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de Waal, F. B. M., Dindo, M., Freeman, C. A., & Hall, M. J. (2005). The monkey in the mirror: hardly a stranger. Proc. Natl. Acad. Sci. U.S.A., 102(32), 11140–11147.
Abstract: It is widely assumed that monkeys see a stranger in the mirror, whereas apes and humans recognize themselves. In this study, we question the former assumption by using a detailed comparison of how monkeys respond to mirrors versus live individuals. Eight adult female and six adult male brown capuchin monkeys (Cebus apella) were exposed twice to three conditions: (i) a familiar same-sex partner, (ii) an unfamiliar same-sex partner, and (iii) a mirror. Females showed more eye contact and friendly behavior and fewer signs of anxiety in front of a mirror than they did when exposed to an unfamiliar partner. Males showed greater ambiguity, but they too reacted differently to mirrors and strangers. Discrimination between conditions was immediate, and blind coders were able to tell the difference between monkeys under the three conditions. Capuchins thus seem to recognize their reflection in the mirror as special, and they may not confuse it with an actual conspecific. Possibly, they reach a level of self-other distinction intermediate between seeing their mirror image as other and recognizing it as self.
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Flack, J. C., Krakauer, D. C., & de Waal, F. B. M. (2005). Robustness mechanisms in primate societies: a perturbation study. Proc Biol Sci, 272(1568), 1091–1099.
Abstract: Conflict management mechanisms have a direct, critical effect on system robustness because they mitigate conflict intensity and help repair damaged relationships. However, robustness mechanisms can also have indirect effects on system integrity by facilitating interactions among components. We explore the indirect role that conflict management mechanisms play in the maintenance of social system robustness, using a perturbation technique to 'knockout' components responsible for effective conflict management. We explore the effects of knockout on pigtailed macaque (Macaca nemestrina) social organization, using a captive group of 84 individuals. This system is ideal in addressing this question because there is heterogeneity in performance of conflict management. Consequently, conflict managers can be easily removed without disrupting other control structures. We find that powerful conflict managers are essential in maintaining social order for the benefit of all members of society. We show that knockout of components responsible for conflict management results in system destabilization by significantly increasing mean levels of conflict and aggression, decreasing socio-positive interaction and decreasing the operation of repair mechanisms.
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Brosnan, S. F., Schiff, H. C., & de Waal, F. B. M. (2005). Tolerance for inequity may increase with social closeness in chimpanzees. Proc Biol Sci, 272(1560), 253–258.
Abstract: Economic decision-making depends on our social environment. Humans tend to respond differently to inequity in close relationships, yet we know little about the potential for such variation in other species. We examine responses to inequity in several groups of chimpanzees (Pan troglodytes) in a paradigm similar to that used previously in capuchin monkeys (Cebus apella). We demonstrate that, like capuchin monkeys, chimpanzees show a response to inequity of rewards that is based upon the partner receiving the reward rather than the presence of the reward alone. However, we also found a great amount of variation between groups tested, indicating that chimpanzees, like people, respond to inequity in a variable manner, which we speculate could be caused by such variables as group size, the social closeness of the group (as reflected in length of time that the group has been together) and group-specific traditions.
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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.
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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. (2000). Primates--A natural heritage of conflict resolution. Science, 289(5479), 586–590.
Abstract: The traditional notion of aggression as an antisocial instinct is being replaced by a framework that considers it a tool of competition and negotiation. When survival depends on mutual assistance, the expression of aggression is constrained by the need to maintain beneficial relationships. Moreover, evolution has produced ways of countering its disruptive consequences. For example, chimpanzees kiss and embrace after fights, and other nonhuman primates engage in similar “reconciliations.” Theoretical developments in this field carry implications for human aggression research. From families to high schools, aggressive conflict is subject to the same constraints known of cooperative animal societies. It is only when social relationships are valued that one can expect the full complement of natural checks and balances.
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Parish, A. R., & De Waal, F. B. (2000). The other “closest living relative”. How bonobos (Pan paniscus) challenge traditional assumptions about females, dominance, intra- and intersexual interactions, and hominid evolution. Ann N Y Acad Sci, 907, 97–113.
Abstract: Chimpanzee (Pan troglodytes) societies are typically characterized as physically aggressive, male-bonded and male-dominated. Their close relatives, the bonobos (Pan paniscus), differ in startling and significant ways. For instance, female bonobos bond with one another, form coalitions, and dominate males. A pattern of reluctance to consider, let alone acknowledge, female dominance in bonobos exists, however. Because both species are equally “man's” closest relative, the bonobo social system complicates models of human evolution that have historically been based upon referents that are male and chimpanzee-like. The bonobo evidence suggests that models of human evolution must be reformulated such that they also accommodate: real and meaningful female bonds; the possibility of systematic female dominance over males; female mating strategies which encompass extra-group paternities; hunting and meat distribution by females; the importance of the sharing of plant foods; affinitive inter-community interactions; males that do not stalk and attack and are not territorial; and flexible social relationships in which philopatry does not necessarily predict bonding pattern.
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Judge, P. G., de Waal, F. B., Paul, K. S., & Gordon, T. P. (1994). Removal of a trauma-inflicting alpha matriline from a group of rhesus macaques to control severe wounding. Lab Anim Sci, 44(4), 344–350.
Abstract: Wounding in an 83-member group of rhesus macaques (Macaca mulatta) housed at the Yerkes Regional Primate Research Center Field Station became excessive to the point that intervention was deemed necessary. When observations indicated that three females from the group's alpha matriline were principally responsible for the wounding, the matriline (N = 7) was removed from the group. This study was conducted to document an atypical pattern of wounding in this group and to evaluate the effectiveness of removal as a procedure for controlling injuries. The aggression rates of 21 adult subjects and the wounds of all group members were recorded before and after the removal procedure and compared with those in a similar-sized group. Removing the alpha matriline did not alter aggression rates in the group or the rank order among the remaining matrilines. Aggression rates in the experimental group were also not significantly different from those in the comparison group before or after the removal. With the alpha matriline present, wounding levels in the group were significantly higher than those in the comparison group. After removal of the matriline, the frequency of wounds decreased significantly to levels similar to those of the comparison group. The pattern of excess wounding attributed to the extracted alpha females was idiosyncratic, involving removal of large patches of skin from the hindquarters of adult females or removal of the distal portion of the fingers, toes, or tail from juveniles.(ABSTRACT TRUNCATED AT 250 WORDS)
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