Effects of Social Environment on Play in Squirrel Monkeys: Resolving Harlequin's Dilemma

Yearling-male squirrel monkeys (Saimiri sciureus) were paired with play partners either matched or different from them in age and/or sex. The frequency of play was maintained at equally high levels with both matched and mismatched partners. However, play between mismatched partners was marked by an unusually high incidence of nondirectional wrestling, a less threatening form of playfighting, and an increase in role reversal, suggesting that youngsters had considerable flexibility in the use of alternate strategies to sustain play activity with partners who were atypical for them. These results also attest to the robustness of play and its importance in the behavioral repertoire. The technique of restricting youngsters to atypical play companions is suggested as an effective means of manipulating play, with minimal disruption of other social experiences, for long-term studies of the developmental impact of social play.     

Sex differences in the termination of social play in cats

Sixteen measures of social play between kittens were found to decrease significantly between 12 and 16 weeks of age. Behaviour was little influenced by the presence of the mother during this time. Although males' rates of behaviour did not differ from females' rates when the sexes were compared regardless of group composition, males from all-male groups played at higher rates than females from all-female groups. Females' rates of behaviour declined as the number of male companions decreased. Males' behaviour was little influenced by the number of females in the group. Females with male companions behaved more like males than did females with no male companions, but this was not because they directed behaviour to brothers at higher rates than to sisters.     

Social development during the first twenty weeks in the cotton-top tamarin (Saguinus o. oedipus)

Fourteen infant cotton-top tamarins from five captive family groups were studied during the first 20 weeks of life. We examined the roles of parents and of siblings of different ages as caregivers and as social companions for the infants, as well as the effects of group composition and group size on these roles. Parents and adult siblings played similar active roles in infant caregiving during the first few weeks after birth, with males showing a greater involvement than females. In groups without other offspring the two parents showed equal infant caregiving behaviour, whreas fathers carried more than mothers did in groups with offspring present. Although infants received more caregiving from males, they showed a preference for contact and proximity with their mothers. No sex differences were found in infant behaviour or in the direction of behaviour by caregivers toward infants. Twins showed greater amounts of both social and solitary play in weeks 15–20 than did singletons, and there was no effect of the number of older siblings on the amount of play shown by an infant.     

Diving-for-Food: A New Model to Assess Social Roles in a Group of Laboratory Rats

In an experimental situation called “diving for food”, groups of laboratory rats are tested in an aquarium where they have to dive and swim under water to reach the single source of food located at the other end. A behavioural differentiation appears: some rats — the carriers — dive to get food and others — non-carriers — stay in the cage and feed by stealing. We examine whether carrier and non-carrier profiles can be considered as social roles, defined as supraindividual features dependent on the social context. Carrier/non-carrier differentiation resulted in all groups tested. Individually tested, almost all rats can get food by diving and swimming. Differentiation also occurred in groups that had been previously trained alone in the device, and in groups whose members had all been carriers or non-carriers exclusively in a preliminary stage. As the access-to-food behaviour of a rat having to cope with the diving-for-food situation is settled by its social environment, we consider that the present experimental model is promising for the study of interactions between the individual and the social structure.     

Zebra finches are bolder in an asocial, rather than social, context

Despite an expanding interest in animal personalities, the influence of social interactions and sex differences on individual differences in behaviour remains poorly understood. Using the social zebra finch (Taeniopygia guttata), we tested for behavioural differences in exploration of a novel environment and objects, between individuals of both sexes in relation to a social context; the presence of three male companions, three female companions or no companion birds. We predicted that the presence of conspecific companions should result in focal birds reacting to novelty by exploring more extensively because the companion birds contribute to anti-predator vigilance behaviour and because social isolation often causes behavioural inhibition in social species. We found that exploratory behaviour of focal individuals was significantly reduced in the presence of conspecific companions, irrespective of the companion's sex. Moreover, we found a weak trend towards females being more exploratory than males, irrespective of the social context. These results demonstrate the importance of considering the social context in animal personality studies and of exploring sex differences in personalities.     

