Circadian and Ultradian Time Patterns in Human Behavior: Part 2: Social Synchronisation During the Development of the Infant's Diurnal Activity-Rest Pattern

Individually developing patterns of activity-rest rhythms in infants and the influence of environmental factors in the tuning and synchronisation of parent-infant pairs have important implications for the health of both infant and parents. After discharge from the hospital newborn infants are exposed to patterned influences of maternal and environmental regularities of a family's daily life resulting in varying degrees of social synchronisation. Actigraphic monitoring was used in this longitudinal study to examine how activity patterns of the entire family agree or disagree with each other, and how the infant entrains to the environment. Activity data of 12 families (father, mother and infant) were continuously recorded using non-invasive Actiwatch units. Recordings of parental activity started at the beginning of the 37th week of gestation, and were continued in parallel with the infants' recordings in three series of 21 days each until four months after birth: 1st to 3rd week, 7th to 9th week and 13th to 15th week of life. Fast Fourier transformation and cross correlation techniques were used to determine frequencies of each family member and to quantify the synchronisation of activity between parents and infants. To elucidate differences in social synchronisation between human cultures, synchronisation of a Melanesian family was additionally compared. Results showed the existence of corresponding ultradian frequencies in the activity patterns of mother-infant pairs at 1, 2 and 4 months. Increases in the synchronisation of parental activity were found from prenatal to postnatal and for mother-infant pairs from the first to the second month. Synchronisation between mother and infant always exceeded that of father and infant. Transient mono-, bi- or polyphasic activity patterns emerged in the infants immediately after birth. Good correspondence of mother-infant activity patterns during the early postnatal period was correlated with a rapid development of an entrained daily pattern in the infant.     

Evolution and sudden infant death syndrome (SIDS)

This paper and its subsequent parts (Part II and Part III) build on an earlier publication (McKenna 1986). They suggest that important clinical data on the relationship between infantile constitutional deficits and microenvironmental factors relevant to SIDS can be acquired by examining the physiological regulatory effects (well documented among nonhuman primates) that parents assert on their infants when they sleep together. I attempt to show why access to parental sensory cues (movement, touch, smell, sound) that induce arousals in infants while they sleep could possibly help one of many different subclasses of infants either to override certain kinds of sleep-induced breathing control errors suspected to be involved in SIDS or to avoid them altogether. I do not suggest that solitary nocturnal sleep “causes” SIDS, that all parents should sleep with their infants, or that traditional SIDS research strategies should be abandoned. However, using evolutionary data, I do suggest that an adaptive fit exists between parent-infant sleep contact and the natural physiological vulnerabilities of the neurologically immature human infant, whose breathing system is more complex than that of other mammals owing to its speech-breathing abilities. This “fit” is best understood, it is argued, in terms of the 4–5 million years of human evolution in which parent-infant contact was almost certainly continuous during at least the first year of an infant’s life. Thus, to dismiss the idea that solitary sleep has no physiological consequences for infants does not accord with scientific facts. James J. McKenna is Associate Professor of Anthropology and Chair of the Department of Sociology and Anthropology at Pomona College. He also has an appointment as an Adjunct Clinical Assistant Professor in the Departments of Pediatrics, Child Psychiatry, and Human Behavior at the University of California, Irvine, School of Medicine. His primary research interests and many of his publications concern aspects of primate parenting and infant development among both human and nonhuman primates. For the past seven years he has been investigating from an anthropological perspective possible environmental correlates of the sudden infant death syndrome (SIDS) and has just finished a preliminary study on the physiological correlates of human parent-infant co-sleeping. His earlier monograph on the subject (cited in this paper) has received much international attention. He and his colleagues (Mosko and Dungy) are the first to have used standard polysomnographic techniques to document simultaneously human parent-infant co-sleeping. He has won three awards for distinguished teaching at Pomona College.     

