Social regulation of electric signal plasticity in male Brachyhypopomus gauderio.

In animal communication, the social context that elicits particular dynamic changes in the signal can provide indirect clues to signal function. Female presence should increase the expression of male signal traits relevant for mate-choice, while male presence should promote the enhancement of traits involved in male-male competition. The electric fish Brachyhypopomus gauderio produces a biphasic electric pulse for electrolocation and communication. Pulse amplitude predicts the signaler's body size while pulse duration predicts circulating androgen levels. Males enhance pulse amplitude and duration when the numbers of males and females increase simultaneously. Here we tested the relative effects of female presence and male presence on male signal enhancement, and whether the size of the male competitor affected this enhancement. We found that male presence drives the enhancement of both pulse amplitude and second phase duration, independently of the size of the male competitor. Female presence induces the enhancement of pulse duration, but not pulse amplitude. These data suggest that males probably attend to information about a competitor's body size coded by pulse amplitude and attend to aggressiveness coded by a competitor's pulse duration, both potential predictors of fight outcome. Females may be primarily concerned about information on reproductive condition coded by pulse duration.

Cellular mechanisms of developmental and sex differences in the rapid hormonal modulation of a social communication signal.

Some gymnotiform electric fish species rapidly modify their electric signal waveforms by altering the action potential (AP) waveforms of their electrocytes, the excitable cells that produce the electric organ discharge (EOD). The fish Brachyhypopomus gauderio alters EOD amplitude and pulse duration as a social signal in accordance with the prevailing social conditions, under the dual regulation of melanocortins and androgens. We show here that B. gauderio uses two distinct cellular mechanisms to change signal amplitude, and its use of these two mechanisms varies with age and sex of the signaler. EOD amplitude and waveform are regulated by the coordinated timing and shaping of two APs generated from two opposing excitable membranes in each electrocyte. The two membranes fire in sequence within 100 µs of each other with the second AP being broader than the first. We have shown previously that mature males increase EOD amplitude and duration when melanocortin peptide hormones act directly on electrocytes to selectively broaden the second AP and increase the delay between the two APs by approximately 25 µs. Here we show that females selectively broaden only the second AP as males do, but increase amplitude of both APs with no change in delay between them, a previously unreported second mechanism of EOD amplitude change in B. gauderio. Juvenile fish broaden both APs and increase the delay between the APs. Cellular mechanisms of EOD plasticity are therefore shaped during development, presumably by sex steroids, becoming sexually dimorphic at maturity.

Neuromodulation of the agonistic behavior in two species of weakly electric fish that display different types of aggression.

Agonistic behavior has shaped sociality across evolution. Though extremely diverse in types of displays and timing, agonistic encounters always follow the same conserved phases (evaluation, contest and post-resolution) and depend on homologous neural circuits modulated by the same neuroendocrine mediators across vertebrates. Among neuromodulators, serotonin (5-HT) is the main inhibitor of aggression, and arginine vasotocin (AVT) underlies sexual, individual and social context differences in behavior across vertebrate taxa. We aim to demonstrate that a distinct spatio-temporal pattern of activation of the social behavior network characterizes each type of aggression by exploring its modulation by both the 5-HT and AVT systems. We analyze the neuromodulation of aggression between the intermale reproduction-related aggression displayed by the gregarious Brachyhypopomus gauderio and the non-breeding intrasexual and intersexual territorial aggression displayed by the solitary Gymnotus omarorum. Differences in the telencephalic activity of 5-HT between species were paralleled by a differential serotonergic modulation through 1A receptors that inhibited aggression in the territorial aggression of G. omarorum but not in the reproduction-related aggression of B. gauderio. AVT injection increased the motivation towards aggression in the territorial aggression of G. omarorum but not in the reproduction-related aggression of B. gauderio, whereas the electric submission and dominance observed in G. omarorum and B. gauderio, respectively, were both AVT-dependent in a distinctive way. The advantages of our model species allowed us to identify precise target areas and mechanisms of the neuromodulation of two types of aggression that may represent more general and conserved strategies of the control of social behavior among vertebrates.

