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PubMed1987

A microprocessor system for the digital synthesis of pulsed and continuous discharges of electric fish (or animal vocalizations).

Traditionally, the sensitivity of electric fish to electric stimuli has been investigated using a variety of electronically generated signal functions, for example sine, square, and sawtooth waves (both continuous and pulsed). None of these resemble the species- or sex-specific electric organ discharges (EODs) of fish. Therefore, EODs recorded on magnetic tape were used in open-loop playback experiments. However, as shown in the present paper, the waveforms of EODs reproduced from magnetic tape are distorted, and cannot be manipulated with speed and accuracy as to frequency or pulse repetition rate by feedback from the experiment. Good EOD imitations are generated by our microprocessor-based system for the digital synthesis of EODs which is controlled by a small 'personal computer'. Digitized EOD samples edited by computer are output by the fast digital-to-analogue converter of the microprocessor system. This system was devised for the synthesis of pulse-type (discontinuous) and wave type (continuous) EODs with continuous, on-line frequency and amplitude control. The rate of digital synthesis is 500 kHz in pulse-type EODs, and 250-500 kHz in wave type EODs at 0.06-0.24 Hz frequency resolution, depending on the repetition frequency of the synthesized wave EOD. The present paper describes the steps in EOD synthesis, and indicates some applications which benefit from the playback of high-fidelity imitations of natural EODs at amplitudes and frequencies selected by the experimenter, or automatically controlled by feedback from the experiment in closed-loop applications.

Analog-Digital Conversion↗
PubMedDate not supplied

Effect of drugs that alter alertness and emotionality on the novelty response of a weak electric fish, Gymnotus carapo.

Weak field electric fish respond to alerting stimuli with a transient increase in the frequency of electric organ discharge (novelty response). In an attempt to demonstrate the influence of different degrees of alertness and of emotionality on the novelty response of Gymnotus carapo, we studied the variations in the magnitude of this response induced by the application of an electric stimulus to the water of the experimental box using a pair of electrodes, before and after intramuscular injections of d-amphetamine (1-2 and 4 mg/kg), sodium pentobarbital (10-20 and 30 mg/kg), diazepam (1-2 and 4 mg/kg), beta-carboline (2 mg/kg), and saline. After d-amphetamine injection the animal presented increased somatic motility but no changes in electric organ baseline firing rate or in response to the alerting electric stimulus. Sodium pentobarbital induced a partial loss of posture and a reduction of fin and operculum movements, as well as a reduction of baseline firing rate and of the response to the alerting electric stimulus, with frequent interruptions in electric organ firing. Beta-carboline caused increased motility, but no changes in basal firing rate or in response to the alerting stimulus. Diazepan-injected fishes remained still throughout the experiment, and some of those threated with the higher dose (4 mg/kg) presented interruptions on electric organ discharges in response to stimulation but no change on baseline firing rate. The data suggest that a reduction of the degree of alertness by the barbiturate and a decrease in emotionality and/or stress by the benzodiazepine interfere with the novelty response. The possible site of action of the drugs is discussed.

Animals↗
PubMed2000

Auditory discrimination in a sound-producing electric fish (Pollimyrus): tone frequency and click-rate difference detection.

Pollimyrus adspersus is a fish that uses simple sounds for communication and has auditory specializations for sound-pressure detection. The sounds are species-specific, and the sounds of individuals are sufficiently stereotyped that they could mediate individual recognition. Behavioral measurements are presented indicating that Pollimyrus probably can make species and individual discriminations on the basis of acoustic cues. Interclick interval (ICI; 10-40 ms) and frequency (100-1400 Hz) discrimination was assessed using modulations of the fish's electric organ discharge rate in the presence of a target stimulus presented in alternation with an ongoing base stimulus. Tone frequency discrimination was best in the 200-600-Hz range, with the best threshold of 1.7% +/- 0.4% standard error at 500 Hz (or 8.5 Hz +/- 1.9 SE). The just noticeable differences (jnd's) were relatively constant from 100 to 500 Hz (mean 8.7 Hz), then increased at a rate of 13.3 Hz per 100 Hz. For click trains, jnd's increased linearly with ICI. The mean jnd's for 10- and 15-ms ICI were both 300 micros (SE= 0.8 ms at 10-ms ICI, SE= 0.11 ms at 15-ms ICI). The jnd at 20-ms ICI was only 1.1 ms +/- 0.25 SE.

Animals↗
PubMed2004

Amazonian ecology: tributaries enhance the diversity of electric fishes.