Parent-infant bed-sharing behavior

An evolutionarily informed perspective on parent-infant sleep contact challenges recommendations regarding appropriate parent-infant sleep practices based on large epidemiological studies. In this study regularly bed-sharing parents and infants participated in an in-home video study of bed-sharing behavior. Ten formula-feeding and ten breast-feeding families were filmed for 3 nights (adjustment, dyadic, and triadic nights) for 8 hours per night. For breast-fed infants, mother-infant orientation, sleep position, frequency of feeding, arousal, and synchronous arousal were all consistent with previous sleep-lab studies of mother-infant bed-sharing behavior, but significant differences were found between formula and breast-fed infants. While breast-feeding mothers shared a bed with their infants in a characteristic manner that provided several safety benefits, formula-feeding mothers shared a bed in a more variable manner with consequences for infant safety. Paternal bed-sharing behavior introduced further variability. Epidemiological case-control studies examining bed-sharing risks and benefits do not normally control for behavioral variables that an evolutionary viewpoint would deem crucial. This study demonstrates how parental behavior affects the bed-sharing experience and indicates that cases and controls in epidemiological studies should be matched for behavioral, as well as sociodemographic, variables.     

Evolution and the sudden infant death syndrome (SIDS)

Postnatal parent-infant physiological regulatory effects described in the previous paper (Part I) are viewed here as being biologically contiguous with events that occur prenatally, preparing and sensitizing the fetus to the average microenvironment into which the infant is expected, based on its evolutionary past, to be born. Following McKenna (1986), evidence (some of which is circumstantial) is presented concerning fetal hearing and fetal amniotic liquid breathing as they are affected both by maternal cardiovascular blood flow sounds in the uterus and by fluctuating maternal blood sugar levels. These data are linked in turn to the infant’s postulated postnatal responsivity to parental sensory cues, including auditory and vestibular respiratory cues that may assist infants as they “learn” to breathe and, for some, to resist a SIDS event. Data on the respiratory and vocalizing behavior of normal and hearing-impaired persons are used to show that not all forms of human breathing are innate; some forms develop with experience. These data reveal how human infants learn, for example, to coordinate higher and lower brain respiratory nuclei in the context of learning initially to cry with intent and purpose and later to speak. Voluntary, cortex-based breathing emerges at the same time that infants are most likely to die from SIDS, between 2 and 4 months of age. This switch between voluntary and involuntary breathing during both sleep (while dreaming) and wake cycles, which depends on the integration of higher cortical and lower brain stem nuclei, is complex and is possibly the basis of the human species’ unique susceptibility to SIDS—a syndrome as yet unrecognized in other species. These human infant vulnerabilities, including delayed maturity, can explain in part why natural selection ought to favor increased infant sensitivity to parental sensory cues provided by a caregiver—stimuli available in the evolving parental care environment that included parent-infant co-sleeping for more than 4–5 million years of human evolution.     

Growth and Investment in Hominin Life History Evolution: Patterns, Processes, and Outcomes

The transitions from apes to lineages allied to humans are marked by shifts in the allocation of parental effort, associated with discontinuous changes in rates of infant and juvenile growth both prenatally and postnatally. Here, I assess growth and life history characteristics of apes within a general mammalian / primate paradigm, using time and energy expenditure as 2 fundamentals that covary with infant survival and success probabilities. I suggest that these survival probabilities depend on the quality, amount, and timing of parental care allocated to infants. Growth to birth, growth to weaning, and growth to reproductive onset are partitioned as separate periods within a life history on the basis of comparative mammalian data. Growth problems such as sexual dimorphism can be incorporated into an investment perspective by assessing when and how sex-specific parental care affects growth rates and the onset of reproduction. I compare features of the hominoid life history with developmental rates for hominin lineages as seen in dentition, and the fossil record of body and brain size changes over time. The links between parental effort and allocation of care to infant growth and survival generate speculative scenarios of sex-specific parental care allocation; I then explore hominin social evolution?—mating system and childhood—?for the lineages thought to lead to modern Homo, and for those that coexisted with ancestors of Homo.     

