Androgen receptors rapidly modulate non-breeding aggression in male and female weakly electric fish (Gymnotus omarorum).

The South American weakly electric fish, Gymnotus omarorum, displays territorial aggression year-round in both sexes. To examine the role of rapid androgen modulation in non-breeding aggression, we administered acetate cyproterone (CPA), a potent inhibitor of androgen receptors, to both male and females, just before staged agonistic interactions. Wild-caught fish were injected with CPA and, 30 min later, paired in intrasexual dyads. We then recorded the agonistic behavior which encompasses both locomotor displays and emission of social electric signals. We found that CPA had no discernible impact on the levels of aggression or the motivation to engage in aggressive behavior for either sex. However, CPA specifically decreased the expression of social electric signals in both males and female dyads. The effect was status-dependent as it only affected subordinate electrocommunication behavior, the emission of brief interruptions in their electric signaling ("offs"). This study is the first demonstration of a direct and rapid androgen effect mediated via androgen receptors on non-breeding aggression. Elucidating the mechanisms involved in non-breeding aggression in this teleost model allows us to better understand potentially conserved or convergent neuroendocrine mechanisms underlying aggression in vertebrates.

Estrogenic influences on agonistic behavior in teleost fishes.

Teleost fishes show an extraordinary diversity of sexual patterns, social structures, and sociosexual behaviors. Sex steroid hormones are key modulators of social behaviors in teleosts as in other vertebrates and act on sex steroid receptor-containing brain nuclei that form the evolutionarily conserved vertebrate social behavior network (SBN). Fishes also display important differences relative to tetrapod vertebrates that make them particularly well-suited to study the physiological mechanisms modulating social behavior. Specifically, fishes exhibit high levels of brain aromatization and have what has been proposed to be a lifelong, steroid hormone dependent plasticity in the neural substrates mediating sociosexual behavior. In this review, we examine how estrogenic signaling modulates sociosexual behaviors in teleosts with a particular focus on agonistic behavior. Estrogens have been shown to mediate agonistic behaviors in a broad range of fishes, from sexually monomorphic gonochoristic species to highly dimorphic sex changers with alternate reproductive phenotypes. These similarities across such diverse taxa contribute to a growing body of evidence that estrogens play a crucial role in the modulation of aggression in vertebrates. As analytical techniques and genomic tools rapidly advance, methods such as LC-MS/MS, snRNAseq, and CRISPR-based mutagenesis show great promise to further elucidate the mechanistic basis of estrogenic effects on social behavior in the diverse teleost lineage.

Agonistic interactions and social behaviors in the Saharan Dorcas gazelle (Gazella dorcas neglecta): Using social network analysis to evaluate relationships and social structure in captive male groups.

Social interactions, including agonistic behavior, are very important for the management and welfare of individuals forming groups in captivity. One of the main concerns for the stability and durability of adult male groups is a noticeable level of intraspecific aggression. This study comprises a Social Network Analysis to illustrate social structure in different groups of captive Saharan Dorcas gazelles (Gazella dorcas neglecta). Our main objectives were to examine the relationship between agonistic, affiliative, and association networks and their reciprocity, assessing also whether the agonistic networks can conform to a linear dominance hierarchy. For these purposes, we recorded the behavior of 23 adult males organized in five herds, three composed only of Saharan Dorcas gazelle males and two mixed herds in which there were also Mhorr gazelle males (Nanger dama mohor). Observations were carried out during 295 h through scan sampling. We found no correlation between the affiliative and association networks in any group, although there was a significant correlation between the agonistic and association networks in mixed-species groups which was not present in single-species groups. Overall, there was no consistent reciprocity in either affiliative nor agonistic networks and none of the agonistic networks showed a linear structure. These results indicate that affiliative behavior in Saharan Dorcas gazelles offers distinctive and valuable information about the bonds between individuals, however, their dominance structure is far more complex than previously thought. As information provided by affiliative and proximity behaviors is different in this species, we suggest considering affiliative interactions to stablish affinity between individuals. Evaluating different social behaviors and not only agonistic interactions in later studies, is also recommended to develop a more accurately daily management in zoos that guarantee group stability and individuals' welfare, which will improve the conservation of captive populations.

