Strong Competition Does Not Always Predict Play Asymmetry: The Case of South American Sea Lions (Otaria flavescens)

Although play fighting has been studied for over a century in both human and non‐human animals, quantitative data on marine mammals are still scarce. Here, we investigated play fighting in South American sea lions (Otaria flavescens), one of the most sexually dimorphic species with an extreme polygynous mating system, high levels of both intra‐ and inter‐sexual competition. All these features make South American sea lions a good model species to test some predictions on play fighting. Our results indicate play is restricted to juveniles, being inhibited among adults, and as to be expected in a species that shows a high degree of sexual dimorphism, it is mainly expressed in males. Even though playful interactions were punctuated by competitive behaviours, animals played in a highly symmetric way and were able to adjust their competitive playful interactions in a flexible manner and so reduce the risk of escalation to a minimum level. They were highly selective in their choice of playmates by limiting the number of players per session and playing more with age‐matched companions and friends. All these factors taken together are probably at the basis of the low risk of escalation recorded during the study. This result is predictive of a high ability and motivation of these animals to engage in play behaviour which can have a possible role not only in the acquisition of dominance status, but also in establishing and maintaining social relationships, an unexpected role in a so highly competitive species.     

Drosophila melanogaster females change mating behaviour and offspring production based on social context

In Drosophila melanogaster, biological rhythms, aggression and mating are modulated by group size and composition. However, the fitness significance of this group effect is unknown. By varying the composition of groups of males and females, we show that social context affects reproductive behaviour and offspring genetic diversity. Firstly, females mating with males from the same strain in the presence of males from a different strain are infecund, analogous to the Bruce effect in rodents, suggesting a social context-dependent inbreeding avoidance mechanism. Secondly, females mate more frequently in groups composed of males from more than one strain; this mitigates last male sperm precedence and increases offspring genetic diversity. However, smell-impaired Orco mutant females do not increase mating frequency according to group composition; this indicates that social context-dependent changes in reproductive behaviour depend on female olfaction, rather than direct male-male interactions. Further, variation in mating frequency in wild-type strains depends on females and not males. The data show that group composition can affect variance in the reproductive success of its members, and that females play a central role in this process. Social environment can thus influence the evolutionary process.     

Social ontogeny and behavioural diversity: consequences for Bighorn sheep Oris canadensis inhabiting desert and mountain environments

The development of social behaviour was studied in three natural populations of Bighorn sheep ( Ovis canadensis ). Individuals in all populations exhibited the same behavioural repertoires, but the utilization of specific behaviours among both infants and adults differed between populations. It was suggested that differences in the adult utilization of behaviour patterns result in part from behaviours used during infancy. In turn, the social and physical environments affected the development of subsequent behaviour patterns in sheep older than lambs.
Desert sheep (in southern California) used mounting behaviours often, presumably because they matured sexually at an early age. Sheep in a more northern environment (British Columbia) used different behaviour patterns more frequently perhaps as a consequence of interactions experienced in larger bands. Additionally, playful interactions were reduced due to physical hazards in the desert, but they were at least nine times as frequent in the northern study population. Play in large groups resulted in the utilization of more different kinds of behaviour patterns. These behavioural findings are interpreted ecologically as consequences of inhabiting environments that differ socially and physically.     

Schooling,dusk flight and dance: social organisations as amplifiers of individual quality?

