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Oceans absorb a huge part of the atmospheric heat, leading to the rise in water temperature. Reefs are among the most affected ecosystems, where the complex behavioral repertoire of fishes is usually an indicator of environmental impacts. Here, we examined whether temperature (28 and 34°C) and habitat complexity (high and low) interact to affect the agonistic behavior (mirror test) of the dusky damselfish (Stegastes fuscus), a key species in Brazilian reefs because of its gardening capacity and territorial behavior. Higher temperatures altered basal behavior in both high and low-complexity conditions. Fish kept at 28°C under the high-complexity condition were more aggressive than those at a higher temperature (34°C) and in a low-complexity condition, which also exhibited lower dispersion. Our data show that changes in behavior of coral reef fish is associated to fluctuations in environmental conditions. Thus, it is important to implement management or conservation strategies that could mitigate global change effects.
Pubertal testosterone programs the level of aggressive behavior displayed by male Syrian hamsters during resident-intruder interactions. To further explore the pubertal programming of adult male agonistic behaviors, the current study investigated the formation, stability, and maintenance of dominant-subordinate relationships in males that either did (T@P) or did not (NoT@P) experience testicular hormones during adolescent development. NoT@P males were gonadectomized prepubertally and T@P males were gonadectomized in adulthood. Four weeks after gonadectomy, all males received testosterone-replacement. Two weeks later, two males of the same hormonal history were given a 60 min introductory trial in a neutral arena, followed immediately and again 24h later by three 5-min trials. During the introductory trial, each male was deemed dominant, subordinate, or no-status. Brains were collected 1h after the last trial and sections were stained for Fos-immunoreactivity. Dominant T@P males flank marked more frequently than subordinate and no-status T@P males; this difference was not found in NoT@P males. NoT@P males showed an increase in the number of offensive postures the day after the first series of tests, whereas T@P males did not. Dominant T@P males had significantly more Fos expression than no-status T@P males in anterior cingulate cortex; this relationship was not observed in NoT@P males. Additionally, dominant T@P males had higher Fos expression than dominant NoT@P males in lateral septum. Thus, pubertal testosterone does not organize the formation or stability of male-male relationships, but does program the behavioral strategies used to maintain these relationships over time and the neural correlates of status.
The enzyme aromatase, responsible for the conversion of C19 androgens to C18 estrogens, exists as two paralogue copies in teleost fish: Cyp19a1a mostly expressed in the gonads, referred as gonadal aromatase, and Cyp19a1b, mostly expressed in the brain, accordingly known as brain aromatase. The neural localization of Cyp19a1b is greatly contained within the social behavior network and mesolimbic reward system in fish, suggesting a strong role of estrogen synthesis in the regulation of social behavior. In this work we aimed to analyze the variation in cyp19a1b expression in brain and pituitary of males of a highly social cichlid, Cichlasoma dimerus (locally known as chanchita), and its relation with inter-individual variability in agonistic behavior in a communal social environment. We first characterized chanchita's cyp19a1b mRNA and deduced amino acid sequence, which showed a high degree of conservation when compared to other teleost brain aromatase sequences, and its tissue expression patterns. Within the brain, Cyp19a1b was solely detected at putative radial glial cells of the forebrain, close to the brain ventricles. We then studied the relative expression levels of cyp19a1b by Real Time PCR in the brain and pituitary of males of different social status, territorial vs. non-territorial, and its relationship with an index of agonistic behavior. We found that even though, brain aromatase expression did not differ between types of males, pituitary cyp19a1b expression levels positively correlated with the index of agonistic behavior. This suggests a novel role of the pituitary in the regulation of social behavior by local estrogen synthesis.
Chinese mitten crab (Eriocheir sinensis) as a commercially important species is widely cultured in China. However, E. sinensis is prone to agonistic behavior, which causes physical damage and wastes energy resources, negatively impacting their growth and survival. Therefore, understanding the regulatory mechanisms that underlie the switching of such behavior is essential for ensuring the efficient and cost-effective aquaculture of E. sinensis. The 5-HT2B receptor is a key downstream target of serotonin (5-HT), which is involved in regulating animal behavior. In this study, the full-length sequence of 5-HT2B gene was cloned. The total length of the 5-HT2B gene was found to be 3127 bp with a 236 bp 5'-UTR (untranslated region), a 779 bp 3'-UTR, and a 2112 bp open reading frame encoding 703 amino acids. Phylogenetic tree analysis revealed that the 5-HT2B amino acid sequence of E. sinensis is highly conserved with that of Cancer borealis. Using in vitro co-culture and luciferase assays, the miR-143 targets the 5-HT2B 3'-UTR and inhibits 5-HT2B expression was confirmed. Furthermore, RT-qPCR and Western blotting analyses revealed that the miR-143 mimic significantly inhibits 5-HT2B mRNA and protein expression. However, injection of miR-143 did not decrease agonistic behavior, indicating that 5-HT2B is not involved in the regulation of such behavior in E. sinensis.