Fighting cichlids: An integrated multimodal analysis to understand female and male aggression in Cichlasoma dimerus.

Aggression has been historically linked to males and androgen levels and, even if females from different species also display aggressive behavior, female aggression is still widely understudied. The aim of the present work is to disentangle how sex differences in social plasticity can be explained by sex steroid hormone levels, gonadal state and/or morphometric characteristics. In this context, we performed intrasexual dyadic encounters to identify social plasticity after acquiring a winner or loser status in males and females of Cichlasoma dimerus. This integral analysis suggests that the reproductive and hormonal variables analyzed explain the behavioral variation among winner and loser males and females, and that there are significant differences between sexes and contest outcome when individual morphometric variables are excluded from the analysis. Interestingly, there are no sex differences in aggressive and submissive behaviors, and clustering into winners and losers is mainly explained by specific behavioral displays, such as bites, chases, approaches, passive copings, and escapes. Correlation heatmaps show a positive correlation between estradiol with aggression and a negative correlation with submission, suggesting estrogens may have a dual role regulating agonistic behavior. Finally, these results suggest that size difference can help to understand aggression in females but not in males, and that assessment of the opponent's body size is important to understand aggression also before the initiation of the contest in both sexes. Overall, this study constitutes an integral approach adding insights into the importance of reproductive and hormonal variables to understand social plasticity in males and females.

Social factors drive vocal exchanges in red-capped mangabeys (Cercocebus torquatus).

Of all the calls made by non-human primates, the function of short-distance contact calls has largely remained to be determined. These calls are the most frequent in the repertoire and are most often exchanged between individuals in a non-random way. To our knowledge, no study has ever examined how vocal exchanges are structured in red-capped mangabeys (Cercocebus torquatus), a semi-terrestrial monkey living in the African forest with a complex semi-tolerant/semi-despotic social system. Our goal was to assess the organization of contact call exchanges in this species and their relationship with individual and social factors such as age, affinity, and hierarchy. Therefore, we observed several captive groups of red-capped mangabeys and collected data on vocal behavior, as well as grooming, agonistic behavior, and spatial proximity. We defined a vocal exchange in this species as a series of contact calls made by two or more individuals within a maximum inter-caller interval of 2 s. At the individual level, the higher the individual's hierarchical rank, the less they initiated exchanges. Furthermore, the most socially integrated individuals had a longer average response time than the less integrated ones. At the dyadic level, preferred exchange partners were individuals often observed near one other or individuals most distant in age. Also, the further apart two individuals were in the dominance hierarchy, the shorter the response time. Our results support both the social bonding hypothesis and a modulating key role of the dominance hierarchy on the social use of contact calls, which is in line with the social style of this species.

Iridescent scales signal male fighting ability to access females in the jewelled splitfin Xenotoca variata.

Jewelled splitfin males (Xenotoca variata) possess multi-colour iridescent scales (speckles) on the flanks. This study tested the hypothesis that the number of speckles could be a good proxy for predicting fighting ability in contests for access to females. The experiments consisted in observing and recording males' agonistic behaviour, courtship displays and mating attempts in mixed-sex groups. The data were analysed and presented based on a Bayesian approach, which revealed that the density (cm-2 ) of speckles was positively correlated with the increase observed in the proportional frequency of attacks. Similarly, the density of speckles was positively associated with the increase observed in the frequency of courtship behaviour and with the frequency of mating attempts. Male-male aggressions drastically diminished (by eightfold) when females were removed from the observation tanks. These results indicate that the number of speckles is a consistent predictor of successful access to females and therefore, speckled males are likely to be able to gain more mating opportunities. This finding highlights the ubiquitous role iridescent colours play in visual signalling.

