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de Waal, F. B. M., & Davis, J. M. (2003). Capuchin cognitive ecology: cooperation based on projected returns. Neuropsychologia, 41(2), 221–228.
Abstract: Stable cooperation requires that each party's pay-offs exceed those available through individual action. The present experimental study on brown capuchin monkeys (Cebus apella) investigated if decisions about cooperation are (a) guided by the amount of competition expected to follow the cooperation, and (b) made instantaneously or only after a period of familiarization. Pairs of adult monkeys were presented with a mutualistic cooperative task with variable opportunities for resource monopolization (clumped versus dispersed rewards), and partner relationships (kin versus nonkin). After pre-training, each pair of monkeys (N=11) was subjected to six tests, consisting of 15 2 min trials each, with rewards available to both parties. Clumped reward distribution had an immediate negative effect on cooperation: this effect was visible right from the start, and remained visible even if clumped trials alternated with dispersed trials. The drop in cooperation was far more dramatic for nonkin than kin, which was explained by the tendency of dominant nonkin to claim more than half of the rewards under the clumped condition. The immediacy of responses suggests a decision-making process based on predicted outcome of cooperation. Decisions about cooperation thus take into account both the opportunity for and the likelihood of subsequent competition over the spoils.
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Bonnie, K. E., & de Waal, F. B. M. (2006). Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees. Primates, 47(1), 27–34.
Abstract: Handclasp grooming is a unique social custom, known to occur regularly among some, but not all populations of chimpanzees (Pan troglodytes). As with other cultural behaviors, it is assumed that this distinctive grooming posture is learned socially by one individual from another. However, statistical comparisons among factors thought to influence how a behavior spreads within a group have never, to our knowledge, been conducted. In the present study, the origination and spread of handclasp grooming in a group of captive chimpanzees was followed throughout more than 1,500 h of observation over a period of 12 years. We report on the frequency, bout duration, and number and demography of performers throughout the study period, and compare these findings to those reported for wild populations. We predicted that dyads with strong affiliative ties, measured by time spent in proximity to and grooming one another, were likely to develop a handclasp grooming partnership during the study period. A quadratic assignment procedure was used to compare correlations among observed frequencies of grooming and proximity with handclasp grooming in all possible dyads within the group. As predicted, the formation of new handclasp grooming dyads was positively correlated with the rate of overall grooming and proximity within a dyad. In addition, in nearly all dyads formed, at least one individual had been previously observed to handclasp groom. We concluded that affiliation and individual experience determines the transmission of handclasp grooming among captive chimpanzees.
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Brosnan, S. F., & de Waal, F. B. M. (2005). Responses to a simple barter task in chimpanzees, Pan troglodytes. Primates, 46(3), 173–182.
Abstract: Chimpanzees (Pan troglodytes) frequently participate in social exchange involving multiple goods and services of variable value, yet they have not been tested in a formalized situation to see whether they can barter using multiple tokens and rewards. We set up a simple barter economy with two tokens and two associated rewards and tested chimpanzees on their ability to obtain rewards by returning the matching token in situations in which their access to tokens was unlimited or limited. Chimpanzees easily learned to associate value with the tokens, as expected, and did barter, but followed a simple strategy of favoring the higher-value token, regardless of the reward proffered, instead of a more complex but more effective strategy of returning the token that matched the reward. This response is similar to that shown by capuchin monkeys in our previous study. We speculate that this response, while not ideal, may be sufficient to allow for stability of the social exchange system in these primates, and that the importance of social barter to both species may have led to this convergence of strategies.
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de Waal, F. B. M. (2005). How animals do business. Sci Am, 292(4), 54–61. |
de Waal, F. B. M. (2008). Putting the Altruism Back into Altruism: The Evolution of Empathy. Annu Rev Psychol, 59(1), 279–300.
Abstract: Evolutionary theory postulates that altruistic behavior evolved for the return-benefits it bears the performer. For return-benefits to play a motivational role, however, they need to be experienced by the organism. Motivational analyses should restrict themselves, therefore, to the altruistic impulse and its knowable consequences. Empathy is an ideal candidate mechanism to underlie so-called directed altruism, i.e., altruism in response to anothers's pain, need, or distress. Evidence is accumulating that this mechanism is phylogenetically ancient, probably as old as mammals and birds. Perception of the emotional state of another automatically activates shared representations causing a matching emotional state in the observer. With increasing cognition, state-matching evolved into more complex forms, including concern for the other and perspective-taking. Empathy-induced altruism derives its strength from the emotional stake it offers the self in the other's welfare. The dynamics of the empathy mechanism agree with predictions from kin selection and reciprocal altruism theory.
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Plotnik, J., Nelson, P. A., & de Waal, F. B. M. (2003). Visual field information in the face perception of chimpanzees (Pan troglodytes). Ann N Y Acad Sci, 1000, 94–98.