Cooperation among Norway Rats: The Importance of Visual Cues for Reciprocal Cooperation,and the Role of Coercion

Some animals reciprocate help, but the underlying proximate mechanisms are largely unclear. Norway rats (Rattus norvegicus) have been shown to cooperate in a variant of the iterated prisoner's dilemma paradigm, yet it is unknown which sensory modalities they use. Visual information is often implicitly assumed to play a major role in social interactions, but primarily nocturnal species such as Norway rats may rely on different cues when deciding to reciprocate received help. We used an instrumental cooperative task to compare the test rats' propensity to reciprocate received help between two experimental conditions, with and without visual information exchange between social partners. Our results show that visual information is not required for reciprocal cooperation among social partners because even when it was lacking, test rats provided food significantly earlier to partners that had helped them to obtain food before than to those that had not done so. The mean decision speed did not differ between the two experimental conditions, with or without visual information. Social partners sometimes showed aggressive behaviour towards focal test individuals. When including this in the analyses to assess the possible role of aggression as a trigger of cooperation, aggression received from cooperators apparently reduced the cooperation propensity, whereas aggression received from defectors increased it. Hence, in addition to reciprocity, coercion seems to provide additional means to generate altruistic help in Norway rats.     

Learning rules for social foragers: implications for the producer-scrounger game and ideal free distribution theory

In population games, the optimal behaviour of a forager depends partly on courses of action selected by other individuals in the population. How individuals learn to allocate effort in foraging games involving frequency-dependent payoffs has been little examined. The performance of three different learning rules was investigated in several types of habitats in each of two population games. Learning rules allow individuals to weigh information about the past and the present and to choose among alternative patterns of behaviour. In the producer-scrounger game, foragers use producer to locate food patches and scrounger to exploit the food discoveries of others. In the ideal free distribution game, foragers that experience feeding interference from companions distribute themselves among heterogeneous food patches. In simulations of each population game, the use of different learning rules induced large variation in foraging behaviour, thus providing a tool to assess the relevance of each learning rule in experimental systems. Rare mutants using alternative learning rules often successfully invaded populations of foragers using other rules indicating that some learning rules are not stable when pitted against each other. Learning rules often closely approximated optimal behaviour in each population game suggesting that stimulus-response learning of contingencies created by foraging companions could be sufficient to perform at near-optimal level in two population games.     

A preliminary study on the function of food begging in Sichuan snub-nosed monkeys (Rhinopithecus roxellana): challenge to begging for nutritional gain

Several hypotheses have been developed to explain what benefits a donor may gain from sharing food with another individual, with nutritional gain assumed to be the sole benefit for the beggar. Recently, it has been proposed that begging behaviour serves a social function in non-human primates. In this study, the nutritional-gain assumption was again challenged based on observations on a captive group of Sichuan snub-nosed monkeys (Rhinopithecus roxellana), or golden snub-nosed monkeys. The major findings from this study are that (1) beggars sometimes left their own branches or passed by available branches to beg for similar food from other individuals, (2) beggars occasionally ignored branches that were acquired by begging and (3) food begging occurred more frequently in the all-male unit after the social rank had changed between 2 individuals in this unit. Overall, these preliminary findings suggest that some begging behaviours in captive golden snub-nosed monkeys were not driven by nutritional gain only; instead, we propose that these begging behaviours could be interpreted as attempts at deriving social benefits.     

Development of Social Behaviour in Four Litters of Dogs (Canis familiaris)

The development of social behaviour in 4 litters of dogs was observed without interfering with the puppies from birth to 8 weeks of age. Direct and continuous observation was combined with video recording. Three of the litters were observed during one session of 2 h once a week, and the fourth litter during one session of 40 min twice a week. Social interactions were divided into 1) investigation of litter mates (licking, sniffing or investigating orally), 2) social play, 3) and interactions in which agonistic elements (dominance postures, threats, bites or submission) were displayed. The different forms of social interactions appeared for the first time when the puppies were between 14 and 21 days of age. Social investigation appeared first and was followed by play and agonistic interactions. From week 5, differences between the puppies in the tendency to initiate social play and agonistic interactions emerged. Generally, within the litters individual differences were consistent over weeks 6-8 (positive correlations between weeks), whereas the tendency in the puppies during these weeks were negatively correlated with those of week 3 (play) or weeks 3 and 4 (agonistic interactions), indicating a rebound effect for both play and agonistic behaviour. No significant correlations, however, were found for social investigation. More often than expected males played or engaged in agonistic with other males, whereas these behaviours occurred less often than expected between females. Both males and females, however, preferred male partners for agonistic interactions. No sex differences were found in the direction of social investigation. Agonistic behaviour was often responded to by play and play was often responded to by agonistic behaviour, and the results indicated that before 8 weeks of age differences in social behaviour between the puppies were already established.     