Daily agonistic interactions of mice are an effective experimental approach to elucidate the molecular mechanisms underlying the excitation of the brain neurons and the formation of alternative social behavior patterns. An RNA-Seq analysis was used to compare the ventral tegmental area (VTA) transcriptome profiles for three groups of male C57BL/6J mice: winners, a group of chronically winning mice, losers, a group of chronically defeated mice, and controls. The data obtained show that both winners and defeated mice experience stress, which however, has a more drastic effect on defeated animals causing more significant changes in the levels of gene transcription. Four genes (Nrgn, Ercc2, Otx2, and Six3) changed their VTA expression profiles in opposite directions in winners and defeated mice. It was first shown that Nrgn (neurogranin) expression was highly correlated with the expression of the genes involved in dopamine synthesis and transport (Th, Ddc, Slc6a3, and Drd2) in the VTA of defeated mice but not in winners. The obtained network of 31 coregulated genes, encoding proteins associated with nervous system development (including 24 genes associated with the generation of neurons), may be potentially useful for studying their role in the VTA dopaminergic neurons maturation under the influence of social stress.
Stressful events promote several neuroendocrine and neurotransmitter changes that might contribute to the provocation of psychological and physical pathologies. Perhaps, because of its apparent ecological validity and its simple application, there has been increasing use of social defeat (resident-intruder) paradigms as a stressor. The frequency of stress-related psychopathology is much greater in females than in males, but the typical resident-intruder paradigm is less useful in assessing stressor effects in females. An alternative, but infrequently used procedure in females involves exposing a mouse to a lactating dam, resulting in threatening gestures being expressed by the resident. In the present investigation we demonstrated the utility of this paradigm, showing that the standard resident-intruder paradigm in males and the modified version in females promoted elevated anxiety in a plus-maze test. The behavioral effects that reflected anxiety were more pronounced 2 weeks after the stressor treatment than they were 2 hr afterward, possibly reflecting the abatement of the stress-related of hyper-arousal. These treatments, like a stressor comprising physical restraint, increased plasma corticosterone and elicited variations of norepinephrine and serotonin levels and turnover within the prefrontal cortex, hippocampus and central amygdala. Moreover, the stressor effects were exaggerated among mice that had been exposed to a chronic or subchronic-intermittent regimen of unpredictable stressors. Indeed, some of the monoamine changes were more pronounced in females than in males, although it is less certain whether this represented compensatory changes to deal with chronic stressors that could result in excessive strain on biological systems (allostatic overload).
As a commercially important species, the Chinese mitten crab (Eriocheir sinensis) has been cultured for a long time in China. Agonistic behavior often causes limb disability and requires much energy, which is harmful to the growth and survival of crabs. In this paper, we divided crabs into a control group (control, no treatment) and an experimental group (fight, agonistic behavior after 1 h) and then collected the thoracic ganglia (TG) to extract RNA. Subsequently, we first used a deep sequencing approach to examine the transcripts of microRNAs (miRNAs) and messenger RNAs (mRNAs) in E. sinensis displaying agonistic behavior. According to the results, we found 29 significant differentially expressed miRNAs (DEMs) and 116 significant differentially expressed unigenes (DEGs). The DEMs esi-miR-199a-5p, esi-let-7d, esi-miR-200a, and esi-miR-200b might participate in the regulation of agonistic behavior by mediating neuroregulation and energy metabolism. Focusing on the transcripts of the mRNAs, the renin-angiotensin system (RAS) Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway might be involved in the regulation of agonistic behavior through glucose metabolism as this pathway was significantly enriched with DEGs. Besides, an integrated analysis of the miRNA and mRNA profiles revealed that the retinoid X receptor (RXR) was also involved in visual signal transduction, which was important for agonistic behavior. In addition, four vital agonistic behavior-related metabolic pathways, including the cAMP signaling, MAPK, protein digestion and absorption, and fatty acid metabolism pathways, were significantly enriched with the predicted target unigenes. In conclusion, the findings of this study might provide important insight enhancing our understanding of the underlying molecular mechanisms of agonistic behavior in E. sinensis.
