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Although cooperating individuals may gain benefits, they risk being cheated by their allies. Therefore, to understand the persistence of cooperation by mutual benefits, it is important to document both the potential benefits of cooperation and the risks of cheating. We experimentally examined evidence of cooperation in the ocellated wrasse (Symphodus ocellatus), a fish species with males that exercise three reproductive strategies: satellite and sneaker males attempt to parasitically spawn in the nests of nesting males. Nesting males have been hypothesized to tolerate satellites as a result of satellite aid in defense against sneaker males and help in courting females. Here, we provide evidence that cooperation between reproductive competitors may arise in response to sperm competition from other males. After we experimentally removed satellites, nesting males had more sneakers at their nest and experienced a higher rate of sneaking when they spawned (increased sperm competition risk). When we experimentally distracted nesting males to prevent their acting aggressively toward satellites, satellites spawned with females and ate eggs in the nest. Our results suggest that nesting males and satellites gain fitness benefits of reduced direct reproductive and sperm competition through cooperation. However, nesting males risk losing paternity, while satellites experience increased aggression and must invest in aggression themselves (i.e., they act aggressively toward sneakers).
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When reproductive competitors tolerate or cooperate with one another, they may gain particular benefits, such as collectively guarding resources or attracting mates. Shared resources may be those essential to reproduction, such as a breeding site or nest. Using the tessellated darter, a species where males but not females compete over potential nest sites, we examined site use and sharing under controlled conditions of differing competitor density. Sharing was observed even when competitor density was low and individuals could have each occupied a potential nest site without same-sex sharing. Males were more likely to share a nest site with one other when the difference in size between them was larger rather than smaller. There was no evidence that female sharing was dependent on their relative size. Fish were generally more likely to use and share larger sites, in accordance with the greater relative surface area they offered. We discuss how one or both sharing males may potentially benefit, and how male sharing of potential nest sites could relate to female mating preferences. Tessellated darter males are known to provide alloparental care for eggs but this occurs without any social contact between the alloparent and the genetic father of the young. Thus, the suggestion that they may also share sites and maintain social contact with reproductive competitors highlights the importance of increased focus on the potential complexity of reproductive systems.
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Agreement on the causes of sexual offending has eluded scholars because of an emphasis on studying proximate causes—as with crime in general, there are numerous variables that correlate with sexual offending. A more consilient view of sexual offending includes an evolutionary approach that attempts to answer ultimately why such behavior exists. To guide this research, Camilleri (The Oxford handbook of evolutionary perspectives on violence, homicide, and war, New York, 173–196) proposed a typology of sexual offenders that classifies offending based on two dimensions: (1) whether the behavior is an adaptation, by-product, or disorder and (2) if the mechanism is obligate or facultative. Categories resulting from this typology seem to capture the variability of sexual offending behavior. This chapter mostly reviews evidence of how sexual offending could function as an adaptation, draws links from nonhuman behavior literature, and briefly reviews alternative explanations, including by-products and disorders.
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While extensive research has focused on how social interactions evolve, the fitness consequences of the neuroendocrine mechanisms underlying these interactions have rarely been documented, especially in the wild. Here, we measure how the neuroendocrine mechanisms underlying male behaviour affect mating success and sperm competition in the ocellated wrasse (Symphodus ocellatus). In this species, males exhibit three alternative reproductive types. “Nesting males” provide parental care, defend territories and form cooperative associations with unrelated “satellites,” who cheat by sneaking fertilizations but help by reducing sperm competition from “sneakers” who do not cooperate or provide care. To measure the fitness consequences of the mechanisms underlying these social interactions, we used “phenotypic engineering” that involved administering an androgen receptor antagonist (flutamide) to wild, free-living fish. Nesting males treated with flutamide shifted their aggression from sneakers to satellite males and experienced decreased submissiveness by sneaker males (which correlated with decreased nesting male mating success). The preoptic area (POA), a region controlling male reproductive behaviours, exhibited dramatic down-regulation of androgen receptor (AR) and vasotocin 1a receptor (V1aR) mRNA following experimental manipulation of androgen signalling. We did not find a direct effect of the manipulation on male mating success, paternity or larval production. However, variation in neuroendocrine mechanisms generated by the experimental manipulation was significantly correlated with changes in behaviour and mating success: V1aR expression was negatively correlated with satellite-directed aggression, and expression of its ligand arginine vasotocin (AVT) was positively correlated with courtship and mating success, thus revealing the potential for sexual selection on these mechanisms. © 2018 John Wiley & Sons Ltd