Abstract: Evidence for a visual field advantage (VFA) in the face perception of chimpanzees was investigated using a modification of a free-vision task. Four of six chimpanzee subjects previously trained on a computer joystick match-to-sample paradigm were able to distinguish between images of neutral face chimeras consisting of two left sides (LL) or right sides (RR) of the face. While an individual's ability to make this distinction would be unlikely to determine their suitability for the VFA tests, it was important to establish that distinctive information was available in test images. Data were then recorded on their choice of the LL vs. RR chimera as a match to the true, neutral image; a bias for one of these options would indicate an hemispatial visual field advantage. Results suggest that chimpanzees, unlike humans, do not exhibit a left visual field advantage. These results have important implications for studies on laterality and asymmetry in facial signals and their perception in primates.
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de Waal, F. B. M. (2003). Animal communication: panel discussion. Ann N Y Acad Sci, 1000, 79–87. |
de Waal, F. B. M. (2003). Darwin's legacy and the study of primate visual communication. Ann N Y Acad Sci, 1000, 7–31.
Abstract: After Charles Darwin's The Expression of the Emotions in Man and Animals, published in 1872, we had to wait 60 years before the theme of animal expressions was picked up by another astute observer. In 1935, Nadezhda Ladygina-Kohts published a detailed comparison of the expressive behavior of a juvenile chimpanzee and of her own child. After Kohts, we had to wait until the 1960s for modern ethological analyses of primate facial and gestural communication. Again, the focus was on the chimpanzee, but ethograms on other primates appeared as well. Our understanding of the range of expressions in other primates is at present far more advanced than that in Darwin's time. A strong social component has been added: instead of focusing on the expressions per se, they are now often classified according to the social situations in which they typically occur. Initially, quantitative analyses were sequential (i.e., concerned with temporal associations between behavior patterns), and they avoided the language of emotions. I will discuss some of this early work, including my own on the communicative repertoire of the bonobo, a close relative of the chimpanzee (and ourselves). I will provide concrete examples to make the point that there is a much richer matrix of contexts possible than the common behavioral categories of aggression, sex, fear, play, and so on. Primate signaling is a form of negotiation, and previous classifications have ignored the specifics of what animals try to achieve with their exchanges. There is also increasing evidence for signal conventionalization in primates, especially the apes, in both captivity and the field. This process results in group-specific or “cultural” communication patterns.
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Preston, S. D., & de Waal, F. B. M. (2002). Empathy: Its ultimate and proximate bases. Behav Brain Sci, 25(1), 1–20; discussion 20–71.
Abstract: There is disagreement in the literature about the exact nature of the phenomenon of empathy. There are emotional, cognitive, and conditioning views, applying in varying degrees across species. An adequate description of the ultimate and proximate mechanism can integrate these views. Proximately, the perception of an object's state activates the subject's corresponding representations, which in turn activate somatic and autonomic responses. This mechanism supports basic behaviors (e.g., alarm, social facilitation, vicariousness of emotions, mother-infant responsiveness, and the modeling of competitors and predators) that are crucial for the reproductive success of animals living in groups. The Perception-Action Model (PAM), together with an understanding of how representations change with experience, can explain the major empirical effects in the literature (similarity, familiarity, past experience, explicit teaching, and salience). It can also predict a variety of empathy disorders. The interaction between the PAM and prefrontal functioning can also explain different levels of empathy across species and age groups. This view can advance our evolutionary understanding of empathy beyond inclusive fitness and reciprocal altruism and can explain different levels of empathy across individuals, species, stages of development, and situations.
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de Waal, F. B. M., & Luttrell, L. M. (1988). Mechanisms of social reciprocity in three primate species: Symmetrical relationship characteristics or cognition? Ethology and Sociobiology, 9(2–4), 101–118.
Abstract: Agonistic intervention behavior was observed in captive groups of chimpanzees (Pan troglodytes), rhesus monkeys (Macaca mulatta), and stumptail monkeys (M. arctoides). Reciprocity correlations of interventions were determined while removing from the data the effects of several symmetrical relationship characteristics, that is, matrillineal kinship, proximity relations, and same-sex combination. It was considered likely that if significant reciprocity persisted after controlling for these characteristics, the reciprocity was based on cognitive mechanisms. Statistical significance was tested by means of recently developed matrix permutation procedures. All three species exhibited significant reciprocity with regard to beneficial interventions, even after controlling for symmetrical traits. Harmful interventions were, however, reciprocal among chimpanzees only. This species showed a “revenge system”, that is, if A often intervened against B, B did the same to A. In contrast, both macaque species showed significantly inversed reciprocity in their harmful interventions: if A often intervened against B, B rarely intervened against A. Further analysis indicates that the strict hierarchy of macaques prevents them from achieving complete reciprocity. Compared to chimpanzees, macaques rarely intervene against higher ranking group members. The observed contrast can be partially explained on the basis of differences in available space, as indicated by a comparison of indoor and outdoor living conditions for the chimpanzee colony. Yet, even when such spatial factors are taken into account, substantial behavior differences between chimpanzees and macaques remain.
Keywords: Reciprocity; Agonistic intervention; Cognition; Chimpanzees; Macaques
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