Arginine vasopressin (AVP) regulates aggression in male Syrian hamsters. In this study, we used radioligand receptor autoradiography to examine whether changes in agonistic behavior following acute and repeated social defeat are accompanied by changes in AVP V1a receptor binding. Social defeat produced high levels of submissive behavior and a loss of territorial aggression when hamsters were subsequently tested with a novel intruder, and repeated agonistic encounters produced similar behavioral changes in subordinates. AVP V1a receptor binding was not reduced by acute social defeat but was affected by repeated agonistic encounters. Dominants had significantly more AVP V1a receptor binding in lateral portions of the ventromedial hypothalamus (VMHL) than did their subordinate opponents, but subordinates were no different from controls. In contrast, receptor binding did not differ in most other brain regions examined. The changes in receptor binding appear to be independent of testosterone levels, as testosterone levels did not differ among dominants, subordinates, and controls. Our results suggest that changes in AVP V1a receptors do not account for the changes in agonistic behavior produced by acute social defeat but AVP V1a binding in the VMHL correlates with, and may modulate, the behavioral changes that occur following repeated experiences of victory.
Social status and resource availability can strongly influence individual behavioral responses to conspecifics. In European starlings, males that acquire nest sites sing in response to females and dominate other males. Males without nest sites sing, but not to females, and they do not interact agonistically with other males. Little is known about the neural regulation of status- or resource-appropriate behavioral responses to conspecifics. Opioid neuropeptides are implicated in birdsong and agonistic behavior, suggesting that opioids may underlie differences in the production of these behaviors in males with and without nest sites. Here, we examined densities of immunolabeled mu-opioid receptors in groups of male starlings. Males that defended nest boxes dominated other males and sang at higher rates when presented with a female than males without nest boxes, independent of testosterone concentrations. Multiple regression analyses showed nest box ownership (not agonistic behavior or singing) predicted the optical density of receptor labeling in the medial bed nucleus of stria terminalis, paraventricular nucleus, ventral tegmental area and the medial preoptic nucleus. Compared to males without nest boxes, males with nest boxes had lower densities of immunolabeled mu-opioid receptors in these regions. Singing additionally predicted the area covered by labeling in the ventral tegmental area. The results suggest that elevated opioid activity in these regions suppresses courtship and agonistic behavioral responses to conspecifics in males without nest boxes. The findings are consistent with a dynamic role for opioid receptors in adjusting social behavior so that it is appropriate given the resources available to an individual.
Winning aggressive disputes is one of several experiences that can alter responses to future stressful events. We have previously tested dominant and subordinate male Syrian hamsters in a conditioned defeat model and found that dominant individuals show less change in behavior following social defeat stress compared to subordinates and controls, indicating a reduced conditioned defeat response. Resistance to the effects of social defeat in dominants is experience-dependent and requires the maintenance of dominance relationships for 14days. For this study we investigated whether winning aggressive interactions increases plasma testosterone and whether repeatedly winning increases androgen receptor expression. First, male hamsters were paired in daily 10-min aggressive encounters and blood samples were collected immediately before and 15min and 30min after the formation of dominance relationships. Dominants showed an increase in plasma testosterone at 15min post-interaction compared to their pre-interaction baseline, whereas subordinates and controls showed no change in plasma testosterone. Secondly, we investigated whether 14days of dominant social status increased androgen or estrogen alpha-receptor immunoreactivity in brain regions that regulate the conditioned defeat response. Dominants showed more androgen, but not estrogen alpha, receptor immuno-positive cells in the dorsal medial amygdala (dMeA) and ventral lateral septum (vLS) compared to subordinates and controls. Finally, we showed that one day of dominant social status was insufficient to increase androgen receptor immunoreactivity compared to subordinates. These results suggest that elevated testosterone signaling at androgen receptors in the dMeA and vLS might contribute to the reduced conditioned defeat response exhibited by dominant hamsters.