Neotropical rivers support a diverse array of endemic taxa, including electric fishes of the order Gymnotiformes. A comprehensive survey of the main channels of the Amazon River and its major tributaries (>2000-kilometer transect) yielded 43 electric fish species. Biogeographical analyses suggest that local mainstem electric fish diversity is enhanced by tributaries. Mainstem species richness tends to increase downstream of tributary confluences, and species composition is most similar between tributaries and adjacent downstream mainstem locations. These findings support a "nodal" or heterogeneous model of riverine community organization across a particularly extensive and diverse geographical region.

Animals↗
PubMed2006

Sodium channel genes and the evolution of diversity in communication signals of electric fishes: convergent molecular evolution.

We investigated whether the evolution of electric organs and electric signal diversity in two independently evolved lineages of electric fishes was accompanied by convergent changes on the molecular level. We found that a sodium channel gene (Na(v)1.4a) that is expressed in muscle in nonelectric fishes has lost its expression in muscle and is expressed instead in the evolutionarily novel electric organ in both lineages of electric fishes. This gene appears to be evolving under positive selection in both lineages, facilitated by its restricted expression in the electric organ. This view is reinforced by the lack of evidence for selection on this gene in one electric species in which expression of this gene is retained in muscle. Amino acid replacements occur convergently in domains that influence channel inactivation, a key trait for shaping electric communication signals. Some amino acid replacements occur at or adjacent to sites at which disease-causing mutations have been mapped in human sodium channel genes, emphasizing that these replacements occur in functionally important domains. Selection appears to have acted on the final step in channel inactivation, but complementarily on the inactivation "ball" in one lineage, and its receptor site in the other lineage. Thus, changes in the expression and sequence of the same gene are associated with the independent evolution of signal complexity.

Amino Acid Sequence↗
PubMed1996

Neuronal circuitry for comparison of timing in the electrosensory lateral line lobe of the African wave-type electric fish Gymnarchus niloticus.

An African wave-type electric fish, Gymnarchus, compares timing on the order of microseconds of sensory feedback from from its high-frequency (approximately 400 Hz) electric organ discharges (EODs) received at different parts of its body surfaces. This capability is essential for and demonstrated by the jamming avoidance response (JAR). The organization of the timing comparison mechanisms was identified in the electrosensory lateral line lobe (ELL) in the hindbrain by field potential, extra- and intracellular recordings, and intracellular labeling with biotinylated agents. Timing of phase of the EOD feedback is carried by action potentials of S-type primary afferent fibers that project to the inner cellular layer (ICL) of the medial zone of the ELL and to the giant neurons in the ELL. The giant neurons bilaterally project to the ICL, where neurons sensitive to phase differences between different parts of the body occur. Although sensitive to dynamic phase changes of several microseconds, these differential-phase-sensitive neurons showed adaptation to steady-state changes of phase difference over a wide range (greater than +/- 100 microseconds) and continued to respond to small modulations after the mean difference was shifted. Gymnarchus and an independently evolved South American electric fish, Eigenmannia, exhibit nearly identical JARs and share a rather complex but identical set of computational algorithms for JAR. This study showed that one of the computational steps, the timing comparison between body surfaces, occurs in the hindbrain in Gymnarchus, in contrast to the midbrain in Eigenmannia. Thus, similar systems with a similar overall function may have evolved differently in different genera by assigning a subfunction to different substructures within the brain.

Afferent Pathways↗
PubMed1993

Weakly electric fish as model systems for studying long-term steroid action on neural circuits.

Weakly electric fish generate electric organ discharges (EODs) that are species-specific and often sexually-dimorphic. The waveform or frequency of an EOD can be altered by treating a fish with sex steroid hormones. The EOD is controlled by a few discrete nuclei in the brainstem and spinal cord and a muscle-derived electric organ. The organizational simplicity and steroid-sensitivity of the electromotor system make it a premier system for investigating how sex steroids modulate behavior, neural circuitry, and ion channels. In addition, the diversity of EOD patterns in the many species of electric fish provide a wealth of material in which to examine the evolution of sexual dimorphisms in the nervous system.

Animals↗
PubMed2000

Molecular systematics of the African electric fishes (Mormyroidea: teleostei) and a model for the evolution of their electric organs.

