and recently uncovered functions and put them into context by describing the more “classical” functions of feedback in the electrosensory system. We further highlight the parallels between the electrosensory system and other systems as well as outline interesting future directions.

Environmental signals act as input and are processed across successive stages in the brain to generate a meaningful behavioral output. However, a ubiquitous observation is that descending feedback projections from more central to more peripheral brain areas vastly outnumber ascending feedforward projections. Such projections generally act to modify how sensory neurons respond to afferent signals. Recent studies in the electrosensory system of weakly electric fish have revealed novel functions for feedback pathways in that their transformation of the afferent input generates neural firing rate responses to sensory signals mediating perception and behavior. In this review, we focus on summarizing these novel and recently uncovered functions and put them into context by describing the more "classical" functions of feedback in the electrosensory system. We further highlight the parallels between the electrosensory system and other systems as well as outline interesting future directions. [ABSTRACT FROM AUTHOR]

Mormyrid weakly electric fish produce electric organ discharges (EODs) for active electrolocation and electrocommunication. These pulses are emitted with variable interdischarge intervals (IDIs) resulting in temporal discharge patterns and interactive signaling episodes with nearby conspecifics. However, unequivocal assignment of interactive signaling to a specific behavioral context has proven to be challenging. Using an ethorobotical approach, we confronted single individuals of weakly electric Mormyrus rume proboscirostris with a mobile fish robot capable of interacting both physically, on arbitrary trajectories, as well as electrically, by generating echo responses through playback of species-specific EODs, thus synchronizing signals with the fish. Interactive signaling by the fish was more pronounced in response to a dynamic echo playback generated by the robot than in response to playback of static random IDI sequences. Such synchronizations were particularly strong at a distance corresponding to the outer limit of active electrolocation, and when fish oriented toward the fish replica. We therefore argue that interactive signaling through echoing of a conspecific's EODs provides a simple mechanism by which weakly electric fish can specifically address nearby individuals during electrocommunication. Echoing may thus enable mormyrids to mutually allocate social attention and constitute a foundation for complex social behavior and relatively advanced cognitive abilities in a basal vertebrate lineage. [ABSTRACT FROM AUTHOR]

Latency-dependent responses to artificial electric stimuli in 11 species of the Nilotic mormyrids were studied. In total, more than 350 poststimulus time histograms from 132 individuals were analyzed. All species studied exhibited the latency-dependent responses. Both main types of responses - phase locking and phase avoidance behavior or the preferred latency response (PLR) and preferred latency avoidance (PLA) - were found in eight species ( Brevimyrus niger, Hippopotamyrus pictus, Marcusenius cyprinoides, Mormyrus kannume, M. hasselquistii, Petrocephalus bane, P. bovei and 'Pollimyrus' petherici), while only PLR were recorded from the remaining three species ( Hyperopisus bebe, Mormyrops anguilloides and Mormyrus caschive). In eight species both types of behavior were found in fish of the same sex. In four species both types of behavior were found both in females and males. Finally, in four species both types of behavior were revealed in the same individual (in total, there were eight such individuals). In some individuals of different species the atypical responses with substantially increased latencies (compared to the typical PLR and PLA) were observed. The possible role of the phase locking and phase avoidance behavior in the mormyrid electrolocation and communication is discussed. [ABSTRACT FROM AUTHOR]

Some weakly electric fish can use electric signals to interact and communicate with each other in dark and complex underwater environments where traditional underwater communication fails. In our previous work, we developed a bio-inspired electrocommunication system (BECS) that serves as an effective alternative to traditional methods in this challenging underwater scenario performing communication at a speed of approximately 1200 bps (bits per second) within approximately 3 m. In this study, a novel underwater wireless communication system (BECS-II) is proposed to upgrade the BECS with much better performance. We first propose theoretical and simulation models for electrocommunication, including the effects of the angular frequency and electrode impedance. A custom-made digital communication system is employed in BECS-II to improve the anti-interference ability and channel capacity of the BECS. In addition, a novel circuit optimization strategy was used to develop a customized circuit to enhance the transmitting and receiving capabilities of the BECS-II. Dual-frequency communication is proposed to meet the communication demands of different tasks by taking inspiration from the task allocation and evolution mechanisms of weakly electric fish. The experimental results showed that BECS-II outperformed BECS in high-frequency mode at both the communication speed (approximately 20 kbps) and distance (approximately 10 m), whereas in low-frequency mode, it extended the communication range by transmitting data up to a distance of approximately 20 m at a speed of approximately 200 bps. A substantial increase in the communication distance can expand the robot motion space in a group and improve group flexibility., (© 2023 IOP Publishing Ltd.)