One striking aspect of certain types of animal assemblages, for example fish schools or bird flocks, is the level of synchrony and spatial regularity that group members achieve. Although the evolutionary mechanisms leading to the formation of animal aggregations appear to be understood reasonably well, the evolution of spatial regularity and the high levels of synchrony that typify the groups in which certain animals move are less clear. Traditional explanations have generally focused on benefits gained during interspecific interactions, particularly the improvement of antipredator responses, or have suggested aero- or hydrodynamic advantages during locomotion. However, since the latter benefits of structural regularity may be largely rejected on theoretical grounds, and because many examples of spatially regular, synchronous groupings – such as dusk-flying flocks of some birds – may occur in the absence of predators, we suggest that these behaviours may not be explained solely in terms of locomotory efficiency or performance in predator-prey interactions. Instead, we suggest that the maintenance of regular spatial positions and the level of synchrony achieved within certain social groups may reveal honest information about an individual's neurosensory or locomotory performance, and that these behaviours may have evolved as amplifiers of individual quality. The evolution of such behaviour therefore need not have occurred as a result of interspecific interactions, but could have happened in the arena of conspecific evaluation.     

Playing it cool: Characterizing social play,bout termination,and candidate play signals of juvenile and infant Tibetan macaques(Macaca thibetana)

Play behaviors and signals during playful interactions with juvenile conspecifics are important for both the social and cognitive development of young animals.The social organization of a species can also influence juvenile social play. We examined the relationships among play behaviors, candidate play signals, and play bout termination in Tibetan macaques(Macaca thibetana) during juvenile and infant social play to characterize the species play style. As Tibetan macaques are despotic and live in groups with strict linear dominance hierarchies and infrequent reconciliation, we predicted that play would be at risk of misinterpretation by both the individuals engaged in the play bout and by those watching, possibly leading to injury of the players. Animals living in such societies might need to frequently and clearly signal playful intent to play partners and other group members to avoid aggressive outcomes. We gathered video data on 21 individually-identified juvenile and infant macaques(one month to five years of age)from the Valley of the Wild Monkeys, Mt. Huangshan,China. We used all-occurrence sampling to record play behaviors and candidate play signals based on an ethogram. We predicted that play groups would use multiple candidate play signals in a variety of contexts and in association with the number of audience members in proximity to the players and play bout length. In the 283 playful interactions we scored,juvenile and infant macaques used multiple body and facial candidate play signals. Our data showed that juvenile and infant Tibetan macaques use a versatile repertoire of play behaviors and signals to sustain play.     

The use of vocal signals in the social play of barbary macaques

Field studies in various species ofMacaca (Cercopithecidae) provided evidence for specific visual displays that typically accompany playful interactions. The aim of our study was to examine whether and when playing individuals would use auditory displays, i.e. vocalizations that often occur during social play as well. The study was conducted on a population of semi-free Barbary macaques (Macaca sylvanus) with a special focus on the composition and dynamics of playful wrestling (synonymous term: ‘rough-and-tumble play’). Analyses of dyadic encounters between subadult males allowed us to distinguish five types of playful behaviours and three types of vocalizations. The latter were clearly linked to encounters where effects of visual signals were impaired, e.g. during close body contact. During wrestling, vocalizations tended to increase in the beginning of an encounter, whereas the last seconds of wrestling often showed a decline in vocalization rate. Our results allowed us to conclude that these vocalizations may supplement or in many cases even substitute interactional effects of visual signals, e.g. the ‘play face.’     

A descriptive study of social development in the rat (Rattus norvegicus)

The social interactions between juvenile rat pups (Rattus norvegicus) were observed daily between 21 and 55 days of age. The data were analysed for developmental changes, sex differences, play-partner preferences, and dominance relations. The results revealed clear patterns in the development of sexual and agonistic behaviours in the rat. This development refers to changes in either the manner in which animals interact or the behaviours involved in social play, but not to the topography of specific behaviours. Development toward the adult pattern of social behaviour was temporally associated with the period of sexual maturation (i.e. between days 36 and 50).     