We present a new molecular phylogeny for 41 species of African mormyroid electric fishes derived from the 12S, 16S and cytochrome b genes and the nuclear RAG2 gene. From this, we reconstruct the evolution of the complex electric organs of these fishes. Phylogenetic results are generally concordant with earlier preliminary molecular studies of a smaller group of species and with the osteology-based classification of Taverne, which divides the group into the Gymnarchidae and the Mormyridae, with the latter including the subfamilies Petrocephalinae (Petrocephalus) and Mormyrinae (all remaining taxa). However, we find that several genera previously recognized by Taverne are non-monophyletic. Within the Mormyrinae, the genus Myomyrus is the sister group to all the remaining taxa. Other well-supported clades within this group are recovered. A reconstruction of electrocyte evolution on the basis of our best-supported topology suggests that electrocytes with penetrating stalks evolved once early in the history of the mormyrids followed by multiple paedomorphic reversals to electrocytes with non-penetrating stalks.

Animals↗
PubMed2004

Sodium-dependent plateau potentials in electrocytes of the electric fish Gymnotus carapo.

The weakly electric fish Gymnotus carapo emits a triphasic electric organ discharge generated by muscle-derived electrocytes, which is modified by environmental and physiological factors. Two electrode current clamp recordings in an in vitro preparation showed that Gymnotus electrocytes fired repetitively and responded with plateau potentials when depolarized. This electrophysiological behavior has never been observed in electrocytes from related species. Two types of plateaus with different thresholds and amplitudes were evoked by depolarization when Na(+)-dependent currents were isolated in a K(+)- and Ca(2+)-free solution containing TEA and 4-AP. Two electrode voltage clamp recordings revealed a classical fast activating-inactivating Na+ current and two persistent Na(+)-dependent currents with voltage-dependencies consistent with the action potential (AP) and the two plateaus observed under current clamp, respectively. The three currents, the APs and the plateaus were reduced by TTX, and were absent in Na(+)-free solution. The different Na(+)-dependent currents in Gymnotus electrocytes may be targets for the modifications of the electric organ discharge mediated by environmental and physiological factors.

Action Potentials↗
PubMed2003

Serotonin modulates the electric waveform of the gymnotiform electric fish Brachyhypopomus pinnicaudatus.

The gymnotiform electric fish Brachyhypopomus pinnicaudatus communicates with a sexually dimorphic electric waveform, the electric organ discharge (EOD). Males display pronounced circadian rhythms in the amplitude and duration of their EODs. Changes in the social environment influence the magnitudes of these circadian rhythms and also produce more transient responses in the EOD waveforms. Here we show that injections of serotonin produce quick, transient, dose-dependent enhancements of the male EOD characters similar to those induced by encounters with another male. The response to serotonin administered peripherally begins 5-10 min post injection and lasts approximately 3 h. The magnitude of the response to serotonin is tightly associated with the magnitude of the day-to-night swing of the circadian rhythm prior to injection. Taken together these findings suggest that the male's social environment influences his response to serotonin by altering the function of some part of the downstream chain between the serotonin receptors and the ion channels involved in control of the EOD waveform. Although chronic activation of serotonin circuitry is widely known to elicit subordinate behavior, we find that 5-HT initially increases a dominance signal in these fish. These findings are consistent with the emerging view that serotonin facilitates different adaptive responses to acute and chronic social challenge and stress.

Animal Communication↗
PubMed2002

Drawing a spark from darkness: John Walsh and electric fish.

John Walsh's research on electric fish, carried out between 1772 and 1775, proved fundamental for demonstrating that electricity might be involved in animal physiology, and, moreover, in favouring a period of great progress in both the physiology and physics of electrical phenomena. However, Walsh is hardly known to modern neuroscientists and is largely neglected by science historians also. One of the reasons for this neglect is that he never published his 'crucial experiment', that is the production of a spark from a discharge of the electric eel.

Animals↗
PubMed2002

Drawing a spark from darkness: John Walsh and electric fish.

John Walsh's research on electric fish, carried out between 1772 and 1775, proved fundamental for demonstrating that electricity might be involved in animal physiology, and, moreover, in favouring a period of great progress in both the physiology and physics of electrical phenomena. However, Walsh is hardly known to modern neuroscientists and is largely neglected by science historians also. One of the reasons for this neglect is that he never published his 'crucial experiment', that is the production of a spark from a discharge of the electric eel.

Animals↗
PubMed1986

Jamming avoidance in electric fish and frogs: strategies of signal oscillator timing.

Repetitive spontaneous calling in frogs and electric signalling in pulse-type electric fishes are driven by neural pacemakers. Minimization of signal overlap between neighbors is achieved in frogs and certain electric fishes by mutual entrainment, requiring cycle-by-cycle adjustment of pacemaker interval. Other fishes, which require a regular electric organ discharge, are more constrained in their ability to avoid jamming. Entrainment may also serve to reciprocally place signals in a temporal 'blindspot' of the receiver, the period of elevated sensory threshold following signal production, to reduce aggressive interactions.

