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1.
Weakly electric fish produce a communication signal, the electric organ discharge, that is species specific, and in many species, sexually dimorphic. Because the neural circuit that controls the electric organ discharge is relatively simple, it is an excellent model in which to study both the biophysical mechanisms underlying a rhythmic behavior and the neuroendocrine control of a sexually dimorphic behavior. By studying the effects of ion channel blockers on neurons in the medullary pacemaker nucleus, I pharmacologically characterized three ionic currents that influence the pacemaker rhythm, and thus electric organ discharge frequency, in the gymnotiform fish, Apteronotus leptorhynchus. These currents included a tetrodotoxin-sensitive sodium current; a potassium current that was sensitive to 4-aminopyridine; and a calcium current that was sensitive to nickel and cadmium, but resistant to specific blockers of L-, N-, P-, and Q-type calcium currents. The pharmacological profiles of the ionic currents in the pacemaker nucleus are similar to those of ionic currents involved in pacemaking in other neuronal oscillators. Because these ionic currents dramatically influence pacemaker firing frequency, which is directly related to electric organ discharge frequency, these ionic currents are likely targets of steroid hormone action in producing sexual dimorphisms in electric organ discharge frequency. Additional studies are needed to determine how these ionic currents interact to generate the electric organ discharge rhythm and to investigate the possibility that sexual dimorphism in the electric organ discharge results from the actions of gonadal steroids on these ionic currents. Accepted: 3 June 1999  相似文献   

2.
At the southern boundary of gymnotiform distribution in America. water temperature changes seasonally, and may be an environmental cue for the onset of breeding. In this study, we aim to describe the role of temperature upon electric organ discharge waveform in Gymnotus carapo, order Gymnotiformes, family Gymnotidae, and to analyze its interactions with the effects of steroid hormones. The effects of water temperature within its natural range were explored using different protocols. All fish tested had temperature-sensitive electric organ discharge waveforms: the amplitude of the last head-negative component (V4) decreased as temperature increased. Rate increases elicited by electrical stimulation had similar but smaller effect on waveform. Temperature sensitivity is a peripheral phenomenon that depends on the conductivity of the aquatic media. We found hormonal-dependent changes in the electric organ discharge waveform not previously described in this species. The amplitude and duration of V4 increased after testosterone administration. Both testosterone treatment and acclimation by sustained temperature at 27-28 degrees C (environmental simulation of breeding conditions) induced a decrease in temperature sensitivity. As in the related species Brachyhypopomus pinnicaudatus, our data strongly suggest interactions between temperature sensitivity of the electric organ discharge waveform and sexual maturity that might be crucial for reproduction.  相似文献   

3.
The mechanisms that trigger the onset of the breeding season depend on geographical latitude. At the edge of Gymnotiform distribution in America, variations in day length and water temperature are likely cues to initiate breeding. In this study we aim to clarify the role of temperature and the interaction between temperature and hormonal state upon electric organ discharge waveform. In breeding ponds, we measured naturally occurring changes of water temperature and of electric organ discharge waveform during two 48-h periods in a sample of identified mature males and females of Brachyhypopomus pinnicaudatus. Water temperature, day-night cycle, and sexual maturity each modified electric organ discharge waveform. Temperature sensitivity was also evaluated in the laboratory in adult sexually-differentiated individuals, adult non-differentiated fish, juveniles, and testosterone-treated fish. Our data strongly suggest an interaction between the effects of temperature and steroid hormones upon electric organ discharge waveform. High temperature (30 °C) induced a significant decay of head negative phase amplitude in temperature-sensitive fish. This sensitivity was observed in physiological conditions that coincide with low levels of steroid hormones: juveniles and adult fish kept in captivity at 20–21 °C. Conversely, temperature resistance was observed in mature fish in the breeding habitat and was induced by testosterone treatment and by captivity at 27–28 °C. Accepted: 23 May 1999  相似文献   

