Behavioral Neuroendocrinology Research Articles

Exposure to polystyrene microplastics reduces sociality and brain oxytocin levels through the gut-brain axis in mice

The rising global prevalence of microplastics (MPs) has highlighted their diverse toxicological effects. The oxytocin (OT) system in mammals, deeply intertwined with social behaviors, is recognized to be vulnerable to environmental stressors. We hypothesized that MP exposure might disrupt this system, a topic not extensively studied. We investigated the effects of MPs on behavioral neuroendocrinology via the gut-brain axis by exposing adolescent male C57BL/6 mice to varied sizes (5 μm and 50 μm) and concentrations (100 μg/L and 1000 μg/L) of polystyrene MPs over 10 weeks. The results demonstrated that exposure to 50 μm MPs significantly reduced colonic mucin production and induced substantial alterations in gut microbiota. Notably, the 50 μm-100 μg/L group showed a significant reduction in OT content within the medial prefrontal cortex and associated deficits in sociality, along with damage to the blood-brain barrier. Importantly, blocking the vagal pathway ameliorated these behavioral impairments, emphasizing the pivotal role of the gut-brain axis in mediating neurobehavioral outcomes. Our findings confirm the toxicity of MPs on sociality and the corresponding neuroendocrine systems, shedding light on the potential hazards and adverse effects of environmental MPs exposure on social behavior and neuroendocrine frameworks in social mammals, including humans.

Gender minority stress and diurnal cortisol profiles among transgender and gender diverse people in the United States

The field of behavioral neuroendocrinology has only begun to explore the lived experiences of transgender and gender diverse (TGD) people exposed to stigma. In light of escalating attacks and legislation targeting TGD people in the United States, it is crucial to examine the physiological pathways through which gender minority stressors become embodied, impact health, and contribute to health inequities. The Trans Resilience and Health Study included baseline data collection from fall 2019 to spring 2020 from a sample of 124 TGD people, reflecting a diversity of gender identities (e.g., trans masculine, trans feminine, and nonbinary) and ages (range = 19–70 years old; M = 34.10), living in Michigan, Nebraska, Oregon, and Tennessee. These analyses examine experiences of gender-related enacted stigma in association with hypothalamic-pituitary-adrenal (HPA)-axis functioning. Among those experiencing the highest levels of enacted stigma, findings show a blunted cortisol awakening response and sluggish daily decline that resulted in elevated concentrations at bedtime compared to those experiencing less enacted stigma. These results of flattened diurnal activity are consistent with an emergent literature on discrimination as a social determinant of potential stress pathophysiology. In contrast, community connectedness was associated with a larger, more dynamic cortisol awakening response. These findings emphasize the importance of incorporating gender-minority stress and resilience measures when studying HPA-axis functioning among TGD people.

Contribution of birds to the study of sexual differentiation of brain and behavior

Behavioral neuroendocrinology has largely relied on mammalian models to understand the relationship between hormones and behavior, even if this discipline has historically used a larger diversity of species than other fields. Recent advances revealed the potential of avian models in elucidating the neuroendocrine bases of behavior. This paper provides a review focused mainly on the contributions of our laboratory to the study of sexual differentiation in Japanese quail and songbirds. Quail studies have firmly established the role of embryonic estrogens in the sexual differentiation of male copulatory behavior. While most sexually differentiated features identified in brain structure and physiology result from the different endocrine milieu of adults, a few characteristics are organized by embryonic estrogens. Among them, a sex difference was identified in the number and morphology of microglia which is not associated with sex differences in the concentration/expression of neuroinflammatory molecules. The behavioral role of microglia and neuroinflammatory processes requires further investigations. Sexual differentiation of singing in zebra finches is not mediated by the same endocrine mechanisms as male copulatory behavior and “direct” genetic effect, i.e., not mediated by gonadal steroids have been identified. Epigenetic contributions have also been considered. Finally sex differences in specific aspects of singing behavior have been identified in canaries after treatment of adults with exogenous testosterone suggesting that these aspects of song are differentiated during ontogeny. Integration of quail and songbirds as alternative models has thus expanded understanding of the interplay between hormones and behavior in the control of sexual differentiation.