Behavioral ecology, endocrinology and signal reliability of electric communication.

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

Food restriction promotes signaling effort in response to social challenge in a short-lived electric fish.

Vertebrates exposed to stressful conditions release glucocorticoids to sustain energy expenditure. In most species elevated glucocorticoids inhibit reproduction. However individuals with limited remaining reproductive opportunities cannot afford to forgo reproduction and should resist glucocorticoid-mediated inhibition of reproductive behavior. The electric fish Brachyhypopomus gauderio has a single breeding season in its lifetime, thus we expect males to resist glucocorticoid-mediated inhibition of their sexual advertisement signals. We studied stress resistance in male B. gauderio (i) by examining the effect of exogenous cortisol administration on the signal waveform and (ii) by investigating the effect of food limitation on androgen and cortisol levels, the amplitude of the electric signal waveform, the responsiveness of the electric signal waveform to social challenge, and the amount of feeding activity. Exogenous cortisol administration did reduce signal amplitude and pulse duration, but endogenous cortisol levels did not rise with food limitation or social challenge. Despite food limitation, males responded to social challenges by further increasing androgen levels and enhancing the amplitude and duration of their electric signal waveforms. Food-restricted males increased androgen levels and signal pulse duration more than males fed ad libitum. Socially challenged fish increased food consumption, probably to compensate for their elevated energy expenditure. Previous studies showed that socially challenged males of this species simultaneously elevate testosterone and cortisol in proportion to signal amplitude. Thus, B. gauderio appears to protect its cortisol-sensitive electric advertisement signal by increasing food intake, limiting cortisol release, and offsetting signal reduction from cortisol with signal-enhancing androgens.

Circadian and social cues regulate ion channel trafficking.

Electric fish generate and sense electric fields for navigation and communication. These signals can be energetically costly to produce and can attract electroreceptive predators. To minimize costs, some nocturnally active electric fish rapidly boost the power of their signals only at times of high social activity, either as night approaches or in response to social encounters. Here we show that the gymnotiform electric fish Sternopygus macrurus rapidly boosts signal amplitude by 40% at night and during social encounters. S. macrurus increases signal magnitude through the rapid and selective trafficking of voltage-gated sodium channels into the excitable membranes of its electrogenic cells, a process under the control of pituitary peptide hormones and intracellular second-messenger pathways. S. macrurus thus maintains a circadian rhythm in signal amplitude and adapts within minutes to environmental events by increasing signal amplitude through the rapid trafficking of ion channels, a process that directly modifies an ongoing behavior in real time.

Mosquitoes Exposed to Sunlight in Florida Are Capable of Making Vitamin D3.

BACKGROUND/AIM: This study aimed to determine whether mosquitoes exposed to natural sunlight are able to produce vitamin D3 Materials and Methods: Mosquito eggs obtained from adult Aedes aegypti mosquitoes were raised to adulthood. Approximately 1,200 male and female mosquitoes were divided among six cages. Four cages were placed on the roof on sunny days, 10-Sep-2021 and 12-Oct-2021. Two of the cages were exposed to sunlight from 8:30-18:00 and brought inside during rainstorms in the midday that lasted approximately 2 hours, while the other two cages were placed in deep shade and served as a control. After the exposure, the mosquitoes were collected and frozen at -80°C. The mosquitoes were thawed and then extracted in methanol followed by high-performance liquid chromatography analysis. RESULTS: The mosquitoes that were exposed to sunlight produced vitamin D3 whereas the mosquitoes kept in deep shade and not exposed to any sunlight did not produce significant vitamin D3 Conclusion: Mosquitoes exposed to sunlight have the ability to produce vitamin D3 Therefore, mosquitoes exposed to sunlight provide an excellent dietary source of vitamin D3 for insect eating vertebrates including birds and especially bats that avoid sun exposure and require vitamin D for their skeletal health.