Male parenting and response to infant stimuli in the common marmoset (Callithrix jacchus)

Callithrix jacchus infants are raised in complex family environments where most members participate in rearing the young. Many studies examining male parental behavior have focused on the carrying of infants with observations made within the family context. However, interference from family members can make it difficult to assess the father's motivation to care for infants. Our goals were to develop a testing paradigm for determining an individual's response to infant stimuli separate from family influences, compare a male's motivation to respond to an infant stimulus outside the family with his paternal behavior within the family, to compare responses to infant stimuli of parentally experienced versus inexperienced males and finally to develop a reproducible and standardized method of testing male responsiveness to infant stimulus that could serve to evaluate hormonal manipulations. Fifteen experienced common marmoset fathers were evaluated using three different measures of parental behavior: (1) instantaneous scan sampling, (2) continuous focal sampling in the family, and (3) continuous focal sampling of males presented with four infant stimuli: familiar and unfamiliar infants, familiar and unfamiliar infant vocalizations. Six parentally inexperienced males (non-fathers) served as controls. Males that carried the most in the family were typically the same males that responded most to the infant vocalization tests. Experienced fathers did not differ in their latency to enter the stimulus cage for any of the four infant stimuli response tests while inexperienced males took significantly longer to enter the stimulus cage. In addition, fathers expressed a greater frequency of infant-directed behavior than did the inexperienced males during the unfamiliar infant and unfamiliar vocalization tests. These studies show that experienced male marmosets are highly motivated to interact with infant stimuli and that there is interindividual variability in response to infant vocalizations. Testing males outside of the family allows for a clear assessment of male's interest in infant stimuli in both parentally experienced fathers and inexperienced males.     

Where Will the Baby Sleep? Attitudes and Practices of New and Experienced Parents Regarding Cosleeping with Their Newborn Infants

An evolutionary perspective on human infant sleep physiology suggests that parent-infant cosleeping, practiced under safe conditions, might be beneficial to both mothers and infants. However, cosleeping is not part of mainstream parenting ideology in the United States or the United Kingdom, and little evidence is available to indicate whether, and under what circumstances, parents sleep with their newborn infants. We present data from an anthropological investigation into the practices and attitudes of new and experienced parents of newborn infants regarding parent-infant sleeping arrangements in a community in the northeast of England. Despite not having contemplated cosleeping prior to the birth, new parents in our sample found it to be a convenient nighttime caregiving strategy, and one which was practiced regularly. Infants slept with both their parents, some being habitual all-night cosleepers, but commonly beginning the night in a cnb and sleeping with their parents for several hours following the early morning feed, [infant sleep, newborn, cosleeping, new parents]     

Focus: The Science of Stress: Social Fear in US Infants: The Roles of Hair and Salivary Cortisol

Elevated social fear in infancy poses risk for later social maladjustment and psychopathology. Hair cortisol concentration (HCC), an index of cumulative cortisol exposure, and diurnal salivary cortisol slope, a biomarker of acute stress regulation, have been associated with social fear behaviors in childhood; however, no research has addressed their relations in infancy. Elucidating potential biomarkers of infant social fear behaviors, as well as environmental factors associated with these biomarkers, may grant insights into the ontogeny of fear behaviors that increase risk for internalizing and externalizing psychopathologies later in life. The current study used multiple linear regression to examine if infant HCC, infant diurnal cortisol slope, and income-to-needs ratios (ITN) were differentially associated with observed social fear responses to a Stranger Approach task at 12 months. Using a sample of 90 infants (M_age = 12.26m, SD = 0.81m, 50% female), results indicated that increased infant HCC was associated with increased distress vocalizations during the Stranger Approach task, while steeper diurnal cortisol slope was associated with fewer distress vocalizations. Ordinary least squares path analyses did not reveal group differences between economically strained and non-strained infants in how cortisol measures and social fear responses related. Findings underscore very early psychobiological correlates of fearfulness that may increase risk for fear-related disorders and adverse mental health symptomology across childhood.     