Leveraging individual power to improve racial equity in academia

Academia in the United States continues to grapple with its longstanding history of racial discrimination and its active perpetuation of racial disparities. To this end, universities and academic societies must grow in ways that reduce racial minoritization and foster racial equity. What are the effective and long-lasting approaches we as academics should prioritize to promote racial equity in our academic communities? To address this, the authors held a diversity, equity, and inclusion (DEI) panel during the Society for Behavioral Neuroendocrinology 2022 annual meeting, and in the following commentary synthesize the panelists' recommendations for fostering racial equity in the US academic community.

No Compelling Evidence that Self-Reported Personality Traits Explain Basal Testosterone and Cortisol’s Associations with Status-Relevant Behavior

ObjectiveA goal of behavioral neuroendocrinology is to understand how basal hormone levels relate to behavior. Studies of human participants sometimes measure self-reported personality traits, in addition to or instead of direct behavioral observation. Although personality traits often predict their respective behaviors, whether personality explains hormone-behavior relationships remains unclear.MethodsWe obtained data from eight previous studies (total N = 985) that examined baseline testosterone and cortisol as predictors of status-relevant behavior (competitiveness, dominance, risk-taking, aggression, affiliation, and social status). We tested whether the previously reported hormone-behavior relationships are mediated by self-reported personality traits (e.g., trait dominance, prestige, extraversion). As a secondary research question, we also tested whether trait dominance moderated the testosterone-behavior relationships.ResultsAs expected, self-reported personality traits often predicted status-relevant behaviors, but there was little evidence that traits also correlated with basal testosterone or the testosterone × cortisol interaction. Across all eight studies, personality traits did not significantly mediate hormone-behavior relationships. Indeed, the effect sizes of the hormone-behavior relationships were robust to the inclusion of personality traits as covariates. Further, we did not find strong or consistent evidence that trait dominance moderates the testosterone-behavior association.ConclusionResults suggest that basal testosterone and cortisol predict status-related behavior independent of self-reported personality. We discuss how these results may have broader implications for the physiological mechanisms by which testosterone and cortisol influence behavior, a process that could be unconscious and automatic. We also discuss alternative explanations, limitations, and future directions.

Methods and considerations for the use of hormonal contraceptives in rat models of neurobehavior

Hormonal contraceptives (HCs), prescribed to millions of women around the world, alter the ovarian hormonal cycle resulting in neurobehavioral changes in HC users. Human epidemiological and experimental data has characterized some of these effects with oftentimes conflicting or irreproducible results, reflecting a dearth of research considering different compositions, routes of administration, or time-courses of HC use. Non-human animal research can model these effects and help elucidate the underlying mechanisms by which different HCs modulate neurobehavioral outcomes. Still, animal models using HCs are not well-established. This may be because the pharmacological profile of HCs – including the metabolism, receptor binding affinity, and neuromodulatory effects – is dynamic and not always clearly translatable between animals and humans. The current review addresses these issues and provides basic methods and considerations for the use of HCs in animal models of neurobehavior to help advance the field of behavioral neuroendocrinology and inform decisions regarding to women’s health.

CRISPR-Cas9 editing of the arginine–vasopressin V1a receptor produces paradoxical changes in social behavior in Syrian hamsters

Studies from a variety of species indicate that arginine–vasopressin (AVP) and its V1a receptor (Avpr1a) play a critical role in the regulation of a range of social behaviors by their actions in the social behavior neural network. To further investigate the role of AVPRs in social behavior, we performed CRISPR-Cas9–mediated editing at the Avpr1a gene via pronuclear microinjections in Syrian hamsters (Mesocricetus auratus), a species used extensively in behavioral neuroendocrinology because they produce a rich suite of social behaviors. Using this germ-line gene-editing approach, we generated a stable line of hamsters with a frame-shift mutation in the Avpr1a gene resulting in the null expression of functional Avpr1as. Avpr1a knockout (KO) hamsters exhibited a complete lack of Avpr1a-specific autoradiographic binding throughout the brain, behavioral insensitivity to centrally administered AVP, and no pressor response to a peripherally injected Avpr1a-specific agonist, thus confirming the absence of functional Avpr1as in the brain and periphery. Contradictory to expectations, Avpr1a KO hamsters exhibited substantially higher levels of conspecific social communication (i.e., odor-stimulated flank marking) than their wild-type (WT) littermates. Furthermore, sex differences in aggression were absent, as both male and female KOs exhibited more aggression toward same-sex conspecifics than did their WT littermates. Taken together, these data emphasize the importance of comparative studies employing gene-editing approaches and suggest the startling possibility that Avpr1a-specific modulation of the social behavior neural network may be more inhibitory than permissive.