Avian vocalizations are common examples of the complex signals used by animals to negotiate during agonistic interactions. In this study, we used two playback experiments to identify agonistic signals in a songbird species with several acoustically complex songs and calls, the veery. In the first experiment, we compared veery singing behavior in response to simulated territorial intrusions including playback of three variations of veery song: 1) song alone as a control, 2) songs with added whisper calls, and 3) songs with introductory notes removed. In the second experiment, we used multimodal stimuli including songs, whisper calls and songs with introductory notes removed, along with a robotic veery mount. Focal males readily responded to all of the playback stimuli, approached the speaker and/or robotic mount, and vocalized. Male veeries gave more whisper calls, and sang more songs without the introductory note in response to all types of playback. However, veeries responded similarly to all types of stimuli presented, and they failed to physically attack the robotic mount. These results indicate that rival veeries use two different types of novel vocalizations: whisper calls and songs lacking the introductory note as agonistic signals, but do not allow us to discern the specific functions of these two vocalizations.
In this study we present evidence that 20-hydroxyecdysone (20E) affects agonistic behavior in male American lobsters and that male and female animals differ in their response to the hormone. Thirty-minute staged fights were conducted between large males exposed either to artificial seawater (ASW) or 20E and small, anosmic opponents. The nephropores of both combatants were blocked. Fights were videotaped and quantitatively analyzed for aggressive, defensive and avoidance behaviors using an ethogram in which behaviors are ranked according to aggressiveness. Unlike female lobsters, exposing male lobsters to 20E did not increase their aggressive behavior; however, there was a marginally significant trend toward an increase in defensive behaviors with a lower aggressive content than in their ASW-exposed counterparts. The opponents of 20E-exposed animals performed significantly more aggressive behaviors than their counterparts. In fights with 20E-exposed animals, the overall aggressive intensity of the fight was increased and the animals performed a greater number of avoidance behaviors. Unlike the effects of 20E on females, where exposure to 20E caused an increase in overall agonistic arousal, males only exhibited a change in frequency of their behaviors. These findings suggest that while 20E affects both males and females in agonistic encounters, the nature of the effect is different for the two sexes.
Several neurobiological mechanisms are implicated in the formation of selective pair bonds in socially monogamous mammals, however much less is known about the mechanisms that underlie the long-term behavioral maintenance of these bonds. In prairie voles (Microtus ochrogaster), agonistic behavior that contributes to pair bond maintenance are regulated by dopamine activity at D1-like receptors (D1R) within the mesocorticolimbic system. Evidence suggests D1Rs similarly regulate the behavioral components of pair bond maintenance in socially monogamous titi monkeys (Callicebus cupreus); however, evaluation with behavioral pharmacology is necessary to evaluate this hypothesis. In the current study we evaluated the role of D1Rs in behavioral components of pair bond maintenance in captive male titi monkeys (N = 8). We administered two doses of a D1R selective antagonist, SCH23390, (0.1 mg/kg, 0.01 mg/kg) or saline vehicle to male titi monkeys and presented pairs with a simulated intruder monkey via the use of a mirror stimulus. The non-reflective back of the mirror stimulus was used for control sessions. We video recorded responses to the five-minute stimulus presentations and later scored for arousal and agonistic behaviors relevant to mate guarding as well as affiliative behavior between the pair mates. We also conducted a locomotor assessment to evaluate the potential side effect for SCH23390 of impaired locomotion. Finally, we collected blood samples at the end of each session to assay for plasma cortisol responses. We found evidence of locomotor impairment only with the high dose of SCH23390, and therefore analyses were conducted comparing only test sessions where low dose SCH23390 and saline were administered. With saline administration, males displayed more agonistic behavior via back arching and tail lashing as well as restraining their female partners when viewing the mirror compared to the back of the mirror. D1R antagonist treatment attenuated these agonistic behaviors indicative of mate guarding when males viewed the mirror. Results also indicated that this reduction in agonistic behavior occurred without evidence of overall behavioral blunting or generally reduced social interest. Likewise changes in agonistic behavior were not driven by differences in HPA activity across testing sessions. Mate-directed affiliative behavior, including lip smacks and approaches to female partners, were not altered by D1R antagonist treatment. Dyadic social contact was higher with D1R antagonist treatment, but this was due to a reduction in contact termination by the treated males, which was typically followed by an approach or arousal display to the simulated intruder. These results provide further evidence that D1R activity regulates mate guarding behaviors in titi monkeys and suggests that the dopamine system plays a similar role in the agonistic behavioral components of pair bond maintenance behavior in non-human primates and rodents.
