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.     

The coevolution of juvenile play–fighting and adult competition

Although play–fighting is widespread among juvenile mammals, its adaptive significance remains unclear. It has been proposed that play is beneficial for developing skills to improve success in adult contests (motor‐training hypothesis), but the links between juvenile play–fighting and adult aggression are complex and not well understood. In this theoretical study, we investigate the coevolution between juvenile play–fighting and adult aggression using evolutionary computer simulations. We consider a simple life history with two sequential stages: a juvenile phase in which individuals play–fight with other juveniles to develop their fighting skills; and an adult phase in which individuals engage in potentially aggressive contests over access to resources and ultimately mating opportunities, leading to reproductive success. The simulations track genetic evolution in key traits affecting adult contests, such as the level of aggression, as well as juvenile investment in play–fighting, capturing the coevolutionary feedbacks between juvenile and adult decisions. We find that coevolution leads to one of two outcomes: a high‐play, high‐aggression situation with highly aggressive adult contests preceded by a prolonged period of juvenile play–fighting to improve fighting ability, or a low‐play, low‐aggression situation in which adult contests are resolved without fighting and there is minimal investment in play–fighting before individuals mature. Which of these outcomes is favoured depends on the mortality costs and on the type of societal structure: societies with strong reproductive skew, favouring monopolization of resources, show high levels of adult aggression and high investment in juvenile play–fighting, whereas societies with low reproductive skew have both low adult aggression and low levels of play–fighting. A review of empirical evidence, particularly in the primate genus Macaca, highlights some limitations of our model and suggests that other, complementary functional explanations are needed to account for the full range of competitive and cooperative forms of play–fighting. Our study illustrates the power of evolutionary simulations to shed light on the long‐standing puzzle of animal play.     

Play vocalizations of squirrel monkeys (Saimiri sciureus)

Vocalizations occurring during play bouts were studied in 2 pairs of young male squirrel monkeys. Two main call types, one having four variants, are described. Vocalization rate varied with the type of ongoing behavior and with play bout duration, an association considered to be indicative of motivation to play. Structural differences also varied with bout duration and longer bouts had longer and more complex calls. In addition to providing direct information about motivation, vocalizations may have a metacommunicative function in alerting adult group members (not play partners) to the presence of playful activity.     

Social and solitary play in a colony of vervet monkeys (Cercopithecus aethiops)

This study describes and defines play in a laboratory colony of vervet monkeys (Cercopithecus aethiops). Play is divided into 26 units of behavior and the frequencies of these behaviors are recorded. Analysis shows that sex, age, and dominance have effects on some play behaviors. The play behavior of each age/sex group within the colony is described both quantitatively and qualitatively. Adult females are characterized by reinforcement of the play of immature monkeys, adult males by rough play with older male juveniles, four-year-old females and three-year-old males by stimulation of play in young infants, and males in general by a preference to play within their peer group. It is suggested that modified repetition of behaviors, diversified interactions, and innovative behaviors, are important qualities of play, and are essential to the adaptive plasticity of behavior in primates.     

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.     

The development of different types of play in gazelles: implications for the nature and functions of play

The development of play behaviour in Cuvier's gazelle, Gazella cuvieri, is described and compared with that of other species. Play was sub-divided into four different types: locomotor play, sexual play, play-fighting and object play. Developmental trends in the types considered were different, supporting the hypothesis that play is a heterogeneous category. Locomotor play started at high frequencies, and accounted for a major proportion of total play, but both measures declined from early months onwards. By contrast, sexual play and play-fighting were infrequent shortly after birth, increased thereafter reaching peaks between 4 and 6 months, and then decreased. The relative importance of these play types steadily increased until they became the predominant types of play. Object play remained infrequent at all ages, and accounted for a small proportion of total play. Young calves played mostly alone, but the proportion of social play increased with age, until all play became social. Before month 4 calves directed more invitations to play and sexual play than they received. Other peers were the major play partners of calves, followed by adult females. Calves directed a higher proportion of invitations and sexual play to adult females and males than they received from either class. These results are discussed in relation to other aspects of the calves' behavioural and social development, and it is argued that they are best explained if play is considered to have immediate benefits.     