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The molecular mechanisms underlying phenotypic plasticity are not well understood. Identifying mechanisms underlying alternative reproductive tactics (ARTs) in species for which the behavioural and fitness consequences of this variation are well characterized provides an opportunity to integrate evolutionary and mechanistic understanding of the maintenance of variation within populations. In the ocellated wrasse Symphodus ocellatus, the behavioural phenotypes of three distinct male morphs (sneakers, satellites and nesting males), which arise from a single genome, have been thoroughly characterized. To determine the neuroendocrine and genomic mechanisms associated with discrete phenotypic variation and ARTs in S. ocellatus in their natural environment, we constructed a whole-brain de novo transcriptome and compared global patterns of gene expression between sexes and male morphs. Next, we quantified circulating cortisol and 11-ketotestosterone (11-kt), mediators of male reproductive behaviours, as well as stress and gonadal steroid hormone receptor expression in the preoptic area, ventral subpallial division of the telencephalon and dorsolateral telencephalon, critical brain regions for social and reproductive behaviours. We found higher levels of 11-kt in nesting males and higher levels of cortisol in sneaker males relative to other male morphs and females. We also identified distinct patterns of brain region-specific hormone receptor expression between males such that most hormone receptors are more highly expressed in satellites and nesting males relative to sneakers and females. Our results establish the neuroendocrine and molecular mechanisms that underlie ARTs in the wild and provide a foundation for experimentally testing hypotheses about the relationship between neuromolecular processes and reproductive success. Copyright © 2016 John Wiley & Sons Ltd.
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Due to the difficulty of simultaneously assessing variation in individual physiology, behaviour and fitness, we often know little about the mechanistic basis of life-history trade-offs and fitness variation. It is similarly challenging to examine how physiological variation in one individual has cascading fitness consequences for others in the social environment. Using a wild-living fish (ocellated wrasse, Symphodus ocellatus), we manipulated a neuropeptide pathway associated with courtship, aggression and parental care in vertebrates (arginine vasotocin, AVT) and directly examined the behavioural and fitness consequences. Nesting males injected with the AVT antagonist increased their paternal care, resulting in increased hatching of offspring and increased reproductive success of all individuals that mated at his nest. By directly examining physiology, behaviour and reproductive success, we revealed how a small change in individual physiology has clear and direct fitness consequences for multiple individuals. © 2019 The Association for the Study of Animal Behaviour
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In species with internal fertilization, females can favour certain males over others, not only before mating but also within the female's reproductive tract after mating. Here, we ask whether such directional post-mating (that is, cryptic) female mate choice can also occur in species with external fertilization. Using an in vitro sperm competition experiment, we demonstrate that female ovarian fluid (ovarian fluid) changes the outcome of sperm competition by decreasing the importance of sperm number thereby increasing the relative importance of sperm velocity. We further show that ovarian fluid does not differentially affect sperm from alternative male phenotypes, but generally enhances sperm velocity, motility, straightness and chemoattraction. Under natural conditions, female ovarian fluid likely increases the paternity of the preferred parental male phenotype, as these males release fewer but faster sperm. These results imply females have greater control over fertilization and potential to exert selection on males in species with external fertilization than previously thought possible.
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Discrete variation in reproductive behavior and physiology is observed in diverse taxa. Although it is known that most within-sex alternative reproductive tactics arise as a consequence of phenotypic plasticity, relatively little is known about differential neural gene expression among plastic alternative reproductive phenotypes. In the ocellated wrasse Symphodus ocellatus, males exhibit one of three alternative tactics (nesting, satellite, and sneaker) within a reproductive season, but switch tactics between years. Satellites and sneakers spawn parasitically in dominant (nesting) males' nests, but only nesting males provide parental care. Nesting and satellite males show transient cooperative defense of nests against sneakers. Here, we analyze circulating sex steroid hormone levels and neural gene expression profiles in these three male phenotypes and in females. 11-ketotestosterone (but not testosterone) was highest in nesting males, while estradiol was highest in females. Brain transcriptomes of satellites and females were most similar to each other and intermediate to nesting and sneaker males. Sneakers showed more total expression differences, whereas nesting males showed higher magnitude expression differences. Our findings reveal the surprising extent to which neural gene expression patterns vary across reproductive tactics that vary in a number of social traits, including aggression, territoriality, and cooperation, providing important insights into the molecular mechanisms that may underlie variation in cooperative and reproductive behavior. © 2014 Blackwell Verlag GmbH.