Human‐directed social behaviour in dogs shows significant heritability

Through domestication and co‐evolution with humans, dogs have developed abilities to attract human attention, e.g. in a manner of seeking assistance when faced with a problem solving task. The aims of this study were to investigate within breed variation in human‐directed contact seeking in dogs and to estimate its genetic basis. To do this, 498 research beagles, bred and kept under standardized conditions, were tested in an unsolvable problem task. Contact seeking behaviours recorded included both eye contact and physical interactions. Behavioural data was summarized through a principal component analysis, resulting in four components: test interactions, social interactions, eye contact and physical contact. Females scored significantly higher on social interactions and physical contact and age had an effect on eye contact scores. Narrow sense heritabilities (h²) of the two largest components were estimated at 0.32 and 0.23 but were not significant for the last two components. These results show that within the studied dog population, behavioural variation in human‐directed social behaviours was sex dependent and that the utilization of eye contact seeking increased with age and experience. Hence, heritability estimates indicate a significant genetic contribution to the variation found in human‐directed social interactions, suggesting that social skills in dogs have a genetic basis, but can also be shaped and enhanced through individual experiences. This research gives the opportunity to further investigate the genetics behind dogs' social skills, which could also play a significant part into research on human social disorders such as autism.     

An investigation of individual, age and sexual differences in the play of Rattus norvegicus (Mammalia: Rodentia)

Eight litters of Rattus norvegicus of the PVG/C hooded strain were observed in order to investigate the roles of individuality, sexuality and age in playful behaviour.
Three measures of social play, namely, initiations, numbers of play bouts and numbers of playful elements performed were all found to be positively correlated. The incidence of social and solitary play were also positively correlated so that a concept of "playfulness" appeared to be justifiable. Playfulness showed no correlation with the general level of non-playful locomotor activity.
Male rats were more playful than females but only minor sex-related differences were found to occur in the sequential organisation of play.
Older rats showed longer play bouts and a higher degree of temporal organisation. Considerable individual differences in playfulness were recorded and in single sex litters playful rats were found to play together.
The majority of opponents, in playfighting, alternated between the roles of attacker and defender.     

Affiliative function of the silent bared-teeth display in moor macaques (Macaca maurus): Further evidence for the particular status of sulawesi macaques

The silent bared-teeth display is described in a captive group of Moor macaques (Macaca maurus). It occurs with either closed or open mouth and is observed in both affinitive and playful interactions. It does not appear to convey specific information about the social status of partners and should be viewed as a signal advertising the emitter's peaceful intentions. This is consistent with what is known for other species of macaques on Sulawesi island.     

Play and social dynamics in a captive herd of gelada baboons (Theropithecus gelada)

Here, we provide quantitative data concerning adult and immature play distribution in geladas discussing the results in the light of their social dynamics. Sex differences in immature play does not seem to reflect sex differences in adult role; in fact, frequency, modality (C-play and LR-play), and motivation (measured by play initiations) to play did not vary according to the different sex-class combinations. The occurrence of adult-adult play suggests that geladas are characterized by an apparent more than actual rigidity in their social relationships and this is particularly evident in females which show peculiar traits in the use of social play. Specifically, adult females played with other adult females as much as with immature ones, thus suggesting their high motivation to engage in such interactions. This adult play behavioural pattern is typical of species showing relaxed/fluid more than despotic/rigid social relationships. Such assumption is also supported by the positive correlation observed between play and affinitive behaviours (grooming and agonistic support) found within each age-category (adult females and immature subjects). This evidence, together with the lack of correlation between aggressive contacts and social play, suggests that geladas use play for social assessment purposes and/or to increase their social affiliation levels.     

An investigation of playful behaviour in Rattus norvegicus and Mus musculus (Mammalia)

The playful behaviour of laboratory rats ( Rattus norvegicus ) was investigated in litters of five individuals with the mother present; parallel observations were made on mice ( Mus musculus ). Seven mixed litters containing four young rats and a young mouse fostered at birth were also observed.
Solitary play was recorded in both species and took a similar form but social play was only observed in rats. In rats, solitary play frequently preceeded social play.
The behavioural elements involved in the social play of Rattus norvegicus were described, and the majority of these were the playful equivalent of adult agonistic behaviour elements. These social play elements were found to be organized into definite sequences which differed from those of adult aggression. Each behavioural element was found to act as a social releaser.
Young mice did not respond playfully to social play from a rat litter mate; mice were less attractive to rats as playmates in comparison with fellow rats.     