Animals have evolved flexible strategies that allow them to evaluate and respond to their social environment by integrating the salience of external stimuli with internal physiological cues into adaptive behavioral responses. A highly conserved social decision-making network (SDMN), consisting of interconnected social behavior and mesolimbic reward networks, has been proposed to underlie such adaptive behaviors across all vertebrates, although our understanding of this system in reptiles is very limited. Here we measure neural activation across the SDMN and associated regions in the male brown anole (Anolis sagrei), within both reproductive and agonistic contexts, by quantifying the expression density of the immediate early gene product Fos. We then relate this neural activity measure to social context, behavioral expression, and activation (as measured by colocalization with Fos) of different phenotypes of 'source' node neurons that produce neurotransmitters and neuropeptides known to modulate SDMN 'target' node activity. Our results demonstrate that measures of neural activation across the SDMN network are generally independent of specific behavioral output, although Fos induction in a few select nodes of the social behavior network component of the SDMN does vary with social environment and behavioral output. Under control conditions, the mesolimbic reward nodes of the SDMN actually correlate little with the social behavior nodes, but the interconnectivity of these SDMN components increases dramatically within a reproductive context. When relating behavioral output to specific source node activation profiles, we found that catecholaminergic activation is associated with the frequency and intensity of reproductive behavior output, as well as with aggression intensity. Finally, in terms of the effects of source node activation on SDMN activity, we found that Ile8-oxytocin (mesotocin) populations correlate positively, while Ile3-vasopressin (vasotocin), catecholamine, and serotonin populations correlate negatively with SDMN activity. Taken together, our findings present evidence for a highly dynamic SDMN in reptiles that is responsive to salient cues in a social context-dependent manner.
Chronic agonistic interactions promote the development of experimental psychopathologies in animals: a depression-like state in chronically defeated mice and the pathology of aggressive behavior in the mice with repeated wins. The abundant research data indicate that such psychopathological states are associated with significant molecular and cellular changes in the brain. This paper aims to study the influence of a 20-day period of agonistic interactions on the expression patterns of collagen family genes encoding the proteins which are basic components of extracellular matrix (ECM) in different brain regions of mice using the RNA-Seq database. Most of differentially expressed collagen genes were shown to be upregulated in the hypothalamus and striatum of chronically aggressive and defeated mice and in the hippocampus of defeated mice, whereas downregulation of collagen genes was demonstrated in the ventral tegmental areas in both experimental groups. Aberrant expression of collagen genes induced by chronic agonistic interactions may be indicative of specific ECM defects in the brain regions of mice with alternative social experience. This is the first study demonstrating remodeling of ECM under the development of experimental disorders.
Complex behavioral phenotyping techniques are becoming more prevalent in the field of behavioral neuroscience, and thus methods for manipulating neuronal activity must be adapted to fit into such paradigms. Here, we present a head-mounted, magnetically activated device for wireless optogenetic manipulation that is compact, simple to construct, and suitable for use in group-living mice in an enriched semi-natural arena over several days. Using this device, we demonstrate that repeated activation of oxytocin neurons in male mice can have different effects on pro-social and agonistic behaviors, depending on the social context. Our findings support the social salience hypothesis of oxytocin and emphasize the importance of the environment in the study of social neuromodulators. Our wireless optogenetic device can be easily adapted for use in a variety of behavioral paradigms, which are normally hindered by tethered light delivery or a limited environment.
In recent years, computer vision has contributed significantly to the study of farm animal behavior. In complex environments such as commercial farms, however, the automated detection of social behavior and specific interactions between animals can be improved. The present study addresses the automated detection of agonistic interactions between caged animals in a complex environment, relying solely on computer vision. An automated pipeline including group-level temporal action segmentation, object detection, object tracking and rule-based action classification for the detection of agonistic interactions was developed and extensively validated at a level unique in the field. Comparing with observations made by human observers, our pipeline reaches 77% precision and 85% recall using a 5-min tolerance interval for the detection of agonistic interactions. Results obtained using this pipeline allow to construct time-dependent socio-matrices of a group of animals and derive metrics on the dominance hierarchy in a semi-automated manner. Group-housed breeding rabbits (does) with their litters in commercial farms are the main use-case in this work, but the idea is probably also applicable to other social farm animals.