The cross-generation transmission of oxytocin in humans

Animal studies demonstrated that the neuropeptide oxytocin (OT), implicated in bond formation across mammalian species, is transmitted from mother to young through mechanisms of early social experiences; however, no research has addressed the cross-generation transmission of OT in humans. Fifty-five parents (36 mothers and 19 fathers) engaged in a 15-min interaction with their infants. Baseline plasma OT was sampled from parents and salivary OT was sampled from parents and infants before and after play and analyzed with ELISA methods. Interactions were micro-coded for parent and child's socio-affective behavior. Parent and infant's salivary OT was individually stable across assessments and showed an increase from pre- to post-interaction. Significant correlations emerged between parental and infant OT at both assessments and higher OT levels in parent and child were related to greater affect synchrony and infant social engagement. Parent-infant affect synchrony moderated the relations between parental and infant OT and the associations between OT in parent and child were stronger under conditions of high affect synchrony. Results demonstrate consistency in the neuroendocrine system supporting bond formation in humans and other mammals and underscore the role of early experience in shaping the cross-generation transmission of social affiliation in humans.     

Longitudinal observations of care and development of infant titi monkeys (Callicebus moloch)

The purpose of this study was to describe patterns of infant care and development in family groups of the monogamous titi (Callicebus moloch). Three infant titis were observed with their natal family groups over the first six months of life. Field observations of extensive male involvement with infants were confirmed. Adult males were clearly the infants' primary social companions, and infants spent more time in contact with adult males at all ages than with mothers or older siblings. However, mothers and siblings also carried infants at times and interacted with them in affiliative ways (e.g., grooming, nuzzling, play). Mothers often invited nursing, but otherwise it was infants who primarily initiated transfers between carriers and approaches to parents, reminding us that the infants' own activities and their effects on caregivers should not be overlooked in considerations of the patterns of infant care. This is particularly important for those species exhibiting extensive parental involvement by males. It is suggested that substantial male care of young titi infants leads to several important consequences for the infant's social development, including the development of a stronger attachment to the father than the mother.     

Social behavior of infant and mother Japanese Macaques (Macaca fuscata): Effects of kinship,partner sex,and infant sex

The behavioral interactions of 22 infant and mother Japanese macaques with other group members were studied. Focal-animal observations were made from the time of each infant’s birth until 1 year of age. Infants and mothers both displayed exceedingly strong preferences for associating with matrilineal kin and, specifically, for female kin. The degree of genetic relatedness was positively correlated with levels of spatial proximity, contact, grooming, aggression, and play. Overall frequencies of interactions with nonkin were very low, and partner sex was not an important factor in interactions with nonkin. There were no significant differences between male and female infants in interactions with kin versus nonkin. There was only one significant difference between male and female infants in interactions with males versus females: female infants showed stronger preferences for initiating proximity with females over males than did male infants. Because mothers provide the focal point for infant interactions during the first year of life, we compared the behavior of infants and mothers. Mothers were the recipients of more social interactions than were infants, mothers engaged in more grooming than did infants, and infants engaged in more social play than did mothers. These findings are only partially consistent with kin-selection theory, and the inadequacies of studying matrilineal kin discrimination to test kin selection are reviewed. The near-absence of infant sex differences in associations with social partners suggests that although maternal kin other than the mother are important to infant socialization, they probably do not contribute to the development of behavioral sex differences until after the first year of life.     

Circadian and Ultradian Time Patterns in Human Behaviour: Part 1: Activity Monitoring of Families from Prepartum to Postpartum

Time patterns of activity-rest rhythms during and after pregnancy are increasingly recognised as important factors for the well-being and health of young families. This longitudinal study examined activity-rest patterns of couples during late pregnancy and subsequently the alterations in the periodic structure of parental and neonatal time patterns during the first four months after birth. Part I concentrates on the effects of late pregnancy and birth to the mother's rest-activity patterns and those of the father and, after birth, what time pattern the infant developed. Part II attempts to clarify how activity patterns of the entire family agree or disagree with each other and investigates how the infant synchronises with the environment that includes the process of parent-infant interaction. Activity data of, so far, seven families (father, mother and child) were continuously recorded using non-invasive Actiwatch units. Recordings of parental activity started at the beginning of the 37th week of gestation, and were continued in parallel with the infants' recordings in three series of three weeks each until four months after birth: 1st to 3rd week, 7th to 9th week and 13th to 15th week of life. In a standardised diary, record was kept of household routines, parental activities, type of feeding, initiation of sleep or waking up. Activity data of seven non-pregnant women were collected and used as a control. Irregular nocturnal activity epochs occurred frequently in pregnant women and were absent in non-pregnant women. Period lengthenings and shortenings of the circadian rhythms appeared in both parents from prepartum to postpartum. Activity at night increased from prepartum to postpartum in mothers and fathers. Three infants showed a marked circadian rhythm between day 3 and 14 after birth. All seven infants showed a predominant circadian rhythm between day 8 and 19 after birth. The onset of daytime activity of mothers and their infants corresponded well to each other. Postpartum frequency spectra of parents and child always had some ultradian components in common. Time patterns of activity-rest rhythms of couples and parents are shown to be altered during and after pregnancy and we suggest that the infants' adaptation to the environment begins during the first week that includes the process of mother-infant interaction.     