Animal Communication↗
PubMed1976

Electric signals and schooling behavior in a weakly electric fish, Marcusenius cyprinoides L. (Mormyriformes).

Field recordings of electric organ discharges and catches of Marcusenius cyprinoides showed that these electric fish form groups and move about in schools. The role electric organ discharges in group cohesion was investigated by comparing interactions in groups of intact and operated, electrically silent fish. The absence of electric organ dischares reduced locomotor activity and resulted in the disappearance of two behaviors: parallel lineup and single file swimming. Electric signals are considered part of a schooling mechanism that aids the fish in maintaining group cohesion in their turbid enviornment and during migration at night.

Animals↗
PubMedDate not supplied

Emergence of temporal-pattern sensitive neurons in the midbrain of weakly electric fish Gymnarchus niloticus.

Sensitivity of neurons in the torus semicircularis of a weakly electric fish, Gymnarchus niloticus, to two stimulus parameters that are critical for its behavior the jamming avoidance response was examined. The first parameter is the sign of frequency difference between discharge frequencies of fish's own electric organ and that of a neighbor's. The second parameter is the spatial orientation of neighbor's electric field. Whereas neuronal ambiguity of frequency coding for different orientations of neighbor's electric field is predicted, unambiguous JAR occurs at the behavioral level. Most neurons in the torus semicircularis showed sensitivity to the sign of frequency difference. Although a small number of neurons showed preference to a consistent sign of the frequency difference, the coding of the sign of frequency differences was found to be ambiguous with a highly variable pattern of responses for different orientations in most of neurons.

Action Potentials↗
PubMed1982

Structural similarities between acetylcholine receptors from fish electric organs and mammalian muscle.

Acetylcholine receptors from fish electric organ tissue and mammalian muscle were compared by peptide mapping. The alpha subunits from receptors of Torpedo californica and Electrophorus electricus electric organ tissue were digested with V8 protease and the resulting fragments separated on polyacrylamide gels and stained for protein or for carbohydrate. 125I-Labeled alpha subunits of acetylcholine receptors from Electrophorus electric organ tissue and bovine muscle were digested with V8 protease, and the resulting fragments were also separated on polyacrylamide gels. Intact receptors from both the fish electric organs and mammalian muscle were labeled with [4-(N-maleimido) benzyl]tri[3H]-methylammonium iodide which binds specifically to the acetylcholine binding site on alpha subunits, and the isolated alpha subunits were subjected to the same peptide mapping procedure. The fragment patterns produced were stained for protein and fluorographed to identify active site containing polypeptides. None of these peptide mapping approaches revealed extensive homologies between alpha subunits. Intact and V8 proteolyzed sodium dodecyl sulfate denatured receptors from Torpedo and Electrophorus electric organs and bovine muscle were electrophoretically transferred to diazophenyl thioether paper and probed with antisera to Torpedo receptor subunits and two monoclonal antibodies. Unique fragment patterns were obtained with each antisubunit serum. A fragment of the same size was derived from the beta subunit of each acetylcholine receptor and was shown to specifically bind the same monoclonal antibody in each case. These results indicate that only in the beta subunits from all of the species examined is a large sequence nearly identical. However, it is likely that corresponding receptor subunits from receptors of all of these species have extensive structural homologies.

Animals↗
PubMed2006

Structure and sexual dimorphism of the electrocommunication signals of the weakly electric fish, Adontosternarchus devenanzii.

Electrocommunication 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.

Animal Communication↗
PubMed1990

Functional analysis of sexual dimorphism in an electric fish, Hypopomus pinnicaudatus, order Gymnotiformes.

Hypopomus pinnicaudatus, an electric fish, has a marked sexual dimorphism in its tail filament. Sexually mature males have long, 'feathered' tails as compared with females. The sexual dimorphism emerges when a fish reaches about 110 mm total length. Mature males have larger electrocytes which are more widely spaced and more numerous than those in mature females. The biphasic electric organ discharge (EOD) is longer in males than in females. The peak-to-peak amplitude of the male's EOD is weaker than a female's of the same total length. The weaker discharge is unexpected given the increase in size and number of electrocytes. It is suggested that the reduction in EOD amplitude is a consequence of the increase in EOD duration among males. Further, female choice probably played a role in the evolution of long duration EODs among males, and males may have secondarily grown long tails to compensate for the loss in active space that would otherwise accompany a weaker EOD.

Agonistic Behavior↗