4.
Many species of electric fish show diurnal or socially elicited variation in electric organ discharge amplitude. In Sternopygus macrurus, activation of protein kinase A by 8-bromo-cAMP increases electrocyte sodium current magnitude. To determine whether the behavioral plasticity in electric organ discharge amplitude is controlled by electrocyte biophysical properties, we examined whether the effects of phosphorylation on ion currents in the electric organ translate directly into electric organ discharge changes. We injected the electric organ of restrained fish with 8-bromo-cAMP and monitored the electric organ discharge. The effect of protein kinase A activation on electrocyte action potentials was examined in isolated electric organ using two-electrode current clamp. Electric organ discharge and action potential amplitude and pulse duration increased in response to 8-bromo-cAMP. Pulse and action potential duration both increased by about 25%. However, the increase in electric organ discharge amplitude (approximately 400%) was several-fold greater than the action potential amplitude increase (approximately 40%). Resting membrane resistance decreased in electrocytes exposed to 8-bromo-cAMP. We propose that in the Thevenin equivalent circuit of the electric organ a moderate increase in action potential amplitude combined with a decrease in internal resistance produces a greater voltage drop across the external resistance (the water around the fish), accounting for the large increase in the externally recorded electric organ discharge.  相似文献   

5.
Summary This paper reports the finding of a sexual dimorphism in the electric organ discharge of the weakly electric fishPollimyms isidori. The difference in the head-positive components of the waveform (Fig. 1) was found in 95% of adults. Juvenile fish always possessed female type discharges. Fourier analysis (Fig. 3) revealed significant sex differences in the power spectra of the discharges, indicating that specific phase information is perhaps unnecessary for sexual identification. The results are discussed with reference to our current knowledge of coding of electrocommunication signals in the electrosensory system.Abbreviation EOD electric organ discharge  相似文献   

6.
Weakly electric fish in the genus Sternopygus emit a sinusoidal, individually distinct, and sexually dimorphic electric organ discharge (EOD) that is used in electrolocation and communication. Systemically applied androgens decrease EOD frequency, which is set by a medullary pacemaker nucleus, and increase pulse duration, which is determined by the cells of the electric organ (the electrocytes), in a coordinated fashion. One possibility is that androgens broaden the EOD pulse duration by acting on the pacemaker neurons, thereby effecting a change in pacemaker firing frequency, and that the change in EOD pulse duration is due to an activity-dependent process. To determine whether androgens can alter pulse duration despite a stable pacemaker nucleus firing frequency, we implanted small doses of dihydrotestosterone in the electric organ. We found that androgen implants increased EOD pulse duration, but did not influence EOD frequency. In addition, using immunocytochemistry, we found that electrocytes label positively with an androgen receptor antibody. While it is not known on which cells androgens act directly, together these experiments suggest that they likely act on the electrocytes to increase EOD pulse duration. Since pulse duration is determined by electrocyte action potential duration and ionic current kinetics, androgens may therefore play a causative role in influencing individual variation and sexual dimorphism in electrocyte electrical excitability, an important component of electrocommunicatory behavior.  相似文献   

7.
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, males 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 electric 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. Accepted: 18 February 1998  相似文献   

8.
Serotonin (5-HT) is an indirect modulator of the electric organ discharge (EOD) in the weakly electric gymnotiform fish, Brachyhypopomus pinnicaudatus. Injections of 5-HT enhance EOD waveform "masculinity", increasing both waveform amplitude and the duration of the second phase. This study investigated the pharmacological identity of 5-HT receptors that regulate the electric waveform and their effects on EOD amplitude and duration. We present evidence that two sets of serotonin receptors modulate the EOD in opposite directions. We found that the 5HT1AR agonist 8-OH-DPAT diminishes EOD duration and amplitude while the 5HT1AR antagonist WAY100635 increases these parameters. In contrast, the 5HT2R agonist alpha-Me-5-HT increases EOD amplitude but not duration, yet 5-HT-induced increases in EOD duration can be inhibited by blocking 5HT2A/2C-like receptors with ketanserin. These results show that 5-HT exerts bi-directional control of EOD modulations in B. pinnicaudatus via action at receptors similar to mammalian 5HT1A and 5HT2 receptors. The discordant amplitude and duration response suggests separate mechanisms for modulating these waveform parameters.  相似文献   