Testosterone and 11-ketotestosterone have different regulatory effects on electric communication signals of male Brachyhypopomus gauderio.

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.

Tight hormonal phenotypic integration ensures honesty of the electric signal of male and female Brachyhypopomus gauderio.

Hormones mediate sexually selected traits including advertisement signals. Hormonal co-regulation links the signal to other hormonally-mediated traits such that the tighter the integration, the more reliable the signal is as a predictor of those other traits. Androgen administration increases the duration of the communication signal pulse in both sexes of the electric fish Brachyhypopomus gauderio. To determine whether the duration of the signal pulse could function as an honest indicator of androgen levels and other androgen-mediated traits, we measured the variation in sex steroids, signal pulse duration, and sexual development throughout the breeding season of B. gauderio in marshes in Uruguay. Although the sexes had different hormone titres and signal characteristics, in both sexes circulating levels of the androgens testosterone (T) and 11-ketotestosterone (11-KT) were strongly related to signal pulse duration. Consequently, signal pulse duration can serve as an honest indicator of circulating androgens in males and females alike. Additionally, through phenotypic integration, signal pulse duration also predicts other sexual traits directly related to androgen production: gonad size in males and estradiol (E2) levels in females. Our findings show that tight hormonal phenotypic integration between advertisement signal and other sex steroid-mediated traits renders the advertisement signal an honest indicator of a suite of reproductive traits.

Energetic cost of communication.

Communication signals may be energetically expensive or inexpensive to produce, depending on the function of the signal and the competitive nature of the communication system. Males of sexually selected species may produce high-energy advertisement signals, both to enhance detectability and to signal their size and body condition. Accordingly, the proportion of the energy budget allocated to signal production ranges from almost nothing for many signals to somewhere in excess of 50% for acoustic signals in short-lived sexually selected species. Recent data from gymnotiform electric fish reveal mechanisms that regulate energy allocated to sexual advertisement signals through dynamical remodeling of the excitable membranes in the electric organ. Further, males of the short-lived sexually selected species, Brachyhypopomus gauderio, trade off among different metabolic compartments, allocating energy to signal production while reducing energy used in other metabolic functions. Female B. gauderio, by contrast, do not trade off energy between signaling and other functions. To fuel energetically expensive signal production, we expect a continuum of strategies to be adopted by animals of different life history strategies. Future studies should explore the relation between life history and energy allocation trade-offs.

Social competition affects electric signal plasticity and steroid levels in the gymnotiform fish Brachyhypopomus gauderio.

Sexually-selected communication signals can be used by competing males to settle contests without incurring the costs of fighting. Steroid regulation of these signals can render them as reliable indicators of a male's physiological state. We investigated how plasticity in electrocommunication signals is driven by social competition for mates, mediated by steroid hormones, and subject to the effects of past social experience. We measured the electric waveform's amplitude and duration and steroid hormone levels of male gymnotiform electric fish (Brachyhypopomus gauderio) following week-long periods of social isolation, and low or high social competition. To quantify the effect of social history on the modulation of the electric signal, six groups of six males experienced all three social conditions but in different order. We found that males differentially modulate their electric signals depending on the order they experienced these conditions. Thus, past social interactions affect both present and future social electric signals. Cortisol levels and the amplitude of the electric signal appeared to track the intensity of competition, while androgen levels and the duration of the electric signal only responded to the presence (low and high competition) or absence (isolation) of a social environment (low and high androgens respectively). In addition, cortisol levels were related to the body size of the males at high social competition. Taken together, these findings suggest that the capacity of males to modulate their signals in response to social competition is regulated by steroids.

Androgens enhance plasticity of an electric communication signal in female knifefish, Brachyhypopomus pinnicaudatus.