Respective influence of extrinsic and intrinsic factors on the age-related decrease of thymic secretion.

The serum level of a circulating thymic factor (FTS) described in our laboratory diminishes in mice after adult thymectomy and with age. However, thymuses from old mice, when grafted into young adult thymectomized recipients lacking circulating FTS, can still partially restore the circulating FTS level of the recipients, whereas newborn thymuses are less efficient in restoring the serum level of FTS in old recipients than in young adult thymectomized recipients. Taken together, our results suggests that in addition to an intrinsic deficiency of the thymic secretion, "environmental" factors play a role in the disappearance of circulating FTS with age, the more so since we observed in old mouse sera factors inhibiting the in vitro biologic activity of FTS, factors that are absent from young mouse sera.     

Retinoic acid in the development, regeneration and maintenance of the nervous system

Retinoic acid (RA) is involved in the induction of neural differentiation, motor axon outgrowth and neural patterning. Like other developmental molecules, RA continues to play a role after development has been completed. Elevated RA signalling in the adult triggers axon outgrowth and, consequently, nerve regeneration. RA is also involved in the maintenance of the differentiated state of adult neurons, and disruption of RA signalling in the adult leads to the degeneration of motor neurons (motor neuron disease), the development of Alzheimer's disease and, possibly, the development of Parkinson's disease. The data described here strongly suggest that RA could be used as a therapeutic molecule for the induction of axon regeneration and the treatment of neurodegeneration.     

Play in Harbour seals (Phoca vitulina)

Harbour seals in a breeding colony at Miquelon exhibited 13 distinct types of play behaviour, 80% of which was solitary locomotor play. Most play was displayed by yearlings and juveniles, though a significant amount of adult play occurred as well. Mothers and pups rarely played. Yearlings played socially as often as alone, but seals of other age classes played by themselves most often. Only social play resembled adult activities. It is suggested that the function of mammalian play has been obscured by its similarity to adult behaviour.     

Homeobox gene expression in the intestinal epithelium of adult mice.

Using a polymerase chain reaction-based strategy, we have detected the expression of nine different homeobox genes in adult mouse intestine. Included among these are the recently described intestine-specific Cdx-1 gene and a new, related gene, Cdx-2. Southern blot experiments show that Cdx-2 is present in a single copy in the mouse genome. Of several adult mouse tissues assayed, intestine was the only one that contained detectable levels of Cdx-2 mRNA. Expression of all nine homeobox genes in different regions of the intestine was quantitated by RNase protection analysis, which revealed a unique expression profile for each gene. These observations suggest that homeobox gene expression may play an important role in cellular differentiation in the adult intestine.     

The social behavior of adult male-infant pairs of Rhesus macaques in a laboratory environment

Adult males in nonhuman primate societies exhibit a wide range of behaviors toward immature members of their social group, ranging from virtually continuous nurturance to systematic infanticide. Among macaques, rhesus monkey males in the wild display perhaps the least degree of paternalistic behavior toward conspecific infants. But adult male-infant rhesus attachments have not as yet been explored in a controlled longitudinal study, even though the mother-infant bond has received careful attention for several decades. In the present study two pairs of adult male and infant rhesus macaques were housed together without the mother for a period of at least seven months. To minimize the probability of physical harm to the infants, one adult male was present at the birth of the infant with whom it was paired and the other was visually familiarized with the infant for several weeks before being paired with it. Both pairs have shown much less ventral-ventral contact than mother-infant pairs and the infants have rarely attempted to gain nipple contact. The female infant now initiates and maintains ventral-ventral contact much more frequently than the male infant in spite of the higher degree of aggression directed toward her by the male. The adults frequently groom their infants and exhibit varying degrees of both tolerance and aggression. Rough-and-tumble play appeared quite early in both pairs and is much more vigorous than mother-infant play. Recently the male-male pair has exhibited reciprocal mounting and thrusting. Comparisons are made with data obtained in an earlier study of mother-infant pairs conducted under nearly identical experimental circumstances. The effects of separating the adult males and infants at seven months are also described.     