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Male–male competition is a well-known driver of reproductive success and sexually selected traits in many species. However, in some species, males work together to court females or defend territories against male competitors. Dominant (nesting) males sire most offspring, but subordinate (satellite) males are better able to obtain fertilizations relative to unpartnered males. Because satellites only gain reproductive success by sneaking, there has been much interest in identifying the mechanisms enforcing satellite cooperation (defense) and reducing satellite sneaking. One such potential mechanism is outside competition: unpartnered satellites can destabilize established male partnerships and may force partnered satellites to restrain from cheating to prevent the dominant male from replacing them with an unpartnered satellite. Here, we manipulated perceived competition in the Mediterranean fish Symphodus ocellatus by presenting an “intruding” satellite male to established nesting and satellite male pairs. Focal satellite aggression to the intruder was higher when focal satellites were less cooperative, suggesting that satellites increase aggression to outside competitors when their social position is less stable. In contrast, nesting male aggression to the intruder satellite increased as spawning activity increased, suggesting that nesting males increase their defense toward outside competitors when their current relationship is productive. We found no evidence of altered spawning activity or nesting/satellite male interactions before and after the presentation. These results collectively suggest that response to outside competition is directly linked to behavioral dynamics between unrelated male partners and may be linked to conflict and cooperation in ways that are similar to group-living species.
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Background:Little is known about the adoption by athletic administrators (AAs) of exertional heat illness (EHI) policies, and the corresponding facilitators and barriers of such policies within high school athletics. This study describes the adoption of comprehensive EHI policies by high school AAs and explores factors influencing EHI policy adoption.Hypothesis:We hypothesized that <50% of AAs would report adoption of an EHI policy, and that the most common facilitator would be access to an athletic trainer (AT), whereas the most common barrier would be financial limitations.Study Design:Cross-sectional.Level of Evidence:Level 4.Methods:A total of 466 AAs (82.4% male; age, 48 ± 9 years) completed a validated online survey to assess EHI prevention and treatment policy adoption (11 components), as well as facilitators and barriers to policy implementation. Access to athletic training services was ascertained by matching the participants? zip codes with the Athletic Training Locations and Services Project. Policy adoption, facilitators, and barriers data are presented as summary statistics (proportions, interquartile range (IQR)). A Welch t test evaluated the association between access to athletic training services and EHI policy adoption.Results:Of the AAs surveyed, 77.9% (n = 363) reported adopting a written EHI policy. The median of EHI policy components adopted was 5 (IQR = 1,7), with only 5.6% (n = 26) of AAs reporting adoption of all policy components. AAs who had access to an AT (P = 0.04) were more likely to adopt a greater number of EHI-related policies, compared with those without access to an AT. An AT employed at the school was the most frequently reported facilitator (36.9%).Conclusion:Most AAs reported having written EHI policy components, and access to an AT resulted in a more comprehensive policy.Clinical Relevance:Employment of an AT within high school athletics may serve as a vital component in facilitating the adoption of comprehensive EHI policies.
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Early life neglect increases risk for the development of psychopathologies during childhood and adulthood, including depression and anxiety disorders. We recently reported epigenetic changes in DNA derived from saliva in three genes predicted depression in a cohort of maltreated children: DNA-binding protein inhibitor ID-3 (ID3), Glutamate NMDA Receptor (GRIN1), and Tubulin Polymerization Promoting Protein (TPPP). To validate the role of these genes in depression risk, secondary analyses were conducted of gene expression data obtained from medial prefrontal cortex (mPFC) tissue of mice subjected to a model of maternal neglect which included maternal separation and early weaning (MSEW). Anxiety and depression-like phenotype data derived using the elevated plus maze (EPM) and forced swimming test (FST), respectively, were also available for secondary analyses. Behavioral tests were conducted in MSEW and control adult male mice when they were between 65 and 80days old. ID3, GRIN1 and TPPP gene expression in the mPFC were found to significantly predict behavioral differences in the EPM and FST. These results further support the role of these genes in the etiology of depressive and anxiety phenotypes following early life stress. Copyright © 2016. Published by Elsevier B.V.