493 The use of visual arts to teach complex ideas on non-binary sex/gender traits and identities.

OBJECTIVES/GOALS: Recent scientific discoveries show that human sex determination and differentiation is a spectrum of developmental processes and that sex/gender traits and identities not always fit binarity. This study aims to determine whether the visual arts can effectively transmit these complex scientific ideas on sex and gender variance. METHODS/STUDY POPULATION: A one-hour lecture for undergraduate students enrolled in a behavioral neuroendocrinology course included 17th century Spanish paintings and representative work from contemporary LGBTQIA+ artists in photography. Pre and post self-evaluation was conducted through five multiple choice questions. Chi Square test was employed for statistical analysis, which required elimination of no responses from item analysis. Statistical significance was defined as p value < 0.05. Three artistic images and two scientific images were included in the questions. Acquisition of content-specific knowledge on diverse body configurations, differences of sex development (DSD), transsexuality, histopathology of the gonad and neuroanatomy of the hypothalamus was evaluated. RESULTS/ANTICIPATED RESULTS: Twenty students completed the online pre- and post-test. We found that a 17th century Spanish painting was effective at making the distinction between transsexuality and intersexuality (p < 0.0005). Similarly, an artistic image that reinterprets the Vitruvius Man was effective for presenting the argument that diverse body configurations does not imply clinical pathology (p < 0.0001). Last, a scientific image showing the histopathology of an ovotestis was effective for introducing undergraduate students to diagnostic criteria for DSD (p < 0.02). No significant differences were found in the use of an artistic photograph depicting a young female to male transsexual individual nor in the use of a scientific image showing the neuroanatomical localization of the hypothalamus. DISCUSSION/SIGNIFICANCE: The learning of complex scientific concepts on human sex determination and differentiation can be affected by preconceived ideas, values, and attitudes towards sex and gender variance. The visual arts can provide a familiar ground of understanding between teachers and learners to transform such preconceptions.

Uncovering the seasonal brain: Liquid chromatography-tandem mass spectrometry (LC-MS/MS) as a biochemical approach for studying seasonal social behaviors

Many animals show pronounced changes in physiology and behavior across the annual cycle, and these adaptations enable individuals to prioritize investing in the neuroendocrine mechanisms underlying reproduction and/or survival based on the time of year. While prior research has offered valuable insight into how seasonal variation in neuroendocrine processes regulates social behavior, the majority of these studies have investigated how a single hormone influences a single behavioral phenotype. Given that hormones are synthesized and metabolized via complex biochemical pathways and often act in concert to control social behavior, these approaches provide a limited view of how hormones regulate seasonal changes in behavior. In this review, we discuss how seasonal influences on hormones, the brain, and social behavior can be studied using liquid chromatography-tandem mass spectrometry (LC-MS/MS), an analytical chemistry technique that enables researchers to simultaneously quantify the concentrations of multiple hormones and the activities of their synthetic enzymes. First, we examine studies that have investigated seasonal plasticity in brain-behavior interactions, specifically by focusing on how two groups of hormones, sex steroids and nonapeptides, regulate sexual and aggressive behavior. Then, we explain the operations of LC-MS/MS, highlight studies that have used LC-MS/MS to study the neuroendocrine mechanisms underlying social behavior, both within and outside of a seasonal context, and discuss potential applications for LC-MS/MS in the field of behavioral neuroendocrinology. We propose that this cutting-edge technology will provide a more comprehensive understanding of how the multitude of hormones that comprise complex neuroendocrine networks affect seasonal variation in the brain and behavior.

A consideration of brain networks modulating social behavior

A primary goal of the field of behavioral neuroendocrinology is to understand how the brain modulates complex behavior. Over the last 20 years we have proposed various brain networks to explain behavioral regulation, however, the parameters by which these networks are identified are often ill-defined and reflect our personal scientific biases based on our area of expertise. In this perspective article, I question our characterization of brain networks underlying behavior and their utility. Using the Social Behavior Network as a primary example, I outline issues with brain networks commonly discussed in the field of behavioral neuroendocrinology, argue that we reconsider how we identify brain networks underlying behavior, and urge the future use of analytical tools developed by the field of Network Neuroscience. With modern statistical/mathematical tools and state of the art technology for brain imaging, we can strive to minimize our bias and generate brain networks that may more accurately reflect how the brain produces behavior.

