AMER. ZOOL., 14:289-294 (1974). The Ontogeny of Play within a Society: Preliminary Analysis G. W. MEIER AND V. D. DEVANNEY Department of Psychiatry, University of Nebraska, College of Medicine, Omaha, Nebraska 68131 SYNOPSIS. Data were collected on play behaviors occuring within a single troop or rhesus monkeys in two environments. Observations were regular and covered a total of almost 2 years. The three categories of play—object, activity, and social play—tended to occur together, typically within the same recording session. Thus, the attending conditions— including satiation and maturity—which permitted the occurrence of one category of play behavior, permitted the occurrence of the other categories as well. Maturity seemed to account for the appearance of a particular behavior within a category (e.g., manipulation rather than touching; climbing rather than active hanging; romping and wrestling rather than contact or touching). The social context, determined by such factors as maternal dominance, the identity of nearby animals, and the overall social tension of the troop, and reflected in the extent of inhibitory control of the target animals by their mothers, seemed to determine the frequency with which these elements in the infant monkey's repertoire could be displayed during any given period in the ontogeny of the individual. Satiation and immaturity remain as the most provocative conditions—but least promising explanations—for mammalian play. Together with the recognized irregularity of play behavior—at least irregular in the sense that unpredictability of occurrence continues to confound systematic experimentation—these conditions remain the stalwart characteristics of play research (Loizos, 1967). Being unable to specify the natui-e of satiation or of maturity save in terms of feeding schedules or of "clocktime," we have elected to describe the ontogenesis of play behavior in infants born to a laboratory simulation of a natural troop of monkeys, rhesus macaques, and have focused on the development of social play under the conditions of routine existence in their environment which included the mothers and other adults, siblings and other older, yet juvenile members, and age-peers (see Rosenblum, 1961; Kaufmann, 1966). Despite a familiarity with the group which began some 5 years before these observations were made, we accepted the erratic ocThis research was supported in part by NIH grants HD-00370 and HD-00973 from NICHHD. currence over any epoch of time and have aspired to sketch only the course or pattern of the development of play behaviors rather than to define ages of initial occurrences or transitions, or to provide absolute values of the quantities of such behaviors and their proportions of the total behavioral display for any given age. METHOD For this report, we will consider only data collected during two phases in the existence of the primate troop of rhesus monkeys (Macaca mulatto.) which are our subjects. All were derived from an extended family group moved from Cayo Santiago, Puerto Rico, in November 1965 to Nashville, Tennessee (Meier, 1969). During the first phase, the group consisting of 7 adults (1 male, 6 females), 3 sub-adults (2 males, 1 female), 11 yearlings and juveniles (4 males, 7 females), and 4 infants (1 male, 3 females) resided in an inside-outside facility with a large outside cage and a much smaller inside room on top of one of the instructional buildings on the campus of George Peabody College. During the second phase, 289 290 G. W. MEIER AND V. D. DEVANNEV the troop, now moved to Omaha, Nebraska (as of early February 1972), consisted of 15 adults (7 males, 8 females), 2 sub-adults (2 females), 10 yearlings and juveniles (2 males, 8 females), and 6 infants (4 males, 2 females). At this time the group lived in quarters—two inside rooms and two somewhat smaller outside cages—on top of one of the buildings at the Medical Center of the University of Nebraska. Under both conditions, the animals had free access to all parts of the facilities at all times. In the first environment, the animals were fed only in the inside room on a self-paced schedule determined by individual patterns of operation of food-delivery system which was available to them at all hours (Meier et al., 1969; Meier, 1971). In the second, the animals were fed in one room only during the midmorning hours (before 1000 hours) by the placement of large quantities of food for easy access. In the first situation, we observed the animals through one-way vision windows strategically placed at sites adjacent to the animal quarters. In the second, we observed the animals through the open doorways to the inside rooms or, as necessary, from the outside area adjacent to the outside cages. The procedures for the actual recording of behavior under the two conditions were almost identical: a check-list, time-sampling technique was used whereby the observer indicated for each 15-sec epoch, for each daily 10-min session, the presence of selected behaviors in each of seven categories (proximity, posture, locomotion, facial display, social manipulation, object manipulation, and vocalization). The particular behaviors recorded differed appropriately by age of animal observed. One check-list was used for the first 2 months following birth; a second was used thereafter (Devanney, 1973). Some variation occurred on the latter as necessitated by the establishment of the check-list for the older age. In each instance, the observer (the authors or, during