Mother-Infant Relations in Free-Ranging Chimpanzees

Growing Independence, Conflict and Learning in Mother-Infant Relations in Free-Ranging Chimpanzees Author(s): Hedwig H. C.
van de Rijt-Plooij and Frans X. Plooij Reviewed work(s): Source: Behaviour, Vol. 101, No. 1/3 (May, 1987), pp. 1-86 Published by: BRILL Stable URL: http://www.jstor.org/stable/4534590 . Accessed: 17/01/2012 10:07
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.
BRILL is collaborating with JSTOR to digitize, preserve and extend access to Behaviour.
http://www.jstor.org
GROWING INDEPENDENCE, CONFLICT AND LEARNING IN MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

by HEDWIG H. C. VAN DE RIJT-PLOOIJ1) and FRANS X. PLOOIJ2) (Department of Physical Anthropology, University of Cambridge, England and Department of Developmental Psychology, University of Nijmegen, The Netherlands, respectively)
(With 28 Figures) (Acc. 28-IV-1986)
Contents
1. Introduction ........................................
2. General methods ........................................
.......................
.......................
2
5
A longitudinal study on single subjects ............................................. Study area and research centre ....................................................... ....................... Study subjects ........................................ ....................... Sampling ........................................ Observation methods .................................................................... 3. Results ........................................ ....................... Part A: Care of neonates .............................................................. Part B: Conflict over ventro-ventral contact ......................................
5 6 6 7 8 10 10 19
1) The fieldwork was carried out in the Gombe National Park, Tanzania, from May 1971 to March 1973. We are indebted to the authorities of the Tanzanian National Parks for permission to work in the area, and to Dr J. GOODALL providing facilities in the for Gombe Stream Research Center. We wish to thank Dr P. GARLICK, Head of the Department of Physical Anthropology of the University of Cambridge, England, for his constant sympathy, advice and encouragement. We acknowledge with gratitude Prof. F. J. M6NKSand Prof. C. van LIESHOUT for permitting us to use the facilities of the Department of Developmental Psychology of the University of Nijmegen, The Netherlands. We are grateful to Prof. G.
Dr M. J. A. SIMPSON Mr F. VELDMAN for their stimulating and constructive and Sr. discussions. The analysis of the data was partly supported by a grant to the first author from the "Fonds Doctor Catharine van Tussenbroek" in the Netherlands. 2) The second author received grant nr 56-83 from the Netherlands Organization for the Advancement of Pure Research (ZWO). Corresponding address: Gemeentelijk Pedotherapeutisch Inst., Amsterdams Pedologisch Centrum, IJsbaanpad 9, 1076 CV Amsterdam, The Netherlands.
P. BAERENDS, ROON, Prof. R. A. HINDE, Prof. A. KORTLANDT, DrJ. M. BAERENDS-VAN
H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ 35 41 62 62 63 66 68 73 79 80
Part C: First excursions ................................................................ Part D: Conflict over body contact .................................................. 4. G eneral discussion .......................................................................... Labile periods at specific ages ........................................................ Labile periods and mother-infant conflict .......................................... Regression and maturation ............................................................ Developmental steps, new types of learning and conflict periods ............ Developmental steps and attachment ................................................ ........................ Summary ........................................ References ........................................................................................
1. Introduction Up to now, we know less about conditions which facilitate normal development than we know about the adverse effects of factors such as marital discord, parental rejection, felt maternal incompetence, institutional upbringing (RUTTER, 1978a, 1980a). Not all insecurity, children have to be damaged by such "bad" circumstances and reasons why they do not may be found not only in the characteristics of the individuals themselves, but also in the balance of good and bad influences they experience as they develop (RUTTER, 1978b). This is all the more reason to know more about the conditions which facilitate normal development. Knowledge of these conditions during infancy seems to deserve highest priority, because this age-span may be a 'sensitive period' as far as certain aspects are concerned. The idea of 'sensitive periods' in development implies that certain events at a particular developmental phase have a greater effect than do the same events at any other phase (Bateson, 1979, 1983). It also implies that certain events have to occur during the 'sensitive' period if normal development is to proceed thereafter. There are few studies which suggest the existence of clearcut 'sensitive' periods in human development (RUTTER, 1979). However, it seems that the development of attachment in early childhood could be an exception to
these findings (RUTTER, 1980b).
A condition which facilitates normal development is a continuing mother-child relationship. Over the years the evidence for the importance of such a relationship has grown considerably (BOWLBY,1969,
1973, 1980; DOUGLAS, 1975; RUTTER, 1981).
One aspect of this relationship is physical contact between mother and child. The occurrence of such contact is not restricted to the nonhuman primate mother-infant relationship. The class of mammals can be
MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES
divided in 4 groups according to the type of contact regulation between mother and offspring: nesting, cacheing, carrying and following. There are convincing arguments to assume that man, together with the
monkeys and apes, belongs to the carrying type (BLURTONJONES, 1972).
Different aspects of the role of body contact in development have been shown. The necessity of body contact for the healthy development of infant chimpanzees and rhesus monkeys is impressively demonstrated by
HARLOW(1958), HARLOW& HARLOW(1965), MASON (1968), and DAVENPORT (1979). The comforting aspect of contact in anxiety-provoking situations has been established experimentally (HARLOW & ZIMMERMAN,
1959). HINDE and coworkers have experimentally established that brief mother-infant separations of 1-2 weeks still have behavioural consequences two years later (HINDE & SPENCER-BOOTH, 1971; HINDE, 1977).
Authors who take a cross-cultural view of human infancy have addressed the importance of body contact (how much time, by whom, and in what position) with respect to the nipple access, the potential for interaction
and so on, e.g. FREEDMAN(1974) and KONNER (1976).
As the infant grows older, he begins to crawl and walk away from mother further and to stay away for a longer timespan. This growing independence has been described not only for monkeys (HINDE, 1974a)
and apes such as gorillas (FOSSEY, 1979), orang-utans (HORR, 1977) and 1968; CLARK, 1977; NICOLSON, chimpanzees (VAN LAWICK-GOODALL, 1977; PUSEY, 1978a, 1980), but also for man (SHIRLEY & POYNTZ, 1941; RHEINGOLD & ECKERMAN, 1970). Thus, it may be expected that a better
understanding of the processes at work during the development of contact- and distance regulation in free-living chimpanzees, ultimately will be of benefit for such a study in man. This is all the more true, if one knows that there have been very few nonlaboratory observations of
human attachment behaviour during the last 10 years (BRETHERTON,
1985). The longer this state of affairs lasts, the more difficult it becomes to assimilate new data or to design incisive research, according to the same author. "Longitudinal studies are wanted that chart in more detail the epigenetic pathways that produce the patterning of attachment relationships" (BRETHERTON,1985). Also MAIN & GOLDWYN(1984) stress the
need for a detailed description of the interactional processes underlying the formation of normal as opposed to pathogenetic attachment relationships. Our study aims to provide such a description. Of course, in order to understand such processes in the human mother-infant relationship one must study this relationship. An ethological study such as ours can only contribute to human studies by providing methods of observing and
H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ
analysing behaviour, and by providing hypotheses that can be tested

(BARNETT, 1977).
Many researchers have stressed the usefulness of conceptualizing the

mother-infant dyad as a system (BOWLBY, 1969; SANDER, 1969; PETERFREUND, 1971; THOMAN, 1975; BATESON, 1976; ALS, 1979). Positive as
well as negative feedback was shown to occur in rhesus-monkey motherinfant interactions (HINDE, 1983). The study to be reported in this article can be viewed upon as taking such an approach to chimpanzee motherinfant contact- and distance regulation. In taking a systems approach, we assume that the mother-infant dyad behaves as an organized whole (WATZLAWICK et al., 1967) characterized by mutual modification of each other's behaviour in response to feedback. In this study we aim to assess the influence of changes in the behaviour of mother or infant on the growing independence of the infant. We will follow the infants' development towards greater independence in individual chimpanzee mother-infant pairs when mothers and infants vary and when they contribute to changes in the mother-infant system. We will look at the history of mother-infant relationship in terms of the effects of changes in the mother's and infant's behaviour, and the way in which the occurrence of these changes affects what happens next. To be able to do so, we will search for age-changes in mother-infant contactand distance regulations and in profiles of maternal behaviours concerning the qualities of each 'make' and 'break' of contact. We expect that changes in mother's and infant's behaviour are associated with: 1. A jump in the independence of the infant. 2. A marked change in the quality of the infant's independence.
This article is mainly organized according to the infants' age and increasing distance from its mother. In section 2 (General Methods) the study area, the mother-infant pairs, the way of sampling, and the observation methods are described. In section 3 the results are presented, divided in four parts. Part A is centered around the perinatal period. First it is shown that an inexperienced primiparous mother has to learn how to handle her newborn baby. Then the consequences of changes in maternal caregiving activities for the baby's growing independence are described. Part B is centered around the age of 5 months. At this age first maternal rebuffs were aimed at warding the baby off the nipple and off the belly. Attention is paid to the consequences of these maternal changes for the onset of ventral riding and eating solid food. Part C is centered around the age of 8 months. Here the changes in the mother-infant relationship surrounding the onset of frequent excursions are described. Part D is centered around 10-11 and 18 months. At these ages regressive behaviour in the infant preceded maternal rebuffs aimed at breaking body contact. Attention is paid to the consequences of these changes in the mother-infant dyad for the infant's growing independence.
The results of each part are discussed. The paper ends with a general discussion (section 4) of the consequences of the findings for the better understanding of the processes involved in the growing independence of free-living chimpanzees.
2. General methods
2.1. A longitudinal
study on single
subjects.
It is our opinion that the study of developmental processes (such as the ones involved in the growing independence) needs research strategies which are somewhat different from the usual designs. This opinion is based on two arguments. First, in the naturalistic setting, like the present one, it is virtually impossible to apply the experimental design involving control- and experimental groups. In the Gombe National Park only one or two babies were born each year. These were observed longitudinally at frequent intervals (section 2.4). Secondly, DENENBERG (1978, 1982) discussed the limitations of experimental and correlational procedures using group data for the understanding of the behavioural development of single individuals and suggested procedures that allow the researcher to determine meaningful statistical parameters for individual subjects. This single subject research design involves the following assumptions. Subjects are studied longitudinally in a naturalistic or semi-naturalistic setting. The observations are sufficiently detailed so that quantitative values can be derived, such as frequency of occurrence of events or percentage of time during which an event occurred. The number of subjects is limited but sufficient to allow a test for generality. The kind of statistical parameters we looked for in our subjects are the following: Patterns over time in individual graphs. Having derived quantitative values from detailed observations, these will be plotted against age. Then, patterns over time in these graphs will be assessed. Within-subjectcomparison. "Natural experiments" occur, which can be used to advantage, provided that infants are observed frequently and for long periods. By comparing certain quantitative values of a number of observation sessions preceding and following the natural experiments it can be examined whether and how patterns in these values over time associate with these natural experiments. A statistical testfor generality. According to DENENBERG (1978) two of the arguments leveled against the use of single be readily tested and easily rejected. and chance relationship-can subjects-uniqueness Each infant can be looked upon as being a separate experiment. If the same pattern of parameters found in one infant is also found in others, this will be sufficient to establish that the phenomenon generalizes beyond one subject. DENENBERG (1978) states that the occurrence of the pattern in 2 out of 50 cases is sufficient to reject the null hypothesis at the .05 level. The null hypothesis is that a particular pattern does not exist in the population. This implies that, if we find the same pattern in all 2 or 3 infants whom we are able to compare at any age, it is all the more convincing that such pattern has some degree of generality. Of course, to define the degree of generality more accurately, it is necessary to study large numbers of infants. However, even if a study population with sufficient suitable subjects were available, it could not be done economically without a hypotheses (1982) puts it: "...you have to have generating study preceding it. Or, as DENENBERG it in order to get started." The present study will generate hypotheses for further study on larger numbers, because this study enables one to define the questions to be asked more specificly in order to obtain data more economically.
centre. 2.2. Study area and research The study was carried out in the Gombe National Park. The park lies on the shore of lake Tanganyika in northwestern Tanzania. The general structure of the park, the climate, flora and fauna have been described by VANLAWICK-GOODALL (1968), CLUTTONBROCK(1972, 1975) and WRANGHAM (1975). The research centre has been described by VANLAWICK-GOODALL (1967, 1968, 1973). In 1960 GOODALLinitiated research at Gombe and free-ranging chimpanzees (Pan troglodytes schweinfurthii) have been studied since. An artificial feeding area was set up in 1962. The disadvantages of the feeding methods used at that time and the change in the manner of feeding after 1968 have been described concluded that size and form of the comby WRANGHAM (1974). In 1975, WRANGHAM munity range and the importance of natural food sources were not greatly influenced by the artificial feeding schedule in the years 1971-1973. We conducted our study from May 1971 till March 1973. Additionally, PLOOIJ(1984) suggested that the influence of the artificial feeding on the association between mother-offspring pairs was minimal if not absent in those years.
+ WK WL
PS + PF FF + FD
4
15 20
ML+ GM PL + PT
month 5
7 10
10 15 20
24 24
25
Fig. 1. Age spans of the mother-offspring pairs. 2.3. Study subjects. 2.3.1. The mother-offspring pairs. We studied five mothers with offspring under 24 months of age. The age-spans over which each individual was studied are given in Fig. 1. Two pairs had a third individual associating with them for 100% of their time. These and other personal data relating to the study-subjects are presented in Table 1.
TABLE 1. The study subjects

Mother Winkle Passion Fifi Melissa Pallas Initial WK PS FF ML PL PAIR MOTHEROFFSPRING Initial Offspring Wilky Prof Freud Gremlin Plato WL PF FD GM PT Sex m m m f m Birthdate 21-10-1972 25/28-10-1971 21/24- 5-1971 14/23-11-1970 7- 9-1970 Goblin GB m juvenile 1Pom PM f juvenile I Sibling MEMBER FAMILY DEPENDENT Sex Initial Age-class
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
FF + FD was the only pair known to have extended family ties. These are described in Table 2.
TABLE 2. Family ties of FF + FD

Kin of FD Grandmother Uncle Uncle Uncle Name Florence*) Flint*) Figan Faben Initial FLO FT FG FB Age class Adult Juvenile Adult Adult
*) Flo and FT died when FD was 15/16 months old, resp.
2.3.2. Age classes. The terms baby, infant, juvenile, adolescent and adult are used throughout the text. They refer to the following age-classes: baby 0-6 months, infant l/2-5 years, juvenile 5-9 years, adolescent 9-14 years, adult > 14 years. Except for the terms baby and infant, use of the terms is the same as VANLAWICK-GOODALL's (1973). The age-classes 'baby' and 'infant' were distinguished by PLOOIJ (1984) on the basis of major behavioural changes that were observed. 2.4. Sampling. Each mother-offspring pair was observed monthly around the day when the baby or infant was 0.5, 1.5, 2.5, etc. months old. During and 'observation session' the pair was followed on foot at a mean distance of 5-10 m. All pairs usually tolerated being followed. When a mother really wanted to be on her own, she could easily leave her human followers behind, by moving only a little faster through the often dense and thorny undergrowth. Each mother-offspring pair was 'followed' until 300 min of 'good observation' were obtained. Periods of 'bad observation' occurred when it was impossible to see mother and/or offspring, due to the nature of the field situation. The frequency and the duration of such 'bad observations' periods varied with the location of the chimpanzees and with the seasons. During the wet season all trees were in leaf and the grass was high as a result of which 'bad observation' periods were frequent and long lasting. To be able to collect 300 min of 'good observation' each mother-offspring pair had to be followed for 2-3 days. Usually a 300 min sample would require 2 days of following. A split-half analysis showed that this sample size provides reliable results (VAN DE RIJT-PLOOIJ, 1982). Observations were dictated into a cassette-tape with the help of an audio cassetterecorder (Sony 1 1A).
2.5. Observation
methods.
Some of the steps involved in building a conceptual framework for the analysis of social systems have been specified by HINDE (1983). The main characteristics of HINDE'S approach are the following. 1. The study of social behaviour must begin with a solid descriptive base. Description of social behaviour in physical terms (i.e. the form of communicatory signals) is usually embedded in descriptions by consequence of longer episodes. The physical, social and
temporal contexts in which the social behaviour is embedded should be described as well. The description of the interactions include what the individuals are doing together (content), how they are doing it (qualifiers), who initiates and who breaks off. The patterning of interactions over time leads to qualifiers that are difficult to deduct from isolated interactions. 2. Social behaviour must be studied at several, different levels, including those of individual behaviour, interactions, relationships and social structure. Each of these levels may show emergent properties not relevant at lower levels. 3. Dyadic relations are of central importance. On the one hand, there is the 'dialectic' between the personality of the individuals and the relationship, through the medium of interactions. On the other hand, the relationship is embedded in a social group or -system which is made up of a network of dyadic, triadic and higher order relationships. In this paper we focus mainly on the mother-infant interactions. This is a simplification. Although we are fully aware of this, we feel this is justified, since PLOOIJ (1984) has shown that a mother-baby pair is alone for approximately 90% of the time. 4. The four types of explanation of the observed phenomena (namely: causal-, functional-, developmental-, and evolutionary explanation) are in principle of equal interest. The details of the application of this approach to our study is extended upon in the next few subsections. 2.5.1. Distance betweenmotherand offspring. It has been shown that the growing independence of the offspring is reflected in the distance between mother and offspring (RHEINGOLD & ECKERMAN, 1970; HINDE, 1971, 1974a; ALTMANN, 1980). The following distance categories, following those used in HINDE & SPENCER-BOOTH'S (1967) observation system, were used: I = Ventral: the infant is ventral to another individual if most of its weight is supported by that individual and the greater part of its body is in front of and encompassed by the arms and thighs of the other individual. "Ventral" includes sitting on the ground between the thighs and facing the other individual. II = Not-ventral: this includes any other position of the infant on the body of another individual (when most of its weight is supported by that individual), when it is not ventral. E.g. on the other individual's back, head, lower-arms and lower-legs, and outer surface of the upper-arms and -legs. III = In contact: most of the infant's weight is supported by the ground or vegetation, but the infant's body is touching the other individual's body. Excluded are the times when the infant is sitting in between the thighs of another individual and facing that other individual (then the infant is said to be ventral). IV = Within arm's reach: the infant is within arm's reach of another individual if it is out of contact of that other individual and if the other can touch the infant by extending his or her arm. This distance varies a little with the circumstances, but usually it is approximately 1.5 meters. V = Within five meters: the infant is within five meters but out of arm's reach of another individual. VI = Within fifteen meters: another individual is over five meters but less than fifteen meters away from the infant. VII = Visible: another individual is visible but over fifteen meters away from the infant. VIII = Out of visibility: another individual is out of visibility but closely around the infant. For instance behind a bush from the point of view of the infant.
The distance between mother and offspring was scored in terms of these distance categories. A time-scale with 15-sec time-markersas smallest units was used. The percentage of time that was spent in each category was assessed from these original records through a scan-sampling technique, which allows adequate sampling of durations, so long as the bout lengths of the behaviour involved are long relative to the sampling interval (SIMPSON& SIMPSON, 1977). Comparing
sampling using the 15 sec time markers with the assessments using progressively longer
the assessment resulting from the scan-
time-units it was shown that a 5-min interval provides reliable results (VAN DE RIJTPLOOIJ,1982).
This distance measuring system could be used immediately upon arrival in the Research Centre, because the subjects studied were well habituated. In this study these distance measurements will be the starting point for further analysis of the contact- and proximity regulations of both partners over time. These will be described in the next few paragraphs.
2.5.2.
Regulation of contact and proximity.
Distance measures alone do not help to explain the roles of mother and offspring in maintaining proximity. The relative contributions by mother and offspring to this maintenance can be assessed by employing a function that takes into account the proportion of approaches or leavings over a given distance that are due to one partner or the other (HINDE & ATKINSON, 1970). Roles in the maintenance of ventro-ventral contact can under 'Observation methods' in parts B, C and D of the results in section 3.
also be assessed (HINDE & WHITE, 1974). More details of these functions will be given
2.5.3.
Maternal punishment.
These functions, in their turn, cannot help to answer the question whether, for instance, detectable aggression directed by the mother towards her infant plays a decisive role in the growing independence of the infant. In order to answer this question, observations are needed of the circumstances in which the approaches and leavings occurred. For example, whether or not a mother signals before leaving her infant may be relevant information about the act of leaving. Similarly, whether or not her infant whimpers when his mother leaves him could determine whether or not his mother waits when the infant next approaches. Furthermore, the quality of a mother's action, such as whether she breaks contact by shoving her baby aside or by punching him could be important extra information. In order to be able to record the circumstances in which the acts of approaching and leaving (or making and breaking contact) occurred, the observation system should enable one to record as much of the behaviour of mother and infant as possible. Such a system was developed during a pilot-study and is described and illustrated by PLOOIJ (1984). Additionally, the circumstances were defined in terms of certain combinations of the behaviour-elements as these preceded, were associated with or followed an approach (or make) or leave (or break). These definitions are given in the sections entitled 'Observation methods' of the appropriate parts of the results. The only point that remains to be mentioned here is that the circumstancesare defined in such a way that the behavioural elements used in the definitions may change over age without the circumstances changing name. For instance, a mother may gather her 12 month old infant before travelling, whereas she may signal her 18 month old to approach her. In both cases the circumstance 'Retrieval' is scored. A similar approach was
10
advocated by SROUFE & WATERS(1977), in their study of the quality of attachment relationships.
3. Results Part A: Care of neonates Through the first 2 months a chimpanzee baby is extremely helpless and dependent upon his mother, according to PLOOIJ (1984). He described how a multiparous mother, after becoming physically handicapped, failed to hold her baby efficiently, as a result of which the baby died. Since maternal behaviour is of such vital importance in these early months, we describe the (prenatal- and postnatal-) part played by the mother. Furthermore, we want to know at what age the baby is able to carry his own weight by clinging to his mother. As a result, the baby's helplessness should diminish. This should have important consequences for maternal caregiving activities. Therefore, maternal behaviours related to the initiation, maintainance and breaking of ventro-ventral contact will be studied from birth towards month 6. In doing so, the individual differences in both babies and mothers will be considered.