Sex differences in play among western lowland gorilla (Gorilla gorilla gorilla) infants: implications for adult behavior and social structure

Mammalian play is believed to improve motor skills as well as facilitate the development of social relationships. Given the marked sexual dimorphism in gorilla body size and the role assumed by the male in protecting the group from conspecifics and predators, the motor-training hypothesis of play predicts that male infants should exhibit higher frequencies of social play than female infants, and that males should prefer to play with other males. Given that adult female gorillas are strongly attracted to adult breeding males and form only weak social bonds with unrelated adult females, the social-relationship hypothesis of play predicts that female infants should prefer to play with males. These hypotheses were tested in a 22-month study of 12 gorilla infants, aged between 0-5 years, living in three zoological parks in Chicago and Atlanta. Consistent with the hypotheses, male infants played more than female infants did, and both male and female infants preferred to play with males rather than with females. These findings suggest that sex differences in play in the great apes and other primates can be predicted by the characteristics of adult behavior and social structure above and beyond the patterns of sex-biased dispersal or coalition formation with same-sex kin.     

Epithelial integrins

The integrin family was originally described as a family of adhesion receptors, utilized by cells for attachment to and migration across components of the extracellular matrix. Epithelial cells in adult tissues are generally stationary cells, but these cells nevertheless express several different integrins. This review will discuss the evidence that integrins on epithelial cells are also likely to function as signaling molecules, allowing these cells to detect attachment or detachment, and changes in the local composition of ligands. Signals initiated by integrins appear to modulate epithelial cell differentiation, proliferation, survival, and gene expression. Because the local concentration of integrin ligands is altered by injury, inflammation, and remodeling, signals initiated through integrins are likely to play important roles in the responses of epithelial cells to each of these processes.     

Topography and electrical activity of peripheral neurons in the abdomen of the tsetse fly (Glossina) in relation to abdominal distension

ABSTRACT. The detailed morphology of the abdominal nervous system of adult male and female Glossina morsitans is described. Staining with methylene blue revealed about twenty peripheral neurons distributed in a relatively constant pattern in each abdominal segment. Suction electrode recordings were made of the electrical activity generated by these neurons. Three pairs per segment present on the ventral body wall responded to mechanical stimulation, and are assumed to play a primary role in monitoring abdominal distension. Also present in each segment is a pair of dorsal longitudinal stretch receptors which require a greater intensity of mechanical stimulation to respond than do the ventral body wall neurons. A proposal as to how the various mechanically responsive receptors are involved in monitoring abdominal distension is presented.     

Vigilance during play in squirrel monkeys

Play by young squirrel monkeys (Saimiri boliviensis) may put them and other troop members at risk for predation because youngsters are noisy, separated from adults, and not vigilant when at play. In a study using separated groups of adults and 1-year-old juveniles caged outdoors, we found that adult female squirrel monkeys become more vigilant during periods of spontaneous play among juveniles. This behavioral response could be obtained with auditory cues (play vocalizations) alone. Five times as much vigilance activity was directed toward an area from which threat or disturbance was likely to come as was directed toward the juveniles themselves. These results suggest both an adaptive, compensatory increase in adult vigilance during play and a function for play vocalizations. Additional functions for play vocalizations remain to be investigated.     

The Development of Play in the South American Fur Seal

Play behaviour in the juvenile South American fur seal (Arctocephalus australis) is described for the period of dependence on the mother (up to 32 months). Play does not appear to be a unitary category of behaviour, but may be separated into four distinct types: terrestrial social and solitary play and swimming social and solitary play. These four types show different development profiles, and appear to serve different functions. Terrestrial solitary play and the two swimming play forms appear to be juvenile specializations, the first to enable very young seals to gain motor skills rapidly in order to avoid dangers from conspecifics, the second two to enhance antipredator behaviour and foraging ability during the gradual transition to independence. In contrast, social play may be best thought of as gaining social skills necessary in adult life, prior to dispersal post-weaning. Juvenile fur seals which do not wean in their first year show an upsurge in social play which is not seen in the other play types. This upsurge coincides temporally with the peak of social play of the new cohort, and is presumably elicited by them.     