Early play may predict later dominance relationships in yellow-bellied marmots (Marmota flaviventris)

Play has been defined as apparently functionless behaviour, yet since play is costly, models of adaptive evolution predict that it should have some beneficial function (or functions) that outweigh its costs. We provide strong evidence for a long-standing, but poorly supported hypothesis: that early social play is practice for later dominance relationships. We calculated the relative dominance rank by observing the directional outcome of playful interactions in juvenile and yearling yellow-bellied marmots (Marmota flaviventris) and found that these rank relationships were correlated with later dominance ranks calculated from agonistic interactions, however, the strength of this relationship attenuated over time. While play may have multiple functions, one of them may be to establish later dominance relationships in a minimally costly way.     

Social modulation of learned behavior by dopamine in the basal ganglia: Insights from songbirds

Dysfunction of the dopaminergic system leads to motor, cognitive, and motivational symptoms in brain disorders such as Parkinson’s disease. The basal ganglia (BG) are involved in sensorimotor learning and receive a strong dopaminergic signal, shown to play an important role in social interactions. The function of the dopaminergic input to the BG in the integration of social cues during sensorimotor learning remains however largely unexplored. Songbirds use learned vocalizations to communicate during courtship and aggressive behaviors. Like language learning in humans, song learning strongly depends on social interactions. In songbirds, a specialized BG–thalamo-cortical loop devoted to song is particularly tractable for elucidating the signals carried by dopamine in the BG, and the function of dopamine signaling in mediating social cues during skill learning and execution. Here, I review experimental findings uncovering the physiological effects and function of the dopaminergic signal in the songbird BG, in light of our knowledge of the BG–dopamine interactions in mammals. Interestingly, the compact nature of the striato-pallidal circuits in birds led to new insight on the physiological effects of the dopaminergic input on the BG network as a whole. In singing birds, D1-like receptor agonist and antagonist can modulate the spectral variability of syllables bi-directionally, suggesting that social context-dependent changes in spectral variability are triggered by dopaminergic input through D1-like receptors. As variability is crucial for exploration during motor learning, but must be reduced after learning to optimize performance, I propose that, the dopaminergic input to the BG could be responsible for the social-dependent regulation of the exploration/exploitation balance in birdsong, and possibly in learned skills in other vertebrates.     

Characterization of behavioural responses in different test contexts after a single social defeat in male golden hamsters (Mesocricetus auratus)

The objective of the present study was to characterize behavioural responses of male hamsters in each of three test contexts after they had experienced either a single social defeat or a neutral encounter. In experiment 1, hamsters were observed in a familiar social context (i.e., their home cages), and defeated males displayed different amounts of time and submissive behaviours towards a known opponent than a novel intruder, whereas males in the neutral-encounter groups did not show such differences. In experiment 2, in an unfamiliar social context (i.e., a Y-maze), defeated males generated submissive behaviours and fear memory towards a known opponent that they re-encountered 5-min and 24-h after the defeat. The formation of long-term memory was interrupted by an injection of anisomycin (210 mg/kg). In experiment 3, in a non-social, anxiogenic context, hamsters that had previously had different social experiences did not demonstrate additional anxiety in an elevated plus-maze, with the exception of males that had previously experienced repeated social defeats. Our data suggested that hamsters’ behavioural changes following defeat are context-dependent and stimulus-specific. The experience of a single social defeat is sufficient to regenerate submissive behaviours and fear memory when reencountering a known opponent.     