A.1. A.l.1.
Subjects
and methods.
Subjects. During my study two babies were born. The multiparous mother PS (Passion) gave birth to a son PF (Prof). The primiparous mother WK (Winkle) gave birth to a son WL (Wilky). At the beginning of the study, the primiparous mother FF (Fifi) had a baby son of one week of age, named Freud (FD). WK + WL was the only pair that was observed intensively shortly before and after birth. Therefore, the description of pre- and early post-natal maternal behaviour will be restricted to this pair. As our field study started when FD was one week old, the observations on this baby were not as detailed as the observations on the other two babies. Consequently, FF + FD's curve of initiating and maintaining ventro-ventral contact is not plotted in Fig.
3.
A.1.2.
Methods.
At the beginning of her pregnancy WK was very shy. To be certain that she would tolerate humans following her closely and looking at her when she had her baby, fieldassistents followed her more and more often until they were with her daily, 15 days before she delivered her baby. The following behavioural measures were used.
11
Frequency of effort grunts. The number of 'effort grunts' per 100 mins of good observation while the observer was within a distance of 5 m. An 'effort-grunt' is an explosive expel of air resulting when breath, after being held while making some physical effort, is released. May be vocalized. Mouth and lips are slightly open. PLOOIJ (1984) showed that the frequency of 'effort grunts' can be used as a measure of the activity level of a baby.
Securing. The percentage of the total number of location-changes that were preceded by pressing the baby into the ventro-ventral position. Thereafter, the mother may support or hold him there. A location-change was defined as the mother changing from one body-position to the other (e.g. from sitting to standing) or from a body-position to a way of locomoting. The decision to analyse this measure was taken after having observed WK + WL on their first day: on occasions, when WK moved after having remained for some time in one position, the baby immediately fell back and screamed. That very day WK learned to 'secure' the baby before moving (subsection A.2.2.5). After having made these observations on WK + WL and looking at the data collected from other pairs studied before, it struck us that every mother, when she 'secured' her baby, did so especially before moving. When they did so their babies seemed to have less difficulty in maintaining their ventral position afterwards.
A.2. Observations. A. 2.1. Thefinal days of pregnancy. Until shortly before WK gave birth, no changes were observed in the distance she travelled in a day, nor was she seen to avoid large mixed groups (males, receptive females and mothers with infants). Two days before parturition she began to avoid social activity, being alone except when she spent a short time near one mother-infant pair. WK was alone throughout the final days of her pregnancy. Moreover, she was far less active on these last days (see PLOOIJ, 1984). She walked slowly and rested often. The day before she gave birth, she slept repeatedly. Five times she was seen to make a day-nest and altogether she slept for a total of almost two hours during the day. At 17.55, WK built a nest in a palmtree. The place she chose for it was surrounded by dense crowns from adjacent trees. It was remarkable that this nest was much larger than those chimpanzees usually make. Somewhere between 18.00 that day and 12.20 the next day the baby (WL) was born. A.2.2. Mother with newbornbaby. The birth was not observed. From 6.00 until 10.00 no Inovement or sounds were observed coming from the nest. Then there was a slight
12
movement. At 12.20 a faint baby-scream was heard when WK shifted weight. Because the nest was built in dense vegetation it was difficult to find a point from which to make the observations. From 13.00 onwards WK and the baby could be observed from the crown of a tree about 15 m away.
A.2.2.1.
The umbilical cord and afterbirth.
The baby's body seemed dry and the attached umbilical cord was about 50 cm long. For the first 1 hour and 19 mins of observation, before she fell asleep, WK licked the baby and the umbilical cord continuously, and she sucked at the end of the umbilical cord. Once she picked leaves and ate them during the sucking sessions. She also bit the umbilical cord close to the baby's body. At no time was the baby not almost completely encircled by mother's body, legs and arms. The afterbirth was not seen. It would have been, had WK thrown it out of her nest after 6.00. When WK, later that day, left the place of birth, the nest and environment was searched thoroughly, but nothing was found. As soon as WK woke up after 33 mins, she continued licking and sucking the umbilical cord. She also began to bite and chew on the cord, and she pulled at it with her incisors and canines. As the day wore on biting and chewing the umbilical cord replaced licking and sucking more and more. Bouts of chewing on a quantity of leaves after biting and chewing the umbilical cord became more frequent. Often WK licked her fingers. Sessions of biting and chewing the umbilical cord, chewing on leaves and finger licking were seen until sunset. The umbilical cord was gone by the next morning.
Mother paid attention to baby. At 16.23 of the baby's first day a striking session in which WK was seen to inspect her baby was observed. During this session WK held the back of his head firmly in her right hand, while she touched his body-parts with quick grooming-like movements. Special attention was paid to the face. While looking at parts of this face intensively, she turned his head in all possible directions. At the end of the session WK looked at and touched the baby's genitals. Inspection was interrupted by short embraces of the baby. When WK stopped her inspection the baby was seen to root. WK did not help him by pressing him to her breast. That day sucking was not observed. It was remarkable that grooming sessions were absent, though over the day quick grooming movements on the baby's head and shoulders became more frequent. However, the contexts in which these movements were seen reflected so clearly mother's conflict between using her arms to carry the baby and using them to locomote and climb, that it seems likely that they were primarily displacement activities (KORTLANDT, 1940; A.2.2.2.
TINBERGEN, 1940).
A.2.2.3.
Mother avoided social contacts.
On the day of parturition WK left the birth-nest for the first time at 15.38. She did not come down to the ground but stayed in the crowns of the adjacent trees where she ate palmnuts and figs. At sunset she made a new nest within 15 m of the birthnest where she and the baby spent the night.
13
During the second day, WK also spent most of her time around the place of birth. Six times she made a nest, every time within 15 m of the birthnest. She was seen asleep regularly and left the nest only for short intervals when she fed. That day it rained almost continuously and WK collected the water from her hair to drink. On WL's birthday, as on most other days, 'panthoot' calls which are individually distinctive (VANLAWICKGOODALL,1968) were to be heard all around. Although WK listened to all the calls and she was able to recognize the calling individuals, she never responded by calling or approaching them as she would have done a few days earlier. Also in the afternoon of the second day, chimpanzees were heard to 'panthoot' in the distance. Judging from the increased vocalizations from one direction a large group was gradually forming. In the afternoon, three mothers with infants came into sight of WK and her baby. When they were more than 20 m away they all stopped to look intensely at the new pair-one mother and her infant peering continuously for more than half an hour before travelling on. WK did not seem to pay attention to anyone, but gave loud 'panthoots' as a reaction to vocalizations from the group in the distance. After the three pairs left, WK began to travel in the same direction, following the three mothers, but at such a distance as not to be seen by them. By this time, WK seemed to have become uneasy in our presence, so we increased our distance to over 50 m and lost her. Human observerswho were present with the group that called from a distance reported that although the three mother-infant pairs who had passed WK + WL in the afternoon joined this group later. WK did not so in the first 7 days of WL's life (our own observations, and observations by Bauer, van der Stoep and Wrangham). However, on the baby's sixth day, WK + WL were again near the group. Although WK could just hear the others, she did not join in social activities with them. After day 7, WK began to interact socially with other. A.2.2.4. The baby was helpless. Because the baby was covered by WK's arms and legs almost continuously, it was impossible to observe details. For example, although we saw the infant suck for the first time on day 2, this could have happened before. In the two days during which the baby was observed, he was rarely seen to move and most times these were only slight movements. Once, the baby was seen to move more vigorously when he flexed and stretched one leg. In so doing, he progressed a little on his mother's body. Through the first two days of life the baby often seemed distressed. Whenever his mother moved he lost close contact with her and gave 'infant-screams'. He seemed unable to hold onto his mother without assistance. Whenever WK shifted weight he fell back and away from her. While WK carried him, he was often resting in her hand with his own hands relaxed and open. Seldom did he grasp her hair when being 'supported'. A.2.2.5. Mother adjusted herself to the baby and maintained ventro-ventral contact. Over the first day, WK gradually learned to secure the baby's ventral position (by pressing the baby into the ventro-ventral position and holding him there) before moving (see definition in subsection A. 1.2). At first WK did not secure her baby in any of these ways, but her progress seemed to be guided by the baby's utterances of distress. In the birthnest the baby was heard to scream as soon as WK moved. WK immediately stopped all movements and pressed the baby against her. Over the first hours WK increasingly more often secured the baby in the ventro-ventral contact position before moving. Next, similar adaptations were seen in the context of climbing and walking and every occurrence of WL's 'distress' corrected WK when she omitted to secure him. By the end of the second day, WK always secured her baby in ventro-ventral contact before moving
off.
14 A.2.2.6.
Inexperienced mother. From observations made on WK's first days, it became obvious that the degree of maternal efficiency increased with experience. WK 'supported' and 'carried' the baby but she appeared to have difficulty doing these things while walking and climbing. She found keeping her balance while walking along horizontal branches especially difficult. Often she was seen to take an extra hold of branches while she moved slowly through the trees making sure of each new hand or foothold before relinquishing the last. The following example illustrates her conflict and the unusual way she solved her problem. WK approached a horizontal branch while carrying the baby in the ventral position. When she put her hand on the branch she immediately took her hand back and sat down. She looked repeatedly at the baby and the branch while making a few quick grooming movements on the baby's head (this sequence was repeated three times). Then WK scratched herself, rubbed her nose and got up while holding WL against her ventrum with her right hand. The baby did not grip at all. She extended this hand (i.e. the hand with the baby still on it) slightly in front of her and in this manner she balanced to the other side. Here she sat down immediately and restored ventro-ventral contact. This was the only time WK was seen to solve her problem by breaking ventro-ventral contact. Later that day she used a different type of locomotion when moving over horizontal branches: the 'sloth-position' (VAN LAWICK-GOODALL, 1968). By hanging underneath the branch she could use both hands and feet to move along the branch while her baby was supported by her ventrum. It is notable that VANLAWICK-GOODALL (1968) mentions that the 'sloth-position' is used by infants under 2 years of age when they are still insecure when moving in branches. She presumed that the sloth position enables them to cling on more safely.
A. 2.3. From 'securing'to breakingthe ventro-ventral position. In this section, we will illustrate how maternal 'securing' changes over age in two mother-baby pairs: WK + WL and PS + PF. On the one hand, it may be expected that the changes depend on the baby's growing ability to make and maintain ventro-ventral contact for himself. Also individual differences between the babies may play a role in the amount of 'securing' they receive. On the other hand, the individual differences between the mothers may influence their amount of 'securing' over age. Therefore, we will first describe the differences between both babies and mothers and compare them with more babies and mothers. A. 2.3.1. Individual differences.
Gombe chimpanzee babies could not be removed from their mothers, nor could they be touched. Therefore, their individual differences could not neonatal assessment scale (1973) as was be assessed with the BRAZELTON done for captive chimpanzee babies by TURNEY(1978). PLOOIJ (1984)
15
described a way to establish individual differences in newborns in their natural environment. All newborns uttered 'effort-grunts' and they always and only did so when they were moving or exerted muscle effort. By comparing the frequency of 'effort-grunts' of two babies, he found a clear difference in the percentage of time each baby spent moving. Additionally, the baby who spent more time moving appeared stronger in that his 'effort-grunts' were given more loudly and clearly. If WL's frequency of effort-grunts is compared with these frequencies of the other two babies PF and FD, it becomes clear that WL was the least active baby (Fig. 2). Moreover, the effort-grunts were soft and weak (i.e. pants rather than grunts). These results suggest that newborns can vary considerably in strength. Therefore, it is understandable that some newborns could support themselves for short periods while others not at
all (VAN LAWICK-GOODALL, 1968; NICOLSON, 1977).
Within each baby's own limits he is able to secure ventro-ventral contact himself: he can hold on to his mother as much as his strength permits. A mother, especially if she is experienced, should be able to adjust herself within a wider range of limits. She can avoid physical and social situations potentially leading to the kinds of strenuous activity that could interfere with her holding the baby onto her body. She can also adapt her
2Fig.
2.
Individual
differences
in
frequency
of
effort-grunts.
50
180
Fig. 2. Individual differences in frequency of effort-grunts.
16
mode of locomotion in a variety of ways to support her baby in difficult situations. Here we consider the question: how far does she allow herself to adjust to her baby? Two of the three mothers, WK and PS, were very different in this respect. WK appeared exceptionally cautious. When walking, she supported and carried her baby almost continually both with exaggerated flexion of her thighs and by keeping one hand under his back and shoulders. She walked slowly and sat down often. When sitting she craddled the baby almost invariably, keeping her knees bent up, one arm or hand behind his shoulders, and her feet close together under his rump. On the other hand, PS was seen to allow the baby to lie on his back on the ground for several minutes and she allowed him to release his handand/or footgrips when she removed her supporting hand while walking. It is apparent from VAN LAWICK-GOODALL (1968) that PS behaved in the same way towards the baby's elder sister Pom (PM). Moreover, this author mentioned that PS's behaviour was least efficient (compared to three other mothers) despite the fact that PM was not her first baby (the first baby had disappeared soon after birth). In conclusion, of all Gombe mothers observed to date, PS has been least solicitous, while WK was at the other extreme.
100 90 t80 70 60 50. \ 3, \ \ \
20 .
10t
1 month 2
If
3
\;
44 5
PS
6
Fig. 3. The decrease of the % of securing over age.
A.2.3.2.
Changes in 'securing' over age. Fig. 3 shows the percentages of 'securing' made by WK + WL (months 1-4) and PS + PF (months 1-6). In line with the descriptions of the individual differences in both babies and mothers (see above) is the finding that WK + WL's amount of 'securing' was higher from months 1-4 then was PS + PF's. Despite these individual differences, both curves show the same sharp decrease between months 2-3.
17
A. 2.3.3.
Breaking ventro-ventral contact. Two of the three mothers were observed to break ventro-ventral contact deliberately in order to transport their baby in the dorsal position: PS and FF. When PF was 3-5 days old, PS pushed him up to her back for the first time. Perhaps it is because he could not cling properly that she was not seen to repeat these trials.
While walking, PS pushed PF sideways with her left arm. PF clung to her side, his right hand and foot gripping the hairs on her back. PS had to hold him in that position with the back and side of her left arm. When she sat down PF lay across her left thigh facing the ground. PS stood up, walked and pushed him all the way up on her back with her left arm and hand. PF showed distress, PS sat down with her hand behind her holding PF up. PF slipped down however and clung onto her side. Then PF rooted against her side and called out in distress. PS pulled him a little way towards her ventrum. PF now screamed and PS pulled him to the full ventral position.
When PF was 8 weeks old, PS tried again to transport him in the dorsal position. Although it still appeared difficult for the baby to hold himself in that position, PS persisted. From this age onwards, PF could be seen regularly on PS's back during travel.
PS turned her back towards PF, put her hand under his rump and pushed him up (scooping) onto her back from the ground as she moved off. She scooped him in similar ways several times during that day, but PF invariably slithered down her side. Within a week PF was able to maintain the dorsal position for a while before slipping off.
At the age from which PS was seen to transport PF regularly in the dorsal position, FF was seen to initiate dorsal riding for the first time. FF carried FD dorsal for over thirty paces when FD was 8 weeks old
(WRANGHAM, pers. comm.). FF lifted FD head first, over her shoulder whilst she was sitting. Then she stood up with FD facing her rump. As she walked she lowered her head and this gave FD more support. For the first few paces FF kept her hand near FD and she seemed to walk more slowly than usual. FD stayed dorsal and did not show distress. He looked back for the whole time and did not change his position. After about 20 m she sat down and took FD ventral.
Though seemingly successful, FF was not seen to lift FD dorsal again until he was 13 weeks old. FF started to try to transport FD in the dorsal position at a later age than PS did. From 13 up to 19 weeks of age, FD was carried dorsal increasingly more often. At this age FD was able to hold himself on his mother properly with both hands and feet. Similar observations were made for PF from week 12 onwards.
18
It was notable, however, that both PF and FD, after having been lifted onto mother's back, were not yet able to change their position. They clung tightly to the mother in whatever position they happened to be. As a result, they were regularly carried in a reversed position and looked back for the whole travel-session. A.3. Discussion. It is quite likely that WK consumed the afterbirth and we saw her eat the umbilical cord. Although many primate mothers consume the placenta and umbilical cord soon after expulsion (POIRIER,1977), not all chimpanzee mothers are reported to do so. For instance, VAN
LAWICK-GOODALL (1968)
reported that one mother (Melissa)
carried her
newborn baby (Goblin) around while the umbilical cord and afterbirth were still attached. These became black and fell off without having been eaten. POIRIER'S (op. cit.) suggestion that "placental consumption might satisfy the mother's appetite for a few days" would be supported if mothers who did not eat the afterbirth ate more and sooner after the birth. However, detailed observations on feeding behaviour of several more mothers are not yet available. WK was hardly seen to eat for 3 days. The cleaning and licking of the neonate, which WK did extensively, is not uncommon among non-human primates (BRANDT & MITCHELL, 1971) and was even reported to occur in some human cultures (LINDBURGH & HAZELL, 1972). It may be one of the factors binding the mother to her infant, as has been shown for some non-primate mammals
(PoIRIER, 1977).
The session during the first day in which WK was seen to pay special attention to her baby (touching body parts, looking at the face, inspecting the genitals) may be another factor playing its role in early mother-infant attachment. Similar behaviours have been reported for humans during the first hours after birth (KENNELL al., 1975). The importance of the et quick formation of a strong attachment of the mother to her baby becomes all the more clear if one considers the initial inefficiency of this primiparous mother in securing ventro-ventral contact. It took this mother 2 days to solve the problems she met in combining locomotion with maintaining ventro-ventral contact efficiently. During these two days, it often seemed that the baby was at risk of falling far enough out of a tree to be killed. Once a mother was able to maintain ventro-ventral contact efficiently, the most marked change in neonatal caregiving was the sharp decrease
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
19
in maternal 'securing' between month 2 and 3 in the two mother-baby

pairs observed. STEINBACHER'S (1941) very detailed descriptions on one
chimpanzee mother-baby pair in captivity also reveal that the baby was not able to maintain the ventro-ventral position for himself until he was almost 3 months old. We propose that the mothers forced their babies to carry their own weight by clinging to the mother's hair. This is illustrated in the actual breaking of ventro-ventral contact by both mothers and the initiation of dorsal riding. Although both mothers initiated dorsal riding at the age of 8 weeks (one tried earlier but failed), only one mother continued forcing her baby to ride dorsal once in a while whereas the other did not. The baby that was not forced to ride dorsal was not seen to do so until one month later, when his mother tried again. These observations are supported by the independent data of STEINBACHER (op. cit.). He observed the first careful trials to break ventroventral contact in the captive pair from the baby's 7th week onwards. He describes the gentle way with which the mother tried and, next, her failure to complete a first break, because the baby did not 'cooperate' but kept clinging to the ventral side. The mother did not force a break, but from this age onwards she was seen to repeat her trials increasingly often. From month 4 onwards, the babies we studied 'cooperated' for the first time when their mothers forced them into the dorsal position. The mother who stopped forcing her baby between the age of 2 and 3 months succeeded in the beginning of month 4, because now her baby no longer struggled. The baby who was forced into the dorsal position earlier (8 weeks), not only stopped struggling during month four but also ceased to whimper (PLOOIJ,1984). This suggests that a 3 months old baby no longer feels insecure when being put in the dorsal position. Part B: Conflict over ventro-ventral contact
Two mother-baby pairs, with 5 month-old babies, were observed. Both mothers were 'aggresive' to their babies. 'Aggression' between mother and infant has been described in captive chimpanzees (YERKES,1943). In the wild, however, 'maternal aggression' towards infants under three years of age has not yet been reported. With older infants, it is rare for
a mother to show aggression (VAN LAWICK-GOODALL, 1968; CLARK,
1977). The maternal rebuffs towards the 5 months-old babies occurred only in the context of ventro-ventral contact, and they were aimed at warding the baby off the nipple and off the belly.
20
In this part we will describe the role of mother-infant conflict in the context of ventro-ventral contact. To be able to do so we will first examine the mother-baby interaction prior to the onset of the maternal rebuffs. For this purpose the changing role of both partners in regulating ventro-ventral contact will be examined during the age range of months 1-6. Our analysis is continued beyond month 6 to check whether motherinfant conflict over ventro-ventral contact continued.
and methods. B.1. Subjects B.1.1. Subjects. In this chapter, data on two mother-infant pairs with male offspring will be analysed and compared. Freud (FD) with his mother Fifi (FF) and Prof (PF) with his mother Passion (PS) were observed monthly from 1-20 and 1-15 months of age, respectively. B.1.2. Observation-methods.
The following behavioural measures were used: On the nipple. The number of times the baby took the nipple in his mouth per 300 mins of good observation. This measure does not take into account the length of each bout of nipple-contact. Time in ventro-ventral contact. Percentage of total number of 5-minute scan-samples that the pair was in ventro-vental contact (see section 2: General Methods). The relative roles in making and breaking. The relative contributions of each of the partners in 'breaking' ventro-ventral contact is the proportion of total 'breaks' due to the infant (%BKi) and the mother (%BKm). The relative contributions of the partners in initiating ventral contact ('makes' or %MKi and %MKm) were assessed in the same way (HINDE, 1974a). Control of ventro-ventral contact. The percentage of all mother-infant 'makes' which were due to the infant's behaviour minus the percentage of all mother-infant 'breaks' which were due to the infant (% MKi%BKi) is regarded as a measure of the infant's control of ventro-ventral contact in this paper. A negative value is regarded as indicating that the mother is taking the primary role in maintaining ventro-ventral contact, and a positive value is showing that the infant is taking the primary role in maintaining ventro-ventral contact. The properties of this measure are discussed further by HINDE & ATKINSON (1970) and HINDE & WHITE (1974). The final five measures are concerned with the qualities of the 'makes' and 'breaks'. They take into account the circumstances in which a 'make' or 'break' occurred. They are frequencies per 300 mins of good observation. Retrieval. A retrieval was recorded whenever a mother turned towards or went to her infant who was out of ventral contact, picked it up and pressed it ventral. A retrieval was also recorded whenever a mother signalled to her infant who was out of contact to make ventral contact. The signals were: looking at, reaching for, grooming leaf, leg bending, grunting and lowering back. The difference between this measure and the next measure is that
21
in this measure interaction between the pair-members was omitted once contact was made. Attention. Immediately after ventro-ventral contact was made (whether by the infant or by the mother), the mother attended to the infant by playing, kissing, embracing, and looking at. The following two contexts in which 'grooming' occurred were also recorded under 'attention'. First, maternal grooming when the infant made intention movements for breaking ventro-ventral contact again, and second, maternal grooming when the infant started sucking immediately upon making ventro-ventral contact. Grooming. The mother groomed the infant immediately after ventral contact was made by either herself or the infant. Rejection. The following three contexts were considered to belong to rejection. The mother prevented her infant from making ventro-ventral contact. The mother refused to interact with the infant after the infant made ventro-ventral contact and initiated the interaction. The mother broke ventro-ventral contact immediately after the infant made this contact. Aggression. The number of times the mother broke ventro-ventral contact by slapping, punching, biting, pushing or stamping on the infant.