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.     

Physiological state influences the social interactions of two honeybee nest mates

Physiological state profoundly influences the expression of the behaviour of individuals and can affect social interactions between animals. How physiological state influences food sharing and social behaviour in social insects is poorly understood. Here, we examined the social interactions and food sharing behaviour of honeybees with the aim of developing the honeybee as a model for understanding how an individual's state influences its social interactions. The state of individual honeybees was manipulated by either starving donor bees or feeding them sucrose or low doses of ethanol to examine how a change in hunger or inebriation state affected the social behaviours exhibited by two closely-related nestmates. Using a lab-based assay for measuring individual motor behaviour and social behaviour, we found that behaviours such as antennation, willingness to engage in trophallaxis, and mandible opening were affected by both hunger and ethanol intoxication. Inebriated bees were more likely to exhibit mandible opening, which may represent a form of aggression, than bees fed sucrose alone. However, intoxicated bees were as willing to engage in trophallaxis as the sucrose-fed bees. The effects of ethanol on social behaviors were dose-dependent, with higher doses of ethanol producing larger effects on behaviour. Hungry donor bees, on the other hand, were more likely to engage in begging for food and less likely to antennate and to display mandible opening. We also found that when nestmates received food from donors previously fed ethanol, they began to display evidence of inebriation, indicating that ethanol can be retained in the crop for several hours and that it can be transferred between honeybee nestmates during trophallaxis.     

Affiliation and social discrimination produced by brief exposure in day-old domestic chicks.

Discrimination between individual strangers and companions was examined in day-old domestic chicks. In one experiment, pecking rates at companions and strangers were observed in pairwise bouts after 16 hr of cohabitation. The discriminability of strangers and companions was varied by means of pre-hatch colouring. Reliable discriminations between individual strangers and companions emerged as early as the first minute of the encounter. Discriminative cues provided by artificial colouring were found not to be necessary in establishing social discrimination. In a second experiment, undyed chicks were housed in pairs for 1, 4 or 16 hr. Half of the pairs lived in cages that separated companions by a wire screen, and half were housed in undivided cages. Observations of pecking in four-way bouts confirmed previous findings and demonstrated that the opportunity to peck during exposure may be a necessary condition in producing social discrimination. Antecedent conditions that lead to the development of affiliative bonds simultaneously appear to establish social discrimination.     

Effects of physical and social experiences and octopamine receptor agonist on fighting behavior of male crickets Velarifictorus aspersus (Orthoptera: Gryllidae)

Fighting commonly occurs among animals and is very important for resolving conflicts between conspecific individuals over limited resources. The plasticity of fighting strategies and neurobiological mechanisms underlying fighting behavior of insects are not fully understood. In the present study, we examined whether physical and social experiences affected the aggressiveness of males of the cricket Velarifictorus aspersus Walker, and whether an octopamine (OA) receptor agonist could affected the aggressiveness of males exposed to different experiences. We found that flight and winning a fight significantly enhanced male aggressiveness, while losing a fight significantly suppressed male aggressiveness, consistent with the findings of existing studies on other cricket species. We also found that female presence had a stronger enhancing effect on male aggressiveness than flight or winning a fight. These findings demonstrated that physical and social experiences can affect the fighting behavior of male V. aspersus. Topical application of a 0.15?M solution of an OA receptor agonist (chlordimeform, CDM) significantly increased male aggression level, suggesting that OA may play an important role as a neuromodulator in controlling fighting behavior of males of this species. Despite displaying a significantly higher aggression level (level 5 or 6), CDM-treated losers did not escalate to physical combat, while fights between courting males usually resulted in physical escalation. It is likely that fighting behavior is only partly regulated by OA, and additional regulatory pathways may be involved in achieving physical combat.     