Impact of temperature-induced sex reversal on behavior and sound production in Nile tilapia (Oreochromis niloticus).

In some fish species, sex is determined by the combination of genetic and environmental factors. In most species concerned, extreme temperatures during the sensitive period of sex differentiation drives masculinization, independently of the female sex chromosomes. In Nile tilapia (XY male heterogamety), XX juveniles exposed to high temperatures (>32 °C) can masculinize and become phenotypical males (neomales). Whether these neomales exhibit a different behavior than XY males remains however unclear. Sex reversal being naturally relevant, we investigated the agonistic behavior of neomales during dyadic fights and the preference of gravid females for one of the two male types. We quantified the behavior, size of the nest, hormone circulating levels (testosterone, 11-ketotestosterone and cortisol) and sound production of the two male types in both contexts. Independently of the individual they face, neomales seem to display more aggressive behaviors than XY males but often fail to become dominant. Agonistic interactions were mainly silent, suggesting that sounds are unnecessary for the establishment of social hierarchy. Although males and neomales produce different kinds of sounds when facing a gravid female, the female does not exhibit a preference. Overall, no differences were observed for hormone circulating concentrations between the two male types. We suggest that the sex chromosomes and/or the sex reversal procedure may have differently shaped the brain of neomales, resulting in differences in the expression of behavior.

Environmental Enrichment Differentially Activates Neural Circuits in FVB/N Mice, Inducing Social Interaction in Females but Agonistic Behavior in Males.

Environmental enrichment induces behavioral and structural modifications in rodents and influences the capability of mice to cope with stress. However, little is understood about hippocampal neurogenesis and the appearance of social/agonistic (aggressive) behavior upon activation of different neuronal circuits in FVB/N mice. Thus, in this study we hypothesized that environmental enrichment differentially regulates neurogenesis, neural circuit activation and social/agonistic behavior in male and female FVB/N mice. We explored the (1) neurogenic process as an indicative of neuroplasticity, (2) neuronal activation in the limbic system, and (3) social behavior using the resident-intruder test. On postnatal day 23 (PD23), mice were assigned to one of two groups: Standard Housing or Environmental Enrichment. At PD53, rodents underwent the resident-intruder test to evaluate social behaviors. Results revealed that environmental enrichment increased neurogenesis and social interaction in females. In males, environmental enrichment increased neurogenesis and agonistic behavior. Enriched male mice expressed higher levels of agonistic-related behavior than female mice housed under the same conditions. Neural circuit analysis showed lower activation in the amygdala of enriched males and higher activation in enriched females than their respective controls. Enriched females also showed higher activation in the frontal cortex without differences in male groups. Moreover, the insular cortex was less activated in females than in males. Thus, our results indicate that environmental enrichment has different effects on neuroplasticity and social/agonistic behavior in FVB/N mice, suggesting the relevance of sexual dimorphism in response to environmental stimuli.

Impact of intraspecific variation in teleost fishes: aggression, dominance status and stress physiology.

Dominance-based social hierarchies are common among teleost fishes. The rank of an animal greatly affects its behaviour, physiology and development. The outcome of fights for social dominance is affected by heritable factors and previous social experience. Divergent stress-coping styles have been demonstrated in a large number of teleosts, and fish displaying a proactive coping style have an advantage in fights for social dominance. Coping style has heritable components, but it appears to be largely determined by environmental factors, especially social experience. Agonistic behaviour is controlled by the brain's social decision-making network, and its monoaminergic systems play important roles in modifying the activity of this neuronal network. In this Review, we discuss the development of dominance hierarchies, how social rank is signalled through visual and chemical cues, and the neurobiological mechanisms controlling or correlating with agonistic behaviour. We also consider the effects of social interactions on the welfare of fish reared in captivity.

Agonistic behaviour of wild eels and depressed survival and growth of farmed eels in mixed rearing experiments.