Circadian and Ultradian Time Patterns in Human Behaviour: Part 1: Activity Monitoring of Families from Prepartum to Postpartum

Time patterns of activity-rest rhythms during and after pregnancy are increasingly recognised as important factors for the well-being and health of young families. This longitudinal study examined activity-rest patterns of couples during late pregnancy and subsequently the alterations in the periodic structure of parental and neonatal time patterns during the first four months after birth. Part I concentrates on the effects of late pregnancy and birth to the mother's rest-activity patterns and those of the father and, after birth, what time pattern the infant developed. Part II attempts to clarify how activity patterns of the entire family agree or disagree with each other and investigates how the infant synchronises with the environment that includes the process of parent-infant interaction. Activity data of, so far, seven families (father, mother and child) were continuously recorded using non-invasive Actiwatch units. Recordings of parental activity started at the beginning of the 37th week of gestation, and were continued in parallel with the infants' recordings in three series of three weeks each until four months after birth: 1st to 3rd week, 7th to 9th week and 13th to 15th week of life. In a standardised diary, record was kept of household routines, parental activities, type of feeding, initiation of sleep or waking up. Activity data of seven non-pregnant women were collected and used as a control. Irregular nocturnal activity epochs occurred frequently in pregnant women and were absent in non-pregnant women. Period lengthenings and shortenings of the circadian rhythms appeared in both parents from prepartum to postpartum. Activity at night increased from prepartum to postpartum in mothers and fathers. Three infants showed a marked circadian rhythm between day 3 and 14 after birth. All seven infants showed a predominant circadian rhythm between day 8 and 19 after birth. The onset of daytime activity of mothers and their infants corresponded well to each other. Postpartum frequency spectra of parents and child always had some ultradian components in common. Time patterns of activity-rest rhythms of couples and parents are shown to be altered during and after pregnancy and we suggest that the infants' adaptation to the environment begins during the first week that includes the process of mother-infant interaction.     

Evolution and the sudden infant death syndrome (SIDS)

This paper extends the evolutionary and developmental research model for SIDS presented in previous articles (McKenna 1990a, 1990b). Data from variety of fields were used to show why we should expect human infants to be physiologically responsive in a beneficial way to parental contact, one form of which is parent-infant co-sleeping. It was suggested that on-going sensory exchanges (touch, movement, smell, temperature, etc.) between co-sleeping parent-infant pairs might diminish the chances of an infantile cardiac-respiratory crisis (such as those suspected to occur in some SIDS cases). In this article we review recent epidemiological data and sleep research findings on SIDS to show how they relate to evolutionary and cross-cultural perspectives. Results of a preliminary study of the co-sleeping behavior of mother-infant pairs indicate that, with respect to sleep, arousal, and respiratory patterns, co-sleeping mother-infant pairs affect each other in potentially important ways. We suggest specifically that co-sleeping may shorten periods of consolidated sleep among young infants by causing them to arouse more frequently. Moreover, we suggest that partner-induced arousals might help the infant to confront sleep crises more competently. In the long run, these arousals might prevent the premature emergence of prolonged (adultlike) sleep bouts from which some infants have difficulty arousing—especially during a breathing pause or apnea.     