a brief period in the first phase, an experienced colleague) entered in a shorthand code in the appropriate cell on the prepared form the behaviors of the target animal (always), its mother (usually), and its siblings, age-peers, and unrelated others in the troop (occasionally). Observations were made daily in the first phase from the day of birth to 10 to 12 months of age; in the second phase they were made daily from the day of birth to 60 days of age and then twice weekly until 6 to 8 months of age. In both instances, they were usually made in the morning hours after the maintenance activities (and the feeding, as well, in the second phase) were complete. The target animals were all of the infants born to the group during the particular phase indicated; these were the 4 animals, 1 male and 3 females, born in 1970 and the 6 animals, 4 males and 2 females, born in 1973. (All were delivered between March and October of the year indicated. Our intent in presenting these data from widely separate times and places is to show the fundamental stability of the developmental-social patterns of infantile play behaviors within a primate troop whose composition closely approximated the feral situation (Lindburg, 1971). In short, our emphasis is on the pertinence of these behaviors to the social environment rather than to other, physical aspects. The comments which follow are preliminary in nature in that they were based upon an analysis which is still incomplete. RESULTS In the analysis of the checklist data, the behaviors were categorized according to the nature of the play orientation: (i) toward an object (Object Play); (ii) toward another member of the troop (Social Play); and (iii) toward "free" activity (Activity Play). These were counted and summed for each session in a manner which indicated the exclusive nature of the categories (i.e., no two categories could be scored for the same epoch). In all cases the mother's proximity was also recorded and an attempt was made to score the arousal level of the group, 1-5, during the period of observation. The appearance of all three types of play occurred at about the same time—when the infants were about 2 weeks of age—as shown in Figure 1. (All illustrative material 291 ONTOGENY OF PLAY IN SITU three patterns which appeared to be temporally distinguishable. The first appeared during the first postnatal month when the infants were 2 to 3 weeks of age. This occurred in close proximity to the mother (in a ventral-ventral position, typically) and consisted of manipulation of the mother by the infant although sometimes the particular interaction was mutual and the mother manipulated the infant alternately as it did her. (We did keep a separate tally of play behaviors initiated by the mother and others directed to the infant.) The manipulation by the infant was subdued, without vocalization or vigor, typified by the manual exploration or displacement of some part of the mother's body: face, ears, fingers, arms. This behavior was usually of FIC. 1. Frequencies of object, activity, and social short duration—it rarely persisted beyond play behaviors computed as totals over 10-day blocks from birth to 300 days of age. Data from LB/EX. a single 15-sec epoch—although it could occur several times within a single session. is taken from the analysis of a single pair, The frequency of occurrence remained low, LB/EX. The mother, LB, was multiparous but stable, during this month and into the and high-ranking. The infant, EX, was a third postnatal month at which time it female born October 3, 1971. The trends diminished but continued until about the portrayed are believed to be representative seventh month when it showed an even of all the pairs observed weighing, of course, sharper decline. The second pattern was somewhat more the influence of maternal parity and domrigorous than the first and like the first, ocinance and infant gender.) Object play most often occurred within 1 m of the mother until the middle of the fourth month, at which time it began to occur increasingly at distances greater than 1 m from the mother. By the age of 5 months the frequency of object play at these greater distances surpassed the frequency of object play close to mother (see Fig. 2). Activity play was the most frequently recorded type of play behavior, perhaps because our definition was so all-encompassing. It began to occur occasionally by the sixth week at a distance from the mother greater than 1 m, occurred in equal proportions both near and far from the mother until the end of the third month, then became more frequent in situations distant l!0 270 from the mother. However, like object play it continued to appear in situations of FIG. 2. Frequencies of object play behaviors as regreater proximity (see Fig. 3). lated to mother-infant distance, computed as totals Observations of social manipulations and over 30-day blocks from birth to 300 days of age. interactions of these rhesus infants revealed Data from LB/EX. 160 AGE IN DAYS On vcnlrum Off vcntiam, within 1 m . . . . Beto.d I m AG! IK DAYS 292 G. W. MEIER AND V. D. DEVANNEY then wrestling with chasing, romping, teasing usually, but not necessarily, with agepeers—was first recorded during the second _ O« vlltna : . Oil Vlltfla, »itkii 1 • ,' at a very low frequency. Thereafter month . l.roid I m i it increased markedly in frequency of observation until the animal was about 6 months of age and then remained stable. With this third type of play the initiator of action