Familiarity Breeds Tolerance: the Development of Social Stability in Flocking Siskins (Carduelis spinus)

Behavioural integration associated with the fusion of two flocks is analyzed in captive siskins (Carduelis spinus) by quantifying changes in social behaviour with time since joining. In general there was an increase in the incidence of tolerant behaviour, supplanting attacks and hopping withdrawals with time since fusion of the flocks. However, the number of displays and flights showed the opposite, negative, trend. Taking dominance status into account, the greatest change in behaviour with time since joining is an increase in tolerance by dominants of new flock companions. Factorial analysis of correspondences was used to study how different birds changed their behaviour with time since joining a flock. This analysis showed that the introduction of new birds did not disrupt relationships with familiar birds, and that residents are dominant in interactions with the incoming new flock companions. The analysis also demonstrated that relationships within the new flock had stabilized 20 days after the flocks had joined. The characteristics of the socially integrated group of siskins are quite similar to those described by Rohwer & Ewald (1981) in their shepherds hypothesis: dominants tolerate their subordinates feeding in close proximity, offering them a profitable feeding area, but also supplant them to obtain food; both dominants and subordinates benefit from being in a flock. As a consequence, constant changes of membership in flocks is costly not only because birds lose dominance status, but also the advantages of clear dominant and subordinate roles.     

Explaining social learning of food preferences without aversions: an evolutionary simulation model of Norway rats

Norway rats (Rattus norvegicus) transmit preferences for novel foods socially by smelling each other's breath. However, rats fail to learn aversions, acquiring a preference even if the rat whose breath they smell has been poisoned. Rats can distinguish between sick and healthy conspecifics and social learning of both preferences and aversions is present in other species - hence it is unclear why rats cannot learn aversions socially. We constructed an evolutionary simulation in which a population of rats foraged from a central location, exploiting food sites that could contain edible or toxic foodstuffs. We examined the relationship between toxin lethality and selection for individual versus social learning and discrimination between sick and healthy conspecifics in order to allow learning of both preferences and aversions. At low lethality levels individual learning was selected for and at intermediate levels we found social learning of both preferences and aversions. Finally, given high lethality levels the simulated rats would employ social learning but failed to learn aversions, matching the behaviour of real rats. We argue that Norway rats do not learn aversions socially because their environment may contain only highly lethal toxins which make interaction with a sick conspecific an extremely rare event.     

Behavioural and hormonal responses of male rhesus monkeys introduced to females in the breeding and non-breeding seasons.

Six adult male rhesus monkeys were introduced individually to an all-female group for 10 days during the mating season. The initial aggressive responses of the females were rapidly replaced by positive social behaviour, and each male achieved alpha status and had access to social and sexual partners. A repetition of this paradigm in the non-breeding season produced significantly more female aggression, and no male attained high rank or engaged in sexual or other social behaviour. Male testosterone levels rose following introduction to the females in both seasons, but were significantly higher during the breeding season. Hormonal levels following removal from the females suggest a complex interplay between social, sexual and seasonal variables and recent social experiences. The differences in female social behaviour with newly introduced males, as a function of season, suggest an explanation for the seasonal limitation of male troop transfers.     

On the feeding and social behaviour of the laying hen

This paper describes a study of the influences of early husbandry conditions, social attraction and social rank on various aspects of the feeding behaviour of laying hens.Birds were raised in flocks of 10, 60 or 500. Groups of 3 birds, selected from flocks of the same size, were then housed in pens. Some groups consisted of hens raised in the same flock, and some of birds raised in different flocks. The feeding and agonistic behaviour of each group at each of 5 types of feeder was observed and compared with the behaviour shown when the birds had free access to a 1-m long food trough. Each of the 5 feeders offered the same area of feeding space, but differed in its partitioning and spatial distribution. One feeder had a single unpartitioned feeding space. The other 4 feeders had 3 partioned feeding spaces which were adjacent, or separated by distances of 10, 20 or 40 cm, respectively.For the 5 feeders, total feeding times and lengths of feeding bouts were greatest, and the number of feeding bouts least, when the feeding space was unpartitioned. Synchrony of feeding behaviour was low when the feeding space was unpartitioned or the partitioned spaces adjacent, but was comparable to that at the 1-m long food trough when the distance between partitioned feeding spaces was 10 cm or greater. When feeding space was partitioned, the likelihood that 2 birds would eat together at the same site increased with the distance between feeding space. Dominant birds always exhibited the longest feeding bouts and greatest total feeding times, but were less likely to feed in the same space as another bird, and exhibited less synchrony of feeding behaviour than subordinates. The size of the flock in which the birds were raised, and whether or not the birds in a group had been raised together or apart, had no clear effect on behaviour.These results indicate that, within the limits of this experiment, early husbandry conditions do not influence behaviour shown during feeding in later life, and that social attraction has a greater influence on the feeding behaviour of hens than is generally assumed. In view of this latter finding, it is postulated that in attempting to determine the requirements of laying hens for feeding space, attention must be paid to social attraction as well as to competition at the feeder.     