Prefrontal asymmetry and parent-rated temperament in infants

Indicators of temperament appear early in infancy and remain relatively stable over time. Despite a great deal of interest in biological indices of temperament, most studies of infant temperament rely on parental reports or behavioral tasks. Thus, the extent to which commonly used temperament measures relate to potential biological indicators of infant temperament is still relatively unknown. The current experiment examines the relationship between a common parental report measure of temperament--the Infant Behavior Questionnaire-Revised (IBQ-R)--and measures of frontal EEG asymmetry in infants. We examined associations between the subscales of the IBQ-R and frontal EEG asymmetry scores recorded during a combined series of neutral attentional and putatively emotional recording conditions in infants between 7 and 9 months of age. We predicted that approach-related subscales of the IBQ-R (e.g., Approach, Soothability) would be related to greater left prefrontal asymmetry, while withdrawal-related subscales (e.g., Distress to Limitations, Fear, Falling Reactivity, Perceptual Sensitivity) would be related to greater right prefrontal asymmetry. In the mid- and lateral-frontal regions, Approach, Distress to Limitations, Fear, Soothability, and Perceptual Sensitivity were generally associated with greater left frontal activation (rs≥.23, ps<0.05), while only Falling Reactivity was associated with greater right frontal activation (rs≤-.44, ps<0.05). Results suggest that variability in frontal EEG asymmetry is robustly associated with parental report measures of temperament in infancy.     

Babies sleeping with parents: case-control study of factors influencing the risk of the sudden infant death syndrome. CESDI SUDI research group

ObjectiveTo investigate the risks of the sudden infant death syndrome and factors that may contribute to unsafe sleeping environments.DesignThree year, population based case-control study. Parental interviews were conducted for each sudden infant death and for four controls matched for age, locality, and time of sleep.SettingFive regions in England with a total population of over 17 million people.Subjects325 babies who died and 1300 control infants.ResultsIn the multivariate analysis infants who shared their parents'' bed and were then put back in their own cot had no increased risk (odds ratio 0.67; 95% confidence interval 0.22 to 2.00). There was an increased risk for infants who shared the bed for the whole sleep or were taken to and found in the parental bed (9.78; 4.02 to 23.83), infants who slept in a separate room from their parents (10.49; 4.26 to 25.81), and infants who shared a sofa (48.99; 5.04 to 475.60). The risk associated with being found in the parental bed was not significant for older infants (>14 weeks) or for infants of parents who did not smoke and became non-significant after adjustment for recent maternal alcohol consumption (>2 units), use of duvets (>4 togs), parental tiredness (infant slept ⩽4 hours for longest sleep in previous 24 hours), and overcrowded housing conditions (>2 people per room of the house).ConclusionsThere are certain circumstances when bed sharing should be avoided, particularly for infants under four months old. Parents sleeping on a sofa with infants should always be avoided. There is no evidence that bed sharing is hazardous for infants of parents who do not smoke.

Key messsages

• Cosleeping with an infant on a sofa was associated with a particularly high risk of sudden infant death syndrome
• Sharing a room with the parents was associated with a lower risk
• There was no increased risk associated with bed sharing when the infant was placed back in his or her cot
• Among parents who do not smoke or infants older than 14 weeks there was no association between infants being found in the parental bed and an increased risk of sudden infant death syndrome
• The risk linked with bed sharing among younger infants seems to be associated with recent parental consumption of alcohol, overcrowded housing conditions, extreme parental tiredness, and the infant being under a duvet

     

Supply and Demand Determine the Market Value of Access to Infants in the Golden Snub-Nosed Monkey (Rhinopithecus roxellana)

