Biology

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.

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.

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 5HT 1A R agonist 8-OH-DPAT diminishes EOD duration and amplitude while the 5HT 1A R antagonist WAY100635 increases these parameters. In contrast, the 5HT 2 R agonist α-Me-5-HT increases EOD amplitude but not duration, yet 5-HT-induced increases in EOD duration can be inhibited by blocking 5HT 2A/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 5HT 1A and 5HT 2 receptors. The discordant amplitude and duration response suggests separate mechanisms for modulating these waveform parameters.

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, dosedependent 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.

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 sociallyinduced 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.

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 voltagegated 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. Citation: Markham MR, McAnelly ML, Stoddard PK, Zakon HH (2009) Circadian and Social Cues Regulate Ion Channel Trafficking. PLoS Biol 7(9): e1000203.

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.

This article describes the construction and operation of a simple interface to control external devices via the IBM PC/XT/AT’s parallel printer port. The interface is relatively inexpensive and easy to build. It is potentially useful for controlling behavioral experiments.

This study investigated the role of maternal attention in evoking and maintaining jealous behavior in four 11-14 month-old twin infant pairs. One infant from each twin pair completed a BA design with shaping while the other twin concurrently completed a yoked ABA design with shaping. During A phases, maternal attention was made contingent upon the absence of jealous behavior and, during B phases, maternal attention was contingent upon the occurrence of jealous behavior. Results indicate that differential maternal attention to a twin sibling can function as a cue that evokes jealous behavior by the observer twin, and that maternal attention contingent upon the jealous behavior of the observer twin can function as a positive reinforcer to systematically increase the rate of that jealous behavior. Demonstrations of how environmental processes operate to establish and maintain jealous behavior can instruct parents of twins/siblings on how to manage the occurrence of jealousy and (sibling) rivalry.

Two experiments were conducted to investigate whether infant vocalizations can be conditioned by using contingent maternal-imitation and contingent motheresespeech to reinforce the infant's vocalizations. The effects of these two treatments were compared to each other, to baseline, and to a noncontingent/yoked treatment. In the first experiment, seventeen 3-to-6 month old infants participated in a repeated-measures single treatment design (Br A Bz): three 2-min conditions were implemented sequentially in one session: contingent maternal-imitation (Br), noncontingent/yoked (A), and contingent maternal-imitation (Bz). In the second experiment, twenty 3-to-5-month old infants participated in a repeated-measures multiple treatment design (ArBiAzBz). In the first session, four 2-n-tin conditions were implemented sequentially: baseline (Al), continpent maternal-imitation (Br), noncontinpent-voked (AZ), and contingent maternal-imitation (BJ. In the second session, contingent motherese-speech was used instead of contingent maternal imitation. The order of sessions was counterbalanced with half of the infants experiencing the imitation treatment in the first session.

have argued that alcoholics show a differential tolerance for punishment relative to nonalcoholics. However, the studies purporting to support this assertion failed to control for the effects of social factors on alcoholics' tolerance of punishment. The present study compared alcoholics' and nonalcoholics' tolerance for punishment under two conditions. In the private condition, subjects were led to believe that no one would know how they responded to a task involving punishment. In the public condition, it was made clear that the subjects' responses would be known to the experimenter. In the public condition, alcoholic subjects made more punished responses than nonalcoholic subjects. In the private condition, neither group responded in the presence of punishment.

Three experiments were conducted to investigate stimulus relations that might emerge when college students are taught relations between compound sample stimuli and unitary comparison stimuli using match-to-sample procedures. In Experiment 1, subjects were taught nine AB-C stimulus relations, then tested for the emergence of 18 AC-B and BC-A relations. All subjects showed the emergence of all tested relations. Twelve subjects participated in Experiment 2. Six subjects were taught nine AB-C relations and were then tested for symmetrical (C-AB) relations. Six subjects were taught nine AB-C and three C-D relations and were then tested for nine AB-D (transitive) relations. Five of 6 subjects demonstrated the emergence of symmetrical relations, and 6 subjects showed the emergence of transitivity. In Experiment 3, 5 college students were taught nine AB-C and three C-D relations and were then tested for nine equivalence (D-AB) relations and 18 AD-B and BD-A relations. Three subjects demonstrated all tested relations. One subject demonstrated the AD-B and BD-A relations but not the D-AB relations. One subject did not respond systematically during testing. The results of these experiments extend stimulus equivalence research to more complex cases.