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Context: The Athletic Training Locations and Services (ATLAS) Annual Report suggested that athletic trainer (AT) employment status differed based on geographic locale. However, the influence of geographic locale and school size on AT employment is unknown. Objective: To determine if differences existed in the odds of having AT services by locale for public and private schools and by student enrollment for public schools. Design: Cross-sectional study. Setting: Public and private secondary schools with athletics programs. Patients or Other Participants: Data from 20 078 US public and private secondary schools were obtained. Main Outcome Measures(s): Data were collected by the ATLAS Project. Athletic trainer employment status, locale (city, suburban, town, or rural) for public and private schools, and school size category (large, moderate, medium, or small) only for public schools were obtained. The employment status of ATs was examined for each category using odds ratios. Logistic regression analysis produced a prediction model. Results: Of the 19 918 public and private schools with available AT employment status and locale, suburban schools had the highest access to AT services (80.1%) with increased odds compared with rural schools (odds ratio = 3.55 [95% CI = 3.28, 3.85]). Of 15 850 public schools with known AT employment status and student enrollment, large schools had the highest rate of AT services (92.1%) with nearly 18.5 times greater odds (odds ratio = 18.49 [95% CI = 16.20, 21.08]) versus small schools. The logistic model demonstrated that the odds of access to an AT increased by 2.883 times as the school size went up by 1 category. Conclusions: Nationally, suburban schools and large public schools had the greatest access to AT services compared with schools that were in more remote areas and with lower student enrollment. These findings elucidate the geographic locales and student enrollment levels with the highest prevalence of AT services. Ó by the National Athletic Trainers’ Association, Inc
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Several predictions of sperm competition theory are not well supported empirically. One potential reason is that most current theory and empirical research ignore how the social environment influence the temporal dynamics of mating. We propose that understanding these dynamics is key to understanding sexual selection and improving the predictive power of theory. To demonstrate the importance of these dynamics, we quantify how males’ social role, interactions among males, and current social environment influence the timing of mating in Symphodus ocellatus, a species with three alternative male reproductive tactics. Nesting males spawn synchronously with females; sneakers and satellites sneak-spawn with some time delay. Satellites also cooperate with nesting males. We found that satellites have shorter sneak-spawning delays than sneakers, a benefit of their cooperation with nesting males. Sneak-spawning delays decreased with increasing nest activity for sneakers but not for satellites, suggesting that sneakers may benefit from increased sperm competition intensity. Current sperm competition models ignore this potential benefit, which may be why the prediction that males should decrease investment when sperm competition involves more than two males is not well supported. Our study provides insight into mechanisms that drive variation in the timing of spawning, which could explain mismatches between theoretical and empirical results.
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This paper describes how individual female ocellated wrasse Symphodus ocellatus distribute their spawning among males and nests in space and time. It is based on previously collected genetic data of larvae from ten different nests (used to reconstruct half and full-sibling groupings both within and among nests on multiple days) and behavioural data of marked females across the reproductive season. Both the genetic analyses and behavioural observations confirm that female S. ocellatus intentionally engage in multiple mating, by repeatedly spawning at the same nest on different days and at several different nests (up to 12 spawning events over 3 weeks), leading to mixed paternity among her young. The main benefit of such high and intentional multiple mating is probably insurance against brood failure due to nest predation, desertion or poor paternal care by the male. These findings reveal that even in systems where females attempt to avoid male-controlled mixed paternity, they may still engage in intentional multiple mating due to these potential benefits. © 2018 The Fisheries Society of the British Isles
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Sexual selection arising from sperm competition has driven the evolution of immense variation in ejaculate allocation and sperm characteristics not only among species, but also among males within a species. One question that has received little attention is how cooperation among males affects these patterns. Here we ask how male alternative reproductive types differ in testes size, ejaculate production, and sperm morphology in the ocellated wrasse, a marine fish in which unrelated males cooperate and compete during reproduction. Nesting males build nests, court females and provide care. Sneaker males only “sneak” spawn, while satellite males sneak, but also help by chasing away sneakers. We found that satellite males have larger absolute testes than either sneakers or nesting males, despite their cooperative role. Nesting males invested relatively less in testes than either sneakers or satellites. Though sneakers produced smaller ejaculates than either satellite or nesting males, we found no difference among male types in either sperm cell concentration or sperm number, implying sneakers may produce less seminal fluid. Sperm tail length did not differ significantly among male types, but sneaker sperm cells had significantly larger heads than either satellite or nesting male sperm, consistent with past research showing sneakers produce slower sperm. Our results highlight that social interactions among males can influence sperm and ejaculate production. © 2021 by the authors. Licensee MDPI, Basel, Switzerland.