9.
The electric organ discharge of the gymnotiform fish Brachyhypopomus pinnicaudatus is a biphasic waveform. The female's electric organ discharge is nearly symmetric but males produce a longer second phase than first phase. In this study, infrared-sensitive video cameras monitored the position of unrestrained fish, facilitating precise measurement of electric organ discharge duration and amplitude every 2 h for 24 h. Males (n=27) increased electric organ discharge duration by 37 ± 12% and amplitude by 24 ± 9% at night and decreased it during the day. In contrast, females (n=8) exhibited only minor electric organ discharge variation over time. Most of a male's increase occurred rapidly within the first 2–3 h of darkness. Electric organ discharge values gradually diminished during the second half of the dark period and into the next morning. Modulation of the second phase of the biphasic electric organ discharge produced most of the duration change in males, but both phases changed amplitude by similar amounts. Turning the lights off at mid-day triggered an immediate increase in electric organ discharge, suggesting modification of existing ion channels in the electric organ, rather than altered genomic expression. Exaggeration of electric organ discharge sex differences implies a social function. Daily reduction of duration and amplitude may reduce predation risk or energy expenditure. Accepted: 12 September 1998  相似文献   

10.
Skates discharge an electrical current too weak to be used for predation or defense, and too infrequent and irregular to be used for electrolocation. Additionally, skates possess a specialized sensory system that can detect electrical stimuli at the same strength at which they discharge their organs. These two factors are suggestive of a communicative role for the electric organ in skates, a role that has been demonstrated in similarly weakly electric teleosts (e.g., mormyrids and gymnotiforms). There is evidence that the sexual and ontogenetic variations in the electric organ discharge (EOD) in these other weakly electric fishes are linked to morphological variations in electric organs and the electrogenerating cells of the organs, the electrocytes. Little work has been done to examine possible sexual and ontogenetic variations in skate EODs or variations in the electrocytes responsible for those discharges. Electric organs and electrocyte morphology of male and female, and mature and immature little skates, Leucoraja erinacea, are characterized here. Female electric organs were bigger than male electric organs. This is suggestive of a sexually dimorphic EOD waveform or amplitude, which might be used as a sex-specific identification signal during courtship. The shapes of electrocytes that make up the organ were found to be significantly different between mature and immature individuals and, in some cases, posterior membrane surface area of the electrocytes increased at the onset of maturity due to the formation of membrane surface invaginations and papillae. This is evidence that the EOD of skates may differ in its waveform or amplitude or frequency between mature and immature skates, and act as a signal for readiness to mate. This study supports a communicative role during courtship for the weak electric organs of little skates, but studies that characterize skate EOD dimorphisms are needed to corroborate this speculation before conclusions can be drawn about the role the electric organ plays in communication during courtship.  相似文献   

11.
Summary Hypopomus occidentalis is a weakly electric Gymnotiform fish with a pulse-type electric organ discharge (EOD).Hypopomus used in this study were taken from one of the northernmost boundaries of this species, the Atlantic drainage of Panama where the animals breed at the beginning of the dry season (December). In normal breeding populations,Hypopomus occidentalis exhibit a sexual dimorphism in EOD and morphology. Mature males are large with a broad tail and have an EOD characterized by a low peak power frequency. Females and immature males are smaller, having a slender tail and EODs with higher peak power frequencies (Fig. 1). This study describes differences in the EOD and electric organ morphology between breeding field populations of male and femaleHypopomus. Changes in physiology, morphology and EOD shape which may accompany this seasonal change were examined in steroid injected fish, using standard histological and physiological techniques.A group of females were injected with hormones (5-dihydrotestosterone (DHT), estrogen or saline) to assess changes in their morphology and EOD. Animals treated with DHT developed characteristics which mimicked the sexually dimorphic characteristics of a male, while the other groups did not (see Fig. 5). Tissue from the tails of breeding males and females, and females treated with DHT, were sampled to measure the size of the electrocytes in the tail. The broader tail of males and DHT-females is composed of large electrocytes, whereas the slender tail of normal females is composed of smaller electrocytes. Therefore, the increase in the tail width in the female DHT group is caused by an enlargement of the electrocytes in this area.Intracellular recordings from the electrocytes of saline and DHT injected females show a difference in the responses of the rostral faces of the electrocytes from the two groups, which reflect the differences in their EODs. Saline-treated animals had symmetrical EODs (the first and second phase of the EOD were equal in duration and amplitude), while the physiological responses from each face of the electrocytes yielded responses that were similarly equal in duration and amplitude. DHT-treated animals had asymmetrical EODs (the first phase of the EOD was similar to that of saline treated fish and larger in amplitude and shorter in duration than the second phase) and the physiological responses of the electrocytes reflected this asymmetry. The differential recordings across the caudal face were similar to those from saline treated fish, while the responses from the rostral face were longer in duration and smaller in amplitude.These data suggest that the effects of androgens underlie the changes in single electrocytes which produce the sexually dimorphic signals and morphology present in natural breeding populations ofHypopomus occidentalis.  相似文献   