could not be determined easily, if V at all. In the females, some decline in this frequency was noted toward the end of the o first year, whereas for the male infants the S IJO frequency increased slightly, if any change could be reported, at this age. In both the / w male and female infants, the relation of this more complex and temporally prolonged (both between epochs and between sessions) form of play to the larger social context was clear. This play seemed to occur only when the infant was well beyond the mother's reach, at a distance of a meter and more from her. Nearly all of the social play episodes recorded during the first 3 months occurred within a proximity of a meter from the FIG. 3. Frequencies of activity play behaviors as related to mother-infant distance, computed as to- mother. Thereafter, the distance and fretals over 30-day blocks from birth to 300 days of quency increased rapidly (see Fig. 4). age. Data from LB/EX. None of these forms of social interaction which we have called play occurred during epochs containing aggression in the troop, curred within close proximity to the mother even if considerably removed from the tar—either in direct contact or within 1 m of get mother-infant pair. Depending upon her. This pattern followed the first in about the nature—contact versus display—of the a week. This pattern was the manipulation, pulling, and patting of another anim * mal, older than the infant, seated near the / Itpii 1 / mother. Typically this animal was a sibling / 120 (usually female) of the mother or infant or, / sometimes, an age-peer and "friend" of the mother. On several occasions we observed i i these maneuvers with (or by) one of the i i adult males. In those instances, we noted i to i that the mother was high in the dominance $ i hierarchy and the male was a fully grown A offspring or was an occasional consort. • When recorded, this behavior was sometimes unilateral (i.e., the infant played with the other animal or the other animal manipulated the infant), but frequently alternating in direction between the two ani- FIG. 4. Frequencies of social play behaviors as related to mother-infant distance, computed as totals mals involved. over 30-day blocks from birth to 30 days of age. The third pattern—first wrestling and Data from LB/EX. 300 110 J O l III DAYS " Oi viali 0(1 viit «m, w i i h i . 1 m \ M • / •E •- /•• 110 AGI IN OATS ONTOGENY OF PLAY IN SITU aggression and the participation within the bout—adult versus subadults—the effect of the bout could be discerned throughout the session by the quiescence and proximity of the target animals or, even, into succeeding sessions that day with other target pairs. With epochs of general troop equanimity, however, the incidence of this complex form of play was related to distance of separation of mother and infant and to the dominance ranking and parity —correlated dimensions—of the mother. In pairs with primiparous, low-ranking mothers, this form of play did occur within a meter of the mother although more often within 1 to 3 m of her. On the other hand in pairs with multiparous, high-ranking mothers this form of play rarely occurred close to the mother (i.e., within a meter), most often beyond 3 m but, frequently, out of her presence and sight entirely. DISCUSSION In effect these data are a lession in grammar: the relative meaning of "can" and "may." The grammarian will state that "can" indicates ability or capacity whereas "may" indicates possibility or permission. These data indicate the existence of two separate processes, inextricably confounded in a complex social situation such as this, which control the observed incidence of play behavior: the development of the behavioral repertoire of the infant macaque and the vicissitudes of the social control provided by the maternal parent. The first is dictated by factors only dimly known, but which operate to increase the speed, vigor, and precision of individual movements as well as their qualitative variety. The second may be related to idiosyncrasies of the particular mother—age, parity, rearing conditions—but are most assuredly dominated by a larger social context in which the mother-infant pair resides. These last are those aspects which determine social hierarchy but which, more importantly, control social adhesion and modulate social tension. The first process is of a maturational nature defining the moment-tomoment capability of the infant; the second 293 is of a governing nature by which the infant's possibilities to explore a rich physical (including social) environment are controlled. Our data reveal the balance, changing over time, between these two sometimes enhancing, sometimes restricting forces. Further, our data underscore the reality that, in contrast to the common supposition that play is an avenue for the immature animal "to learn social behaviors," the ontogeny of play occurs in a social context within which the infant is already responding as an established member. We have seen that the form and complexity of social play are related, first, to what the infant can do and, second, to the physical distance between the infant and its mother. Whereas the first observations of social play behaviors are of motherinfant interactions which are brief and tentative, successively later interactions are between the infant and siblings and other intimates of the mother and between the infant and age-peers. The first of such behaviors are in close proximity to the mother (even in the ventral-ventral, early