To contribute to improving eel stocking procedures, the survival, growth and behaviour of farmed Japanese eels reared together with wild individuals were evaluated to learn about possible encounters of farmed- and stocked eels with wild eels. Two experiments were conducted to evaluate the (a) effect of the presence of wild eels on survival and growth of farmed eels in experimental ponds and (b) behavioural differences between wild and farmed eels placed in a small tank. In the survival and growth experiment, significantly lower survival and growth rates of farmed eels reared with wild eels were observed compared with farmed eels reared without wild eels (control). In the behavioural observations, the authors observed significantly higher occupation rates of a refuge and more frequent biting for wild eels, indicating a dominance of wild eels over farmed eels. Because wild and farmed eels used in the current experiments belong to the same genetic population, these differences resulted within one generation through the aquaculture rearing process.

Enhancement of synaptic responses in ascending interneurones following acquisition of social dominance in crayfish.

When crayfish have attained dominant status after agonistic bouts, their avoidance reaction to mechanical stimulation of the tailfan changes from a dart to a turn response. Ascending interneurones originating in the terminal ganglion receive sensory inputs from the tailfan and they affect spike activity of both uropod and abdominal postural motor neurones, which coordinates the uropod and abdominal postural movements. Despite the varying output effects of ascending interneurones, the synaptic responses of all interneurones to sensory stimulation were enhanced when they acquired a dominant state. The number of spikes increased as did a sustained membrane depolarizations. Regardless of social status, the output effects on the uropod motor neurones of all interneurones except VE-1 remained unchanged. VE-1 mainly inhibited the uropod opener motor neurones in naive animals, but tended to excite them in dominant animals. Synaptic enhancement of the sensory response of ascending interneurones was also observed in naive animals treated with bath-applied serotonin. However, subordinate animals or naive animals treated with octopamine had no noticeable effect on the synaptic response of their ascending interneurones to sensory stimulation. Thus, enhancement of the synaptic response is a specific neural event that occurs when crayfish attain social dominance and it is mediated by serotonin.

Association of androgens and estrogens with agonistic behavior in the annual fish Austrolebias reicherti.

Agonistic behavior governs the settlement of conflicts among conspecifics for limiting resources. Sex steroids play a critical role in the regulation of agonistic behavior which in turn may produce modulations in hormone titres. In this study we analyzed the association of androgens and estrogens with agonistic behavior in the annual fish Austrolebias reicherti. This native species inhabits temporary ponds that dry out completely during summer, having one of the shortest lifespans among vertebrates. They are highly sexually dimorphic and have a single breeding season during which they reproduce continuously. Here we measured plasma levels of 11-ketotestosterone (11KT) and 17ß-estradiol (E2) in adult males after the resolution of a social conflict and assessed the role of the aromatase conversion of testosterone (T) to E2 in male aggression. Winners had higher levels of 11KT than losers yet; winner 11KT levels did not differ from those of males not exposed to a social challenge. E2 levels did not show differences among winners, losers or control males. However, fights under the aromatase inhibitor Fadrozole were overall less aggressive than control fights. Our results suggest an androgen response to losing a conflict and that the conversion of T to E2 is involved in the regulation of aggressive behavior. Annual fish extreme life history may give new insights on hormone-behavior interactions.

How to alleviate the agony of providing negative feedback: Emotion regulation strategies affect hormonal stress responses to a managerial task.