B.2. Observations. B.2.1. Maternal aggression. In month 5 and 6 respectively, the mothers, FF and PS, were observed to change their behaviour towards their babies drastically by biting, pushing, pulling, pinching, and slapping them repeatedly (Fig. 4). The following example illustrates FF's behaviour towards FD.
When FD was 19 weeks (i.e. month 5) old, FF's behavior towards him changed strikingly. She had always been the more patient mother and was more sensitive to FD's demands than PS was to PF's. Now she sat down to rise again and jump up and down repeatedly. She suddenly acceleratedfrom standing still as if to shake FD loose. FD clung tightly with hands and feet to the ventral side and was not dislodged on these occasions. Alternatively, FF pushed, pulled and bit FD away from her ventral side, while having a full open grin and giving cough threats. Whenever FD lost his grip, he immediately took hold again. Finally, FF had to tear him loose to lift him onto her back. FD tried to struggle back into the ventral position, but FF prevented this by pushing him away backwardswith her elbows. Then FF stood up and walked away while FD was clinging to her back.
Such sequences were seen repeatedly on that day, as were aggressive acts of rejection when FD began to suck. Sometimes these nipple rebuffs were combined with ventro-ventral rejection because FD repeatedly tried to reach the nipple while being pushed away. It was notable that FF was aggressive to FD only when they were in ventro-ventral contact. A second point worth mentioning is that these aggressive encounters all oc-
114
.. Maenlages, Fig 4.. er ab.
Fi.4 atrargreso"o,e
_ig Ma to as .
ay
Fig.4.
ateral
ggresio to er aby
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
23
curred during one morning and were not seen on the near preceding or the other succeeding observation days by us or other observers. This will be discussed further below (subsection B.2.3.1.). PS's aggressive behaviour seemed less dramatic in that she did not push, bite and pinch PF in persistent bouts, but only once in a while. Moreover, the episodes of aggression were not restricted to one morning but occurred over a longer timespan (months 6 and 7). contactand the relativerolesof motherand baby. Controlof ventro-ventral To understand how the mother-infant relationships changed through time, we need to know more than that all mothers sometimes rejected their babies aggressively. First, how did mother and baby influence ventro-ventral contact from birth up to the time the mothers were seen to behave aggressively? We will show that the mother was totally responsible for maintaining ventro-ventral contact in the first month of life. The baby became more and more responsible over the first half year. By the time the mother behaved aggressively, the baby was more responsible than the mother for the first time in its life. This was associated with the onset of dorsal riding and eating solid food. Second, we will show that after month 6 the infant's responsibility for ventro-ventral contact fluctuates with age. The peaks are associated with peaks in maternal 'rejection' and 'aggression' and with dips in maternal 'retrieval', 'attention' and 'grooming'. The measure in Fig. 5 shows the relative contributions of mother and baby to maintaining ventro-ventral contact. The measure shows an increase in control of ventro-ventral contact for both babies from most negative (month 1) to positive (month 6). However, the points in this measure could have been achieved in different ways and on its own the measure does not explain how the baby's responsibility increased from month one to six. The increase could have been due to an increase in 'makes' or to a decrease in 'breaks'. The measures in Table 3 show the relative roles of mother and baby in, respectively, 'making' and 'breaking' ventro-ventral contact from month one to six. In relation to these measures the following four points are worth mentioning. First, during month 1, the babies were initiating all the 'breaks' and the mothers all, or most of, the 'makes'. In combination with Fig. 5 this means that the mothers of both babies were primarily responsible for ventro-ventral contact. Second, the increase over the first six months in both babies' responsibility for the maintenance of ventro-ventral contact (Fig. 5) was due to B.2.2.
24

+40 % - 'BKI MK
+20
+ 10
1'
-40 -50 -60 -70 -80 / /
-100
month
10
12
14
16
18
20
Fig. 5. Baby's role in ventro-ventral contact (%MKi-%BKi) over age.
the increase in their roles in 'making' ventro-ventral contact. Although the measures in Table 3 also show a decrease in 'breaking' ventro-ventral contact, the increase in 'making' this contact was much more pronounced. Third, the increase in 'making' ventro-ventral contact by both babies was such that the mothers were no longer initiating the majority of 'makes' from month 3 (PS) and month 4 (FF) onwards (Table 3). This means that now the babies began to be more responsible for 'making' ventro-ventral contact. It should be noted that, at these very ages, both mothers began transporting their babies regularly in the dorsal position (subsection A.2.3.3.). Fourth, the increase in both babies' responsibility for 'making' ventroventral contact from months 1 to 6 was such that the babies were
TABLE
3. The proportion of total ventro-ventral contact 'makes' and 'breaks' due to the baby
FD %Makes 0 30 40 50
79
Age Month 1 2 3 4
5
PF %Breaks 100 100 96 87

92
%Makes 33 40 63 62
79
%Breaks 100 80 86 88
88
100
87
82
72
25
initiating (nearly) all the 'makes' in month 6 (Table 3). Also, at this age, the measure in Fig. 5 was positive for both babies for the first time. This indicates that now the babies were primarily responsible for ventroventral contact for the first time in their lives.
B.2.3.
The onset of frequent dorsal riding and eating solidfood.
In the previous section we saw that the mother's role in determining the time she and her baby spent in ventro-ventral contact decreased from month one onwards. In the fifth and sixth months the mothers aggressively forced the babies to leave the ventral side. In this section the relation between these extreme rebuffs at this age and the onset of frequent dorsal riding and eating solid food will be described.
From ventral to dorsal transport. FF (who always seemed to us the more sensitive mother) was more persistent in her attempts to shake FD loose (subsection B.2.1.). FF used a variety of means: she slapped, pushed, bit and pinched him. These aggressive incidents occurred in sessions often lasting several minutes. All aggressive sessions occurred during one morning and were not seen the immediately preceding or succeeding observation days by us or other observers. Moreover, after one hour these sessions occurred less frequently and the duration of each decreased rapidly. Thereafter, FF and FD were seen to play again as usual. Later in the day, FF was observed to offer FD an alternative to being pushed away from the ventral side before travelling. FF 'taught' FD the behaviours that were to serve as 'signals' for climbing dorsal. Here follows a description of this observation: B.2.3.1.
FD was walking on a rock (about 50 cm high) while FF was sitting out of contact but within arm's reach. Then FF stood up slowly, turned her back towards him and approached him while flexing her knees slightly and looking back at him with her lowered back closest to him. FD did not cling immediately and FF waited motionless while looking at him. Finally, he clung and FF walked a few paces to turn around and come back to the same rock to return him onto the rock. Over and over she started travel in this way and when FD did not walk onto the rock himself, she placed him there. Finally, FD clung immediately whenever FF 'signalled' by flexing her knees and looking back. Next, FF started the whole procedure again, with FD on the ground. She lowered her back by crouching onto the ground in front of him while looking back at him over her shoulder. Whenever FD seemed not to be looking at FF she would wait and, ultimately, gain his attention by touching him. When touched in this way he always responded by clinging immediately. 'Signalling' in this context was also repeated several times and gradually slight variations were made.
26
FF was very patient with FD. She sat down with him and waited until he tottered off himself. A baby of this age can hardly walk and FF had to wait for several minutes before FD was far enough away to make it worth her while to signal again. On the first day, FF was seen to use different combinations of the following signals: looking at, reaching for, leg bending, grunting, lowering back. Moreover, she used these signals in a variety of situations: when being on the ground on the same height, when being in a tree, when being with other individuals, and when being alone. FD was seen to agree to these signals by climbing dorsal himself. This apparently tedious 'teaching' and 'learning' of signals had advantages to both partners. Having learnt to understand the signals, FD showed that he accepted the dorsal position and FF no longer needed to be aggressive to prevent ventro-ventral contact. PS was never able to prevent PF from making ventro-ventral contact first, and so she continued to push PF round onto her back after he made ventro-ventral contact with her. The onset of eating solid food. In month 5 (18 weeks) FD was seen to chew and ingest food for the first time'). He had obtained this food from FF by taking it away with his mouth. It had been impossible to observe swallowing movements at an earlier age, although FD was seen to chew on objects such as leaves two weeks earlier. The very day that FD was first seen to be pushed, bitten and pulled off the nipple (month 5; 19 weeks), he had then imitated FF's feeding behaviour twice, by taking, chewing and swallowing leaves of the species she was then feeding on. From this week onwards, FD was seen to collect 'food' for himself. FF was seen to inspect the objects which FD had gathered for himself to eat, especially at the end of month 5 (21 B.2.3.2. weeks). Similar results were obtained for PS and PF. Although PF chewed and ingested food in month 4 (15 weeks), it was only in month 5 (18 weeks) that PS allowed him to take some of her food. He was trying to obtain PS's food from month 4 (16 weeks) onwards, but PS invariably kept her food out of his reach. For PS and PF we were able to oberve the frequency of making nipple contact over the age range of 3-9 months. The results in Fig. 6 show an
1) For both babies tooth eruption had commenced in month 4 (between the ages of 13 to 15 weeks). FF had occasionally been seen to inspect FD's jaws from month 3 (week 11) onwards. At the end of month 5 at least 7 incisors were present in both babies.
27
increase in the number of times PF made nipple-contact, reaching a peak in month 6. Controlof ventro-ventral contactand the relativerolesof mother and infant. The onset of dorsal riding does not mean that the infants were never seen in ventro-ventral contact with their mothers thereafter. The measure in Fig. 7 shows that the percentage of time spent in ventro-ventral contact is still relatively high after month 6. From this we may conclude that ventro-ventral contact plays a part in the mother-infant relationship after six months as well. In this section the control of ventro-ventral contact and the relative roles of mother and infant in 'making' and 'breaking' from month six onwards will be examined. B.2.4.
"/300 mins. 50
:30 20 10 /
Fig. 6. Frequency of nipple contact-makes per 300 mins of good observation.
9o
70 % 60 50
-'
, PF
o3
v month 2 4 6 8
-vv 10 12 14 16 18 20
Fig. 7. Amount of ventro-ventral contact over age.
The measure in Fig. 5, showing the relative contribution of mother and infant to maintaining ventro-ventral contact, fluctuates with age. Over the first twenty months that FF + FD were observed, three peaks in the infant's responsibility can be found, respectively in month six, eleven and fifteen. The first two peaks in FD's responsibility for ventro-ventral contact (Fig. 5) are due to increases followed by decreases in his role in 'making'
28
this contact (Tables 3 and 4). This was concluded from the finding that the differences between the 'makes' of the particular months and the 'makes' of the preceding and following months were greater than the differences between the 'breaks'. The third peak in FD's responsibility for ventro-ventral contact, however, was due to mother FF's greater role in 'breaking' ventro-ventral contact. This can be concluded from the finding that the aforementioned differences between the 'breaks' are greater (Table 4).
TABLE
4. The proportion of total ventro-ventral contact 'makes' and 'breaks' due to the infant
FD %Makes 80 74 68 67 93 88 67 88 82 81 85 85 81 72 %Breaks 71 83 96 85 81 82 92 88 72 92 94 100 100 94 %Makes 93 88 76 95 96 94 97 85 83 PF %Breaks 64 69 58 78 74 67 64 83 65
Age Month 7 8 9 10 11 12 13 14 15 16 17 18 19 20
For PS and PF a similar picture arises, the infant reaching peaks in months seven and thirteen. Because data on PS and PF are available until month fifteen only, a possible third peak cannot be shown. However, month fifteen shows an increase towards this possible third peak. The two peaks in PF's responsibility for ventro-ventral contact (months 7 and 13; Fig. 5) are also due to increases followed by decreases in his role in 'making' this contact (Tables 3 and 4). However, the increase in his role in 'making' this contact leading to the second peak (month 13) had occurred in month 10. Thus, his role in 'making' this contact remained high for the 4 months 10 through 13. Notwithstanding the similarities, there exist striking differences between the two pairs. First of all, from month 7 onwards, the measures of PS + PF lie much higher (Fig. 5). It was most remarkable that their
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
29
measure remained positive from month 6-15. This means that it was mainly up to PF to maintain ventro-ventral contact with PS from month 6 onwards. FF + FD's measure, on the other hand, was negative over age except for the three peaks mentioned above. The measures in Table 4 also reveal differences between the two pairs that add to a better understanding of the differences described above. Through months 7-15, mother PS broke contact proportionally more than did mother FF. The making of contact shows a reversed picture. Mother PS made contact proportionally less than did mother FF, except in month 14 when PF seemed ill. In other words, FF played a greater role in initiating ventro-ventral contact than PS did and PS played a greater role in initiating separation than FF did. interaction. Changesin othermeasuresof mother-offspring Measures of maternal retrieval, attention, grooming, rejection and aggression are among the many aspects of the interaction of chimpanzee mother-offspring dyads. Profiles of the progressive changes in these several measures through the infants' first 15 months show that they vary independently to some degree (Fig. 8). However, the changes can also be related to the peaks in the infant's responsibility for ventro-ventral contact. Thus, FD's 6th month and PF's 7th month were times when this measure first peaked after becoming positive. In these months, both mothers almost ceased to retrieve and attend to their infants for the first time. Then retrieval and attention increased again. Perhaps the peak before the dip in retrieval and attention reflected the babies' increased boldness, where the increase after the dip reflected renewed attention from the mothers as the offspring wandered further away (month 8, 9, etc.). The second peaks in the infant's responsiblity for contact (FD's 11th month, PF's 13th month) coincided with maxima in maternal rejection and peaks in maternal aggression. The profiles suggest that the first and second times in which the infants' responsibility for ventro-ventral contact peaked are different in that on the second occasion increases in rejection and aggression were more noticeable, while on the first occasion decreased retrieval and attention were more obvious. The little rejecting and aggressive behaviour that was observed in month 5 (FF) and month 6 (PS) seemed to reflect irritation at what may have been "inappropriate" demands of 5/6 months old babies. Rejection and aggression were only seen to follow attempts to suck and bids for being carried in the ventro-ventral position. Therefore, it seems likely B.2.5.
30
H.
H.
C. VAN DE RIJT-PLOOIJ
& F. X. PLOOIJ
PS+PF
FF+ FD n/300 mins. T 15 10 _5 5 20 months in which the infant's responsibility for ventro-ventral
contact peaked
5RETRIEVAL
ATTENTION
0 1io
_5L __ GRONOMING
15
S0 _A
i
_ REJECTION
1
l 51 month3 4 5 6 7 8 9 10 11 12 13 14
I
AGGRESSION 15
I
1
5 6 7 8 9 10 11 12 month 4
I
13 14 15
Fig. 8. Profiles of maternal behaviour in association with ventro-ventralcontact over age.
that they reflected attempts at promoting the independence of the infants. High rates of rejection were maintained by PS during months 6-9. This was due to her shoving PF aside or pushing him dorsal before she began to walk. In contrast, FF no longer needed to be so rejecting in the context of ventro-ventral contact. In month 5 she offered FD an alternative to being pushed away from the ventral side. She 'taught' FD the behaviours that serve as 'signals' for climbing dorsal and FD was seen to accept the dorsal position. An attempt to explain the differences between FD's and PF's role in ventro-ventral contact over age (Fig. 5) might have to invoke different aspects of the dyads' interactions before and after month 6. Early on, FF's greater contribution in maintaining ventro-ventral contact is reflected in her higher retrieval and attention rates. Later, it may be PS's more frequent rejection that requires PF to play the greater role in maintaining ventro-ventral contact. B.2.6. Decreasesand increasesin time spent in ventro-ventral contact. Fig. 7 shows the percentages of time that each pair was in ventro-ventral contact at different ages. In both pairs this decreased considerably over the first year. For PS + PF the decrease was a smooth one, but FF + FD showed a marked drop in time spent in ventro-ventral contact between
31
month 7 and 8. This marked drop is associated with changes in FD's excursion behaviour following a further developmental step which will be discussed in part C. The general trend in decreasing ventro-ventral contact over age was sometimes reversed, as in month 10 and 17 for FF + FD. Similar results were obtained for PS + PF. This pair increased the time spent in ventroventral contact in months 10/11. The most striking aspect of these findings is that these increases preceded months of further conflict. We return to consider these temporary increases in ventro-ventral contact preceding further conflict in part D. B.3. Discussion. B.3.1. First conflict over nipple contact. Up to now it has been assumed that free-living chimpanzee mothers begin weaning their infants from the nipple by the infant's second year
of life (VAN LAWICK-GOODALL,1968; CLARK, 1977). In this study mother-
infant conflict over nipple contact was first seen in month five. First conflict over nipple contact coincided with changes in the motherinfant interaction. Up to month 5, the mothers were progressively less responsible for ventro-ventral contact (subsection B.2.2., Fig. 5), culminating in a reversal in responsibility for ventro-ventral contact between month 5 and 6. In this month, the infants were primarily responsible for the first time in their lifes. Especially increases in the infants' role in 'making' ventro-ventral contact were responsible for this change (Table 3). Furthermore, first conflict over nipple contact is associated with changes in the physical development. Fig. 9 presents the growth curve (total body weight) as reconstructed from data on captive chimpanzees given by GAVAN (1971). The rapid (linear) growth in the beginning of the curve abruptly slows down in month 5 (week 19). Furthermore, FD's and PF's milkteeth appeared in months 5-7, when conflict over nipple contact was first seen. The first incisors erupted between 13-15 weeks and by 5 months at least 7 incisors were present. These observations are in accord
with the literature. VAN LAWICK-GOODALL (1968) observed the first erup-
tion of milkteeth in four infants between the ages of twelve to seventeen weeks. The first and second upper and lower incisors were seen to erupt before the age of 7 months. Of course, it is well known that physical maturation is accelerated in captivity (PUSEY, 1978b). However, PUSEY presents data on older chimpanzee. It is our guess that physical growth
32
in the wild cannot be too discrepant from that in captivity at the early age we are discussing. In connection with the appearance of the teeth, it is interesting to note POND'S (1977) paper. She compared many mammal species and found that as a rule the milkteeth erupt at weaning (from the nipple). She argues that lactation is an important factor in making possible a delay in the development of the teeth until the skull and jaws are almost fully
15 WEIGHT (kg)
19
weeks
19 weeks 0 1 AGE (years) 2
Fig. 9. Growth curve (total body weight) reconstructed from data on captive chimpanzees given by GAVAN(1971).
grown. An adequate supply of the postnatal, nutricious mothermilk allows rapid growth. Furthermore, PONDhas shown that lactation provides the young with a buffer against the fluctuations in quantity, quality and toxicity of the normal food supply. In addition, the role of lactation in tiding a baby over periods when he is too small or imperfectly developed to feed on the normal food of the adult members of the species is exemplified by JOLLY(1972). The onset of eating solid food in the two babies described in this chapter was observed at the age of 18 weeks, which was shortly before the first period of conflict over nipple contact. These data, again, are indirectly supported by the literature. VAN LAWICK-GOODALL (1968)
33
noticed the first chew and swallow movements between 14-26 weeks. NICOLSON (1977) even saw one female chimpanzee-baby, Aphrodite, chew solid food by 11 weeks. However, from her study it was not clear whether or not the baby swallowed food she chewed on. Furthermore, NICOLSON found that the mother did not allow Aphrodite to take food until she was 18 weeks old. A similar observation was made by STEINBACHER (1941). This author saw that the baby he studied was not allowed to take food until it was 17 weeks old, though he also saw first chewing and swallowing movements three weeks earlier. The first chewing movements in the two babies reported in this chapter were also observed 2-3 weeks before the mothers allowed them to feed. And, finally, many foods are not accessible to harvest nor assimilable before infants are at certain
stages of physical development (ALTMANN, 1980).
First conflict over nipple contact at the age of 5 months was also suggested by HORR (1977) for orang-utans. He reported that at this age infants are heard to whine after the mother had retired to the nest with them, and sometimes this was heard during daytime rest. He suggested that some of this whining was indicative of maternal withholding of the breast.
B.3.2. Distinct periods of conflict over age. The findings in this chapter contradict the assumption that the process of reducing nipple contact in wild chimpanzees is a gradual one (VAN
LAWICK-GOODALL,
1968; CLARK, 1977). The frequency
distributions
of
behaviours over age associated with mother-infant conflict over nipple contact and ventral riding show marked inhomogeneity. Three peaks were found. These are summarized in a diagram (Fig. 10). The first peak (discussed in the former section) was found during months 5-7, the second during months 11-12 and the third during month 15. All three were characterized by an increase in control of ventro-ventral contact by the infants (Fig. 5), and an increase in maternal rejection and aggression (Fig. 8). These peaks corresponded with dips in 'retrieval' and 'attention' by the mother (Fig. 8). Although the first peak was not explicitly mentioned by VAN
LAWICK-GOODALL
(1968),
it can be reconstructed
from her data.