According to a biological market paradigm, trading decisions between partners will be influenced by the current ‘exchange rate’ of commodities (good and services), which is affected by supply and demand, and the trader’s ability to outbid competitors. In several species of nonhuman primates, newborn infants are attractive to female group members and may become a desired commodity that can be traded for grooming within a biological market place. We investigated whether grooming was interchanged for infant handling in female golden snub-nosed monkeys (Rhinopithecus roxellana) inhabiting the Qinling Mountains of central China. R. roxellana exhibit a multilevel social organization characterized by over 100 troop members organized into 6–11 one-male units each composed one adult male and several adult females and their offspring. Behavioral data were collected over the course of 28 months on grooming patterns between mothers with infants less than 6 months old (N = 36) and other adult female troop members. Our results provide strong evidence for the interchange of grooming for access to infants. Grooming for infant access was more likely to be initiated by potential handlers (nonmothers) and less likely reciprocated by mothers. Moreover, grooming bout duration was inversely related to the number of infants per female present in each one-male unit indicating the possibility of a supply and demand market effect. The rank difference between mothers and handlers was negatively correlated with grooming duration. With increasing infant age, the duration of grooming provided by handlers was shorter suggesting that the ‘value’ of older infants had decreased. Finally, frequent grooming partners were allowed to handle and maintain access to infants longer than infrequent groomers. These results support the contention that grooming and infant handling may be traded in R. roxellana and that the price individuals paid for access to infants fluctuated with supply and demand.     

Variations in care for cottontop tamarin, Saguinus oedipus, infants as a function of parental experience and group size

We examined the effects of parental experience and group size on infant care and development. Ten cottontop tamarin families were followed across three consecutive births of offspring to examine differences in survival, quality of care, developmental maturation and physical development for infants in two experimental conditions: (1) in large groups with previous parenting experience and (2) in small groups with little or no parenting experience. Although we found no differences in infant mortality, parents treated infants differently between conditions. However, when we examined the cumulative care provided by parents and sibling helpers together across experimental conditions, there were no differences in the rates of retrieval or rejection of infants. Overall levels of infant transport and food transfers were similar between conditions, with fathers and sibling helpers contributing the bulk of care. Fathers in small groups carried infants and transferred food with infants much more than helpers but, as group size increased, helpers augmented this role. Infant development varied with the experience of the family both between conditions and within a family. The results indicate that infants receive similar levels of care regardless of group size and parenting experience, although the individual contributions of family members change with both factors. Parental experience with previous infants had effects on early infant development, but only for the first sets of infants. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Separation of preadolescents from infant rhesus monkeys

Each of eight infant rhesus monkeys was paired with a preadolescent conspecific for two months and then separated. Four of the infants were mother-reared and four were isolate-reared. Separation responses were compared with data from preseparation and reunion phases of the study for all pairs. The results indicate that (1) although males interact with infants in a parental fashion, preadolescent females show a greater capacity for parental behavior, (2) both preadolescents and normal (mother-reared) infants contribute to the development of a social bond, (3) isolate infants contributed little to the development of a social bond and were relatively less-valued as social partners by their preadolescent cagemates.     

Group and matriline differences in the behaviour of rhesus monkey infants

A ‘Monte Carlo’ method was developed for characterizing matrilines of rhesus monkey infants (Macaca mulatta) born to their mothers within six similar captive social groups, according to behavioural measures, and for characterizing social groups according to their matrilines' median scores. Eight- and 16-week-old infants differed according to their groups, but not according to their matrilines, in measures of their activity in the vicinity of their mothers, and the frequencies with which social companions other than the mother initiated social contacts with them while they were out of contact with their mothers. Sixteen-week-olds differed according to their groups in the frequencies with which their mothers rejected them, and 8-week-olds in the frequencies of the social contacts they initiated with companions other than mother while off their mothers. When tested with their mothers, in a mildly disturbing situation away from other social companions, 52-week-olds differed according to their group of origin in time spent out of contact with their mothers. Different groups were seen as producing infants differing in ‘enterprise’ from their eighth week, in the sense of being ready to initiate social contacts with others while off their mothers at 8 weeks, and of being off their mothers in the test at 52 weeks. Cases where the 8-week measures did not reflect infant enterprise could be explained in terms of other aspects of the relationships involving the infants and their mothers in the social group. In particular, matrilines differed consistently in dominance status, and some mothers also received high levels of aggression from other adults. Mothers receiving high levels of aggression were more responsible for maintaining proximity with their 8-week-old infants, and their 8-week-olds were less often involved in social contacts with others.