Providing negative feedback can be demanding, as it typically requires dealing with multiple negative emotions. The first aim of this study was to transfer this work-related task to a new laboratory protocol and to investigate short-term hormonal changes among feedback providers. The second aim was to test if such hormonal stress responses can be attenuated through a priori instructions on how to regulate emotions. Each of 150 participants (51% women) provided eight saliva samples before, during, and after anticipating and conducting a negative feedback conversation with a professional actor who displayed negative emotional reactions. Participants were divided into four conditions regarding the way they were instructed to regulate their emotions: expressive suppression (keeping a neutral expression); cognitive reappraisal (staying task-oriented and emotionally distanced); affect utilization (moving towards and using emotions); or control condition. By means of three-phase spline growth models, latent growth factors during baseline, stress response, and recovery were specified. Providing negative feedback was followed by significant temporary testosterone decreases as well as cortisol increases. Testosterone (but not cortisol) responses were attenuated when feedback providers had been instructed to either follow a cognitive reappraisal or affect utilization strategy. This study provides evidence that a typical managerial task, that is, having to provide negative feedback, is a testosterone- and cortisol-relevant experience. Down-regulation of an individual's emotional involvement through reappraisal, as well as the newly introduced technique of moving towards and making use of the interaction partner's emotions (affect utilization), revealed consequences in terms of attenuating the testosterone response to stress.

Opposing effects of dopamine on agonistic behaviour in crayfish.

The effects of dopamine on the agonistic behaviour of crayfish were analysed. When dopamine concentrations of 1 µmol l-1 were injected into large crayfish, individuals were beaten by smaller opponents, despite their physical advantage. Injection of 10 µmol l-1 dopamine into small animals increased their rate of winning against larger opponents. Injection of a D1 receptor antagonist prohibited the onset of a 'loser' effect in subordinate animals, suggesting that the inhibitory effect of dopamine on larger animals is mediated by D1 receptors. Similarly, injection of a D2 receptor antagonist prohibited the onset of a 'winner' effect in dominant animals, suggesting that the facilitating effect of dopamine on small animals is mediated by D2 receptors. Since the inhibitory effect of 1 µmol l-1 dopamine was similar to that seen with 1 µmol l-1 octopamine and the facilitating effect of 10 µmol l-1 dopamine was similar to that of 1 µmol l-1 serotonin, functional interactions among dopamine, octopamine and serotonin were analyzed by co-injection of amines with their receptor antagonists in various combinations. The inhibitory effect of 1 µmol l-1 dopamine disappeared when administered with D1 receptor antagonist, but remained when combined with octopamine receptor antagonist. Octopamine effects disappeared when administered with either D1 receptor antagonist or octopamine receptor antagonist, suggesting that the dopamine system is downstream of octopamine. The facilitating effect of 10 µmol l-1 dopamine disappeared when combined with serotonin 5HT1 receptor antagonist or D2 receptor antagonist. Serotonin effects also disappeared when combined with D2 receptor antagonist, suggesting that dopamine and serotonin activate each other through parallel pathways.

Novel wing display and divergent agonistic behaviors of two incipient Psolodesmus damselflies.

Sexual selection via male competition is a strong evolutionary force that can drive rapid changes in competitive traits and subsequently lead to population divergence and speciation. Territorial males of many odonates are known to use their colorful wings as visual signals and to perform agonistic displays toward intruders. Psolodesmus mandarinus dorothea and Psolodesmus mandarinus mandarinus are two parapatrically distributed sister damselflies that share similar ecological characteristics but differ markedly in wing coloration. The wings of P. m. dorothea are mostly clear, whereas those of P. m. mandarinus have a large area of black pigmentation and a central white patch. We investigated whether territorial males of the two damselflies at breeding sites display distinct agonistic behaviors associated with their respective wing colors. Behavioral interactions between territorial and intruder males and their wing kinematics were filmed and analyzed for P. m. dorothea in Lienhuachih of central Taiwan, and P. m. mandarinus in Tianxiyuan and Fusan of northern Taiwan. We observed that the P. m. mandarinus males exhibited a novel set of perched wing displays, which was not only absent in its sister P. m. dorothea but also previously unknown in Odonata. At breeding sites, perched rival males of P. m. mandarinus with pigmented wings exhibited escalating agonistic wing-flapping and wing-hitting displays toward each other. In contrast, territorial males of P. m. dorothea with clear wings engaged only in aerial chase or face-to-face hovering when intruder males approached from the air. These results indicate that the two sister P. mandarinus damselflies diverged behaviorally in territorial contests and support the hypothesis of coadaptation on the basis of wing colors and types of wing movement in Odonata. Our findings further suggest that divergent agonistic wing displays may play a pivotal role in the speciation mechanism of P. mandarinus damselflies. The sequential analyses of behavioral characteristics and progression suggest that P. m. mandarinus damselflies likely use mutual assessment of rivals in territorial contests.