She
showed that the frequency of sucking increased during Flint's sixth month as if bouts of nipple contact were being interrupted more often. As we reported a similar increase in frequency of sucking for baby PF in the month preceding the first peak (Fig. 6), it is possible that the higher
34
frequency of sucking reflects a similar peak in conflict occurring in Flint.

VAN LAWICK-GOODALL (1968) also showed that the mean length of suck-
ing per feed for the infant Flint decreased during the period when the frequency of his feeding increased. This change in mean length of sucking per feed was also reported for two other infants (Pom and Goblin) during month 5 or 6, respectively. This supports the idea that at this age the
babies described by VAN LAWICK-GOODALLwere in conflict with their
mothers over nipple contact as well.
XXX XXX XXX
XX XX XX
X X X
month
5-7
11-12
15
AGE
Fig. 10. Summary diagram of the peaks in conflict.
Although STEINBACHER (1941) did not discuss distinct peaks in conflict over nipple contact as such, his detailed descriptions reflect the three peaks we found. Moreover, these peaks were found at ages similar to those described in this chapter. In month 6 the frequency of sucking increased, whereas the duration of each bout decreased (the first peak). In month 12, it was remarkable that the infant was now pushed away during sucking (the second peak). Changes in the infant's feeding behaviour were found to occur at the ages of the first and second peak in conflict. At the time of the first peak, the infant was allowed to take small lumps of food and pieces that were chewed by mother first. The time of the second peak was the first time when the mother allowed the infant to take lumps of food as large as those she took for herself. Though we could not find a third peak indicating conflict in this study, we found a further change in feeding behaviour at the age of my third peak. From month 16 onwards the infant accepted all the drinks offered to him by the human caretaker. STEINBACHER concluded that the mother had milk completely now and that the infant had to rely stopped supplying upon other sources for his drinking needs from now on. Finally, HORWICH (1974) reported also fluctuations of high frequency of mother-infant contact in 12 species of monkeys. Furthermore, his review of the literature on primates and other mammals supported these findings.
35
B.3.3. Maternal aggression. The present close study of two mother-infant pairs revealed first appearance of aggressive maternal prevention of sucking at an extremely early age (month 5-7). Before, aggressive prevention of sucking was seen
after the age of 3-4 years (VAN LAWICK-GOODALL, 1968; CLARK, 1977).
Because in the earlier studies the individuals were studied less intensively, it is very likely that relatively infrequent behaviours (one mother was aggressive for just a few hours) were missed. We found that aggressive prevention of nipple contact and ventral riding played a decisive role in the changing mother-infant interaction leading to reducing ventro-ventral contact. Part C: First excursions From month 8 onwards, the infant is able to look for his mother and restore body contact even when she is behind his back and thus out of sight. On such occasions the younger infants did not look for their mothers but walked away from her while whimpering (PLOOIJ,1984). The ability to look for mother and find her implies that the infant should now be able to make excursions regularly. These are defined as those periods of time during which the infant is out of body contact with the mother. The idea that the infant is now able to make excursions regularly is supported by the following finding. From month 8 onwards, the infant no longer whimpers when mother breaks body contact (PLOOIJ,1984). However, now he whimpers as soon as mother begins to increase the distance between them. In this part we will describe the changes in the mother-infant relationship surrounding the onset of regular excursions. For this purpose the changing role of mother and infant in regulating body contact will be examined from the age of first observed contact breaks up to month 9.
C.l. C. 1.. Subjects and methods.
Subjects. In this chapter,data on four mother-infant pairs will be analysed. Three pairs contained male offspring: Prof (PF) with his mother Passion (PS), Freud (FD) with Fifi (FF), and Plato (PT) with Pallas (PL). The fourth pair contained female offspring: Gremlin (GM) with her mother Melissa (ML). C.1.2. Observationmethods.
Most of the behavioural measures used in this part carry the same names as the measures used in the foregoing chapter in order to make comparison more easy. The measures are redefined, though, because they are not concerned with ventro-ventral contact alone, but also with other forms of body contact.
36
Body contact. The baby or infant is in any one of the first three distance categories (I = ventral; II = not ventral but on the body; III = in contact; for further definition see section 2: General Methods) to the mother. Time out of body contact. Percentage of total number of 5-min scan-samples that the pair was out of body contact. The relative roles of mother and infant. The relative contributions of each of the partners in 'breaking' body contact is the proportion of total 'breaks' due to the infant (% BKi) and the mother (% BKm). The relative contributions of the partners in initiating body contact ('makes' or %MKi and %MKm) were assessed in the same way (HINDE, 1974a). Control of body contact. The percentage of all mother-infant 'makes' which were due to the infant's behaviour minus the percentage of all mother-infant 'breaks' which were due to the infant (% MKi%BKi) is regarded as a measure of the infant's control of body contact in this thesis. A negative value is regarded as indicating that the mother is taking the primary role in maintaining proximity, and a positive value as showing that the infant is taking the primary role in maintaining proximity. The properties of this measure are discussed further by HINDE & ATKINSON (1970) and HINDE & WHITE (1974).
C.2. Observations. Initial breakingof mother-infant contact. In this section we will give a description of every observed contact break before the infants (FD and PF) were able to walk quadrupedally on their own. FD was first seen out of contact with his mother when he was twelve weeks old. He himself released his grip on his mother when he reached out into the vegetation. Next, FF got up and took FD onto her ventrum in order to travel. Three days later another contact break, also initiated by FD, was observed. FD released his grip on FF as he sat in front of her on the ground facing her. After about thirty seconds he reached out and held onto FF again. It was remarkable that no further contact breaks between FF and FD were observed until FD was 18 weeks old and able to walk. PS + PF were first seen out of contact when PF was 13 weeks old. In this pair the first break was initiated by the mother who broke contact while shifting weight. When PF found himself out of contact he began to move and PS immediately gathered and embraced him. Two weeks later the second break in contact was observed. This time, PS walked away from PF for the first time. PF did not try to follow PS, but immediately showed distress while whimpering and reaching for PS. At once PS returned to gather him up and she then travelled away. The fact C.2.1.
37
that PS tried to make PF follow her would have been very surprising if PF had not that very day been seen to take his first tottering steps. However, at this age he was not seen to do so without holding his mother. The next day, PS was seen to encourage PF to follow her after she broke contact. She walked a few steps away from him, but was behind PF and out of his sight. PF immediately stopped reaching for objects and looked around while shifting weight. PS looked at him for twenty-five seconds, then reached for him, but PF could not see her. He gradually stopped moving. Next, he slowly pursed and pushed his lips forward into the so called pout-face and finally he began to whimper. As a result PS also whimpered, showing a pout-face, and gathered PF to embrace him. When PF was sixteen weeks old he was observed for the first time to initiate a contact break himself. On this occasion, PF let go his grip on PS as he dangled in front of her from a little branch. Then he stopped all movement and then he reached out and took hold of PS again. In the above-described contact breaks walking and following the mother did not play a role. FD and PF were seen to take their first quadrupedal steps when they were respectively 18 and 19 weeks old. C.2.2. First quadrupedal steps and onset of frequent contactbreaks.
In month five both babies could be seen tottering around and near their mothers. The mothers would keep a watchful eye on them and hurry to their rescue on occasion. Moreover, both mothers sometimes followed their infants around when they went on exploratory expeditions. Finally, both mothers would sometimes walk a few steps from their infants and then wait until the latter rejoined them. Fig. 11 shows the percentage of the total number of five-minute scan-samples that each pair was out of contact up to month 9. In month five all three pairs observed showed contact breaks: FF + FD spent 9 % of the time out of contact and PS + PF and ML + GM only 1%. Although FF + FD and PS + PF already broke contact for the first time one month earlier, as was shown in the previous section, these breaks were too infrequent and/or short to affect the measure as shown here. The curves in Fig. 11 show that three out of four infants began making frequent excursions in month 8 (GM) and 9 (PF, PT). The fourth infant (FD) was exceptional in the way that he began to break contact regularly from month 5 onwards, when he was barely able to walk (Fig. 11). We suggest that FD's onset of regular excursions at such an early age was the consequence of a difference in mothering between his mother and the other mothers. These differences will be described in the next section.
38
H.
H. C. VAN DE RIJT-PLOOIJ
40
& F. X. PLOOIJ
FD 30 /
PT
GM 10 / / /
---ot0 month 1 2 3-4 4 5 67 6
' 7 S 9 9
Fig. 11. Amount of time spent out of body contact.
C. 2.3. Styles of mothering. FF's behaviour towards her infant differed from the other mothers in that she directed her behaviour to that of her infant more than the others did
(PLOOIJ & VAN DE RIJT-PLOOIJ, 1974). For example, she would tickle her
infant after he asked to be tickled by taking her hand and showing a playface. She would stop when his attention was drawn by someone or something else. In a similar way, FF was seen to consider FD's wishes to break contact with her from month 5 to 7. When FD broke contact while sitting beside her, she always stayed on the same spot and looked at him until he restored contact. Only when in contact again, she would move or climb towards the next fruit, one meter further. In similar vein, when FD was walking away from her, she was often seen to follow him and stop in front of him, making it possible for FD to restore contact. On the contrary, the other mothers tended to follow and /or restore contact with their infant only after these showed to be distressed. Also, they were seen regularly to shift weight and/or climb towards the next fruit before restoring contact. On such occasions, all 5-7 month old infants began to whimper. It seems likely that experiences involving insecurity have prevented the three infants from breaking contact as frequently as FD did. No mother was seen to change the above described style of mothering as her infant grew older. Despite this observation, all three infants that did not make regular excursions immediately upon being able to walk did so after month 8. Interestingly, also FD, who's onset of regular excursions began at an earlier age, showed abrupt changes in his excursion behaviour in month 8. These will be described now. Changesin excursionbehaviour. First, a marked increase in the frequency of 'makes' and 'breaks' of body contact (the number of excursions) was found in month 8 (Table 5). C.2.4.
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
39
TABLE 5. The frequency
of 'makes'
and 'breaks' of body contact due to
mother or infant (per 300 mins of good observation)

Age month FF + FD makes breaks PS + PF makes breaks
2 3 4 5 6 7 8 9
> no contact-breaks < 1 38 40 25 104 187 1 39 43 25 106 190 2 2 10 10 5 5 (no contact-breaks) 35 33 78 75
TABLE 6.
1.5 % of time that FF + FD were out of contact < m or 1.55 5 m

Distance category no contact < m 1.5 1.5 5 m
Age month
IV
5 8 11 6 7 10 _____________________________ 8 22 9 31 * = increase in the frequency of excursions.
V
1 4 7 2 2
Second, the amount of time spent on excursions did not increase as much as might have been expected from the above mentioned marked increase in the frequency of excursions (Fig. 11), indicating that FD's excursions were shorter after month 8. Third, the measure in Table 6 shows that FD made use of space closer to mother after month 8. This is concluded from the finding that increases were found in FD's amount of time spent within 1.5 m, whereas the time spent in the area between 1.5 and 5 m from mother tended to decrease. Fourth, though a sharp drop in the amount of time spent on excursions was not found in FF + FD, a sharp drop was found in the amount of time spent in ventro-ventral contact (part B, Fig. 7).
40
H.
& F. X. PLOOIJ
Finally, the measure in Fig. 13 shows a dip in FD's responsibility for body contact in the month prior to the changes described above (month 7). From Table 7 we may conclude that this dip was due to an increase in his mother's proportion of 'makes'. The dip was associated with fewer excursions (Table 5) and with a peak in using space more distant away from mother (1.5< 5 m) when on excursion (Table 6). Two of these changes found in FF + FD's relationship can also be found in PS + PF's. Also PF's measure showed a marked increase in the frequency of excursions in month 8 (Table 5). Furthermore, PF also made excursions less frequently prior to the marked increase when he stopped making excursions in month 7 (Table 5). C.3. Discussion. In month 8 and 9 three infants began to make first frequent excursions and one infant (FD), who already made frequent excursions at that age, showed marked changes in his excursion behaviour in month 8. It was most striking that FD's excursion behaviour no longer differed from the behaviour of the infants who began making first frequent excursions after month 7. Now he made shorter excursions (Table 5 and Fig. 11) and he showed a tendency to make use of space closer to mother (Table 6). It was typical for all 7 months old infants we studied to make use of space within arm's reach of their mothers and to 'make' and 'break' contact
frequently. These characteristics are also described by STEINBACHER
(1941) for the captive chimpanzee infant and he too found that these marked changes occurred in month 8. We propose that the infants' frequent but shortlasting excursions within arm's reach of their mothers express the infants' concern with the distance to the mother after month 7. Also PLOOIJ'S (1984) data on the behavioural development of infant chimpanzees suggest that the infant is able to be occupied with the distance relationship between itself and its mother from month 8 onwards. This means that the infant can now break contact, walk away and return to mother, on its own accord. At the same time, PLOOIJ argues, the infant for the first time realizes the limits of its control over its distance to mother. This means that the infant also knows when it may not be able to return to mother on its own accord. As a consequence it shows concern over its distance to mother. In our study the changes in FD's excursion behaviour in month 8 seem to express such first concern. As a consequence we observed that FD stayed closer to mother and stayed away for a shorter timespan. Also
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
41
human infants are concerned with the distance to the mother after the age
of 8 months (MORGAN & RICCIUTI, 1969; BRONSON, 1972), and the nine
months
olds
possess
the
cognitive
ability
to
perceive
distance
arrangements
(LEMPERS et al., 1977).
Our finding that FD's excursion behaviour before the changes in month 8 was comparatively careless indicates that he was not yet concerned over his distance to his mother. In our opinion, 5-7 months old FD had the opportunity to make use of space more distant from mother and stay away for a longer timespan, because his mother prevented distressing situations by following him, passing him and waiting in front of him. The other three infants did not have a mother who always stopped with whatever she was doing, to accommodate the behaviour of her infant. Therefore, it is understandable that the other infants, after having had distressing experiences of losing mother, began making their first frequent excursions only after being able to prevent losing mother by perceiving and controling the distance between themselves and their mother. Finally, we suggest that in month 8 behaviours such as clinging to mother, holding onto her and being distressed are only evoked in circumstances in which the infant realized that it lost control over its newly emerged ability to regulate its distance to mother. It is interesting that the infants were not upset as long as they themselves were responsible for enlarging the distance, but that they only were so when mother did the leaving. This observation is supported by ESCALONA (1968) who pointed out that the phenomenon 'stranger anxiety' is essentially the fear of losing mother and not the 'fear of strangers'. A stranger appearing on the scene is just one of the conditions in which the fear of losing mother expresses itself most clearly. Part D: Conflict over body contact At around 5 months maternal rebuffs were aimed at warding the baby off the mother's ventrum and nipple (part B). However, the mothers did not seem to object when their 5 month old babies made dorsal contact (distance category II), or sat and leaned against them (distance category III). As a consequence, in the same day one of the mothers would push her baby away after it made ventro-ventral contact, and kiss or play with it when it came to sit in contact without going ventral. It was striking that, around 11 months, the mothers were no longer ready to accept any form of contact by their infants and they used
42
aggressive rebuffs to prevent or break body contact for the first time. In doing so they forced the infants to make excursions. In this part we will describe the role of the first mother-infant conflicts over all forms of body contact. To be able to do so we will examine the mother-infant interaction in the context of all forms of body contact before and after month 11.
and methods. D.1. Subjects D. .1. Subjects. Data on four mother-infant pairs will be analysed and compared. Three pairs contained male offspring: Prof (PF) with his mother Passion (PS), Freud (FD) with Fifi (FF), and Plato (PT) with Pallas (PL). The fourth pair contained female offspring: Gremlin (GM) with her mother Melissa (ML). The ages of the offspring varied between 1-26 months. D.1.2. Observationmethods.
All behavioural measures described in the foregoing part (subsection C. 1.2.) will be used in this part as well. These are concerned with the amount of time in and the control of body contact. The following six measures are concerned with the qualities of the 'makes' and 'breaks' of body contact. They take into account the circumstances in which a 'make' or 'break' occurred. They are frequencies per 300 mins of good observation. Retrieval. A retrieval was recorded whenever a mother turned towards or went to her infant who was out of body contact, touched it or picked it up. A retrieval was also recorded whenever a mother signalled her infant who was out of body contact to make this contact. The signals were: looking at, reaching for, grooming leaf, leg bending, grunting, and lowering back. The difference between this measure and the next measure is that in this measure interaction between the pair-members was omitted once body contact was made. Attention. Immediately after body contact was made (whether by the infant or by the mother), the mother attended to her infant by playing, kissing, embracing, and looking at the infant. The following two contexts in which 'grooming' occurred were also recorded under 'attention'. First, maternal grooming when the infant made intention movements for leaving mother again, and second, maternal grooming when the infant started sucking immediately upon making body contact. Meshing. Maternal coordination to and anticipation of infant behaviour in travel situations. Meshing was recorded whenever the mother started travel immediately after body contact or eye contact was made by her infant. Meshing was also recorded whenever the mother waited for the infant to make contact during travel. Excluded was a mere responsiveness on the part of the mother, e.g. when she waited for the infant after the infant showed distress. Grooming. The mother groomed the infant immediately herself or the infant. after body contact was made by either
MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES Rejection. The following three walking away from The motehr refused initiated interaction. this contact.
43
contexts were considered to belong to rejection. The mother kept her infant (travel-situations) after the latter showed vocal distress. to interact with the infant after the infant made body contact and The mother broke body contact immediately after the infant made
Aggression. The mother broke body contact by slapping, punching, biting, pushing, or stamping on the infant.
D.2. Observations. D.2.1. First conflict over body contact. In month 11 and 12, respectively, the mothers FF and PS were observed to change their behaviour towards their infants. Both mothers were never seen before to behave aggressively when their infants made body contact. Now, FF and PS were seen to push their infant forcefully away from them, for instance when these mothers were eating or termite-fishing. However, maternal aggression was not the only method by which mothers controlled their infants at that age. Most striking were the observations of FF (month 11) and PS (months 11 + 12) in finding ways to avoid and prevent the infant from making and/or staying in body contact. This will be illustrated with the following observation on FF + FD.
When FD was 11 months old, he manipulated a leaf while sitting at a distance of 1 m from his mother FF, his grandmother Flo and his juvenile uncle FT. FF picked a leaf and began to groom this leaf. FD immediately approached his mother to look closely at what she was doing while holding her. Next, FF began to groom FD and, when she stopped, FD moved into the ventro-ventral position and began to suck. At the same time, he invited FF to groom him by raising his arm. While being groomed, FD was seen to close his eyes gradually more often and with shorter pauses in between. After 5 mins, FF began to groom more intensively and gradually she groomed/shoved him out of ventro-ventral contact with care. As soon as ventro-ventral contact was broken, FD opened his eyes to close them again. FF groomed FD for 2.5 mins before she stopped. FD immediately opened his eyes and moved into ventro-ventral contact. At once FF groomed FD intensively and FD closed his eyes soon thereafter. 1.5 min later, FF groomed/shoved FD out of ventro-ventral contact once more and continued grooming him for 5 mins. As soon as she stopped and groomed herself, FD opened his eyes and moved into the ventro-ventral position again. This time, FF had to groom FD for about 8 mins before FD closed his eyes. After about 1.5 min, FF carefully shoved FD out of ventro-ventral contact, before she broke contact completely. FD opened his eyes and FF at once made contact by grooming him intensively. FD closed his eyes. 4 mins later, FD rolled onto his back (FF might have groomed/shoved him too forcefully). FD opened his eyes and came into ventro-ventral contact once more. FF groomed FD continuously and FD immediately closed his eyes again. After 30 secs, FF stopped and groomed herself. FD opened his eyes to pull himself up to reach the nipple. All of a sudden, FF pinched her brother FT who happened to sit closeby, and then she groomed FT for about 6 mins. In the meantime, FD closed his eyes and gradually lost nipple contact but was still clinging tightly. FF looked at FD and, next, groomed him again. After 4 mins, FD released
44
his grip on FF. She groomed him for another few seconds, then she stopped. Now, FF moved out of contact very gradually, walked a few steps and sat down to groom herself. 5 mins later, she approached FD who was still asleep, gathered him and began to travel.
From month 11 onwards, FF could be seen to "groom FD to sleep" in this way, before she left him. It was remarkable that these grooming sessions had a different quality; FF groomed more intensively than usual and she was bent closer to FD than usual. Moreover, these sessions lasted longer than the usual ones. After FF left FD behind, she always stayed within 5 m to sit and groom or to sit and look around. It was striking that she seemed unworried when other chimpanzees or baboons were in the vicinity. FD could be asleep while adult male baboons came towards him gradually (as they would when approaching a strange object, or a corpse) and finally bent over him to look at him closely (Fig. 12). FD could go on sleeping even when adult male chimpanzees came between him and his mother (Fig. 12). On no occasion did FD wake up while being separated from FF. Somehow, before he was to wake up, FF always restored contact. Sometimes, she herself woke him up before signalling to climb onto her back before travelling away. FF was the only mother ever seen to break contact after the infant had fallen asleep. To understand how the mother-infant relationships changed through time, we need to know more than that mothers came to avoid and prevent body contact. Were these maternal behaviours not to be predicted? Or were they preceded, months before, by more subtle changes in the maternal regulation of body contact? In order to answer such questions, we need to know more about how mother and infant influenced body contact over age. Controlof body contactand the relativeroles of motherand infant. The measure in Fig. 13 shows the relative contributions of the infants FD and PF in maintaining body contact over age. The measure shows that for both infants the contribution to maintaining contact was negative from month 4 up to month 9 (except for month 5 of PS and PF). This means that the proportion of contacts broken on the infant's initiative was higher than the proportion made by the infant. In this sense, the mothers of both infants were primarily responsible for body contact at this age, before they avoided and prevented body contact. In order to examine whether the increase in responsibility for body contact between months 4 and 11 (FD) or 12 (PF) was primarily due to an increase in 'makes' or to a decrease in 'breaks', Table 7 is presented. D.2.2.
45
Fig. 12. Infant FD asleep while off the body of his mother and a few meters away from her. The mother seemed unworried when a baboon approached or when other chimpanzees sat in between her and the infant.