12.
Sexually dimorphic signaling is widespread among animals and can act as an honest indicator of mate quality. Additionally, differences in signaling and morphology within a sex can be associated with different strategies for acquiring mates. Weakly electric fish communicate via self-generated electrical fields that transmit information about sex, reproductive state, and social status. The weakly electric knifefish Parapteronotus hasemani exhibits sexual dimorphism in body size as well as substantial within-male variation in body size and jaw length. We asked whether P. hasemani exhibits hormonally mediated sexual dimorphism in electrocommunication behavior. We also asked whether males with short versus long jaws differed significantly from each other in morphology, behavior, hormone levels, or reproductive maturity. Males produced longer chirps than females, but other signal parameters (electric organ discharge frequency; chirp rate and frequency modulation) were sexually monomorphic. Pharmacologically blocking androgen receptors in males reduced chirp duration, suggesting that this sexually dimorphic trait is regulated at least in part by the activational effects of androgens. Males sorted into two distinct morphological categories but did not differ in circulating 11-ketotestosterone or testosterone. Short-jawed males and long-jawed males also did not differ in any aspects of signaling. Thus, chirping and high levels of 11-ketotestosterone were reliably associated with reproductively active males but do not necessarily indicate male type or quality. This contrasts with other alternative male morph systems in which males that differ in morphology also differ in androgen profiles and signaling behavior.  相似文献   

13.
14.
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.  相似文献   

15.
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.  相似文献   

16.
The communication signals of electric fish can be dynamic, varying between the sexes on a circadian rhythm and in response to social and environmental cues. In the gymnotiform fish Brachyhypopomus gauderio waveform shape of the electric organ discharge (EOD) is regulated by steroid and peptide hormones. Furthermore, EOD amplitude and duration change on different timescales and in response to different social stimuli, suggesting that they are regulated by different mechanisms. Little is known about how androgen and peptide hormone systems interact to regulate signal waveform. We investigated the relationship between the androgens testosterone (T) and 11-ketotestosterone (11-KT), the melanocortin peptide hormone α-MSH, and their roles in regulating EOD waveform of male B. gauderio. Males were implanted with androgen (T, 11-KT, or blank), and injected with α-MSH before and at the peak of androgen effect. We compared the effects of androgen implants and social interactions by giving males a size-matched male stimulus with which they could interact electrically. Social stimuli and both androgens increased EOD duration, but only social stimuli and 11-KT elevated amplitude. However, no androgen enhanced EOD amplitude to the extent of a social stimulus, suggesting that a yet unidentified hormonal pathway regulates this signal parameter. Additionally, both androgens increased response of EOD duration to α-MSH, but only 11-KT increased response of EOD amplitude to α-MSH. Social stimuli had no effect on EOD response to α-MSH. The finding that EOD amplitude is preferentially regulated by 11-KT in B. gauderio may provide the basis for independent control of amplitude and duration.  相似文献   

17.
Weakly electric gymnotiform fish specialize in the regulation and modulation of the action potentials that make up their multi-purpose electric signals. To produce communication signals, gymnotiform fish modulate the waveforms of their electric organ discharges (EODs) over timescales spanning ten orders of magnitude within the animal’s life cycle: developmental, reproductive, circadian, and behavioral. Rapid changes lasting milliseconds to seconds are the result of direct neural control of action potential firing in the electric organ. Intermediate-term changes taking minutes to hours result from the action of melanocortin peptides, the pituitary hormones that induce skin darkening and cortisol release in many vertebrates. Long-term changes in the EOD waveform taking days to weeks result from the action of sex steroids on the electrocytes in the electric organ as well as changes in the neural control structures in the brain. These long-term changes in the electric organ seem to be associated with changes in the expression of voltage-gated ion channels in two gene families. Electric organs express multiple voltage-gated sodium channel genes, at least one of which seems to be regulated by androgens. Electric organs also express multiple subunits of the shaker (Kv1) family of voltage-gated potassium channels. Expression of the Kv1 subtype has been found to vary with the duration of the waveform in the electric signal. Our increasing understanding of the mechanisms underlying precise control of electric communication signals may yield significant insights into the diversity of natural mechanisms available for modifying the performance of ion channels in excitable membranes. These mechanisms may lead to better understanding of normal function in a wide range of physiological systems and future application in treatment of disease states involving pathology of excitable membranes.  相似文献   