Sex steroids were initially defined by their actions shaping sexually dimorphic behavioral patterns. More recently scientists have begun exploring the role of steroids in determining sex differences in behavioral plasticity. We investigated the role of androgens in potentiating circadian, pharmacological, and socially-induced plasticity in the amplitude and duration of electric organ discharges (EODs) of female gymnotiform fish. We first challenged female fish with injections of serotonin (5-HT) and adrenocorticotropic hormone (ACTH), and with social encounters with female and male conspecifics to characterize females' pre-implant responses to each treatment. Each individual was then implanted with a pellet containing dihydrotestosterone (DHT) concentrations of 0.0, 0.03, 0.1, 0.3, or 1.0 mg 10 g(-1) body weight. We then repeated all challenges and compared each female's pre- and post-implant responses. The highest implant dose enhanced EOD duration modulations in response to all challenge types, responses to male challenge were also greater at the second highest dose, and responses to ACTH challenge were enhanced in females receiving all but the smallest dose (and blank) implants. Alternatively, amplitude modulations were enhanced only during female challenges and only when females received the highest DHT dose. Our results highlight the differential regulation of EOD duration and amplitude, and suggest that DHT enhanced the intrinsic plasticity of the electrogenic cells that produce the EOD rather than modifying behavioral phenotypes. The relative failure of DHT to enhance EOD amplitude plasticity also implies that factors other than androgens are involved in regulating/promoting male-typical EOD circadian rhythms and waveform modulations displayed in social contexts.

Melanocortins regulate the electric waveforms of gymnotiform electric fish.

The hypothalamic-pituitary-adrenal/interrenal axis couples serotonergic activity in the brain to the peripheral regulators of energy balance and response to stress. The regulation of peripheral systems occurs largely through the release of peptide hormones, especially the melanocortins (adrenocorticotropic hormone [ACTH] and alpha melanocyte stimulating hormone [alpha-MSH]), and beta-endorphin. Once in circulation, these peptides regulate a wide range of processes; alpha-MSH in particular regulates behaviors and physiologies with sexual and social functions. We investigated the role of the HPI and melanocortin peptides in regulation of electric social signals in the gymnotiform electric fish, Brachyhypopomus pinnicaudatus. We found that corticotropin releasing factor, thyrotropin-releasing hormone, and alpha-MSH, three peptide hormones of the HPI/HPA, increased electric signal waveform amplitude and duration when injected into free-swimming fish. A fourth peptide, a synthetic cyclic-alpha-MSH analog attenuated the normal circadian and socially-induced EOD enhancements in vivo. When applied to the electrogenic cells (electrocytes) in vitro, only alpha-MSH increased the amplitude and duration of the electrocyte discharge similar to the waveform enhancements seen in vivo. The cyclic-alpha-MSH analog had no effect on its own, but blocked or attenuated alpha-MSH-induced enhancements in the single-cell discharge parameters, demonstrating that this compound functions as a silent antagonist at the electrocyte. Overall, these results strongly suggest that the HPI regulates the EOD communication signal, and demonstrate that circulating melanocortin peptides enhance the electrocyte discharge waveform.

Signal Cloaking by Electric Fish.

Electric fish produce weak electric fields to image their world in darkness and to communicate with potential mates and rivals. Eavesdropping by electroreceptive predators exerts selective pressure on electric fish to shift their signals into less-detectable high-frequency spectral ranges. Hypopomid electric fish evolved a signal-cloaking strategy that reduces their detectability by predators in the lab (and thus presumably their risk of predation in the field). These fish produce broad-frequency electric fields close to the body, but the heterogeneous local fields merge over space to cancel the low-frequency spectrum at a distance. Mature males dynamically regulate this cloaking mechanism to enhance or suppress low-frequency energy. The mechanism underlying electric-field cloaking involves electrogenic cells that produce two independent action potentials. In a unique twist, these cells orient sodium and potassium currents in the same direction, potentially boosting their capabilities for current generation. Exploration of such evolutionary inventions could aid the design of biogenerators to power implantable medical devices, an ambition that would benefit from the complete genome sequence of a gymnotiform fish.