46

%1KI- %BKI +20
10
P FD
-50
month
10
12
14
16
18
20
Fig. 13. Infant's relative contribution in body contact (%MKi-%BKi) over age.
If we neglect the fluctuations in the percentages over age, and if we only look at the differences between months 11 and 4 for FD and the differences between months 12 and 4 for PF, we may conclude that the increase in responsibility for body contact (Fig. 13) was primarily due to an increase in 'makes' (Table 7). The measure in Fig. 13 peaked in month 11 for FD and in month 12 for PF. Remarkably, at exactly those ages the infants experienced TABLE7. The proportion of total body contact 'makes' and 'breaks' due to the infant
Age month 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 FF + FD %Makes %Breaks PS + PF %Makes %Breaks
> no contact-breaks< 50 72 80 36 81 85 82 99 90 80 89 91 85 81 88 88 80 100 93 98 91 91 91 93 89 87 91 93 94 87 82 83 90 91 33 67 93 93 (no contact-breaks) 83 100 81 92 66 83 89 88 86 82 94 86 94 90 79 89 79 92
47
aggressive objection to body contact for the first time (subsection D.2.7). Furthermore, during those months and the one or two months preceding or following them, the measure in Fig. 13 was positive for both infants. This indicates that both infants were primarily responsible for body contact during those months. Rise and drop in time spent in body contact. A rise in the percentage of time spent in contact was found in month 11 for FF + FD and in month 10 for PS + PF (Fig. 14). In these months both infants were primarily responsible for body contact for the first time (Fig. 13). Data on the percentage of time spent in body contact were available for two more pairs: PL + PT and ML + GM (Figs 15 and 16). Their curves are similar to those of FF + FD and PS + PF (Fig. 14). At comparable ages, also these pairs stopped decreasing their time spent in body contact (Figs 15, 16). Like FF + FD and PS + PF, PL + PT rose their percentage spent in body contact in month 11. The pause in decreasing time spent in body contact or rises in time spent in this contact were only temporary, because thereafter all pairs dropped the percentage of time spent in contact sharply. The sharp drop for two pairs (FF + FD and PL + PT) occurred between months 11 and 12 (Figs 14, 15). For ML + GM time in body contact dropped most markedly between months 10 and 11 (Fig. 16). Finally, PS + PF also showed a decrease in time spent in contact between months 11 and 13 (Fig. 14), albeit less markedly than the other pairs. A possible further decrease might have been interrupted by PF's illness in month 14: in this month, time spent in contact increased. D.2.3. Secondrise and drop in time spent in body contact. A second rise and drop were found in all three pairs which were available for comparison after month 16 (Figs 14, 15, 16). However, for all three pairs this drop appeared less markedly than the first one. For FF + FD and ML + GM this drop is shown between month 18 and 19 (Figs 14, 16). Whereas for PL + PT this drop is shown between months 21 and 22 (Fig. 15). After the first drop, time spent in contact remained at about 50 % for three out of four pairs. After the second drop the amount of time spent in contact remained at about 30%. In month 20 GM appeared to be ill and the pair increased their time spent in contact temporarily. The second drop was also preceded by a relatively high percentage of time spent in contact. For FF + FD, the measure shows a local maximum D.2.4.
48
H.
H.
& F. X. PLOOIJ
90
40
20 10 0 month 2
10
12
14
16
18
20
Fig. 14. Amount of body contact over age. * PF appeared to be ill.
001
703o
month
10 8 month 10 6
8 12 10 12 14 16
12 18 16 20
20
Fig. 15. Amount of body contact between L and PT over age.re
1This
can
be
seen
in
Figs
8
h .ont
and
6
19,
10 12
which
14 16 13
show
20 22 24
the
relative
Th921 15 (Figscnbs.
contribu-
Fig. 16. Amount of body contact between ML and GM over age. * GM appeared to be ill.
in month 17/18 (Fig. 14). ML + GM increased their time spent in contact in month 18 (Fig. 16). PL + PT did so for three months from month 1921 (Fig. 15). Finally, one month before the second drop in time spent in body contact, peaks in the infants' responsibility for body contact were found. This can be seen in Figs 17, 18 and 19, which show the relative contribu-
MOTHER-INFANT
RELATIONS
month 1 3 5 \ 7 9
IN FREE-RANGING
11 13 15 17 19 90 8O 70 60
CHIMPANZEES
49
of time in body contact
40 +20 +10 2
731K1-b
BK
30
-10
-20 -30 -40 -50 -60 month 1 3 5 7 9 11 13 15 17 19
Fig. 17. Association between peaks in %MKi-%BKi and peaks in body contact for FD.
month 8 10 12 14 16 18 20 22 24 26
body contact
90 80 70 60 50 40
+20 '10
BKi MKi-%
-10 -20 month 18 20 22 24
Fig. 18. Association between peaks in %MKi-%BKi and peaks in body contact for PT, 4 months before and 4 months after the drop in body contact.
month 5 7 9 11 13 15 17 19 21 23
i^F
%of time in body contact
100
90 80 70 60 50
30
+10
% %MK.- BK. 1 1
-10
month 15 17 19 21 +
TG1 appeared to be ill
Fig. 19. Association between peaks in %MKi- %BKi and peaks in body contact for GM, 4 months before and 4 months after the drop in body contact. * GM appeared to be ill.
50
H.
H.
& F. X. PLOOIJ
tions of mother and infant in maintaining body contact 4 months before and 4 months after the second drop in time spent in body contact. interaction. Changesin othermeasuresof mother-offspring In part B (subsection B.2.5.), we showed that progressive changes in maternal behaviour in the context of ventro-ventral contact were related to the peaks in the infants' responsibility for this contact. Here we are concerned with the relation between maternal changes in the context of body contact and the peaks in the infants' responsibility for this latter contact. Profiles of the progressive changes in retrieval, attention, meshing, grooming, rejection and aggression in the context of body contact over age show that they vary independently to some degree (Figs 20, 21, 22, 23). However, the changes can also be related to the peaks in the infants' responsibility for body contact. For FF + FD (FD's month 11 and 18, Fig. 20) and PL + PT (PT's month 21; Fig. 22) the changes related to the peaks in responsibility (Figs 17, 18) were similar. Thus, in these months, both mothers almost ceased to retrieve, attend, and mesh to their infants. Then, retrieval, attention and meshing increased again. Furthermore, the peaks in responsibility coincided with the first observed maternal rejection and aggression (FD's month 11; Fig. 20) and maxima in maternal rejection (FD's month 18; Fig. 20) and aggression (FD's month 18, Fig. 20, and PT's month 21, Fig. 22). Furthermore, FF + FD's profiles suggest that on the second occasion in which the infant's responsibility for body contact peaked, rejection and aggression were more obvious than on the first occasion. Also, for PS + PF (PF's month 12, Fig. 21) maxima in rejection and first observed maternal aggression coincided with the infant's first peak in responsibility for body contact (Fig. 13). Unlike FF, PS began to reject PF's attempts for body contact from month 9 onwards (Fig. 21). It may be the case that PF needed to play the greater role in maintaining body contact already in month 10 and 11 (Fig. 13), because PS was behaving in a rejecting way. Next, PS also showed dips in retrieval, attention and meshing, but she did so one month prior to the infant's peak in responsibility for body contact (PF's month 11, Fig. 21). For ML + GM (GM's month 18, Fig. 23) no minima in retrieval, attention and meshing were found to coincide and maxima in maternal rejection and aggression were found one month prior to the infant's peak in responsibility for body contact. D.2.5.
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
51
"/300 mins. 30 25 20 15 5 0 0
FF+FD months in which the infant's for contact peaked responsibility
RETR I EVAL
25 15 0 ATTENTION
10
5 0 .MESHING
GROOHING
agg
n ad 0 r ~--
n (
s 2,
21,
2,
2.
s in REJECTIOr
AGGPRESSION
month 8 Fig. 20. Profiles
10
11
12 13 14
15 16
17 18
19 20 over age (FF).
of maternal
behaviour in association with body contact - - - - - Infant grooms mother
For all pairs it is notablethat groomingcan be relatedto maternal and aggression rejection (Figs20, 21, 22, 23). Peaksin maternal groomto coincidewith maximain maternalaggression,exceptfor ing appear
FD's month 18 (Fig. 20). In this month, however, the infant showed a peak in grooming his mother. Grooming may be an alternative for loss of amount of body contact. It may therefore be understandable that no peaks in grooming were found when ventro-ventral contact had to be
52

PS + PF month in which the infant's for contact peaked responsibility
n/300 mins. 40 35 30 25 , 20 15 -
10 I 50
ETRIEVAL
_ATTENTION
n 1 n I
im
rlESHING
iiionth '1
0otI
61A1GrRESSION 9 10 8 6 7
11
12
13
14
15
Fig. 21. Profiles of maternal behaviour in association with body contact over age (PS). -------Infant grooms mother
rejected (Fig. 8), because body contact (except for ventro-ventral contact) was accepted. When all forms of body contact were rejected, the only form of body contact available to the infant was grooming, which may have substituted for the loss of other forms of body contact. Here a more detailed study to find out what body parts in fact were touching when grooming occurred will be of interest. We expect that the spot that was
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
53
PL + PT "/300 mins. 35 30 month in which the infant's for contact peaked responsibility
25
20 15 10 RETRIEVAL
25
5 0
ATTENT
ION
50~
__
~~~~MESHING
15 10 10t 18
20
GROOMING
10 5
-_
REJECTION
15 5 0 month 18 _AGGRESSION 19 20 21 22 23 24 25
Fig. 22. Profiles of maternal behaviour in association with body contact over age (PL).
- - - -Infant grooms mother
groomed was the only physical contact between the partners at the ages at which body contact was reduced. We propose that dips in retrieval, attention, meshing and/or peaks in rejection and aggression (Figs 20, 21, 22, 23) heralded further increases in the independence of the infants. The increases in maternal retrieval, attention and meshing following the dips, and the decreases in rejection
54
H.
H.
& F. X. PLOOIJ
ML+ GM month in which the infant's responsibility for contact peaked /300 mins. 25 20 15 _ 10 IEV RRETR 5 0
AL
0 5
GROOMING ATTGGRESSNTION
month 16 15
17
19
20
21 22MESHING
Fig. 23. Profiles of maternal behaviour in association with body contact over age (ML). - - - - - Infant grooms mother
and aggression following the peaks indicated that the mothers were effective in inducing the onset of further independence in their infants. This is supported by the fact that at the same time sharp drops in the amount of body contact between mother and infant were found (subsections D.2.3. and D.2.4.). Shifts in using space more distantfrom each other. With each drop in body contact all infants made use of space more distant from their mothers. Figs 24, 25, 26 and 27, show the rankorders of the D.2.6.
55
percentage of time spent in each of the six distance-categories as defined in section 2: General Methods. The distribution of the top ranks over age is indicated by the solid black line. The two drops in time spent in body contact are shown in these figures'). After the first drop in body contact, FF + FD were mainly seen out of contact, but within 1.5 m (Fig. 24). PS + PF were still mainly in contact, but no longer mainly in ventro-ventral contact (Fig. 25). Here PF's illness (first seen in month 12 and peaking in month 14) may have been responsible for interrupting the use of space more distant from mother. After the first drop in body contact, PL + PT were mainly seen out of contact but within 5 m away from each other (Fig. 26). Finally, after their first drop, ML + GM were seen more often within 1.5-5 m for three months, though they remained mainly in ventro-ventral contact (Fig. 27). Then a shift toward using space more distant from each other was found. Now the pair was mainly seen within 1.5 m but out of contact. The second drop in the amount of body contact for ML + GM (Fig. 27) and for FF + FD (Fig. 24) was associated with a shift from mainly staying within 1.5 m to mainly using space more than 1.5 m away from each other but remaining within 5 m from each other. The second drop for PL + PT was associated with a shift from mainly staying out of contact but within 5 m to mainly staying between 5-15 m from each other (Fig. 26). increasesin ventro-ventral contact. Temporary A temporary shift back to mainly staying in ventro-ventral contact is found prior to the months in which dips in maternal retrieval, attention, meshing and/or peaks in maternal rejection and aggression were found. We shall call such shifts 'regressive'. The type of regression discussed here has to be sharply distinguished from the pathological type of regression. The latter is characteristic of almost all versions of psychoanalytic theory except BOWLBY'S (BOWLBY,1980). In our analysis regression is associated with spurts in development and refers to the temporary return during early development of behaviours which were shown in an even earlier developmental period. This type of regression is less well known. In contrast to the pathological type, this could be called the normal type of regression. D.2.7.
1) For FF + FD the first drop in body contact was preceded by a drop in the amount of ventro-ventral contact between month 7 and 8 (Fig. 22). This drop was associated with both a shift from mainly staying in 'ventro-ventral contact' toward mainly staying 'incontact', as well as changes in excursion behaviour (subsection C.2.4.).
56

distance cateqories month--1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 within arm' s reach
v -, 1 1 1 1 1 1 .1 2 3 1 2 3 2 3 2 4 1 3 4 4
notentral ventral
in contact
between between 1.5-5m 5-15m
2 3 4 4 2 3 2 2 4 1 1 5 4 5 5 5 5 6 52 5 5 5 4 5 3 1 2 2 3 1 2 3 2 4 2 2 3 2 3 2 2 3 1 1 1 1 1 5 4 4 A 4 4 4 5
1
3 3 4 3 3 3 1 1 1
6 6
5 6 6 2 2 6
A-drop in ventro-ventral contact 1=first drop in contact 2.second drop in contact
Fig. 24. Rankorder of the amount of time spent in each of the six distance-categories FF and FD.
for
Thus, two 'regressive' shifts are found for FF + FD. In month 10 and 17 this pair was mainly seen in ventro-ventral contact (Fig. 24). Both shifts are also shown in Fig. 7, where the measure shows temporary increases in the amount of ventro-ventral contact for these months. For PS + PF the regressive shift can not be shown (Fig. 25), because the pair was mainly in ventro-ventral contact up to the age of 'regression'. However, in this pair the occurrence of 'regression' was revealed in the temporary increase in ventro-ventral contact during month 10/11 (Fig. 7). For ML + GM the 'regressive' shift is found in month 16 (Fig. 27). For PL + PT the shift back towards ventro-ventral contact is not found. However, before mother PL induced the onset of further indepence (leading to the 2nd drop) a 'regressive' shift towards mainly staying in contact was found (Fig. 26). Prior to this 'regressive' shift the pair was mainly further away from each other than were the two other pairs available for com-
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
57
distance categories ,, ventral month 1 2 3 4 5 6 7 8 9 10 11 -4213 14 x 15 1 -2 2 1 1 1 1 1 1 1 1 1 1 1 -.-
notventral
in contact
within arm' s reach
between between 1.5-5m 5-15m
2 3 2 2 2 2 3 2 3 1 3 2 2 3 3 3 4 2 3 -1 4 2 5 4 3 4 4 -4 2 4 3 5 5 4
6 5 4
- - 5
x PFappearedto be ill 1=first drop in contact
Fig. 25. Rankorder of the amount of time spent in each of the six distance-categories PS and PF.
for
parison. It may be that the further a pair is usually away from each other, the less likely they are to return into ventro-ventral contact during the regressive period. In this paper we will not consider PF's (month 14) and GM's (month 20) shift towards ventro-ventral contact, because here the regressive shifts were associated with illness. D.3. Discussion.
Recapitulation: regression,conflict and jump towards greaterindependence. The data presented above show that the decrease over age in time spent in body contact does not proceed smoothly. In each mother-infant pair periods of rapid decrease were followed by steady periods, or even reversals. Two stages in the decrease in body contact can be distinguished. Between months 1012, time spent in body contact decreased sharply and remained at about 50% over the following months (Figs 14, 15, 16). During the infants' second year of life (month 19-21), this measure decreased sharply once more and remained at about 30%. The remainder of the data are summarized in Table 8. They show that two phases precede each sharp decrease in body contact. These phases appear in the following distinct order. D.3.1.
58

distance categories month \ 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 5 6 within arm' s reach 4 1 3 5 5 4 5 4 4 4 6 4 5 6 4 3 2 2 3 5 4 5 3 5 4 5 3 6 5 2 2 3 2 4 2 2 1 1 1, 3 3 3 2 3 1 1 4 1 2 1 3 2 1 2 -1~ 2 2 4 2 2 1 > 3 1 2 3 3 2 2 1 4 3 1 4 6 6 6 5 4 4 1 4
ventral 1
notventral
in contact 2
betweenbetween 5-15m 1.5-5m
1lfirst drop in contact 2=second drop in contact
Fig. 26. Rankorder of the amount of time spent in each of the six distance-categories PL and PT.
for
The first phase concerns regression. Two months before each sharp decrease in time spent in body contact, temporary regressions were found. Regression was expressed, first, in a temporary shift back to mainly staying closer to mother (Figs 24, 25, 26, 27) and, second, in a temporary increase in the amount of ventro-ventral contact. The second phase concerns conflict. One month prior to each sharp decrease in time spent in body contact, conflict between mother and infant over this contact was seen to peak. Conflict was expressed, first, in temporary maxima in the infants' responsibility for body contact (Figs 17, 18, 19) and, second, in peaks in maternal 'rejection' and 'aggression' associated with dips in maternal 'retrieval', 'attention' and 'meshing' (Figs 20, 21, 22, 23). Finally, conflict over body contact was associated with either an increase or a stop in the rate of decrease of time spent in this contact (Figs. 14, 15, 16). Each sharp decrease in body contact itself is part of a third phase which we called a "jump". Except for the sharp decreases already mentioned, shifts in using space more distant from mother occurred (the distance category in which an infant spent most of the time: Figs 24, 25, 26, 27).
Regressionandjumps. The first regressive period occurred at the age of 10-11 months. Also VAN LAWICK-GOODALL'S (1968) data show the occurrence of a regressive period between these ages. She found an increase in ventro-ventral contact and sucking for two chimpanzee infants between month 10-12. D.3.2.
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
59
distance categories month 10
ventral
notventral 3
in contact 2
within arm's reach 4
betweenbetween 1.5-5m 5-15m 5 6
11 12
13 14 15 16 17 18 19 20a 21 22 23 24
4 5
4 4
4
3 2
2 3 1i--1 1
2 2
2
I
L~ 4 2 13 4 3 1 4 6 6 4
2 4 3-4 2 6 5 6 3 5 5
4 3 ---2 3 4 5 5 5 2 3
5 4 5
6 6
5 1 5 2 3 3 4 6
3 1
2 3 2
1 1 1
1=first drop in contact 2=second drop in contact x GM appeared to be ill
Fig. 27. Rankorder of the amount of time spent in each of the six distance-categoriesfor ML and GM.
HORWICH (1974) found that the first regressive period occurs at similar times in development in many more species (man, gibbon, 12 species of monkeys, 2 prosimians, and 2 non-primate mammals) when accounting for the fact that some species develop faster and have a shorter life-span than do others. He calculated the ratio of age at first regression to age at sexual maturity and found that in all these species the initial regression occurs between the first 7-17% of the infancy period. The finding that regression was not observed to be associated with the jumps in the growing independence we found at earlier ages (2, 5 and 7 months) not only holds for mammals, but for another non-mammalian species as well. KORTLANDT (1955, p. 238) found that the first jumps in progression towards greater independence in cormorants were not preceded by regression. He tried to explain why, and suggested that "regression will not occur until a certain height and complexity of organization have been attained. At the lower levels during ascending development, the building stones will naturally fit in more easily than at the higher ones". And he continues that, "in order to accomplish
60 TABLE
8. Distinct phases in two interstage periods in the context of body contact

Month prior to phase 3 -2 -1 Data
Phases
1. Regression 2. Conflict
3. Jump
Increase in ventro-ventral contact. Shift back to mainly staying closer. Stop in decrease or temporary increase in amount of body contact. Peak in infants' responsibility for body contact. Peaks in maternal rejection and aggression. Dips in maternal retrieval, attention and meshing. Sharp drop in amount of body contact. Shift in using space more distant from each other.
integration in spite of this, a remodelling or reshuffling of the building stones will be necessary". This may require a revival of infantile forms of behaviour, or regression, followed by a reorganization of low level behaviour. A second regressive period, which we found in months 16-17, was reported for many more species. HORWICH (1974) described up to four of high motor-infant contact in man, gibbon, 12 species regressive peaks of monkeys, 2 prosimians and 2 non-primate mammals. There is a general dampening effect so that each regressive peak is lower than the preceding one. We might have found more than two periods if our study had not stopped shortly after the second one. Conflict andjumps. We suggest that the mother recognizes that her infant is 'ready' to reorganize his behaviour, that he is capable of more, upon which she forces him to do so. In the next three paragraphs we will extend upon this. Around 11 months chimpanzee infants are able to perform 'programs' (PLOOIJ,1984). A program is defined to be a complex sequence of motor patterns and relations between them, as part of a goal-directed activity. Examples of programs are nestbuilding, termite fishing (VAN LAWICKGOODALL, 1968) and eating. For example, eating can involve locomoting towards a fruit, picking or bit-pulling it off the branch, biting a piece off the fruit and chewing and swallowing it. D.3.3.
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
61
Just as the infant is able to perform programs for the first time, so should he be able to perceive that the behaviour of his mother (or others) is organized in programs. On a par with the above given examples, also travel behaviour of a chimpanzee mother can be considered to be a program, for instance. It can involve climbing out of a tree, walking down a slope, jumping over a small stream, walking up the opposite slope of the valley, climbing along a vine into the next tree where the chimpanzee mother sits down, stops the travel program and starts the next eating program. If the infant is able to perceive such programs, he should be able to recognize from this age onwards that his mother has just finished a travel-program and started an eating-program or termite-fishing program which gives him ample time to start an excursion. It is interesting that around 11 months of age mother-infant conflict was observed only in the context of such 'programs'. For instance, mother would push her infant away during termite-fishing: he should know that she is in the midst of a program and that there is no need for restoring contact. The reverse is also true. Mothers were observed for the very first time to start travelling without signalling this first to the infants. One may speculate that in doing so the mothers are implicitly saying to their infants: you should know that I am about to finish my termitefishing program and to start travel and you are able to return to me on youw own accord. In similar vein, in month 5-7 (FF + FD and PS + PF) conflict only occurred in the context of nipple contact and ventral riding (subsection B.2.3.). These findings indicate that the mother does not promote the infant's independence as a whole in each conflict peak, but that she does so only for a particular aspect in guiding him each time to master yet another type of independence.