The behavioral neuroendocrinology of dopamine systems in differently reared juvenile male rhesus monkeys (Macaca mulatta)

Dopamine (DA) is a critical neuromodulator of behavior. With propensities for addiction, hyper-activity, cognitive impairment, aggression, and social subordinance, monkeys enduring early maternal deprivation evoke human disorders involving dopaminergic dysfunction. To examine whether DA system alterations shape the behavioral correlates of adverse rearing, male monkeys (Macaca mulatta) were either mother-reared (MR: N = 6), or separated from their mothers at birth and nursery-reared (NR: N = 6). Behavior was assessed during 20-minute observations of subjects interacting with same- or differently-reared peers. Cerebrospinal fluid (CSF) biogenic amines, and serum testosterone (T), cortisol (CORT), and prolactin (PRL) were collected before and after pharmacologic challenge with saline or the DA receptor-2 (DRD2) antagonist Raclopride (RAC). Neuropeptide correlations observed in MR were non-existent in NR monkeys. Compared to MR, NR showed reduced DA tone; higher basal serum T; and lower CSF serotonin (5-HT). RAC increased PRL, T and CORT, but the magnitude of responses varied as a function of rearing. Levels of PRL significantly increased following RAC in MR, but not NR. Elevations in T following RAC were only significant among MR. Contrastingly, the net change (RAC CORT – saline CORT) in CORT was greater in NR than MR. Finally, observations conducted during the juvenile phase in a novel play-arena revealed more aggressive, self-injurious, and repetitive behaviors, which negatively correlated with indexes of dopaminergic tone in NR monkeys. In conclusion, early maternal deprivation alters brain DA systems, and thus may be associated with characteristic cognitive, social, and addiction outcomes.

Vocal and Electric Fish: Revisiting a Comparison of Two Teleost Models in the Neuroethology of Social Behavior.

The communication behaviors of vocal fish and electric fish are among the vertebrate social behaviors best understood at the level of neural circuits. Both forms of signaling rely on midbrain inputs to hindbrain pattern generators that activate peripheral effectors (sonic muscles and electrocytes) to produce pulsatile signals that are modulated by frequency/repetition rate, amplitude and call duration. To generate signals that vary by sex, male phenotype, and social context, these circuits are responsive to a wide range of hormones and neuromodulators acting on different timescales at multiple loci. Bass and Zakon (2005) reviewed the behavioral neuroendocrinology of these two teleost groups, comparing how the regulation of their communication systems have both converged and diverged during their parallel evolution. Here, we revisit this comparison and review the complementary developments over the past 16 years. We (a) summarize recent work that expands our knowledge of the neural circuits underlying these two communication systems, (b) review parallel studies on the action of neuromodulators (e.g., serotonin, AVT, melatonin), brain steroidogenesis (via aromatase), and social stimuli on the output of these circuits, (c) highlight recent transcriptomic studies that illustrate how contemporary molecular methods have elucidated the genetic regulation of social behavior in these fish, and (d) describe recent studies of mochokid catfish, which use both vocal and electric communication, and that use both vocal and electric communication and consider how these two systems are spliced together in the same species. Finally, we offer avenues for future research to further probe how similarities and differences between these two communication systems emerge over ontogeny and evolution.

Hacia una historia revisada de la teoría organizacional-activacional

En las últimas décadas la neuroendocrinología del comportamiento ha pasado de un área marginal de la incipiente revolución neurocientífica a una disciplina bien consolidada, hasta el punto de convertirse en el abordaje estándar para las diferencias y la diferenciación sexual de la conducta. Tanto en los trabajos de historia general de la endocrinología, como en las reviews de la especialidad y las aproximaciones críticas desde el feminismo académico, se ha generado un racconto según el cual la teoría central de la neuroendocrinología, esto es, la Teoría Organizacional Activacional habría sido fundada en 1959 por W. C. Young y sus colaboradores al descubrir los efectos de la testosterona fetal sobre conejillas de indias hembra. En el presente artículo se propone una revisión de tal historia oficial, comenzando con los trabajos de Arnold Berthold, profundizando en los desarrollos endocrinos de los años veinte y treinta del siglo pasado y revisando las disputas entre Young y Beach en los años previos a la publicación del 59. Con esta revisión, se pretende mejorar el conocimiento histórico de la teoría organizacional activacional, lo cual de forma derivada puede ayudar a iluminar algunas polémicas en torno a las explicaciones biológicas de la sexualidad humana.