The coevolution of play and the cortico-cerebellar system in primates

Primates are some of the most playful animals in the natural world, yet the reason for this remains unclear. One hypothesis posits that primates are so playful because playful activity functions to help develop the sophisticated cognitive and behavioural abilities that they are also renowned for. If this hypothesis were true, then play might be expected to have coevolved with the neural substrates underlying these abilities in primates. Here, we tested this prediction by conducting phylogenetic comparative analyses to determine whether play has coevolved with the cortico-cerebellar system, a neural system known to be involved in complex cognition and the production of complex behaviour. We used phylogenetic generalised least squares analyses to compare the relative volume of the largest constituent parts of the primate cortico-cerebellar system (prefrontal cortex, non-prefrontal heteromodal cortical association areas, and posterior cerebellar hemispheres) to the mean percentage of time budget spent in play by a sample of primate species. Using a second categorical data set on play, we also used phylogenetic analysis of covariance to test for significant differences in the volume of the components of the cortico-cerebellar system among primate species exhibiting one of three different levels of adult-adult social play. Our results suggest that, in general, a positive association exists between the amount of play exhibited and the relative size of the main components of the cortico-cerebellar system in our sample of primate species. Although the explanatory power of this study is limited by the correlational nature of its analyses and by the quantity and quality of the data currently available, this finding nevertheless lends support to the hypothesis that play functions to aid the development of cognitive and behavioural abilities in primates.     

Affiliative,playful, and homosexual interactions of adult female lowland gorillas

Over a period of seven months four wild born adult female lowland gorillas were observed under semi-natural captive conditions. Thirty instances of contact interactions were recorded during 80 hours of observation. These interactions were segregated into categories operationally defined as connoting playful, homosexual, and affiliative activities. During observation periods the animals' labial tumescence was rated and the stage of the cycle was estimated. Metacommunicative signals were involved during playful interactions which resembled social rough-and-tumble play. Sexual interactions included partner positioning, thrusting, and usually were accomplished through ventral-ventral genital approximations. Sexual activity was also significantly related to cycle state. Contact interactions appeared to represent a specification and elaboration of a trend revealed through measures of proximity.     

Locomotor performance and social dominance in male Anolis cristatellus

The proximal mechanisms determining social dominance are not well understood. We used the highly territorial lizard A. cristatellus to test two main hypotheses: (1) that male social dominance is associated with locomotor abilities; (2) that locomotor abilities (maximal performance), as measured in the laboratory, are correlated with behaviour in the field. In the field, we recorded locomotor behaviours and assertion displays, then characterized microhabitat use and thermal relations. In the laboratory, we measured maximum sprint running speed, endurance and morphometric characters, and assessed dominance by pairing males of similar body size in an experimental arena. In 72 of 77 interactions, one lizard (the ‘winner’) was unequivocally determined to be dominant over the other (the ‘loser’). Winners performed more assertion displays than losers before capture and also had higher endurance in laboratory tests. Although contestants were matched for snout-vent length, winners had significantly deeper and wider heads. However, we found no significant differences in field locomotor behaviours, perch or thermal characteristics, head length, or maximal sprint speed. Our findings support those of previous studies, and extend them in several ways. This is the first demonstration that assertion displays in the field are related to both locomotor performance and laboratory-assessed social dominance. Locomotor performance may directly affect social dominance by allowing some males to perform better in dyadic interactions. Alternatively, both locomotor performance and social dominance may be linked to a common underlying mechanism, such as variation in hormone levels, which are known to affect aggression, locomotor performance and morphology.     

同胞侵犯对猞猁幼仔行为的影响

研究了非自由生活的4窝11只猞猁幼仔的玩耍行为、亲和及入侵关系的变化.观察到36-57日龄幼仔的同胞打斗行为,这些打斗减少了这些幼仔之间游戏性接触的频次.这个"打斗期"与游戏行为和食性的变化同步发生.打斗后,幼仔游戏性接触的非对称性以及同胞之间的偏爱更加清楚.这些打斗导致了不同窝的幼仔之间等级结构的建立.