ALTMANN'S
(1980)
discussion
of baboon
mothers
training
their 4-6
months old infants to know when is a good time for being in close contact and when not shows parallels with our discussion of chimpanzee motherinfant conflict in the context of 'programs'. At 4-6 months of age, the baboon infant learns not just that he can be out of contact sometimes, but that he can be out of contact at particular times that are determined by his mother's activity').
1) The age at which the baboon infant learns these things deserves some more attention. In the light of HORWICH'S (1974) finding that the first regressive period occurs at the age of 2-3 months in 12 monkey species (including one baboon species), one would expect such learning to start well before the age of 4 months.
62
Chimpanzee mother-infant conflict is preceded by regression (Table 8). We suggest that this conflict may be triggered by regression. It seems likely that mother is irritated by the return of infantile behaviours which
are a drain on her resources
(CLUTTON-BROCK
et al., 1981).
4. General discussion The major aim of this study was to achieve a better understanding of the processes involved in the growing independence of free living chimpanzees. We looked at the history of mother-infant relationship in terms of how changes in the behavioiur of mother or infant affected the motherinfant dyad as a self regulating homeostatic system. We first discuss the possibility that labile periods at specific ages in the mother-infant relationship are a common feature in development; then the possibility that changes in maternal behaviour in each labile period are responsible for the phenomenon of jumps in the growing independence, provided that the infants are not pushed beyond their maturational abilities; then the possibility that regression, which precedes mother-infant conflict, is associated with maturational changes in the infants; then the possibility that maturational changes which may underly developmental steps trigger mother-infant conflict, and that mother-infant conflict is of vital importance in realizing the infant's developmental potential; and finally the possibility that attachment as a construct exists from birth and takes different forms depending on the proximity involved and the skills available to the infant. periods at specific ages. We found that the development towards greater independence proceeds stepwise and that relatively long lasting stabile periods and relatively short lasting labile periods in the mother-infant relationship alternate. It appears that the onsets of labile periods are found at specific ages for all mother-infant pairs studied (2-3 per age range): the infants being 2, 5, 7, 9-10 and 15-18 months old. Interestingly, PLOOIJ(1984) described the onset of a cluster of new behaviours in infant chimpanzees at the specific ages at which we found the onset of a labile period in the mother-infant 4.1. Labile relationship. Not only chimpanzee infants seem to show labile periods at specific ages, also human infants seem to do so. The resemblance in the sequence of developmental phenomena of both chimpanzee- and human infants is supported by the literature on captive apes. WOOD et al. (1980) and
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
63
REDSHAW (1978) compared cognitive development during the first year in human infants with the cognitive development in chimpanzee- and gorilla infants. Both found that the similarities were pronounced and that all three ape infants follow much the same course of developmentpassing through the same stages in similar order without a systematic difference in the rate of development. CHEVALIER-SKOLNIKOFF made (1977) similar conclusions after her application of a Piagetian model for describing and comparing socialization in monkey-, ape- and human infants. MCCALL et al. (1977) examined the principal components of the California first year and the California preschool tests at various ages and discovered periods in which the character of the components stays more or less the same, and periods in which the components alter considerably. They concluded that the former periods can be considered to correspond to stages (stabile periods) and the latter to stage-intervals (labile periods). Thus, in contrast to previous efforts, stages were found by focusing on developmental change and transition, not continuity and stability. The same approach was used by PLOOIJ (1984), with similar results. with these changes in the composition of the first prinSimultaneously cipal components (developmental function), MCCALLet al. (1977) found changes in the stability of individual differences. The rank order of the individuals remains more stable within a stage than in between stages and dips in the cross age correlations were found at 2, 8, 13, 21 and 30-36 months. The age of the fourth stage-interval (18-21 months) is slightly extended as compared to the age at which a dip in the cross age correlations was found (21 months), because MCCALLet al. (1977) place the end of stage IV at 18 months. Fig. 28 illustrates the concurrence between the ages at which stageintervals were found by MCCALLet al. (1977) for man and the ages at which we found the onset of labile periods in the chimpanzee motherinfant relationship. MCCALL et al.'s second age (8 months), third age (13 months) and fourth age (18-21 months) are slightly later than the ages at which we found the onset of labile periods in chimpanzees (7 months, 9-10 months and 15-18 months). This is not surprising if one considers that man develops at a slightly slower rate than do chimpanzees. 4.2. Labile conflict. periods and mother-infant Because certain data are impossible to collect without interference in the daily routine of free-living chimpanzee mother-infant pairs, we could not
64
H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ 8 13 2 . ___x_____x___ _x__ 30-36 18-21 xxxx ___ ____xxxxxxx_
Man
Chimpanzeee
2 7 10 15-18 ___x____x_xx____xxxx____9
_ > AGE (months)
Fig. 28. The concurrence between the ages at which stage intervals were found in man and the onset of labile periods in chimpanzee.
study how mother-infant conflict is associated with changes in the mother such as alterations in hormone balance, decreasing body weight and decreasing milk production. Indirect evidence for decreasing and subsequently increasing milkproduction can be found in STEINBACHER'S (1941) report on one chimpanzee mother-infant dyad in captivity. The observers concluded that milkproduction was diminishing around the age of 5.5 months from the fact that the baby sucked more frequently and for shorter duration. When they offered extra drinks, the baby accepted these. One month later, the baby refused these drinks and completely relied upon his mother for his drinking needs again. Furthermore, it has been found by FUCHS (1982), MARTIN (1986) and RANDOLPH et al. (1977) that mother's body weight and milkproduction decreased steadily in house mice, cats and cotton rats until the offspring was weaned from the nipple, whereupon body weight increased again. In the present study we focussed on how mother-infant conflict is associated with changes in the behaviour of the infant and we found that periods of mother-infant conflict precede jumps in the development towards greater independence. During conflict the mother forces the infant to use its new abilities and to reorganize its behaviour. We suggest that the phenomena of jumps in independence that we found represent the consequences of mother-infant conflict. In other species maternal rejection is also found to increase the speed with which the infant attains
independence
ALTMANN,
(HANSEN, 1966; HINDE, 1969, 1974a; JENSEN et al., 1973; in a
1980), if the infant is not upset so much that it becomes cling-
ing (HARLOW, 1971; HARLOW et al., 1971). We found that mothers
period of conflict make use of aggression and rejection when teaching their infants new behaviours. We expect that the absence of such maternal contributions might be associated with a more gradual increase in independence. Here a detailed study of infants brought up without a mother-figure would be of interest. It is not yet established whether the mother's actions are needed to attain the infant's independence. HANSEN
65
(1966) found that also the infants reared on inanimate surrogate mothers achieved independence, albeit at a slower rate. We found that mothers are unable to force the infant to perform new behaviours when it is too young for it. In part A of the results we described how PS forced PF to ride on her back when he was only a few days old and how she failed. When PF was two months old, she tried again and succeeded, because now PF was able to hold himself and no longer slipped from her back. Another example occurred when PF was 15 weeks old. Now PS forced PF repeatedly to approach her by standing two meters in front of him. Though PF was observed to make his first step that same day, he was still unable to walk quadrupedally and PS's attempts failed. When she tried again PF was 19 weeks old and she succeeded. Thus, it is futile to force new interaction patterns upon the infant when it is too young and not ready for it. MCGRAW(1945) expresses a similar opinion for human infants. HINDE (1970) illustrated the occurrence of age changes in learning ability in many species and his review of the literature shows that the earliest age at which a certain type of learning is possible is little affected by earlier experience. After the earliest possible age of learning certain specific behaviours, large variation is found in the age of onset of each specific achievement (actual behaviour patterns or proof of learning) worked out by an
individual. For instance, YERKES & TOMILIN (1935) report that although
the usual age of onset for quadrupedal walking in captive chimpanzee babies lies during the fifth month, a spread of 4-12 months occurs. Such delay can be caused, among other things, by rearing practices. RIESEN & KINDER (1952) showed, for instance, that motor development of chimbabies in captivity which are separated from their mothers and panzee raised in a nursery environment is retarded. It should by possible to advance specific achievements beyond the age normally expected, if ample opportunity is afforded (McGRAW, 1945). This is likely if one assumes that the group-average for a specific age of onset is later than the earliest possible age of onset and if normal expectation is based on the group-average. It would be interesting for further research to vary the experiential conditions experimentally while keeping records of independent maturational markers of the offspring, and to see whether specific achievements can be pushed beyond the offspring's maturational abilities (earliest age of onset). BERGER (1979) observed a natural experiment in desert bighorn canadensiscremnobates). Under the extreme ecological condisheep (Ovis tions of the desert the ewes weaned their lambs earlier and more
66
abruptly. Early weaning does not necessarily imply that the offspring was pushed beyond their maturational abilities. Berger suggests that the development in the lambs may have been accelerated during evolution as an adaptation to the desert environment. However, he did not measure independent maturational markers, nor did he know the exact ages of the lambs. We suggest that the development of these lambs was not accelerated, but that the lambs were pushed beyong their maturational abilities as lamb mortality following early weaning was 95%. 4.3. Regression and maturation. We found that from month 10 onwards regressive behaviour precedes a period of mother-infant conflict and we propose that regressive behaviour in this context is associated with maturational changes in the infant.
Several 1955; (MCGRAW, 1945; WERNER, 1948, 1957; KORTLANDT, PETERFREUND, 1971; MOUNOUD, 1976) considered the relation authors
between regressive behaviour and spurts of development. To be able to understand the possible role of regression in development, we need to extend upon each author's view of development. McGRAW (1945) addressed the question of how organization of neural functioning is connected with organization of behaviour. Changes in motor expression are considered to reflect reorganization of the central nervous system. Higher control systems are thought to be nonfunctional at birth and to be superimposed at later, quite specific ages. Despite the emphasis on maturation and the conviction that training in any particular activity before the neural mechanisms have reached a certain state of readiness is futile, McGRAw leaves no doubt about the fact that learning plays an important role in the fundamental process of growth. If ample opportunity is afforded at the proper time (after a neural mechanism has matured), "specific achievements can be advanced beyond the stage normally expected". According to MCGRAW, "periods of transition from one type of neuromuscular organization to another are an inherent part of development and are often characterized by disorganization and confusion. Spurts and regressions are an integral part of organic growth, and there is reason to believe that they also function in the development of complex behavior activities". McGRAw reported a regressive dip in toilet training of one infant at the age of 17.5 months. KUNST'S(1948) data on finger sucking reveals a possible regressive period in human development at the same age (18 months) at which MCCALLet al. (1977) found the fourth interstage period (18-21 months).
67
WERNER (1948) mentioned that reorganization begins with regression to some lower level of functioning. He (1957) states that "whenever development occurs, it proceeds from a state of relative globality and lack of differentiation to a state of increasing differentiation, articulation, and hierarchic integration". He showed that the developmental process may be divided into stages, since some of the reorganizations may take place rapidly (from BALDWIN, 1967). KORTLANDT(1955) showed that the development of cormorants pro-
ceeded in a number of developmental jumps. He argued that development proceeds through the emergence of isolated appetites followed by an ascending integration into a hierarchy. An appetite is defined as "either the performance of a specific activity or the presence of a specific external object or situation which causes the ending of a variable sequence or series of activities leading to this particular activity or situation". The appetites are arranged in levels that successively mature and integrate during ontogeny. The appetites of one level are isolated in the sense that they emerge independently and often at different ages. At a later age these isolated appetites become integrated into larger units and subordinated to appetites of a higher level. This is repeated several times, resulting in a hierarchy of appetites. Finally, KORTLANDT pointed out the what he called 'regenerative aspect' of regression. This aspect becomes understandable if one considers KORTLANDT'S suggestion that when appetites become subordinated to appetites of a higher level, "it may naturally occur that the building stones appear not to fit with each other. Consequently, in order to accomplish integration in spite of this, a remodelling or reshuffling of the building stones will be necessary". In other words, the subordinate appetites (specific activities or external objects) towards which an individual has become conditioned during ontogeny, have to be re-developed. This may require a revival of infantile forms of behaviour, or regression, followed by a reorganization of low level behaviour. PETERFREUND (1971) expresses the point of view that development proceeds through the unfolding, according to a maturational blueprint, of hierarchically arranged, feedback-regulated information processing control systems. He found that with each unfolding of a new level of control systems a process results which implies disordering, regression and recombination of existing learning. According to HORWICH (1974) the fact that the regressive period seems to be a common feature of development in many mammals implies a common genetic base and also normality in development.
68
MOUNOUD (1976), a member of the Genevian school, does not discuss regression as such but mentions a temporary dissociation followed by a recoordination during the change from one stage to another. He is of the opinion that new possibilities (or new 'instruments') of the organism arise through maturation. He argues that these new possibilities lead to new problems being solved. In the light of this, it is interesting to note that Kopp et al. (1974) and UZGIRIS (1973) found relapses in the course of repeated testing. It may well be that such relapses were the consequences of regression. In the opinion of all authors cited above regression is the consequence of maturation and associated with periods of transition from one type of neuromuscular organization to another. Strikingly, these authors have very different backgrounds from developmental psychology, ethology and psycho-analysis to the Genevan school of PIAGET. Our data are in accord with this opinion in that regression in the infant did not follow but precede the period of mother-infant conflict. Thus, it cannot be argued that regression is the consequence of this conflict. It remains difficult, however, to envisage how neuromuscular reorganization is related to regression. A speculative answer to this question is elaborated in the next section. 4.4. Developmental conflict periods. steps, new types of learning and
We propose that infants contribute to changes in the mother-infant system as they change maturationally. Maturational changes in infant chimpanzees must be established independently of their effect on the mother-infant relationship. PLOOIJ (1984) studied in detail the behavioural development of the chimpanzee during its first year of life and recognized 5 steps. Each step represents not so much the emergence of new discrete behaviours as a change in the 'quality' of all the infant's performances. The idea that maturational changes can be detected by looking for qualitative changes over development in behaviour was already mentioned by MCGRAW approach was based on the early ethologists' demonstra(1945). PLOOIJ's tion that entire sequences of behaviour can in principle be described and thereby subjected to analysis. Behaviour was viewed as being made up of "elements" or discrete motor-patterns (among which were the patterns usually called 'fixed action patterns') that integrated into functional sequences. These discrete patterns of behaviour were used in the search for the existence of behaviour control systems (BAERENDS, 1956, 1976;
69
HINDE, 1953). These systems were envisaged to control a group of func-
tionally related activities (MCFARLAND, 1971) in adult organisms. It were these very fundaments of the ethological approach to the analysis of adult behaviour which caused problems in the search for control systems underlying early ontogenetical stages of chimpanzee behaviour. It was impossible to recognize discrete motor-patterns before the age of 5 months (PLOOIJ, 1984). This finding was supported by CONDON'S (1977, 1979) frame by frame analysis of body motion by human neonates. A similar conclusion was reached for a non-primate mammal by FENTRESS (1983): only after the age of 10 days was it easy to identify the stroke types in mice grooming. Only then did these become predictable and 'stereotyped'. The age of 10 days in mice is comparable with the age of 5 months in chimpanzees, since development in the apes is retarded with at least a factor ten. Another way to establish the existence of such control systems was developed (PLOOIJ,1984). Data collection was based on the following argument. If one is looking for evidence concerning the existence of systems controlling early behaviour before the age at which discrete behaviour patterns emerge, it is only logical that such systems, among other things, concern the ontogeny of movement-regulation leading up to the ability to execute discrete behaviour patterns. In such regulation, feedback through proprioception and other perception (of the results of movement outside the body) has been found to play an important role in primates (MARSDEN et al., 1978; POLIT & BIZZI, 1979), in mice (FENTRESS, 1976) and some bird species (BASTOCK et al., 1953; BAERENDS, 1956, 1970). In such feedback a deviation from set-values plays a crucial role (BAERENDS, 1976). As soon as the perceived stimuli conform to the all activity aimed at restoring those values stops. Such stimuli set-values, have been called 'consummatory stimuli' (BASTOCK al., 1953). Thus, et systems may also be thought of as controlling set-values in the perception, effectuated by the behaviour, as a result of which consummatory stimuli can be observed. Consequently, these consummatry stimuli can help in finding the control systems underlying early behaviour. In order to illustrate what such consummatory stimuli look like, the role of thermal and tactile stimuli in chimpanzee neonatal behaviour was described (PLOOIJ, 1984). For instance, a species specific, optimal temperature is discrepant from the optimal temperature, the neonates are active. As soon as the optimal temperature is reached, all activity ceases. Thus, rooting in the direction of a nipple may be guided by a temperature gradient.
70
H.
H.
& F. X. PLOOIJ
TEITELBAUM al. (1983) showed that the same and other, very elemenet
tary (vestibular, kinesthetic and gastric) stimuli are important in the control of normal infant behaviour. Chimpanzee babies of two months and older, however, show overt behaviour which can no longer be directed at obtaining such simple, consummatory stimuli (PLOOIJ, 1984). For instance, finding a nipple through rooting (via a temperature gradient) disappeared. From this age onwards, babies show the so-called 'directed head-turning response' and go straight to the nipple, if they are constantly on the body of the mother. In the latter case they must find their way over the body of the mother in a different manner. Other characteristics must have become important as beacons for finding the nipple. Thus, the question arises what new consummatory stimuli start to rule the behaviour of babies of two months and older? This question is not readily answerable, for these stimuli are, so to speak, hidden. Similar ideas have been expressed by GOLANI (1976) and HOFER(1978, 1981). The latter speaks of 'hidden, regulatory processes'. The former showed that chaos seemed present in the observed behaviour as long as one did not know what was controlled. As soon as one knew, order appeared in et the same overt behaviour. In similar vein, TEITELBAUM al. (1983) showed that understanding of motivation is limited as long as an identifiable control is lacking. PLOOIJ (1984) developed a hierarchical systems control feedback model originally proposed by POWERS (1973) and experimentally tested over the years by COOLS (1985) through chemical stimulation of the brain of adult cats and monkeys. Along with this model, behavioural operational rats, definitions were presented, identifying hierarchical levels of perceptualcognitive-motor competence. These definitions lead to empirical tests for the induction and facilitation of new levels of control. Which type of control operates developmentally depends on maturation level. Which skills and overt behaviours develop actually depends on that type of control interacting with situational-social contexts. PLOOIJ (1984) used three kinds of empirical tests in his search for differences in the qualities of performances. He studied the infants' reaction to disturbances, the speed of concrete behaviours and the degree of variability of overt behaviour. From these analyses he recognized 5 abrupt overall changes in the infants' quality of performances in its first year. He assumes that each overall change in the infant results from the emergence of a new collection of negative feedback control systems in the
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
71
central nervous system. The systems in one such a collection share the same 'type of perception' and the same 'type of control'. In our study we found the onset of four labile periods in the motherinfant relationship during the infant's first year of life, respectively when the infant was 2, 5, 7, and 9-10 months old. At each of these ages PLOOIJ (1984) described a developmental step to occur (Table 9). Additionally, the qualitative changes in the growing independence that took place following a period of mother-infant confict reflected the consequences of We will now describe each developmental step as described by PLOOIJ. our findings together with the behavioural consequences of each new type of perception and control.
TABLE 9. Developmental
Step 1 2 3 4 5 Age (months) 2 3 4.5-5 8 10-11
steps reconstructed
from PLOOIJ (1984)
Type of perception
Configurations Transitions Sequences Relationships Programs
In part A of the results we described the marked drop in maternal 'securing' of the ventro-ventral position after month 2. At the same time we observed that the babies were now able to carry their own weight by clinging to their mothers when they were forced to. This change in maternal behaviour follows the first developmental step as suggested by PLOOIJ (1984). When the baby is able to perceive and control configurationsit can control various body positions (such as clinging to mother) and control the perception of visual- and acoustic patterns (PLOOIJ,1984). In month 5 we found aggressive maternal prevention of nipple contact and ventral riding in favour of gathering solid food for themselves and dorsal riding. These changes in the maternal behaviour follow the third developmental step. When the baby is able to perceive and control sequencesit can make several smooth movements in succession. This means that it can make several steps and thus start walking and climbing. In general, the baby starts to perform various discrete motor patterns of the type which is usually described in the so-called "behaviour repertoire". A further example resulting from controlling sequences is selectivity in behaviour towards other individuals than mother (PLOOIJ,1984). After month 7 the infants are concerned over their distance to mother. All infants show similar excursion behaviour in that they make excursions within arm's reach, while making and breaking contact frequently. These marked changes in excursion behaviour followed the 4th developmental step: the perception and control of relationshipsinvolves, for instance, space-time relationships, cause-effect relationships, but also the logical concept "and" as a relationship. Such relations may exist between the distinct self and another object, relations between the distinct self and another individual, relations between two other individuals, or two objects (PLOOIJ, 1984).