Electric fishes modulate their electric organ discharges with a remarkable variability. Some patterns can be easily identified, such as pulse rate changes, offs and chirps, which are often associated with important behavioral contexts, including aggression, hiding and mating. However, these behaviors are only observed when at least two fish are freely interacting. Although their electrical pulses can be easily recorded by non-invasive techniques, discriminating the emitter of each pulse is challenging when physically similar fish are allowed to freely move and interact. Here we optimized a custom-made software recently designed to identify the emitter of pulses by using automated chirp detection, adaptive threshold for pulse detection and slightly changing how the recorded signals are integrated. With these optimizations, we performed a quantitative analysis of the statistical changes throughout the dominance contest with respect to Inter Pulse Intervals, Chirps and Offs dyads of freely moving Gymnotus carapo . In all dyads, chirps were signatures of subsequent submission, even when they occurred early in the contest. Although offs were observed in both dominant and submissive fish, they were substantially more frequent in submissive individuals, in agreement with the idea from previous studies that offs are electric cues of submission. In general, after the dominance is established the submissive fish significantly changes its average pulse rate, while the pulse rate of the dominant remained unchanged. Additionally, no chirps or offs were observed when two fish were manually kept in direct physical contact, suggesting that these electric behaviors are not automatic responses to physical contact. [ABSTRACT FROM AUTHOR]

Five variables of the electric organ discharge (EOD) were studied in 132 freshly caught individuals of Mormyrus kannume from the Gibe and Gojeb rivers (the enclosed Omo-Turkana system) and Baro River (the White Nile system) in south-western Ethiopia, East Africa. Most individuals exhibited the typical biphasic EODs with an initial head-positive phase followed by a head-negative phase exceeding initial one in the relative amplitude. In three repeated samples from the Gibe, stable differences in the discharge variables between females and males were found, while no sex differences were observed in other samples. In three cases, a positive correlation between the individual size and discharge duration was found: in two male subsamples from the Gibe and in a mixed sample of both sexes from the Gojeb. In most samples studied, some males exhibited the outstandingly long EOD duration. At the same time, no relationships between the discharge variables and reproductive status of individuals were found. The possible influence of individual and intra-individual variations on the results of inter-population and interspecific comparisons of the discharge variables in mormyrids are highlighted. [ABSTRACT FROM AUTHOR]

Mormyrid fishes communicate using pulses of electricity, conveying information about their identity, behavioral state, and location. They have long been used as neuroethological model systems because they are uniquely suited to identifying cellular mechanisms for behavior. They are also remarkably diverse, and they have recently emerged as a model system for studying how communication systems may influence the process of speciation. These two lines of inquiry have now converged, generating insights into the neural basis of evolutionary change in behavior, as well as the influence of sensory and motor systems on behavioral diversification and speciation. Here, we review the mechanisms of electric signal generation, reception, and analysis and relate these to our current understanding of the evolution and development of electromotor and electrosensory systems. We highlight the enormous potential of mormyrids for studying evolutionary developmental mechanisms of behavioral diversification, and make the case for developing genomic and transcriptomic resources. A complete mormyrid genome sequence would enable studies that extend our understanding of mormyrid behavior to the molecular level by linking morphological and physiological mechanisms to their genetic basis. Applied in a comparative framework, genomic resources would facilitate analysis of evolutionary processes underlying mormyrid diversification, reveal the genetic basis of species differences in behavior, and illuminate the origins of a novel vertebrate sensory and motor system. Genomic approaches to studying the evo-devo-neuroethology of mormyrid communication represent a deeply integrative approach to understanding the evolution, function, development, and mechanisms of behavior. [ABSTRACT FROM AUTHOR]

Current underwater communication typically includes acoustic, optical, radio frequency, and magneto-inductive channels. Wireless sensor networks are usually built on these four channels. However, these underwater networks are vulnerable to complex aquatic environments. In nature, weakly electric fish are able to communicate electrically (called electrocommunication), which is 'invisible' to most other animals, to convey information such as species, courtship, and environmental conditions. Inspired by the electrocommunication of weakly electric fish, an artificial electrocommunication system that uses an electric induction (EI) channel has been developed recently. This paper further develops an underwater networking system using the EI channel, which addresses the solutions to collision avoidance and routing problems during electrocommunication networking. In particular, a CSMA/CA-based electrocommunication mechanism was used to solve the collision problem. Then, a single-hop underwater electrocommunication network (UEN) was established. Furthermore, a complex multi-hop UEN was implemented on the basis of the ad hoc on-demand distance vector routing protocol. Theoretical analysis, simulations, and experiments were conducted to demonstrate the effectiveness of the developed UEN. Extensive results show that the UEN holds the potential to serve as a complement to future underwater wireless sensor networks., (© 2022 IOP Publishing Ltd.)