Oxytocin, vasopressin and social behavior in the age of genome editing: A comparative perspective

Behavioral neuroendocrinology has a rich history of using diverse model organisms to elucidate general principles and evolution of hormone-brain-behavior relationships. The oxytocin and vasopressin systems have been studied in many species, revealing their role in regulating social behaviors. Oxytocin and vasopressin receptors show remarkable species and individual differences in distribution in the brain that have been linked to diversity in social behaviors. New technologies allow for unprecedented interrogation of the genes and neural circuitry regulating behaviors, but these approaches often require transgenic models and are most often used in mice. Here we discuss seminal findings relating the oxytocin and vasopressin systems to social behavior with a focus on non-traditional animal models. We then evaluate the potential of using CRISPR/Cas9 genome editing to examine the roles of genes and enable circuit dissection, manipulation and activity monitoring of the oxytocin and vasopressin systems. We believe that it is essential to incorporate these genetic and circuit level techniques in comparative behavioral neuroendocrinology research to ensure that our field remains innovative and attractive for the next generation of investigators and funding agencies.

Forebrain Transcriptional Response to Transient Changes in Circulating Androgens in a Cichlid Fish.

It has been hypothesized that androgens respond to the social interactions as a way to adjust the behavior of individuals to the challenges of the social environment in an adaptive manner. Therefore, it is expected that transient changes in circulating androgen levels within physiological scope should impact the state of the brain network that regulates social behavior, which should translate into adaptive behavioral changes. Here, we examined the effect that a transient peak in androgen circulating levels, which mimics socially driven changes in androgen levels, has on the forebrain state, which harbors most nuclei of the social decision-making network. For this purpose, we successfully induced transient changes in circulating androgen levels in an African cichlid fish (Mozambique tilapia, Oreochromis mossambicus) commonly used as a model in behavioral neuroendocrinology by injecting 11-ketotestosterone or testosterone, and compared the forebrain transcriptome of these individuals to control fish injected with vehicle. Forebrain samples were collected 30 min and 60 min after injection and analyzed using RNAseq. Our results showed that a transient peak in 11-ketotestosterone drives more accentuated changes in forebrain transcriptome than testosterone, and that transcriptomic impact was greater at the 30 min than at the 60 min post-androgen administration. Several genes involved in the regulation of translation, steroid metabolism, ion channel membrane receptors, and genes involved in epigenetic mechanisms were differentially expressed after 11-ketotestosterone or testosterone injection. In summary, this study identified specific candidate genes that may regulate socially driven changes in behavioral flexibility mediated by androgens.

Steroid profiles in quail brain and serum: Sex and regional differences and effects of castration with steroid replacement.

Both systemic and local production contribute to the concentration of steroids measured in the brain. This idea was originally based on rodent studies and was later extended to other species, including humans and birds. In quail, a widely used model in behavioural neuroendocrinology, it was demonstrated that all enzymes needed to produce sex steroids from cholesterol are expressed and active in the brain, although the actual concentrations of steroids produced were never investigated. We carried out a steroid profiling in multiple brain regions and serum of sexually mature male and female quail by gas chromatography coupled with mass spectrometry. The concentrations of some steroids (eg, corticosterone, progesterone and testosterone) were in equilibrium between the brain and periphery, whereas other steroids (eg, pregnenolone (PREG), 5α/β-dihydroprogesterone and oestrogens) were more concentrated in the brain. In the brain regions investigated, PREG sulphate, progesterone and oestrogen concentrations were higher in the hypothalamus-preoptic area. Progesterone and its metabolites were more concentrated in the female than the male brain, whereas testosterone, its metabolites and dehydroepiandrosterone were more concentrated in males, suggesting that sex steroids present in quail brain mainly depend on their specific steroidogenic pathways in the ovaries and testes. However, the results of castration experiments suggested that sex steroids could also be produced in the brain independently of the peripheral source. Treatment with testosterone or oestradiol restored the concentrations of most androgens or oestrogens, respectively, although penetration of oestradiol in the brain appeared to be more limited. These studies illustrate the complex interaction between local brain synthesis and the supply from the periphery for the steroids present in the brain that are either directly active or represent the substrate of centrally located enzymes.