Managing Aedes aegypti populations in the first Zika transmission zones in the continental United States.

The African Zika virus swept across the Pacific, reaching the New World in 2014. In July, 2016, Miami-Dade County, Florida became the locus of the first mosquito-borne Zika transmission zones in the continental United States. Control efforts were guided by the Centers for Disease Control and Prevention, including aerial and truck sprays of adulticides and larvicides. To improve our understanding of how best to fight Zika transmission in an urban environment in the developed world, trap counts of adult Aedes (Stegomyia) aegypti (L.) mosquitoes from the treatment zones were analyzed to determine efficacy of the different insecticide treatments. Analysis revealed that application of four different ester pyrethroid and one non-ester pyrethroid had no statistically significant effect on mosquito counts. Aerial application of naled, a potent organophosphate adulticide, produced significant but short-lived drops in Ae. aegypti counts in the first two applications in the first active transmission zone (Wynwood), then lost some efficacy with subsequent application. In the other active transmission zone (Miami Beach), naled produced no measurable effect in the first three applications, and only a small, transient, and marginally significant reduction in the fourth application. Repeated application of the larvicidal bacterium Bti was accompanied by steady declines of Ae. aegypti populations in both sites. Zika transmission ceased in the first transmission zone, but expanded in the second transmission zone during this period. Specific recommendations are proposed for future treatments of urban mosquitoes.

Circadian rhythms in electric waveform structure and rate in the electric fish Brachyhypopomus pinnicaudatus.

Weakly electric fish have long been known to express day-night oscillations in their discharge rates, and in the amplitude and duration of individual electric organ discharges (EODs). Because these oscillations are altered by social environment and neuroendocrine interactions, electric fish are excellent organisms for exploring the social and neuroendocrine regulation of circadian rhythm expression. Previous studies asserting that these oscillations are circadian rhythms have been criticized for failing to control temperature and randomize feeding regimes, or for running the fish under constant conditions for just 2-3 days. Here we show that the day-night oscillations in the EODs of the neotropical gymnotiform fish Brachyhypopomus pinnicaudatus free-run for over a week under constant photic and thermal conditions, and randomized food provisioning. Sex differences were apparent in strength and magnitude of the circadian oscillations; male oscillations were stronger and larger. All three parameters retain a common oscillation period while differing in the persistence of oscillation strength and magnitude, a difference consistent with proposals by others that declines of behavioral circadian rhythms may result from breakdowns downstream of the central oscillator.

Adrenocorticotropic hormone enhances the masculinity of an electric communication signal by modulating the waveform and timing of action potentials within individual cells.

We report here that melanocortin peptides appear to serve as the mechanism by which weakly electric fish couple socially regulated and stress-regulated brain pathways to unique changes in the intrinsic excitability and action potential waveform of excitable membranes in peripheral cells involved in communication. Gymnotiform electric fish modulate their electric organ discharges (EODs) by reshaping the electric discharges of excitable cells in the periphery. These fish show circadian enhancement of the EOD waveform. They also enhance their EOD waveforms within minutes in response to stressors and changes in the social environment, thus altering the communication value of the signal. Changes in the EOD waveform that occur within minutes result from changes in the discharges of individual electrocytes (microEODs) mediated by the cAMP/protein kinase A (PKA) pathway acting on ion channel kinetics. What activates the cAMP/PKA pathway in electrocytes has not been identified. In vivo injections of the melanocortin peptide adrenocorticotropic hormone (ACTH) increase the amplitude and duration of the electric signal waveform of the gymnotiform Brachyhypopomus pinnicaudatus over the course of 1 h. Applied to single electrocytes in vitro, ACTH increases microEOD amplitude and duration within minutes by differentially modulating the action potentials of the two excitable membranes of the electrocyte and changing the timing of these two spikes. Serotonin modulates the EOD in vivo but has no effect on the microEOD in vitro. The cAMP analog 8-bromo-cAMP mimicked the effects of ACTH, whereas inhibition of PKA by protein kinase A inhibitor 14-22 amide blocked the modulatory effects of ACTH, confirming the role of the cAMP/PKA pathway in microEOD modulation by ACTH.