18.
Weakly electric fish communicate with brief electrostatic field pulses called electric organ discharges (EODs). EOD waveforms are sexually dimorphic in most genera, a condition thought to result from mate choice acting to shape the electric signal's constituent action potentials. We have no direct behavioural evidence that sexual selection by either mate choice or intrasexual competition is responsible for sex differences in the EOD waveforms of electric fish. We explored sexual selection in electric fish by conducting two-choice unforced preference tests with live, unaltered gymnotiform electric fish,Brachyhypopomus pinnicaudatus , which are sexually dimorphic. In the initial test, gravid females selected males over females only when the males were larger than average. Gravid females in later tests preferred larger males to smaller males in a significant majority of those trials in which they showed a preference. In about one-third of those trials, females spawned with their preferred male, confirming their preference. We concluded that passage through the choice apparatus was related to mate choice. The signals of chosen males had larger EOD amplitudes and longer EOD durations. These findings show that femaleB. pinnicaudatus do have a preference for a certain male phenotype. The system requires additional study to dissociate correlated male phenotypic characters to identify which male traits the female prefers. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.   相似文献   

19.
In Sternopygus macrurus, electrocyte action potential duration determines the electric organ discharge pulse duration. Since the electric organ discharge is a sexually-dimorphic behavior under the control of steroid hormones, and because electrocyte action potential durations can range from 3–14 ms, the electrocytes provide a unique opportunity to study how sex steroids regulate membrane excitability. In this study, the voltage-sensitive ionic currents of electrocytes were identified under current- and voltage-clamp as a prelude to further studies on their regulation by sex steroid hormones.Bath application of TTX completely abolished the spike and eliminated an inward current under voltage clamp, indicating that the action potential is due primarily to a sodium current. Calcium-free saline had no effect on spike waveform or voltage-clamp currents, indicating that neither calcium nor calcium-dependent currents contribute to the action potential. Application of potassium channel blocking agents, such as tetraethylammonium and cesium ions, caused changes in the spike which, together with voltage-clamp results, indicate the presence of two potassium currents: an inward rectifier and a classical delayed rectifier. In addition, these cells have a large, presumably voltage-insensitive, chloride current. Differences in one or more of these currents could be responsible for the range of action potential durations found in these cells and for the steroid-mediated changes in spike duration.Abbreviations EOD electric organ discharge - VC voltage clamp - CC current clamp - AP action potential - VI/IV voltage-current/current-voltage  相似文献   

20.
Weakly electric fish such as Sternopygus macrurus utilize a unique signal production system, the electric organ (EO), to navigate within their environment and to communicate with conspecifics. The electric organ discharge (EOD) generated by the Sternopygus electric organ is quasi-sinusoidal and sexually dimorphic; sexually mature males produce long duration EOD pulses at low frequencies, whereas mature females produce short duration EOD pulses at high frequencies. EOD frequency is set by a medullary pacemaker nucleus, while EOD pulse duration is determined by the kinetics of Na+ and K+ currents in the electric organ. The inactivation of the Na+ current and the activation of the delayed rectifying K+ current of the electric organ covary with EOD frequency such that the kinetics of both currents are faster in fish with high (female) EOD frequency than those with low (male) EOD frequencies. Dihydrotestosterone (DHT) implants masculinize the EOD centrally by decreasing frequency at the pacemaker nucleus (PMN). DHT also acts at the electric organ, broadening the EO pulse, which is at least partly due to a slowing of the inactivation kinetics of the Na+ current. Here, we show that chronic DHT treatment also slows the activation and deactivation kinetics of the electric organ's delayed rectifying K+ current. Thus, androgens coregulate the time-varying kinetics of two distinct ion currents in the EO to shape a sexually dimorphic communication signal.  相似文献   

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