72
In month 11 we found aggressive maternal prevention of all forms of body contact in favour of making excursions and staying away for a longer timespan. These changes in maternal behaviour followed the fifth developmental step. When the infant is able to perceive and control programsit is able to understand that a complex sequence of motor patterns and relations between them may be part of a goal-directed activity. For instance, eating is a program. It may involve locomoting towards a fruit, picking it off the branch, biting a piece off the fruit, chewing and swallowing it. However, eating can also be performed in different ways and in different succession of the same motor patterns. Thus, a program is not a list, it is a structure which the infant is now able to perform itself and to recognize when other individuals are performing these (PLOOIJ,1984). It is interesting to note that when this fifth developmental step occurs, we observed mother-infant conflict only in the context of 'programs' (part D of the results). PETERFREUND'S (1971)
thinking
on the relation
between
maturation
and learning (reprogramming) may clarify when, why and how mother changes her behaviour over age. Maturation, as a new source of information, results in stress. PETERFREUND distinguishes two types of sources of stress: external and internal. Maturational change is one example of an internal source of stress. The effects of maturational changes disrupt the so far existing homeostatic conditions. When a baby experiences stress, it operates less efficiently and, consequently, regression is shown. Regression enables a sorting, selecting, and trial- and error process that aims at reducing the stress. However, a baby is hardly able to control the outer world and thus he needs a helping hand to reduce his stress and to restore homeostasis. PETERFREUND points out that a new dynamic equilibrium cannot be reached and maintained without the ministrations of the mother: "mother and infant must adapt to each other. In every response to her infant, the mother conveys a vast amount of information and helps to create a degree of biological order". In this way mother and infant form a selfregulating, homeostatic system in which it seems mother's role to reduce the infant's stress and to restore homeostasis. We propose that mother's behaviour in a conflict period is of vital importance; it is mother's role to (force) teach the infant how to use newly emerged abilities it might not, or not fully, have used otherwise. Thus, what the infant will actually learn depends on the opportunities it receives or the situations it becomes involved in, because each developmental step only creates a new potential for a certain type of perception and control of that perception (PLOOIJ,1984). In such context, conflict between a mother and her infant has its positive sides. During the conflict periods, the mother did not reject the infant as a whole, but only certain aspects of its behaviour. Our data clearly show that the conflict serves to realize the infant's new ability (Parts B-D of the results). We found that the mother contributes to the
73
developmental steps by demanding that the infant reorganizes its behaviour. As soon as the infant reaches a next stage in its independence, the conflict disappears again. Conflict between a mother and her infant in each labile period is not only centered around the infant's growing technical knowledge, but also around demands concerning social behaviour. We found that the specific demands concerning social behaviour in a labile period varied with the new abilities resulting from a developmental step at that age. It appears that over age the infants also learn how to direct their behaviour according to the ongoing behaviour of the other. For instance, during months 11-12, the infants learn when not to interfere with mother's ongoing behaviour or 'programs' (Part D of the results). A similar finding was
reported for yellow baboons by ALTMANN (1980), and HINDE & WHITE
(1974) found that over the same period in which rhesus monkey mothers and infants were spending less and less time together, their behaviour was also becoming better integrated. Thus, each conflict period combined with a developmental step led to a new 'type of learning'. In our opinion, periods of conflict between mother and infant are an integral part of normal development. This does not imply that motherinfant conflict cannot escalate and, in extreme cases, lead to child abuse. The conflict periods may be considered as periods in which the infant and thus the mother-infant system is labile and in a state of transition from one stage to the next. Is is reasonable to assume that the equilibrium in the mother-infant system runs a higher risk of being destroyed in these periods. One can imagine that the amount of mother-infant conflict and maternal aggression varies between being just enough to promote learning and too much to handle. According to PETERFREUND (1971), learning
is not optimal in traumatic situations, because defense mechanisms are activated in the infant. Optimal learning and defense mechanisms are, in a sense, at the opposite ends of a spectrum. In the former, the input of new information tends to be maximal; in the latter, it tends to be minimal. In this respect it is important to realize that a deepened knowledge of normal periods of conflict may lead to a better understanding of how conflict periods become pathogenic and, ultimately, could lead to child abuse. 4.5. Developmental steps and attachment. One of the abrupt changes in the mother-infant interaction is the onset of regular excursions around the age of 7 months. This change was shown to be related to the fourth developmental step.
74
Another abrupt change in the human mother-infant interaction is the emergence of attachment behaviour between weeks 28-30 (BOWLBY, 1973). Attachment behaviour is defined as actively seeking proximity to
a preferred individual above given
(BOWLBY,
1980; RUTTER, 1980b) and is mediated
by homeostatic behavioural systems which become goal-corrected at the

age, according to BOWLBY. This is in line with PLOOIJ's
(1984) view that the emergence of a new type of control systems concerning 'relationships' such as the distance between mother and infant (fourth developmental step) accounts for the appearance of behaviours around 7 months which are grouped under the term attachment in the literature. The association of both the onset of outgoingness and the onset of attachment behaviour to the fourth developmental step seems contradictory. However, both onsets are the two sides of the same coin, being the regulation by the infant of the distance between the infant and its mother. On the one hand, the new ability to perceive and control this distance relationship enables and encourages the infant to go away from mother further and more frequently. On the other hand, the same new ability causes the infant to be concerned when control over this distance is taken away from him (when mother is the one who leaves, for instance). Attachment behaviour is potentially active throughout life (BOWLBY, 1980). Yet, after a certain age, attachment behaviour diminishes and one of the several different processes underlying this change over age is a change in form taken by the behavioural systems mediating attachment behaviour itself (BOWLBY, 1969). This statement is compatible with our view that the fourth step, resulting in among others one system controling the distance between mother and infant, becomes subordinated, at later ages, to systems of at least two more developmental steps: the fifth developmental step (month 10-11) and the sixth developmental step (month 16-19; see part D of the results). statement (quoted above), he was not so much However, in BOWLBY'S thinking of the end of the first year and the second year of life. He remarks that the course which attachment behaviour takes during the second and subsequent years is not very well chronicled (BOWLBY, 1969). Therefore, our detailed account of the two progressive jumps towards greater independence of the chimpanzee infants up to the age of 24 months, resulting from additional levels of control systems added to those regulating distance, may be considered an addition to what is already known. The first of these two jumps is supported by one of BOWLBY's (1969)
75
descriptions. He notes that, during his 1lth or 12th month, the infant becomes able, by noting mother's behaviour, to anticipate her imminent departure. This is in full accord with our notion of the fifth developmental step around that age and the infant's new ability to perceive programs . Our second jump (around 18 months) was not reported by BOWLBY. He reported a jump towards the end of the third year of life (an age period which we did not observe). Then children are usually much better able to accept mother's temporary absence and to engage in play with other children. "In many children the change seems to take place almost abruptly, suggesting that at this age some maturational threshold is
passed" (BOWLBY, 1969). Interestingly, MCCALL et al. (1977) reported
an inter-stage period around the same age (30-36 months). This view of attachment taking different forms between the ages of 836 months, depending on the skills available to the infants (developmental steps), is very much in line with the thinking of attachment as a construct (SROUFE & WATERS, 1977) rather than as particular patterns of
behaviour. This view is also envisaged in the recent reconceptualization of attachment as representation. BRETHERTON (1985) thinks it may be fruitful (for studying the development of so called "internal working models" of the attachment relationship) "to integrate into attachment theory recent advances regarding the development of social-emotional understanding, communication and general representational processes". And a little further: "A theory of human attachment relationships must take into account the set-goals of relevant motivational behavioural systems". This author states that internal working models of attachment must be revised to remain serviceable, especially at an early age when development is rapid. With increasing age, more complex working models come to replace earlier and simpler versions. MAIN et al. (1985) even go so far as to suggest that "Some type of internal working model of specific relationships may be formed in the first months of life". This suggestion is remarkable in the light of the following. Whereas our account of the changes in independence starts from birth, accounts of attachment taking different forms did not up till recently. Despite all the differences between the various concepts of attachment'),
1) The various concepts of attachment are: the trait concept, the social-learning-, the psychobiological-, the psychoanalytic-, the ethological-, and the clinical-developmental et concept. These concepts are discussed in GEWIRTZ (1972), ALLOWAY al. (1977), SROUFE & WATERS(1977), RAJECKI al. (1978) and RUTTER(1980b). et
76
H.
& F. X. PLOOIJ
they shared the view that in human infants some attachment formation process takes place and that attachment has been developed by 8 months. The view of attachment taking different forms depending on the skills available to the infants only applied to older ages. It is possible, in our opinion, to argue that attachment exists from birth onwards and takes different forms depending on the skills available to the baby in a similar way as attachment is supposed to take different forms from month 8 onwards. Furthermore, we suggest that the ages at which each change in form occurs is associated with a developmental step in the infant. Before explaining our view of attachment, we need to give a wider meaning to the words "actively seeking proximity" in the definition of
attachment given by BOWLBY (1980) and RUTTER (1980b): "actively
seeking proximity to a preferred individual". We will do this in the next few paragraphs. There is no need to argue here that human babies younger than 8 months are capable of showing preference for a particular individual. RUTTER (1980b) reviewed the literature and found ample evidence that babies of only a few weeks old are capable of selective responsiveness to a particular individual such as the mother. "Actively seeking proximity" develops late (month 8), according to all attachment theorists. We fully agree, if this phrase is taken to mean "seeking to be within arm's reach" or even "restoring contact". The following discussion concerning this part of the above given definition of attachment centers around the notion that this meaning of the phrase does not take into account that there are other forms of proximity or "closeness" and less obvious ways of seeking or maintaining these. Being in contact with another individual is not unitary, but can mean various forms of "closeness". We were able to study these different forms of "closeness", because chimpanzee mother and infant spent nearly 100% of their time in body contact in the first 7 months of the infant's life. The closest form of contact is being "ventral". This is said to occur when most of the infant's weight is supported by the body of the mother, and when the greater part of its body is in front and facing the mother and encompassed by the arms and thighs of the mother. A less close form of contact is being "not-ventral". This is said to occur when most of the infant's weight is supported by the body of the mother while not being "ventral". This means the infant is on a more peripheral body part of the mother such as her back or her limbs. The least close form of contact is being merely "in-contact". This is said to occur when the infant's weight is not supported by the body of the mother (but by the
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
77
ground or another substrate instead) while the infant's body is still in touch with mother's body. The more dependent (thus younger) a baby is, the closer is the form of contact to which that baby is restricted. For instance, the neonate is 100% of its time in the ventro-ventral position. Only after the age of two months is this contact broken for the first time and can the baby be observed on the mother's back. After the age of 3 months, the baby can be seen regularly off the body of the mother (but "in-contact"). And after the age of 4-4.5 months, the baby can be observed for the first time to break body contact regularly in order to move more freely round the mother while remaining within arm's reach. Thus, if the mother's ventrum is considered to be the centre of the baby's world, this world slowly expands over the first half year from being ventro-ventral, via more peripheral parts of the mother's body towards being out of contact. As long as the baby is totally restricted to the ventro-ventral closeness, is also very limited in the skills it has to its disposal to actively seek this it closeness. The neonate can only "mountaineer" and "root" when trying to reach a nipple. The abilities the baby has to its disposal for actively seeking closeness change with each developmental step (the steps as described in subsection 4.4., Table 9). After the first step (2 months), the baby is able to hold onto mother's hair without her support (part A of the results). After the second step, the baby is able to change any awkward position on the body of the mother (part A of the results). After the third step, the baby can climb into the ventro-ventral position from a less close form of contact (part B of the results). Therefore, actively seeking "closeness" changes form over the first half year with each developmental step in a similar way as actively seeking proximity (attachment) takes different forms thereafter. Realizing this, there seems reason to abandon the concept of attachment as something new having been formed by the age of 8 months. In our view, the emergence of actively seeking proximity (attachment) associated with the fourth developmental step at the age of 8 months, is just another change in form of actively seeking "closeness". In view of this, one may say that attachment exists from the very beginning and changes form over development. This view of attachment is supported by some human literature. First of all, the overall changes in proximity or "closeness" regulation were
confirmed by SORENSEN(1979) in a New Guinea case study. CASSEL & SANDER (1975) concluded from an experimental Secondly, study on
78
human neonates that a primitive form of attachment is already present et during the second week of life. From former work (SANDER al., 1972) a particular caregiving context and a parthey hypothesized that, given ticular set of neonates, by the end of the first week of life the neonate has registered, and is entrained to a specific caregiving "context gestalt". The latter is considered to be characterized for the neonate in terms of the particular unique configuration of sensory, kinaesthetic, and motor cues and their temporal patterning. In order to test this hypothesis, CASSEL & SANDER introduced a perturbation of the specific context gestalt (the wearing of a mask by the mother and the maintaining of silence during the caregiving exchange). This perturbation resulted in striking alterations in neonatal behaviour such as the pattern of visual contact with the mother, thepatternofposturalcontact(rearing back from the mother or visual and postural adjustment away from mother), the pattern of feeding and the transitions to REM and then to NREM sleep. Moreover, increased frequency of distress events and increased crying frequencies
were observed. And thirdly, VAN WULFFTEN PALTHE & HOPKINS (1984)
have shown that for human babies a relationship exists between the maturation of certain neural functions and the first signs of social competence during face-to-face interaction. On a par with what we have shown for chimpanzee babies, practically all human infants they studied attained full postural control at nine weeks. This enabled the babies to coordinate head and eye movements in such a way that they could regulate their looking behaviour efficiently. Combining looking towards the mother with smiling and "pleasure" vocalizations, the infant of 2-3 months of age is perceived as a capable participant in social interaction. Although these authors did not study physical contact regulation per se, one can imagine that in human cultures where infants are carried around, attainment of full postural control has consequences for the contact regulation along similar lines as we have shown for the chimpanzee. Just as reconceptualization of attachment as representation (BRETHERTON, 1985; MAIN et al., 1985) facilitates further research on attachment across the life span beyond infancy, so our view of attachment may extend research in the opposite direction. One may even speculate that for attachment to be present shortly after birth, some sort of representation or "working model" of the mother-infant relationship should have been formed prenatally. This need not be as far-fetched as is may seem at first. After all, GOTTLIEB (1976a, b) found that embryonic auditory contributes in a facilitative way to the preference for the experience maternal call 24 and 48 hours after hatching. Also for the human fetus
MOTHER-INFANT
RELATIONS
IN FREE-RANGING
CHIMPANZEES
79
prenatal determinants of postnatal behavior have been suggested (FEIJOO, 1976; LILEY, 1972; MONTAGU, 1962; SONTAG, 1970; SPELT,
1948; De CASPER & FIFER, 1980). And it seems possible that by the time
of birth human babies have learned their mother's unique combination of auditory, sensory, kinesthetic and motor cues and their temporal patterning.
Summary Several investigators discussed the need to know more about conditions which facilitate normal human development, especially the need for a better understanding of the processes at work during the development towards greater independence in normal and pathogenic human relationships. The study reported in this paper aims to provide a description of the processes at work during the development of contact- and distance regulation for free-living chimpanzee mother-infant relationships. We believe that such an ethological study contributes to human studies by providing methods of observing and analysing behaviour, and by providing hypotheses that can be tested. Our study takes a systems approach to mother-infant contact- and distance regulation. We looked at the history of mother-infant relationship in terms of the effects of changes in the mother's and infant's behaviour on the mother-infant dyad as a self-regulating homeostatic system. A single-subject research design was used, because it has special advantages for the study of developmental processes, not shared by the usual experimental- or correlational designs. We found that the infants' progress towards greater independence proceeds discontinuously in 5 jumps over the first 24 months. With each jump the following drastic changes are found in the quality of the distance-regulation between mother and infant and/or the amount of time spent at a certain distance: 1. After month 2 maternal behaviour related to carrying and supporting the babies in the ventro-ventral position decreased sharply and the babies were forced to cling to the mother themselves. 2. In month 5-6 a period of mother-infant conflict was found in which aggressive maternal rebuffs were aimed at breaking nipple- and ventro-ventral contact. Before this age the mothers were primarily responsible for ventro-ventral contact, and after this age the infants were. A relation was found between the rebuffs and the onset of dorsal riding and eating solid food. These changes in the mother-infant interaction coincided with physical changes such as a change in the speed of growth and the eruption of teeth. The findings are placed in the wider framework of mammalian development. 3. After month 7 an explosion of the frequency of the infants' excursions was found and the infants now made short lasting excursions and remain within arm's reach from their mothers. It is argued that such excursion behavior expresses the infants' concern with the distance to their mothers at this age. 4. Around month 12 and month 18 periods of mother-infant conflict were found in which the aggressive maternal rebuffs were aimed at breaking body contact. Both periods of mother-infant conflict were associated with peaks in the infants' responsibility for body contact and with rises in the amount of time spent in this contact. Both periods of motherinfant conflict were followed by sharp drops in the amount of time spent in body contact, and after each drop time spent in body contact remained at a newly reached level. Furthermore, after both periods of mother-infant conflict all infants made use of space more distant from their mothers. We found that mothers do not promote the infants' independence as a whole in each confict period, but that they do so only for a particular aspect in training the infants how
80
to adapt to other individuals and to the physical outer world when using the new ability. It is argued that mothers recognize the ages that their infants are ready to reorganize their behaviour, upon which they force them to do so. The periods of mother-infant conflict around month 12 and 18 are preceded by periods of regression: temporary shifts back to mainly staying in closer contact (= qualitative regressive shifts) and temporary increases in the amount of time spent in ventro-ventral contact (= quantitative regressive peaks). It is suggested that regressive behaviour is a common feature of normal development after a certain age. We called a period in which the succession of regression and/or conflict and/or jump towards greater independence was found a labile period, as opposed to stabile periods. In the general discussion the following topics are attended to: a) The possibility that "labile periods at specific ages" in the mother-infant relationship are a common feature in normal development. b) The possibility that changes in maternal behaviour in each labile period are responsible for the phenomenon of jumps in the growing independence, provided that the infants are not pushed beyond their maturational abilities. c) The possibility that regression, which precedes mother-infant conflict, is associated with maturational changes in the infant is discussed. Several authors associated regressive behaviour with spurts in development. d) The possibility that maturational changes are associated with developmental steps. Infants contribute to changes in the mother-infant systems as they change maturationally. The latter changes were established independently of their effect on the motherinfant relationship by PLOOIJ (1984). He looked for qualitative changes over development in behaviour and recognized 5 steps in the infant's first year of life. It is discussed that the onsets of labile periods in the mother-infant relationship are associated with the developmental steps. It seems that a developmental step triggers mother-infant conflict, and that mother's behaviour during this conflict is of vital importance in realizing the infant's developmental potential. PETERFREUND'S (1971) thinking on the relation between maturation and learning (reprogramming) suggests mother's helpful contributions to what happens around the times of occurrence of developmental steps. We related our findings to BOWLBY'S (1969, 1973, 1980) attachment theory. The possibility is discussed that attachment takes different forms over age, depending on the proximity involved and the skills available to the infant, and that it is active and present from birth onwards.
References P. & KRAMES,L. (1977). Attachment behavior. Advances in the study of communication and affect, vol. 3. Plenum Press, New York. ALS, H. (1979). Social interaction: Dynamic matrix for developing behavioral organization. - In: New directions for child development, number 4: Social interaction and communication during infancy (I. C. UZGIRIs, ed.). Jossey-Bass Inc., Publ., San Francisco. Harvard University Press, CamALTMANN, J. (1980). Baboon mothers and infants. bridge Massachusetts and London. BAERENDS,G. P. (1956). Aufbau des tierischen Verhaltens. - In: Handbuch der & Zoologie (VIII) (J.-G. HELMCKE G. C. HIRSCH, eds). Berlin: Walter de Gruyter
ALLOWAY, Th., PLINER, & Co.
(1970). A model of the functional organization of incubation behaviour. Behaviour Suppl. 17, p. 263-312. -(1976). The functional organization of behaviour. - Anim. Behav. 24, p. 726-738. BALDWIN, A. L. (1967). Theories of child development. - New York: Wiley.