Due to the challenging communication and control systems, few underwater multi-robot coordination systems are currently developed. In nature, weakly electric fish can organize their collective activities using electrocommunication in turbid water. Inspired by this communication mechanism, we developed an artificial electrocommunication system for underwater robots in our previous work. In this study, we coordinate a group of underwater robots using this bio-inspired electrocommunication. We first design a time division multiple access (TDMA) network protocol for electrocommunication to avoid communication conflicts during multi-robot coordination. Then, we revise a distributed controller to coordinate a group of underwater robots. The distributed controller on each robot generates the required controls based on adjacent states obtained through electrocommunication. A central pattern generator (CPG) controller is designed to adjust the speed of individuals according to distributed control law. Simulations and experimental results show that a group of underwater robots is able to achieve coordination with the developed electrocommunication and control systems., (© 2022 IOP Publishing Ltd.)

This study describes the extrinsic connections of the dorsal telencephalon (pallium) of gymnotiform fish. We show that the afferents to the dorsolateral and dorsomedial pallial subdivisions of gymnotiform fish arise from the preglomerular complex. The preglomerular complex receives input from four clearly distinct regions: 1) descending input from the pallium itself (dorsomedial and dorsocentral subdivisions and nucleus taenia); 2) other diencephalic nuclei (centroposterior, glomerular, and anterior tuberal nuclei and nucleus of the posterior tuberculum); 3) mesencephalic sensory structures (optic tectum, dorsal and ventral torus semicircularis); and 4) basal forebrain, preoptic area, and hypothalamic nuclei. Previous studies have implicated the majority of the diencephalic and mesencephalic nuclei in electrosensory, visual, and acousticolateral functions. Here we discuss the implications of preglomerular/pallial electrosensory-associated afferents with respect to a major functional dichotomy of the electric sense. The results allow us to hypothesize that a functional distinction between electrocommunication vs. electrolocation is maintained within the input and output pathways of the gymnotiform pallium. Electrocommunication information is conveyed to the pallium through complex indirect pathways that originate in the nucleus electrosensorius, whereas electrolocation processing follows a conservative pathway inherent to all vertebrates, through the optic tectum. We hypothesize that cells responsive to communication signals do not converge onto the same targets in the preglomerular complex as cells responsive to moving objects. We also hypothesize that efferents from the dorsocentral (DC) telencephalon project to the dorsal torus semicircularis to regulate processing of electrocommunication signals, whereas DC efferents to the tectum modulate sensory control of movement. J. Comp. Neurol. 520:3338-3368, 2012. © 2012 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

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-ordersignalling 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. [ABSTRACT FROM AUTHOR]

Abstract: When animals are under stress, glucocorticoids commonly inhibit adult neurogenesis by acting through glucocorticoid receptors (GRs). However, in some cases, conditions that elevate glucocorticoids promote adult neurogenesis, and the role of glucocorticoid receptors in these circumstances is not well understood. We examined the involvement of GRs in social enhancement of brain cell addition and aggressive signaling in electric fish, Apteronotus leptorhynchus. In this species, long-term social interaction simultaneously elevates plasma cortisol, enhances brain cell addition and increases production of aggressive electrocommunication signals (“chirps”). We implanted isolated and paired fish with capsules containing nothing (controls) or the GR antagonist, RU486, recorded chirp production and locomotion for 7d, and measured the density of newborn cells in the periventricular zone. Compared to isolated controls, paired controls showed elevated chirping in two phases: much higher chirp rates in the first 5h and moderately higher nocturnal rates thereafter. Treating paired fish with RU486 reduced chirp rates in both phases to those of isolated fish, demonstrating that GR activation is crucial for socially induced chirping. Neither RU486 nor social interaction affected locomotion. RU486 treatment to paired fish had a partial effect on cell addition: paired RU486 fish had less cell addition than paired control fish but more than isolated fish. This suggests that cortisol activation of GRs contributes to social enhancement of cell addition but works in parallel with another GR-independent mechanism. RU486 also reduced cell addition in isolated fish, indicating that GRs participate in the regulation of cell addition even when cortisol levels are low. [Copyright &y& Elsevier]