Fitness correlates of song repertoire size in free-living song sparrows (Melospiza melodia).

Models of sexual selection propose that exaggerated secondary sexual ornaments indicate a male's own fitness and the fitness of his offspring. These hypotheses have rarely been thoroughly tested in free-living individuals because overall fitness, as opposed to fitness components, is difficult to measure. We used 20 years of data from song sparrows (Melospiza melodia) inhabiting Mandarte Island, British Columbia, Canada, to test whether a male's song repertoire size, a secondary sexual trait, predicted overall measures of male or offspring fitness. Males with larger song repertoires contributed more independent and recruited offspring, and independent and recruited grandoffspring, to Mandarte's population. This was because these males lived longer and reared a greater proportion of hatched chicks to independence from parental care, not because females mated to males with larger repertoires laid or hatched more eggs. Furthermore, independent offspring of males with larger repertoires were more likely to recruit and then to leave more grandoffspring than were offspring of males with small repertoires. Although we cannot distinguish whether observed fitness variation reflected genetic or environmental effects on males or their offspring, these data suggest that female song sparrows would gain immediate and intergenerational fitness benefits by pairing with males with large song repertoires.

The evolutionary origins of electric signal complexity.

This study explores the evolutionary origins of waveform complexity in electric organ discharges (EODs) of weakly electric fish. I attempt to answer the basic question of what selective forces led to the transition from the simplest signal to the second simplest signal in the gymnotiform electric fishes. The simplest electric signal is a monophasic pulse and the second simplest is a biphasic pulse. I consider five adaptive hypotheses for the evolutionary transition from a monophasic to a biphasic EOD: (i) electrolocation, (ii) sexual selection, (iii) species isolation, (iv) territory defense, (v) crypsis from electroreceptive predators. Evaluating these hypotheses with data drawn largely from the literature, I find best support for predation. Predation is typically viewed as a restraining force on evolution of communication signals, but among the electric fishes, predation appears to have served as a creative catalyst. In suppressing spectral energy in the sensitivity range of predators (a spectral simplification), the EOD waveforms have become more complex in their time domain structure. Complexity in the time domain is readily discernable by the high frequency electroreceptor systems of gymnotiform and mormyrid electric fish. The addition of phases to the EOD can cloak the EOD from predators, but also provides a substrate for subsequent modification by sexual selection. But, while juveniles and females remain protected from predators, breeding males modify their EODs in ways that enhance their conspicuousness to predators.

Signal modulation as a mechanism for handicap disposal.

Signal honesty may be compromised when heightened competition provides incentive for signal exaggeration. Some degree of honesty might be maintained by intrinsic handicap costs on signalling or through imposition of extrinsic costs, such as social punishment of low quality cheaters. Thus, theory predicts a delicate balance between signal enhancement and signal reliability that varies with degree of social competition, handicap cost, and social cost. We investigated whether male sexual signals of the electric fish Brachyhypopomus gauderio would become less reliable predictors of body length when competition provides incentives for males to boost electric signal amplitude. As expected, social competition under natural field conditions and in controlled lab experiments drove males to enhance their signals. However, signal enhancement improved the reliability of the information conveyed by the signal, as revealed in the tightening of the relationship between signal amplitude and body length. Signal augmentation in male B. gauderio was independent of body length, and thus appeared not to be curtailed through punishment of low quality (small) individuals. Rather, all individuals boosted their signals under high competition, but those whose signals were farthest from the predicted value under low competition boosted signal amplitude the most. By elimination, intrinsic handicap cost of signal production, rather than extrinsic social cost, appears to be the basis for the unexpected reinforcement of electric signal honesty under social competition. Signal modulation may provide its greatest advantage to the signaller as a mechanism for handicap disposal under low competition rather than as a mechanism for exaggeration of quality under high competition.