81
S. BARNETT, A. (1977). Ethology and man: Science or myth? - Develop. Med. Child Neurol. 19, p. 252-264. BASTOCK, M., MORRIS, D. & MOYNIHAN,M. (1953). Some comments on conflict and thwarting in animals. - Behaviour 6, p. 56-84. BATESON, P. P. G. (1976). Rules and reciprocity in behavioural development. - In: & Growing points in ethology (P. P. G. BATESON R. A. HINDE, eds). Cambridge Un. Press, Cambridge. - Anim. Behav. 27, -(1979). How do sensitive periods arise and what are they for? p. 470-486. - (1983). The interpretation of sensitive periods. - In: The behavior of human infants (A. OLIVERIO M. ZAPELLA, & eds). Plenum Press, New York. BERGER, J. (1979). Weaning conflict in desert and mountain bighorn sheep (Ovis - Zeitschr. canadensis): An ecological interpretation. Tierpsychol. 50, p. 188-200. - In: BLURTON JONES, N. (1972). Comparative aspects of mother-child contact. studies of child behaviour (N. BLURTON JONES, ed.). Cambridge Un. Ethological Press, London. - Penguin Books. BOWLBY, J. (1969). Attachment and loss. Volume I. Attachment. - (1973). Idem. Volume II. Separation, anxiety and anger. - The Hogarth Press, London, and the Institute of Psycho-analysis. -(1980). Idem. Volume III. Loss, sadness and depression. - The Hogarth Press, London, and the Institute of Psycho-analysis. BRANDT, E. & MITCHELL, G. (1971). Parturition in primates: behavior related to birth. - In: Primate behavior: Developments in field and laboratory research, vol. 2 (L. ROSENBLUM, ed.). Academic Press, New York. I. BRETHERTON, (1985). Attachment theory: retrospect and prospect. - In: Growing & points of attachment theory and research (I. BRETHERTON E. WATERS, eds). Monographs of the Society for Research in Child Development, serial no. 209, vol. 50, nos 1-2. BRONSON, G. W. (1972). Infants' reaction to unfamiliar persons and novel objects. Monogr. Soc. Research Child Devel. 37, Serial number 148. CASSEL,T. Z. K. & SANDER,L. W. (1975). Neonatal recognition processes and attachment: effects of masking the mother's face at seven days of life. - Paper presented at meeting Soc. Res. Child Devel., Denver, Colorado. CHEVALIER-SKOLNIKOFF, S. (1977). A Piagetian model for describing and comparing socialization in monkey, ape, and human infants. - In: Chevalier-Skolnikoff and Poirier, 1977. -& POIRIER,F. E. (1977). Primate bio-social development: biological, social and ecological determinants. - Garland Publishing, Inc.: New York & London. CLARK, C. B. (1977). A preliminary report on weaning among chimpanzees of the Gombe National Park, Tanzania. - In: Chevalier-Skolnikoff and Poirier, 1977. T. CLUTTON-BROCK, H. (1972). Feeding and ranging behaviour of the red colobus monkey. - Ph.D. thesis, University of Cambridge. - (1975). Feeding behaviour of Colobusguereza in East-Africa. - Folia Primatol. 23, p. 165-207. - , ALBON,S. &GUINNESS, F. (1981). Parental investment in male and female offspring in polygynous mammals. - Nature 289, p. 487-489. CONDON, W. S. (1977). A primary phase in the organization of infant responding behaviour. - In: Studies in mother-infant interaction (H. R. SCHAFFER, ed.). London: Academic Press. - In: Before speech: The begin-(1979). Neonatal entrainment and enculturation. of interpersonal communication (M. BULLOWA, ed.). Cambridge: Cambridge ning University Press. CooLS, A. R. (1985). Brain and behavior: hierarchy of feedback systems and control of
82
- In: & input. Perspectives in Ethology, volume 6 (P. P. G. BATESON P. H. KLOPFER,eds). New York: Plenum Press. DAVENPORT, R. K. (1979). Some behavioral disturbances of great apes in captivity. In: The great apes, perspectives on human evolution, vol. V (D. A. HAMBURG E. & R. McCOWN, eds). The Benjamin/Cummings Publ. Co., Menlo Park, California. DE CASPER, A. J. & FIFER, W. P. (1980). Of human bonding: Newborns prefer their mothers' voices. - Science 208, p. 1174-1176. V. DENENBERG, H. (1978). Paradigms and paradoxes in the study of behavioral development. - In: The origins of the infant's social responsiveness (E. B. THOMAN,ed.). Hillsdale, N.J., Lawrence Erlbaum Assoc. -(1982). Comparative psychology and single-subject research. - In: Single-case research designs (A. E. KAZDIN& A. H. TUMA, eds). San Francisco: Jossey-Bass. DOUGLAS, J. W. B. (1975). Early hospital admissions and later disturbances of behaviour and learning. - Dev. Med. & Child Neurol. 17, p. 456-480. S. ESCALONA, K. (1968). The roots of individuality. - Tavistock Publications. FEIJOO, (1976). Ut conscientia noscatur. - Les Cahier de Sophrologie 13, p. 14-20. J. FENTRESS, J. C. (1976). Behavioral networks and the simpler systems approach. - In: Simpler networks and behavior (J. C. FENTRESS, ed.). Sunderland, MA: Sinauer Associates, Inc. -(1983). Ethological models of hierarchy and patterning of species specific behavior. - In: Handbook of behavioral neurobiology: Volume 6, Motivation (E. SATINOFF & P. TEITELBAUM, eds). New York: Plenum Press. D. The first FOSSEY, (1979). Development of the mountain gorilla (Gorilla gorilla beringez): thirty-six months. - In: The great apes. Perspectives on human evolution, volume V (D. A. HAMBURG E. R. McCOWN, eds). The Benjamin/Cummings Publ. Co., & Menlo Park, California. D. FREEDMAN, G. (1974). Human infancy: An evolutionary perspective. - John Wiley & Sons, New York. FUCHS, S. (1982). Optimality of parental investment: the influence of nursing on reproductive success of mother and female young house mice. - Behav. Ecol. Sociobiol. 10, p. 39-51. GAVAN,J. A. (1971). Longitudinal postnatal growth in chimpanzee. - In: The chimpanzee, vol. 4 (J. BOURNE,ed.), pp. 46-102. Karger, Basel. GEWIRTZ, J. L. (1972). Attachment and dependency. - V. H. Winston, New York. I. (1976). Homeostatic motor processes in mammalian interactions: A GOLANI, choreography of display. - In: Perspectives in ethology (Vol. 2) (P. P. G. BATESON & P. H. KLOPFER,eds). New York: Plenum Press. GOTTLIEB, G. (1976a). Conceptions of prenatal development: behavioral embryology. Psychol. Rev. 83, p. 215-234. - (1976b). The roles of experience in the development of behavior and the nervous system. - In: Neural and behavioral specificity (G. GOTTLIEB, ed.). New York: Academic Press. HANSEN, E. W. (1966). The development of maternal and infant behaviour in rhesus monkeys. - Behav. 27, p. 107-149. HARLOW,H. F. (1958). The nature of love. - Amer. Psychol. 13, p. 673-685. - (1971). The affectional systems. - In: Psychology (H. F. HARLOW, J. L. McGAUGH & R. F. THOMPSON, eds). Albion Press, San Francisco. -- & HARLOW,M. K. (1965). The affectional systems. - In: Behavior of nonhuman & primates, vol. 2 (A. M. SCHRIER,H. F. HARLOW F. STOLLNITZ, eds). Academic Press, New York. & SUOMI, S. J. (1971). From thought to therapy: lessons from a primate , -laboratory. - Amer. Scient. 59, p. 538-549. - &ZIMMERMAN, R. (1959). Affectional responses in the infant monkey. - Science R. 130, p. 421-432.

HINDE, R. A. (1953).
83
Appetitive behaviour, consummatory act, and the hierarchical organization of behaviour, with special reference to the great tit (Parus major). Behaviour 5, p. 189-224. - (1969). Analyzing the roles of the partners in a behavioural interaction - motherinfant relations in rhesus macaque. - Ann. N. Y. Acad. Sc. 159, p. 651-667. -(1970). Animal Behaviour. A synthesis of ethology and comparative psychology. McGraw-Hill, Kogakusha, Ltd., Tokyo. -- (1971). Development of social behaviour. - In: Behavior of non-human primates, & vol. 3 (A. SCHRIER F. STOLNITZ, eds). New York: Academic Press. - (1974a). Biological bases of human social behaviour. - McGraw-Hill: New York. -(1974b). Mother-infant relations in rhesus monkeys. - In: Ethology and psychiatry (N. F. WHITE, ed.). University of Toronto Press, Toronto. -- (1977). Mother-infant separation and the nature of inter-individual relationships: experiments with rhesus-monkeys. - Prc. R. Soc. (London) 196, p. 29-50. - (1983). Primate social relationships. An integrated approach. - Oxford: Blackwell Scientific Publications. -- &ATKINSON, (1970). Assessing the roles of social partners in maintaining mutual S. proximity, as exemplified by mother-infant relations in rhesus monkeys. - Anim. Behav. 18, p. 169-176. -- & SPENCER-BOOTH, (1967). The behaviour of socially living rhesus monkeys in Y. their first two and a half years. - Anim. Behav. 15, p. 169-196. - &-(1971). Effects of brief separation from mothers on rhesus monkeys. Science (N.Y.) 173, p. 111-118. -- &WHITE, L. E. (1974). Dynamics of a relationship: rhesus mother-infant ventroventral contact. - J. Comp. Physiol. Psychol. 86, p. 8-23. HOFER, M. A. (1978). Hidden regulatory processes in early social relationships. - In: & Perspectives in ethology (Vol. 3): Social behavior (P. P. G. BATESON P. H. KLOPFER, eds). New York: Plenum Press. -(1981). The roots of human behavior. An introduction to the psychobiology of early development. - San Francisco: W. H. Freeman and Co. HORR, D. A. (1977). Orang-utan maturation: Growing up in a female world. - In: Chevalier-Skolnikoff and Poirier, 1977. R. H. (1974). Regressive periods in primate behavioral development with HORWICH, reference to other mammals. - Primates 15, p. 141-149. JENSEN, G., BOBBIT, R. &GORDON, A. (1973). Mother's and infant's roles in the development of independence of Macaca nemestrina. - Primates 14, p. 79-88. JOLLY, A. (1972). The evolution of primate behaviour. - MacMillan: New York. - Scient. Amer. 226, p. 74-82. KAGAN, J. (1972). Do infants think? -In: Lewis and (1974). Discrepancy, temperament, and infant distress. Rosenblum, 1974. KENNELL, J. H., TRAUSE, M. A. &KLAUS, M. H. (1975). Evidence for a sensitive period in the human mother. - In: Parent-infant interaction. Ciba Foundation Symposium 33, Elsevier, Amsterdam. KLEIN, M. (1935). A contribution to the psychogenesis of manic depressive states. - In: Love, guilt and reparation and other works, 1921-1945. The Hogarth Press, London, 1975. -(1940). Mourning and its relation to manic-depressive states. In: Love, guilt and reparation and other works, 1921-1945. The Hogarth Press, London, 1975. M. J. (1976). Maternal care, infant behavior, and development among the KONNER, !Kung. - In: Kalahari hunter-gatherers: studies of the !Kung San and their neighbors (R. B. LEE & I. DEVORE, eds). Harvard Un. Press, Cambridge, Mass. A. KoPP, C. B., SIGMAN,M. & PARMELEE, H. (1974). A longitudinal study of sensorimotor development. - Dev. Psychol. 10, p. 687-695. A. (1955). Aspects and prospects of the concept of instinct (vicissitudes of KORTLANDT, the hierarchy theory). - Arch. neerl. Zool. 11, p. 155-284.
84
KUNST, M. S. (1948).
A study of thumb- and finger-sucking in infants. - Psychol. Monogr. 62, p. 1-71. - The National LAWICK-GOODALL, J. VAN (1967). My friends the wild chimpanzees. Geographic Society. -- (1968). The behaviour of free-living chimpanzees in the Gombe Stream Reserve. - Anim. Behav., Monogr. 1, p. 161-311. -(1973). Cultural elements in a chimpanzee community. - Symposium of the IVth International Congress of Primatology 1, p. 144-184. Karger, Basel. LEMPERS, J. D., FLAVELL, E. R. & FLAVELL, J. H. (1977). The development in very young children of tacic knowledge concerning visual perception. -- Gen. Psychol. Monographs 95, p. 3-53.
LEWIS, M. & ROSENBLUM, L. A. (1974). The origins of fear. Wiley and Sons, New
York. LILEY, A. (1972). The fetus as a personality. - Austr. N. Zeal. J. Psychiatry, 6, p. 99-105. LINDBURGH, D. & HAZELL, L. (1972). Licking of the neonate and duration of labor in great apes and man. - Amer. Anthrop. 74, p. 318-325. MCCALL, R. B., EICHHORN, D. H. & HOGARTY, P. S. (1977). Transitions in early mental - Monogr. Soc. Res. Child Devel. 42, No. 171. development. MCFARLAND, D. J. (1971). Feedback mechanisms in animal behaviour. - London: Academic Press. MCGRAW, M. B. (1945). The neuromuscular maturation of the human infant. - Hafner Press, New York, fifth printing, 1974. MAIN, M. & GOLDWYN,R. (1984). Predicting rejection of her infant from mother's representation of her own experience: implications for the abused-abusing - Child abuse & neglect 8, p. 203-217. intergenerational cycle. KAPLAN, N. & CASSIDY, J. (1985). Security in infancy, childhood, and adulthood: --, a move to the level of representation. - In: Growing points of attachment theory & and research (I. BRETHERTON E. WATERS, eds). Monographs of the Soc. Res. Child Devel. 209, vol. 50, nos 1-2. Chicago: The University of Chicago Press. MARSDEN, C. D., MERTON, P. A., MORTON, H. B. & ADAM, J. E. R. (1978). The role of afferent feedback in the regulation of movement. - In: Recent advances in primatology (Vol. 1): Behaviour (D. J. CHIVERS& J. HERBERT,eds). London: Academic Press. MARTIN, P. (1986). An experimental study of weaning in the domestic cat. - Behaviour 99, p. 221-249. MASON, W. A. (1968). Early social deprivation in the nonhuman primates: Implications for human behavior. - In: Biology and behavior: Environmental influences (D. C. GLASS, ed.). Rockefeller Un. Press, New York. MONTAGU,A. (1962). Prenatal Influences. - Springfield, Ill.: Charles C. Thomas. MORGAN, G. A. &RICCIUTI, H. N. (1969). Infants' responses to strangers during the first year. - In: Determinants of infant behaviour, vol. 4 (B. M. Foss, ed.). Methuen, London. P. MOUNOUD, (1976). The development of systems of representation and treatment in the child. - In: Piaget and his school. A reader in developmental psychology (B. INHELDER H. CHIPMAN, eds). Springer Verlag, Berlin. & NICOLSON, N. A. (1977). A comparison of early behavioral development in wild and captive chimpanzees. - In: Chevalier-Skolnikoff and Poirier, 1977. E. PETERFREUND, (1971). Information, systems, and psychoanalysis. An evolutionary biological approach to psychoanalytical theory. - International Universities Press, New York. F. PLOOIJ, X. (1984). The behavioural development of free-living chimpanzee babies and infants. - Norwood, New Jersey: Ablex Publ. Comp. POIRIER,F. E. (1977). Introduction. - In: Chevalier-Skolnikoff and Poirier, 1977.
85
POLIT, A. & BIZZI, E. (1979). Characteristics of motor programs underlying arm movements in monkeys. -J. Neurophysiol. 42, p. 183-194. POND, C. M. (1977). The significance of lactation in the evolution of mammals. Evolution 31, p. 177-199. POWERS,W. T. (1973). Behavior: the control of perception. - Chicago, Ill: Aldine Publishing Co. PUSEY, A. E. (1978a). Age changes in the motor-offspring association of wild chimpanzees. - In: Recent advances in primatology. Volume One: Behaviour (D. J. CHIVERS&J. HERBERT,eds). Academic Press, London. - (1978b). The physical and social development of wild adolescent chimpanzees (Pan Ph.D. thesis, Stanford University. University troglodytes schweinfurthil). Microfilms International, Order no. 7808824. - (1980). Inbreeding avoidance in chimpanzees. - Anim. Behav. 28, p. 543-552. P. RAJECKI,D. W., LAMB, M. E. & OBMASCHER, (1978). Toward a general theory of infantile attachment: A comparative review of aspects of the social bond. - Behav. Brain Sciences 1, p. 417-464. K. RANDOLPH,P. A., RANDOLPH, C., MATTINGLY, & MEAD, M. F. (1977). Energy J. costs of reproduction in the cotton rat, Sigmodon hispidus. - Ecology 58, p. 31-45. REDSHAW, M. (1978). Cognitive development in human and gorilla infants. - J. hum. evol. 7, p. 133-141. RHEINGOLD, H. L. & ECKERMAN, C. 0. (1970). The infant separates himself from his mother. - Science 168, p. 78-83. RIESEN, A. H. & KINDER, E. F. (1952). Postural development of infant chimpanzee. Yale Un. Press, New Haven. RIJT-PLooIJ, H. H. C. VANDE(1982). Mother-infant relations in free-living chimpanzees of the Gombe National Park, Tanzania. - Ph.D. thesis, Cambridge, U.K. RUTTER,M. (1978a). Early sources of security and competence. - In: Human growth and development. Wolfson College lectures 1976 (J. BRUNER& A. GARTON, eds). Clarendon Press, Oxford. - (1978b). Family, area and school influences in the genesis of conduct disorders. In: Aggression and anti-social behaviour in childhood and adolescence. Suppl. J. child psychol. & Psychiatry 1 (L. A. HERSOV M. BERGER, eds). Pergamon Press. & - (1979). Maternal deprivation, 1972-1978: New findings, new concepts, new approaches. - Child Devel. 50, p. 283-305. -- (1980a). Introduction. - In: Scientific foundations of developmental Psychiatry (M. RUTTER, ed.). Heinemann Medical Books, London. -(1980b). Attachment and the development of social relationships. - In: Idem. Maternal deprivation reassessed. Second edition. - Penguin Books. --(1981). SANDER, L. W. (1969). Regulation and organization in the early infant-caretaker system. In: Brain and early behavior (R. J. ROBINSON, ed.). Academic Press, New York. --, JULIA, H. L., STECHLER, G. & BURNS, P. (1972). Continuous 24-hour interactional monitoring in infants reared in two caretaking environments. - Psychosom. Med. 34, p. 270-282. SCHAFFER, H. R. (1974). Cognitive components of the infant's response to strangeness. - In: Lewis & Rosenblum, 1974. SHIRLEY,M. & POYNTZ,L. (1941). The influence of separation from the mother on children's emotional responses. - J. Psychol. 12, p. 251-282. SIMPSON, M. J. A. & SIMPSON, A. E. (1977). One-zero and scan methods for sampling behaviour. - Anim. Behav. 25, p. 726-731. L. SONTAG, W. (1970). Prenatal determinants of postnatal behavior. - In: Fetal growth and development (H. A. WEISMAN& G. KERR, eds). New York: McGraw Hill. E. SORENSON, R. (1979). Early tactile communication and the patterning of human organization: a New Guinea case study. - In: Before speech: the beginning of
86
interpersonal communication (M. BULLOWA, ed.). Cambridge Un. Press, Cambridge. SPELT, D. K. (1948). The conditioning of the human fetus in utero. -J. Exp. Psychol. 38, p. 338-346. SROUFE,L. A. &WATERS,E. (1977). Attachment as an organizational construct. - Child Devel. 48, p. 1184-1199. STEINBACHER, G. (1941). Geburt und Kindheit eines Schimpansen. - Z. Tierpsychol. 4, p. 188-203. TEITELBAUM, Ph., SCHALLER, T. & WHISHAW, I. Q. (1983). Sources of spontaneity in motivated behavior. - In: Handbook of behavioral neurobiology, vol. 6: Motiva& tion (E. SATINOFF Ph. TEITELBAUM, eds). New York: Plenum Press. THOMAN,E. B. (1975). How a rejecting baby affects mother-infant synchrony. - In: Parent-infant interaction. CIBA Foundation Symposium number 33, Elsevier, Amsterdam. C. TREVARTHEN, & HUBLEY, P. (1978). Secondary intersubjectivity: confidence, confiding and acts of meaning in the first year. - In: Action, gesture and symbol: the emergence of language (A. LOCK, ed.). Academic Press, London. TURNEY,T. H. (1978). Human neonatal and infant behavioral assessment scales being applied to chimpanzees. - Laboratory Primate Newsletter 17, p. 14-15. UZGIRIS, I. C. (1973). Patterns of cognitive development in infancy. - Merrill-Palmer Quart. 19, p. 181-204. WATZLAWICK, P., BEAVIN, J. H. &JACKSON, D. D. (1967). Pragmatics of human communication. - Norton, New York. WERNER, H. (1948). Comparative psychology of mental development. Chicago: Follett. - (1957). The concept of development from a comparative and organismic point of view. - In: The concept of development: an issue in the study of human behavior (D. HARRIS, ed.). University of Minnesota Press. K. R. WOOD, S., MORIARTY, M., GARDNER,B. T. & GARDNER, A. (1980). Object permanence in child and chimpanzee. - Anim. learn. & behav. 8, p. 3-9. WRANGHAM, R. W. (1974). Artificial feeding of chimpanzees and baboons in their natural habitat. - Anim. Behav. 22, p. 83-93. -(1975). The behavioural ecology of chimpanzees in Gombe National Park, Tanzania. - Ph.D. thesis, University of Cambridge. WULFFTEN PALTHE, T. VAN &HOPKINS, B. (1984). A developmental approach to the study of early mother-infant interaction. - In: Continuity of neural functions from preto postnatal life (H. F. R. PRECHT,ed.). Oxford: Blackwell Scientific Publications. YERKES,R. M. (1943). Chimpanzees. A laboratory colony. - Yale University Press: New Haven, Connecticut. -& TOMILIN,M. I. (1935). Mother-infant relations in chimpanzee. - J. Comp. Psychol. 20, p. 321-360.

Mother-Infant Relations in Free-Ranging Chimpanzees

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Mother-Infant Relations in Free-Ranging Chimpanzees

Uploaded by

Copyright:

Available Formats

Growing Independence, Conflict and Learning in Mother-Infant Relations in Free-Ranging Chimpanzees Author(s): Hedwig H. C.

GROWING INDEPENDENCE, CONFLICT AND LEARNING IN MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

P. BAERENDS, ROON, Prof. R. A. HINDE, Prof. A. KORTLANDT, DrJ. M. BAERENDS-VAN

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ 35 41 62 62 63 66 68 73 79 80

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

analysing behaviour, and by providing hypotheses that can be tested

Many researchers have stressed the usefulness of conceptualizing the

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

TABLE 1. The study subjects

TABLE 2. Family ties of FF + FD

*) Flo and FT died when FD was 15/16 months old, resp.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

the assessment resulting from the scan-

Regulation of contact and proximity.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

The umbilical cord and afterbirth.

Mother avoided social contacts.

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

Fig. 2. Individual differences in frequency of effort-grunts.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

Fig. 3. The decrease of the % of securing over age.

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

reported that one mother (Melissa)

in maternal 'securing' between month 2 and 3 in the two mother-baby

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

.. Maenlages, Fig 4.. er ab.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

-40 -50 -60 -70 -80 / /

Fig. 5. Baby's role in ventro-ventral contact (%MKi-%BKi) over age.

PF %Breaks 100 100 96 87

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

The onset of frequent dorsal riding and eating solidfood.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

Fig. 6. Frequency of nipple contact-makes per 300 mins of good observation.

Fig. 7. Amount of ventro-ventral contact over age.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

Fig. 8. Profiles of maternal behaviour in association with ventro-ventralcontact over age.

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

19 weeks 0 1 AGE (years) 2

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

1968; CLARK, 1977). The frequency

from her data.

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

frequency of sucking reflects a similar peak in conflict occurring in Flint.

mothers over nipple contact as well.

XXX XXX XXX

Fig. 10. Summary diagram of the peaks in conflict.

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

H. H. C. VAN DE RIJT-PLOOIJ & F. X. PLOOIJ

MOTHER-INFANT RELATIONS IN FREE-RANGING CHIMPANZEES

---ot0 month 1 2 3-4 4 5 67 6

> no contact-breaks < 1 38 40 25 104 187 1 39 43 25 106 190 2 2 10 10 5 5 (no contact-breaks) 35 33 78 75

1.5 % of time that FF + FD were out of contact < m or 1.55 5 m

> no contact-breaks< 50 72 80 36 81 85 82 99 90 80 89 91 85 81 88 88 80 100 93 98 91 91 91 93 89 87 91 93 94 87 82 83 90 91 33 67 93 93 (no contact-breaks) 83 100 81 92 66 83 89 88 86 82 94 86 94 90 79 89 79 92