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 2.5h 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 X 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. [ABSTRACT FROM AUTHOR]

Abstract: Peripheral sensory neurons respond to stimuli containing a wide range of spatio-temporal frequencies. We investigated electroreceptor neuron coding in the gymnotiform wave-type weakly electric fish Apteronotus leptorhynchus. Previous studies used low to mid temporal frequencies (<256 Hz) and showed that electroreceptor neuron responses to sensory stimuli could be almost exclusively accounted for by linear models, thereby implying a rate code. We instead used temporal frequencies up to 425 Hz, which is in the upper behaviorally relevant range for this species. We show that electroreceptors can: (A) respond up to the highest frequencies tested and (B) display strong nonlinearities in their responses to such stimuli. These nonlinearities were manifested by the fact that the responses to repeated presentations of the same stimulus were coherent at temporal frequencies outside of those contained in the stimulus waveform. Specifically, these consisted of low frequencies corresponding to the time varying contrast or envelope of the stimulus as well as higher harmonics of the frequencies contained in the stimulus. Heterogeneities in the afferent population influenced nonlinear coding as afferents with the lowest baseline firing rates tended to display the strongest nonlinear responses. To understand the link between afferent heterogeneity and nonlinear responsiveness, we used a phenomenological mathematical model of electrosensory afferents. Varying a single parameter in the model was sufficient to account for the variability seen in our experimental data and yielded a prediction: nonlinear responses to the envelope and at higher harmonics are both due to afferents with lower baseline firing rates displaying greater degrees of rectification in their responses. This prediction was verified experimentally as we found that the coherence between the half-wave rectified stimulus and the response resembled the coherence between the responses to repeated presentations of the stimulus in our dataset. This result shows that rectification cannot only give rise to responses to low frequency envelopes but also at frequencies that are higher than those contained in the stimulus. The latter result implies that information is contained in the fine temporal structure of electroreceptor afferent spike trains. Our results show that heterogeneities in peripheral neuronal populations can have dramatic consequences on the nature of the neural code. [Copyright &y& Elsevier]

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. [ABSTRACT FROM AUTHOR]

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 (700-1 100Hz) 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 (1 0-300 Hz) sine-wave stimuli; at 30 and 150 Hz, the threshold amplitude for response was 1 mVcm1. 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 gauderlo, but did not respond to the presence of the non-electric fish Carass!us auratus. Fish chirped to low-frequency (150Hz) signals that mimic those of S. macrurus and to EOD playbacks of B. gauderlo. The response to the B. gauderlo playback was reduced when the low-frequency component (<150Hz) was experimentally filtered out. Thus, A. leptorhynchus appears to chirp specifically to the electric signals of heterospecific electric fish, and the low-frequency components of heterospecific EODs significantly influence chirp rate. These results raise the possibility that chirps function to communicate to conspecifics about the presence of a heterospecific fish or to communicate directly to heterospecific fish. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