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Introduction/Purpose Athletic administrators (AA) in U.S. high schools are uniquely positioned to provide guidance and endorsement for the prevention and management of common sport-related catastrophic injuries, such as the development of a spinal cord injury management (SCIM) policy. The purpose of this project was to evaluate AA’s knowledge of comprehensive SCIM policies in U.S. high schools. Secondarily, we aimed to investigate the factors, facilitators, and barriers to the development and adoption of comprehensive policies. Methods An online questionnaire was distributed to 6423 AA working in U.S.-based high school athletics, with 366 included in the final data analysis. The questionnaire evaluated AA’s knowledge of comprehensive SCIM policies addressing components of adoption, documentation, and communication. The questionnaire was developed based on the 2002 “National Athletic Trainers’ Association Position Statement: Acute Management of the Cervical Spine-Injured Athlete.” The questionnaire also inquired about facilitators and barriers to developing and adopting comprehensive policies. Proportions were calculated for policy adoption, and independent-samples t-tests evaluated the influence of athletic training services on comprehensive SCIM policies. Results About half of AA (49.2%, n = 180/366) reported having all recommended components of a written SCIM policy. AA who had access to an athletic trainer (yes = 62.9%, no = 18.1%, unknown = 19%) were more likely to have a comprehensive SCIM policy compared with those without access (80.6% vs 19.4%, χ21 = 7.091, P = 0.008, prevalence ratio = 1.24, 95% confidence interval = 1.03–1.50). The most commonly reported facilitator (49.9%) was “having a medical professional at the school,” and the main barrier (30.2%) was “my school would need more information, resources, assistance, etc.” Conclusions The findings identify areas for improvement in supporting the health and safety of children in high school participating in interscholastic athletics. Future dissemination and implementation research should develop strategies tailored to individual school community and need to improve SCIM policy adoption and implementation.
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Child neglect is the most prevalent form of child maltreatment in the United States, and poses a serious public health concern. Children who survive such episodes go on to experience long-lasting psychological and behavioral problems, including higher rates of post-traumatic stress disorder symptoms, depression, alcohol and drug abuse, attention-deficit/hyperactivity disorder, and cognitive deficits. To date, most research into the causes of these life-long problems has focused on well-established targets such as stress responsive systems, including the hypothalamus-pituitary-adrenal axis. Using the maternal separation and early weaning model, we have attempted to provide comprehensive molecular profiling of a model of early-life neglect in an organism amenable to genomic manipulation: the mouse. In this article, we report new findings generated with this model using chromatin immunoprecipitation sequencing, diffuse tensor magnetic resonance imaging, and behavioral analyses. We also review the validity of the maternal separation and early weaning model, which reflects behavioral deficits observed in neglected humans including hyperactivity, anxiety, and attentional deficits. Finally, we summarize the molecular characterization of these animals, including RNA profiling and label-free proteomics, which highlight protein translation and myelination as novel pathways of interest.
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Background: Approximately 10 to 15% of women consume alcohol (ethanol [EtOH]) during pregnancy in the United States. Even low amounts of EtOH consumption during pregnancy can elicit long-term consequences. Prenatal experience with as few as 3 drinks has been associated with increase problem drinking in adulthood. Such effects are corroborated in rodents; however, the underlying neural adaptations contributing to this effect are not clear. In the current set of experiments, we investigated whether changes in EtOH responding following prenatal EtOH exposure involved kappa opioid receptor activation and expression. Methods: Sprague-Dawley rats were prenatally exposed to low levels of alcohol (1.0 g/kg) during late gestation (gestational days 17 to 20 [GD17-20]) via intragastric intubation of pregnant dams. Following birth, EtOH intake, kappa- and mu-opioid-induced place conditioning, and kappa opioid receptor expression in mesolimbic brain regions were assessed in infant rats (postnatal days 14 to 15 [PD14-15]) that were offspring of dams given EtOH, vehicle, or untreated, during pregnancy. Results: Animals exposed to prenatal alcohol drank more alcohol later in life and exhibited significant changes in the kappa opioid system. While control subjects found kappa opioid activation aversive, animals exposed to EtOH prenatally exhibited either no aversion or appetitive responding. Further analysis revealed that synaptosomal kappa opioid receptor expression was significantly decreased in brain areas implicated in responding to EtOH. Conclusions: Overall, these data suggest that prenatal EtOH affects kappa opioid function and expression and that these changes may be involved in increased drinking later in life. © 2014 by the Research Society on Alcoholism.
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Fish exhibit impaired sensory function and altered behaviour at levels of ocean acidification expected to occur owing to anthropogenic carbon dioxide emissions during this century. We provide the first evidence of the effects of ocean acidification on reproductive behaviour of fish in the wild. Satellite and sneaker male ocellated wrasse (Symphodus ocellatus) compete to fertilize eggs guarded by dominant nesting males. Key mating behaviours such as dominant male courtship and nest defence did not differ between sites with ambient versus elevated CO2 concentrations. Dominant males did, however, experience significantly lower rates of pair spawning at elevated CO2 levels. Despite the higher risk of sperm competition found at elevated CO2, we also found a trend of lower satellite and sneaker male paternity at elevated CO2 Given the importance of fish for food security and ecosystem stability, this study highlights the need for targeted research into the effects of rising CO2 levels on patterns of reproduction in wild fish. Copyright © 2016 The Author(s).
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