Deep-diving beaked whales dive together but forage apart

Echolocating animals that forage in social groups can potentially benefit from eavesdropping on other group members, cooperative foraging or social defence, but may also face problems of acoustic interference and intra-group competition for prey. Here, we investigate these potential trade-offs of sociality for extreme deep-diving Blainville′s and Cuvier''s beaked whales. These species perform highly synchronous group dives as a presumed predator-avoidance behaviour, but the benefits and costs of this on foraging have not been investigated. We show that group members could hear their companions for a median of at least 91% of the vocal foraging phase of their dives. This enables whales to coordinate their mean travel direction despite differing individual headings as they pursue prey on a minute-by-minute basis. While beaked whales coordinate their echolocation-based foraging periods tightly, individual click and buzz rates are both independent of the number of whales in the group. Thus, their foraging performance is not affected by intra-group competition or interference from group members, and they do not seem to capitalize directly on eavesdropping on the echoes produced by the echolocation clicks of their companions. We conclude that the close diving and vocal synchronization of beaked whale groups that quantitatively reduces predation risk has little impact on foraging performance.     

A warmer environment can reduce sociability in an ectotherm

The costs and benefits of being social vary with environmental conditions, so individuals must weigh the balance between these trade-offs in response to changes in the environment. Temperature is a salient environmental factor that may play a key role in altering the costs and benefits of sociality through its effects on food availability, predator abundance, and other ecological parameters. In ectotherms, changes in temperature also have direct effects on physiological traits linked to social behaviour, such as metabolic rate and locomotor performance. In light of climate change, it is therefore important to understand the potential effects of temperature on sociality. Here, we took the advantage of a ‘natural experiment’ of threespine sticklebacks from contrasting thermal environments in Iceland: geothermally warmed water bodies (warm habitats) and adjacent ambient-temperature water bodies (cold habitats) that were either linked (sympatric) or physically distinct (allopatric). We first measured the sociability of wild-caught adult fish from warm and cold habitats after acclimation to a low and a high temperature. At both acclimation temperatures, fish from the allopatric warm habitat were less social than those from the allopatric cold habitat, whereas fish from sympatric warm and cold habitats showed no differences in sociability. To determine whether differences in sociability between thermal habitats in the allopatric population were heritable, we used a common garden breeding design where individuals from the warm and the cold habitat were reared at a low or high temperature for two generations. We found that sociability was indeed heritable but also influenced by rearing temperature, suggesting that thermal conditions during early life can play an important role in influencing social behaviour in adulthood. By providing the first evidence for a causal effect of rearing temperature on social behaviour, our study provides novel insights into how a warming world may influence sociality in animal populations.     

Immediate and Delayed Benefits of Play Behaviour: New Evidence from Chimpanzees (Pan troglodytes)

Evidence for the anticipation of competition at feeding time has been previously documented in both Pan species. Chimpanzees seem to cope with competitive tendency through behavioural mechanisms of tension reduction, and grooming is certainly one of these. Social play and grooming are often matched because they bring animals into close physical contact for long periods, and they have an important role in social cohesion. Our goal was to investigate the occurrence of play behaviour during the pre‐feeding period, before a basic maintenance activity is about to take place, in the chimpanzee colony housed in the ZooParc de Beauval (St Aignan sur Cher, France). The group was composed of 10 adults and nine immature individuals. By scan animal sampling (344 h of observation), we recorded play and grooming interactions in all age‐class combinations during four different periods (pre‐feeding, feeding, post‐feeding, control). We found peak levels of grooming interactions among adults during the pre‐feeding time. A peak frequency at the pre‐feeding time was also found in social play between adults and unrelated immature subjects. This finding suggests that during high tension periods, grooming and play might share similar functions in conflict management. Like grooming, play might have an important role to limit aggression and increase tolerance around food (immediate benefits). Immature animals showed a higher frequency of play in the pre‐feeding than in any other condition (feeding, post‐feeding, and control). During high excitement periods social play probably represents a safe mechanism for immature subjects to test their personal abilities (self‐assessment), the strength/weakness of playmates, and the degree of cooperation/competition with them (social‐assessment). In the light of this new evidence, we can assert that play behaviour is far from being a purposeless activity, at least in the chimpanzee colony under study.