In adult males of the South African weakly electric bulldog fish, Marcusenius pongolensis, the duration of the electric organ discharge (EOD) increases with body size over lifetime. Although there is experimental support for intrasexual selection (male-male competition) having shaped the males' EOD pulse duration in evolution, nothing is known about intersexual selection, such as female choice. Playback of 25 natural male EODs of pulse duration varying from 320 µs (close to the average female value) to 716 µs, to eight female experimental subjects elicited approach, head butts and circling behaviour. The rate of head butts on the dipole electrode model increased significantly with stimulus pulse duration in seven out of eight experimental subjects. In ten experimental female subjects we contrasted the shortest playback pulse with simultaneous playback of one of four longer ones (424, 524, 628 and 716 µs). Pooled responses for all experimental subjects were stronger for the dipole playing back the longer pulse in a pulse pair. The difference in the number of head butts (Δhead butts) that were dealt the two dipoles per 60 s test session increased significantly with the difference in pulse duration (Δpulse duration). The increase followed a significant linear trend (P<0.0001). Similar results were obtained for Δassociation time, Δcircles with head butts, and Δcircles without head butts. These results suggest that a male's reproductive success is enhanced by longer, i.e. more attractive EODs, and that both intra- and intersexual selection must have played a significant role in shaping the EOD of male M. pongolensis. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: In electric fish, Apteronotus leptorhynchus, both long-term social interaction and cortisol treatment potentiates chirping, an electrocommunication behavior that functions in aggression. Chirping is controlled by the diencephalic prepacemaker nucleus (PPn-C) located just lateral to the ventricle. Cells born in adult proliferative zones such as the periventricular zone (PVZ) can migrate along radial glial fibers to other brain regions, including the PPn-C. We examined whether social interactions or cortisol treatment influenced cell addition and radial glia fiber formation by (1) pairing fish (4 or 7 days) or (2) implanting fish with cortisol (7 or 14 days). Adult fish were injected with bromodeoxyuridine 3 days before sacrifice to mark cells that were recently added. Other fish were sacrificed after 1 or 7 days of treatment to examine vimentin immunoreactivity (IR), a measure of radial glial fiber density. Paired fish had more cell addition than isolated fish at 7 days, coinciding temporally with the onset of socially induced increase in chirping behavior. Paired fish also had higher vimentin IR at 1 and 7 days. For both cell addition and vimentin IR, the effect was regionally specific, increasing in the PVZ adjacent to the PPn-C, but not in surrounding regions. Cortisol increased cell addition at 7 days, correlating with the onset of cortisol-induced changes in chirping, and in a regionally specific manner. Cortisol for 14 days increased cell addition, and cortisol for 7 days increased vimentin IR but in a regionally non-specific manner. The correlation between treatment-induced changes in chirping and regionally specific increases in cell addition, and radial glial fiber formation suggests a causal relationship between such behavioral and brain plasticity in adults, but this hypothesis will require further testing. [Copyright &y& Elsevier]

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 500–1000 ms. We hypothesize that this “echo response” serves a communicatory function. [ABSTRACT FROM AUTHOR]

Bulldog fish ( Marcusenius macrolepidotus) generate short (<1 ms) electric-organ discharges (EODs), separated by much longer and highly variable interdischarge intervals (IDIs). We observed overt behaviour and electrical activity during reproductive behaviour in a male and in a female bulldog, and identified IDI patterns with putative signal functions. In contrast to Pollimyrus adspersus and Pollimyrus isidori, in which an elaborate and extended courtship precedes spawning proper, our fish started spawning almost immediately when we allowed the female to enter the male’s territory. The male did not construct a nest, and neither parent provided parental care. The male showed very little aggression towards the intruding female. Fish spawned in bouts near the male’s hiding place, and eggs were scattered by the female’s vigorous tail flips as she left the spawning site, only to return shortly thereafter. During spawning bouts, both fish generated highly stereotyped IDI patterns: the male generated a series of IDIs gradually decreasing from about 200 ms to about 55 ms that was abruptly terminated by a long IDI. The female generated a series of relatively regular IDIs (about 54 ms) that was followed by a marked increase in IDI duration (the probable time of spawning). Finally, a sharp decrease in IDIs to about 20 ms accompanied the female’s sudden escape from the spawning site. In between spawning bouts, both fish generated series of very short IDIs (high discharge rate, HD) that alternated abruptly with very low-rate inter-HD activity (especially in the male). IDIs as short as 9 ms (male) or 11 ms (female) occurred during HD displays. No visible aggression, in fact very little overt behaviour, occurred during these HD displays in both fish. Agonistic interactions between male and female, outside a reproductive context, were similar to those previously described in male pairs, including overt behavioural patterns such as parallel swimming, antiparallel display and attack, as well as HD displays. When not interacting, fish did not generate HD displays. We suggest the HD display is a communication signal in both reproductive and agonistic contexts. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Constraints introduced by signal carriers, pre-receptor mechanisms and receptor transduction are fundamental for shaping the signals used by the brain to build up perceptual images. This review analyses some of these constraints in the electrosensory system of pulse Gymnotids. First, it describes the characteristics and differences of electrolocation and electrocommunication carriers. Second, it analyses the role of electrogenic and non-electrogenic tissues of the fish body in the generation and conditioning of these carriers. Two pre-receptor mechanisms are discussed: (a) the funneling of currents to the perioral region and (b) a Mexican-hat profile involved in edge detection. Finally, some characteristics of the electroreceptor structure and the sensory mosaic are examined. We conclude that there is an electrosensory fovea at the perioral region where a large density and variety of receptors is stimulated by self- and conspecific-generated currents funneled there by non electrogenic tissues. Differences in carrier waveform may be used to distinguish between reafferent and communication signals. [Copyright &y& Elsevier]