The future of sex and gender in psychology: Five challenges to the gender binary.

The view that humans comprise only two types of beings, women and men, a framework that is sometimes referred to as the "gender binary," played a profound role in shaping the history of psychological science. In recent years, serious challenges to the gender binary have arisen from both academic research and social activism. This review describes 5 sets of empirical findings, spanning multiple disciplines, that fundamentally undermine the gender binary. These sources of evidence include neuroscience findings that refute sexual dimorphism of the human brain; behavioral neuroendocrinology findings that challenge the notion of genetically fixed, nonoverlapping, sexually dimorphic hormonal systems; psychological findings that highlight the similarities between men and women; psychological research on transgender and nonbinary individuals' identities and experiences; and developmental research suggesting that the tendency to view gender/sex as a meaningful, binary category is culturally determined and malleable. Costs associated with reliance on the gender binary and recommendations for future research, as well as clinical practice, are outlined. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

A Flat CHAT perspective on transliteracies development

Examining two longitudinal cases of writing development, we argue for a Flat CHAT (cultural-historical activity theory) perspective on transliteracies development that focuses attention on laminated assemblage—how times, spaces, artifacts, and people converge in complex and unfolding trajectories of becoming. The first case focuses on Nora, a post-doctorate biologist working on the behavioral neuroendocrinology of birds' sociality. Based on life-history and text-based interviews, observations, and texts reaching back to elementary school, the analysis explores how a laminated history of semiotic engagements have formed her ways of being-in-the-world as a biologist. Drawing on a larger connective ethnographic study of writing development, the second case focuses on the distributed becoming of an out-of-school organization. Based on interviews with focal young men from the larger study, their peers, and mentors; fieldnotes from two years of observations; and youths' compositions, the analysis traces genesis and disruption across time-space scales. In both cases, we see a host of becomings, where each element of the assemblage is deeply shaped by diverse histories, in intra-active flux (not finalized or isolated) in the moment, and spinning off, changed, into emergent futures.

A personal view on traits useful for success in science: Daniel S. Lehrman Award Lecture

As the 2018 recipient of the Daniel S. Lehrman Lifetime Achievement Award from the Society for Behavioral Neuroendocrinology, I was asked to give a short lecture in the Young Investigator Symposium of the combined Society for Behavioral Neuroendocrinology/International Congress on Neuroendocrinology meeting in Toronto. This lecture focused on one person's thoughts on what it takes to be successful in an academic science career. In this paper, I elaborate on success, on traits that may be useful for success in science, and on where the field of Behavioral Neuroendocrinology may be going.

Oxytocin receptor knockout prairie voles generated by CRISPR/Cas9 editing show reduced preference for social novelty and exaggerated repetitive behaviors

Behavioral neuroendocrinology has benefited tremendously from the use of a wide range of model organisms that are ideally suited for particular questions. However, in recent years the ability to manipulate the genomes of laboratory strains of mice has led to rapid advances in our understanding of the role of specific genes, circuits and neural populations in regulating behavior. While genome manipulation in mice has been a boon for behavioral neuroscience, the intensive focus on the mouse restricts the diversity in behavioral questions that can be investigated using state-of-the-art techniques. The CRISPR/Cas9 system has great potential for efficiently generating mutants in non-traditional animal models and consequently to reinvigorate comparative behavioral neuroendocrinology. Here we describe the efficient generation of oxytocin receptor (Oxtr) mutant prairie voles (Microtus ochrogaster) using the CRISPR/Cas9 system, and describe initial behavioral phenotyping focusing on behaviors relevant to autism. Oxtr mutant male voles show no disruption in pup ultrasonic vocalization, anxiety as measured by the open field test, alloparental behavior, or sociability in the three chamber test. Mutants did however show a modest elevation in repetitive behavior in the marble burying test, and an impairment in preference for social novelty. The ability to efficiently generate targeted mutations in the prairie vole genome will greatly expand the utility of this model organism for discovering the genetic and circuit mechanisms underlying complex social behaviors, and serves as a proof of principle for expanding this approach to other non-traditional model organisms.