Agonistic interactions with asymmetric body size in two adult-age groups of the annual killifish Millerichthys robustus (Miller & Hubbs, 1974).

In this study, the author evaluated two adult age groups of the Mexican rivulus Millerichthys robustus with body size asymmetries to determine the strategies used by an annual killifish during agonistic interactions of different ontogenetic stages. To achieve this goal, the author first characterized the ethogram of agonistic interactions of M. robustus composed of seven behavioural units in males and five behavioural units in females. The author then analysed agonistic interaction strategies used by males and females with body size asymmetries in two groups of different adult ages that represent different ontogenetic stages: (a) just after sexual maturity was reached, at 5 weeks of age, and (b) near natural death, at 24 weeks of age. The agonistic behaviour patterns of M. robustus were compatible with the logic of mutual assessment. Large males had an advantage during their interactions in both age groups, winning all of the encounters. Nonetheless, there was more aggression in 5-week-old fish encounters. In addition, small 24-week-old fish were more aggressive than small 5-week-old fish. These changing strategies may be because of the cost-benefits required during a fight at each ontogenetic stage. In the female encounters, size did not predict winners, as both small and large fish won a similar number of encounters, and some contests remained unresolved regardless of age group. There was a tendency for small females of any age to risk more than males in fights to maintain reproductive fitness.

Serotonin Receptor 1A Variation Is Associated with Anxiety and Agonistic Behavior in Chimpanzees.

Serotonin is a neurotransmitter that plays an important role in regulating behavior and personality in humans and other mammals. Polymorphisms in genes coding for the serotonin receptor subtype 1A (HTR1A), the serotonin transporter (SLC6A4), and the serotonin degrading enzyme monoamine oxidase A (MAOA) are associated with anxiety, impulsivity, and neurotic personality in humans. In primates, previous research has largely focused on SLC6A4 and MAOA, with few studies investigating the role of HTR1A polymorphic variation on behavior. Here, we examined variation in the coding region of HTR1A across apes, and genotyped polymorphic coding variation in a sample of 214 chimpanzees with matched measures of personality and behavior. We found evidence for positive selection at three amino acid substitution sites, one in chimpanzees-bonobos (Thr26Ser), one in humans (Phe33Val), and one in orangutans (Ala274Gly). Investigation of the HTR1A coding region in chimpanzees revealed a polymorphic site, where a C/A single nucleotide polymorphism changes a proline to a glutamine in the amino acid sequence (Pro248Gln). The substitution is located in the third intracellular loop of the receptor, a region important for serotonin signal transduction. The derived variant is the major allele in this population (frequency 0.67), and is associated with a reduction in anxiety, decreased rates of male agonistic behavior, and an increase in socio-positive behavior. These results are the first evidence that the HTR1A gene may be involved in regulating social behavior in chimpanzees and encourage further systematic investigation of polymorphic variation in other primate populations with corresponding data on behavior.

Transcriptional regulatory dynamics drive coordinated metabolic and neural response to social challenge in mice.