Social play in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): Implications for natural social systems and interindividual relationships

This study compares adult play behavior in the two Pan species in order to test the effects of phylogenetic closeness and the nature of social systems on play distribution. The social play (both with fertile and immature subjects) performed by adults did not differ between the two species. In contrast, in bonobos, play levels among fertile subjects were higher than in chimpanzees. Findings regarding levels of undecided conflicts (more frequent in bonobos) and formal submission displays (lacking in bonobos) confirm, in the two colonies under study, that bonobos exhibit "egalitarianism" more than chimpanzees. Some authors emphasized the importance of play-fighting for social assessment when relationships among individuals are not codified and structured according to rank-rules. Indeed, adult bonobos played more roughly than chimpanzees. Moreover, adult bonobos displayed the full play-face at a high frequency especially during rough play sessions, whereas in chimpanzees, the frequency of play signals was not affected by roughness of play. The frequency of social play among bonobo females was higher than in any other sex combinations, whereas no difference was found for chimpanzees. As a matter of fact, social play can be viewed as a balance between cooperation and competition. Among bonobo females, characterized by social competence and affiliation, social play might enhance their behavioral flexibility and increase their socially symmetrical relationships which, after all, are the basis for their egalitarian society.     

Enhanced neurite outgrowth in PC12 cells mediated by connexin hemichannels and ATP

Gap junctions have traditionally been described as transmembrane channels that facilitate intercellular communication via the passage of small molecules. Connexins, the basic building blocks of gap junctions, are expressed in most mammalian tissues including the developing and adult central nervous system. During brain development, connexins are temporally and spatially regulated suggesting they play an important role in the proper formation of the central nervous system. In the current study, connexins 32 and 43 were overexpressed in PC12 cells to determine whether connexins are involved in neuronal differentiation. Both connexin 32 and 43 were appropriately trafficked to the cell membrane following overexpression and resulted in the formation of functional gap junctions. Connexin overexpression was found to cause enhanced neurite outgrowth in PC12 cells treated with nerve growth factor to initiate neuritogenesis. Surprisingly, however, enhanced neurite outgrowth was found to be the consequence of functional hemichannel formation as opposed to traditional intercellular communication. Additional analysis revealed that ATP was released into the media likely through hemichannels and acted on purinergic receptors to cause enhanced neurite outgrowth. Collectively, the results of the current study suggest that connexins may play an important role in neuronal differentiation by non-traditional mechanisms.     

Immunohistological localization of serotonin in the CNS and feeding system of the stable fly Stomoxys calcitrans L. (Diptera: Muscidae)

Serotonin, or 5-hydroxytryptamine (5-HT), plays critical roles as a neurotransmitter and neuromodulator that control or modulate many behaviors in insects, such as feeding. Neurons immunoreactive (IR) to 5-HT were detected in the central nervous system (CNS) of the larval and adult stages of the stable fly, Stomoxys calcitrans, using an immunohistological technique. The location and pattern of the 5-HT IR neurons are described and compared for these two different developmental stages. Anatomical features of the fly feeding system were analyzed in third instar larvae and adult flies using a combination of histological and immunohistological techniques. In third instar larvae, the cibarial dilator muscles were observed within the cibarial pump skeleton and innervated by 5-HT IR neurons in nerves arising from the brain. There were four pairs of nerves arising from the frontal surface of the larval brain that innervate the cibarial pump muscles, pharynx, and muscles controlling the mouth hooks. A strong serotoninergic innervation of the anterior stomatogastric system was observed, which suggests 5-HT may play a role in the coordination of different phases of food ingestion by larvae. Similarly, many 5-HT IR neurons were found in both the brain and the thoracico-abdominal ganglia in the adult, some of which innervate the cibarial pump dilator muscles and the stomatogastric muscles. This is tnhe first report describing neuromuscular structures of the stable fly feeding system. The results reported here suggest 5-HT may play a critical role in feeding behaviors of stable fly larvae and adults.