Two studies investigated the transfer of respondent elicitation through equivalence classes. In Experiment 1, match-to-sample procedures were used to teach 8 subjects two four-member equivalence classes. One member of one class was then paired with electric shock, and one member of the other class was presented without shock. All remaining stimuli were then presented. Using skin conductance as the measure of conditioning, transfer of conditioning was demonstrated in 6 of the 8 subjects. In Experiment 2, similar procedures were used to replicate the results of Experiment 1 and investigate the transfer of extinction. Following equivalence training and conditioning to all members of one class, one member was then presented in extinction. When the remaining stimuli from this class were then presented, they failed to elicit skin conductance. In the final phase of the experiment, the stimulus that was previously presented in extinction was reconditioned. Test trials with other members of the class revealed that they regained elicitation function. These results demonstrate that both respondent elicitation and extinction can transfer through stimulus classes. The clinical and applied significance of the results is discussed.

This experiment investigated whether directly trained covarying functions are necessary for stimulus class formation and transfer of function in humans. Initial class training was designed to establish two respondent-based stimulus classes by pairing two visual stimuli with shock and two other visual stimuli with no shock. Next, two operant discrimination functions were trained to one stimulus of each putative class. The no-shock group received the same training and testing in all phases, except no stimuli were ever paired with shock. The data indicated that skin conductance response conditioning did not occur for the shock groups or for the no-shock group. Tests showed transfer of the established discriminative functions, however, only for the shock groups, indicating the formation of two stimulus classes only for those participants who received respondent class training. The results suggest that transfer of function does not depend on first covarying the stimulus class functions.

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 (μEODs) 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 μEOD 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 μEOD 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 μEOD modulation by ACTH.

This study investigated the efficacy of Group Cognitive Behavioral Therapy (GCBT) in the treatment of a heterogeneous set of anxiety disorders in children and adolescents using a partially nonconcurrent multiple-baseline across groups design with 12 clinically referred youth between 6 and 16 years of age who met DSM-1V criteria for an anxiety disorder. Targeted diagnoses included specific phobia, separation anxiety disorder, social phobia, generalized anxiety disorder, and obsessive-compulsive disorder, with 3 of the children also presenting with school refusal behavior. Duration of baseline for each of the 3 groups varied and ran for 1,2, or 3 weeks. Dependent measures included diagnostic status, daily child and parent ratings of child anxiety severity, and child-and parent-completed questionnaires. Results indicated that GCBT was generally efficacious in reducing anxious symptoms in youth treated in diagnostically heterogeneous groups, and that gains were generally maintained at 6 and 12 month follow-ups. Findings are discussed in terms of their theoretical and practical implications for the efficient treatment of children and adolescents with anxiety disorders.

We describe an operant conditioning apparatus that uses computerized touch-screen technology and is designed for the versatile and highly controlled testing of rats in a potentially wide variety of behavioral paradigms. Although computer-controlled touch-screen systems have been developed for use with pigeons, monkeys, and humans, analogous technologies and methods have not yet been developed for rats. The development of a touch-screen system for rats could enhance the efficiency of behavioral research with rats, and may offer a unique tool for studying animal learning. In the first test of the utility of the apparatus, 3 Sprague-Dawley rats learned to activate the touch screen only after the touch-screen panel was made slightly movable. These animals then learned to discriminate visual stimuli presented on the computer monitor, but only after the food magazine and pellet dispenser were moved to the rear of the chamber opposite the stimulus display and response window. In a test of the utility of the modified apparatus, 6 Long-Evans rats learned to activate the touch screen and learned one of three different simple discriminations using computer-generated, visually presented stimuli. A basic method for training rats to activate the computer touch screen and for visual discrimination training is described. Results show that rats learned to activate the touch screen and discriminate visual stimuli presented on a computer monitor. Potential applications and advantages of the touch-screen-equipped rat operant conditioning chamber are discussed.