Excessive home cage aggression often results in severe injury and subsequent premature euthanasia of male laboratory mice. Aggression can be reduced by transferring used nesting material during cage cleaning, which is thought to contain aggression appeasing odors from the plantar sweat glands. However, neither the composition of plantar sweat nor the deposits on used nesting material have been evaluated. The aims of this study were to (1) identify and quantify volatile compounds deposited in the nest site and (2) determine if nest and sweat compounds correlate with social behavior. Home cage aggression and affiliative behavior were evaluated in 3 strains: SJL, C57BL/6N, and A/J. Individual social rank was assessed via the tube test, because ranking may influence compound levels. Sweat and urine from the dominant and subordinate mouse in each cage, plus cage level nest samples were analyzed for volatile compound content using gas chromatography-mass spectrometry. Behavior data and odors from the nest, sweat, and urine were statistically analyzed with separate principal component analyses (PCA). Significant components, from each sample analysis, and strain were run in mixed models to test if odors were associated with behavior. Aggressive and affiliative behaviors were primarily impacted by strain. However, compound PCs were also impacted by strain, showing that strain accounts for any relationship between odors and behavior. C57BL/6N cages displayed the most allo-grooming behavior and had high scores on sweat PC1. SJL cages displayed the most aggression, with high scores on urine PC2 and low scores on nest PC1. These data show that certain compounds in nesting material, urine, and sweat display strain specific patterns which match strain specific behavior patterns. These results provide preliminary information about the connection between home cage compounds and behavior. Salient compounds will be candidates for future controlled studies to determine their direct effect on mouse social behavior.
Sniffing is a specialized respiratory behavior that is essential for the acquisition of odors [1-4]. Perhaps not independent of this, sniffing is commonly displayed during motivated [5-7] and social behaviors [8, 9]. No measures of sniffing among interacting animals are available, however, calling into question the utility of this behavior in the social context. From radiotelemetry recordings of nasal respiration, I found that investigation by one rat toward the facial region of a conspecific often elicits a decrease in sniffing frequency in the conspecific. This reciprocal display of sniffing was found to be dependent upon the rat's social status in two separate paradigms, with subordinates reliably decreasing their sniffing frequency upon being investigated in the face by dominant rats. Failure of subordinates to decrease their sniffing frequency shortened the latency for agonistic behavior by dominant rats, reflecting that decreases in sniffing serve as appeasement signals during social interactions. Rats rendered unable to smell persisted in displaying reciprocal sniffing behavior, demonstrating the independence of this behavior from olfaction. Oxytocin treatment in rats with established social hierarchies abolished agonistic behaviors and reciprocal sniffing displays. Together, these findings demonstrate that rodents utilize sniffing behaviors communicatively, not only to collect [6, 10-14] but also to convey information.
Among female rats, some individuals show estrus cycle-dependent irritability/aggressive behaviors, and these individual rats may be used as a model for premenstrual dysphoric disorder (PMDD). We wanted to investigate if these behaviors are related to the estrus cycle phase containing moderately increased levels of positive GABA-A receptor-modulating steroids (steroid-PAM), especially allopregnanolone (ALLO), and if the adverse behavior can be antagonized. The electrophysiology studies in this paper show that isoallopregnanolone (ISO) is a GABA-A-modulating steroid antagonist (GAMSA), meaning that ISO can antagonize the agonistic effects of positive GABA-A receptor-modulating steroids in both α1β2γ2L and α4β3δ GABA-A receptor subtypes. In this study, we also investigated whether ISO could antagonize the estrus cycle-dependent aggressive behaviors in female Wistar rats using a resident-intruder test. Our results confirmed previous reports of estrus cycle-dependent behaviors in that 42% of the tested rats showed higher levels of irritability/aggression at diestrus compared to those at estrus. Furthermore, we found that, during the treatment with ISO, the aggressive behavior at diestrus was alleviated to a level comparable to that of estrus. We noticed an 89% reduction in the increase in aggressive behavior at diestrus compared to that at estrus. Vehicle treatment in the same animals showed a minimal effect on the diestrus-related aggressive behavior. In conclusion, we showed that ISO can antagonize Steroid-PAM both in α1β2γ2L and α4β3δ GABA-A receptor subtypes and inhibit estrus cycle-dependent aggressive behavior.
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