Agonistic encounters are powerful effectors of future behavior, and the ability to learn from this type of social challenge is an essential adaptive trait. We recently identified a conserved transcriptional program defining the response to social challenge across animal species, highly enriched in transcription factor (TF), energy metabolism, and developmental signaling genes. To understand the trajectory of this program and to uncover the most important regulatory influences controlling this response, we integrated gene expression data with the chromatin landscape in the hypothalamus, frontal cortex, and amygdala of socially challenged mice over time. The expression data revealed a complex spatiotemporal patterning of events starting with neural signaling molecules in the frontal cortex and ending in the modulation of developmental factors in the amygdala and hypothalamus, underpinned by a systems-wide shift in expression of energy metabolism-related genes. The transcriptional signals were correlated with significant shifts in chromatin accessibility and a network of challenge-associated TFs. Among these, the conserved metabolic and developmental regulator ESRRA was highlighted for an especially early and important regulatory role. Cell-type deconvolution analysis attributed the differential metabolic and developmental signals in this social context primarily to oligodendrocytes and neurons, respectively, and we show that ESRRA is expressed in both cell types. Localizing ESRRA binding sites in cortical chromatin, we show that this nuclear receptor binds both differentially expressed energy-related and neurodevelopmental TF genes. These data link metabolic and neurodevelopmental signaling to social challenge, and identify key regulatory drivers of this process with unprecedented tissue and temporal resolution.

Discrete modulation of anti-predatory and agonistic behaviors by sensory communication signals in juvenile crayfish.

We investigated how the exchange of sensory signals modulates the individual behaviors of juvenile crayfish in an anti-predatory context as well as during intraspecific agonistic encounters. We first compared crayfish housed in total sensory isolation or in pairs with access to chemical and visual cues. After 1 week of housing, we analysed their individual responses to a visual danger signal while they were foraging. We found that crayfish previously housed in pairs with exchange of sensory signals responded to a simulated predator attack predominantly with freezing behavior, whereas animals deprived of all sensory communication mostly responded by performing escape tail-flips. Next, we used the same housing conditions in between repeated fights in pairs of crayfish. Aggressive and submissive behaviors increased in subsequent fights both after total isolation and after exchange of olfactory and visual signals. Thus, unlike responses to simulated predator attacks, intraspecific agonistic behavior was not modulated by exposure to the same sensory signals. However, when we tested the effects of olfactory or visual communication independently, aggression increased dramatically after the exchange of olfactory signals, which also led to a high number of rank reversals in second fights, suggesting a destabilization of the original dominance relationship. Exposure to visual cues during the 1-week separation, however, produced the opposite effect, reducing agonistic behaviors and rank reversals. These findings demonstrate that exchange of sensory signals modulates future anti-predatory decision-making and intraspecific agonistic behaviors discretely, suggesting that the effect of these signals on shared neural circuitry is context dependent.

Both sexes produce sounds in vocal fish species: testing the hypothesis in the pygmy gourami (labyrinth fishes).

In vocal fish species, males possess larger sound-generating organs and signal acoustically with pronounced sex-specific differences. Sound production is known in two out of three species of croaking gouramis (Trichopsis vittata and T. schalleri). The present study investigates sex-specific differences in sonic organs, vocalizing behaviour and sounds emitted in the third species, the pygmy gourami, T. pumila, in order to test the hypothesis that females are able to vocalize despite their less-developed sonic organs, and despite contradictory reports. Croaking gouramis stretch and pluck two enhanced (sonic) pectoral fin tendons during alternate fin beating, resulting in a series of double-pulsed bursts, termed croaking sound. We measured the diameter of the first and second sonic tendon and showed that male tendons were twice as large as in similar-sized females. We also determined the duration of dyadic contests, visual displays, number of sounds and buttings. Sexes differ in all sound characteristics but in no behavioural variable. Male sounds consisted of twice as many bursts, a higher percentage of double-pulsed bursts and a higher burst period. Additionally, male sounds had a lower dominant frequency and a higher sound level. In summary, female pygmy gouramis possessed sonic organs and vocalized in most dyadic contests. The sexual dimorphism in sonic tendons is clearly reflected in sex-specific differences in sound characteristics, but not in agonistic behaviour, supporting the hypothesis that females are vocal.