The variability of the bioelectric field of the electrosensitive catfish, Ictalurus nebulosus, was investigated by recording the potential variation occurring when the fish passed a stationary electrode, and by recording the field of a stationary fish by a 15-electrode array. A good first order approximation of the recorded field of a 20 cm long fish is a dipole dc source with the source and sink about 7 cm apart, carrying a current of about 1 μA in water with a specific resistivity of 3.3 kohm cm. At 5 cm distance from the dipole axis such a source generates an electric potential swing in the order of 50 μV in free space, head negative, tail positive. Superimposed on the basic component are respiration related fluctuations, and fluctuations related to the activity of the alimentary canal, gills, and skin. Novel stimuli, or stressors like investigators approaching the aquarium, evoke sudden increases in field strength, which last about 15 min. Demineralization of the aquarium water causes changes in field strength and reversal of field polarity. The administration of food causes field variations in the vicinity of the anal opening. The bioelectric field shows diurnal fluctuations of 100 μV. The peak is at about 04:00, the dip at 14:00. The fluctuations of the bioelectric field are sufficiently strong and specific to serve as electrical stimuli to other electrosensitive catfish. It is suggested that the field changes allow a simple form of electrocommunication, i.e. inform conspecifics about some physiological properties of the field source. The cellular mechanisms underlying the fluctuations of the bioelectric field are homeostatic processes mediated by ion pumps and ion channels. [Copyright &y& Elsevier]

Brown ghost knife fish, Apteronotus leptorhynchus, continually emit a weakly electric discharge that serves as a communication signal and is sensitive to sex steroids. Males modulate this signal during bouts of aggression by briefly (∼15 ms) increasing the discharge frequency in signals termed “chirps.” The present study examined the effects of short-term (1–7 days) and long-term (6–35 days) male–male interaction on the continuous electric organ discharge (EOD), chirping behavior, and plasma levels of cortisol and two androgens, 11-ketotestosterone (11KT) and testosterone. Males housed in isolation or in pairs were tested for short-term and long-term changes in their EOD frequency and chirping rate to standardized sinusoidal electrical stimuli. Within 1 week, chirp rate was significantly higher in paired fish than in isolated fish, but EOD frequency was equivalent in these two groups of fish. Plasma cortisol levels were significantly higher in paired fish than in isolated fish, but there was no difference between groups in plasma 11KT levels. Among paired fish, cortisol levels correlated positively with chirp rate. To determine whether elevated cortisol can cause changes in chirping behavior, isolated fish were implanted with cortisol-filled or empty Silastic tubes and tested for short-term and long-term changes in electrocommunication signals and steroid levels. After 2 weeks, fish that received cortisol implants showed higher chirp rates than blank-implanted fish; there were no difference between groups in EOD frequency. Cortisol implants significantly elevated plasma cortisol levels compared to blank implants but had no effect on plasma 11KT levels. These results suggest that male–male interaction increases chirp rate by elevating levels of plasma cortisol, which, in turn, acts to modify neural activity though an 11KT-independent mechanism. [Copyright &y& Elsevier]

Brown ghost knife fish, Apteronotus leptorhynchus, produce sexually dimorphic, androgen-sensitive electrocommunication signals termed chirps. The androgen regulation of chirping has been studied previously by administering exogenous androgens to females and measuring the chirping response to artificial electrical signals. The present study examined the production of chirps during dyadic interactions of fish and correlated chirp rate with endogenous levels of one particular androgen, 11-ketotestosterone (11KT). Eight males and four females were exposed to short-term (5-min) interactions in both same-sex and opposite-sex dyads. Twenty-four hours after all behavioral tests, fish were bled for determination of plasma 11KT levels. Males and females differed in both their production of chirps and their ability to elicit chirps from other fish: males chirped about 20–30 times more often than females and elicited 2–4 times as many chirps as females. Among males, chirp rate was correlated positively with plasma 11KT, electric organ discharge frequency, and body size. Combined with results from experimental manipulation of androgen levels, these results support the hypothesis that endogenous 11KT levels influence electrocommunication behavior during interactions between two male fish. [Copyright &y& Elsevier]

Apteronotus 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]