infantile posture); later, the behaviors occur at greater distances; and, ultimately, entirely out of her sight and presence. The first occur in a context which is established and virtually constant, at least through early infancy, and are requisite to the early survival of the infant. The second occur in a context which is also well established antedating the delivery of the infant and reflect instances of such intimacy during previous seasons and peer patterns established during those years. The third, those play behaviors which occur well beyond the mother's arm length and typically involve other infants, are the resultant of the combined relaxation of inhibitory control of the two or more mothers involved. This relaxation —or disinhibition—can take place only when the overall social tension of the troop is low or removed altogether. These behaviors reflect the establishment of entirely new inter-individual patterns within the troop. Our analysis of these developmental trends of play behavior is supported by the observations on social distance made by 294 G. W. MEIER AND V. D. DEVANNEY Jensen and his associates (Jensen et al., 1973) on mother-infant pairs of pig-tailed macaques maintained in environments of differing social complexity. Where the mother and infant were maintained in isolation with minimal physical complexity, the distances were small and changes were slow in coming. Where the two were maintained in isolation with maximal physical complexity, the distances also were maximal and occurred early in postnatal life. Where the two were part of a social troop with other adults and infants, the distances were also small and increased slowly with postnatal age. Wolfheim et al. (1970) have suggested that the last is the outcome of inhibitory control by the mother dictated by the communication between the adults of the troop which, of course, include the mother. This last interpretation is corroborated by the observations of Meier et al. (1971) on a group of rhesus macaques the adults of which had had a bilateral sectioning of the VII th cranial nerve and thereby were rendered unable to make most of the facial expressions typical of the species. Under these conditions of limited communication among the adults, infants born to the group revealed a developmental pattern of physical separation comparable to mother-infant pairs maintained in some degree of physical isolation, e.g., in a playpen setting, but far different from other pairs observed in this communal group since first observed in 1965. The infants of the sectioned mothers moved away from these females into the total group at much earlier ages than observed elsewhere, usually as early as the second month. Another aspect of the data provides further support for the described trends. The three categories of play—object, activity, and social play—tended to occur together, typically within the same recording session. Thus, the attending conditions—including satiation and maturity—which permitted the occurrence of one category of play behavior, permitted the occurrence of the other categories, as well. Maturity seemed to account for the appearance of a particular behavior within a category (e.g., manipulation rather than touching; climbing rather than active hanging; romping and wrestling rather than contact or touching); the social context seemed to determine the frequency with which these elements in the infant's repertoire could be displayed during any session and, presumably, any other instance in the ontogeny of the individual. REFERENCES Devanney, V. D. 1973. A developmental study of operant behavior acquired by infant rhesus monkeys in a social group. Master's thesis. George Peabody College for Teachers, Nashville, Tenn. Jensen, G. D., R. A. Bobbitt, and B. N. Gordon. 1973. Mothers' and infants' roles in the development of independence of Macaca nemestrina. Primates 14:79-88. Kaufmann, J. H. 1966. Behavior of infant rhesus monkeys and their mothers in a free-ranging band. Zoologica 51:17-32. Lindburg, D. G. 1971. The rhesus monkey in North India: an ecological and behavioral study, p. 1106. In L. A. Rosenblum [ed.], Primate behavior: developments in field and laboratory research. Vol. 2. Academic Press, New York, N.Y. Loizos, C. 1967. Play behavior in higher primates: a review, p. 176-218. In D. Morris [ed.], Primate ethology. Morrison and Gibb Ltd., London. Meier, G. W. 1969. The re-establishment of a rhesus social group, p. 66-71. In C. R. Carpenter [ed.], Proceedings of the Second International Congress of Primatology: Behavior. Vol. 1. Karger, Basel, Switzerland. Meier, G. W. 1971. Operant cycles and imitational learning in a social setting. Primates 12:221-227. Meier, G. W., C. E. Izard, and C. Cobb. 1971. Facial display in primate communication. Paper presented to Southeastern Psychological Association, Miami, Florida. Meier, G. W., H. C. Wilcoxon, R. Orlando, and D. G- Paulson. 1969. The development of foodreinforced behaviors in a social situation, p. 143148. In C. R. Carpenter [ed.], Proceedings of the Second International Congress of Primatology: Behavior. Vol. 1. Karger, Basel, Switzerland. Rosenblum, L. A. 1961. The development of social behavior in the rhesus monkey. Ph.D. Diss., University of Wisconsin, Madison. Wolfheim, J. H., G. D. Jensen, and R. A. Bobbitt. 1970. Effects of group environment on the mother-infant relationship in pigtailed monkeys (Macaca nemestrina). Primates 11:119-124.
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