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      Emergent Patterns of Social Affiliation in Primates, a Model

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          Abstract

          Many patterns of affiliative behaviour have been described for primates, for instance: reciprocation and exchange of grooming, grooming others of similar rank, reconciliation of fights, and preferential reconciliation with more valuable partners. For these patterns several functions and underlying cognitive processes have been suggested. It is, however, difficult to imagine how animals may combine these diverse considerations in their mind. Although the co-variation hypothesis, by limiting the social possibilities an individual has, constrains the number of cognitive considerations an individual has to take, it does not present an integrated theory of affiliative patterns either. In the present paper, after surveying patterns of affiliation in egalitarian and despotic macaques, we use an individual-based model with a high potential for self-organisation as a starting point for such an integrative approach. In our model, called GrooFiWorld, individuals group and, upon meeting each other, may perform a dominance interaction of which the outcomes of winning and losing are self-reinforcing. Besides, if individuals think they will be defeated, they consider grooming others. Here, the greater their anxiety is, the greater their “motivation” to groom others. Our model generates patterns similar to many affiliative patterns of empirical data. By merely increasing the intensity of aggression, affiliative patterns in the model change from those resembling egalitarian macaques to those resembling despotic ones. Our model produces such patterns without assuming in the mind of the individual the specific cognitive processes that are usually thought to underlie these patterns (such as recordkeeping of the acts given and received, a tendency to exchange, memory of the former fight, selective attraction to the former opponent, and estimation of the value of a relationship). Our model can be used as a null model to increase our understanding of affiliative behaviour among primates, in particular macaques.

          Author Summary

          Individual primates distribute their affiliative behaviour (such as grooming) in complex patterns among their group members. For instance, they reciprocate grooming, direct it more to partners the higher the partner's rank, use it to reconcile fights and do so in particular with partners that are more valuable. For several types of patterns (such as reconciliation and exchange), a separate theory based on specific cognitive processes has been developed (such as individual recordkeeping, a tendency to exchange, selective attraction to the former opponent, and estimation of the value of a relationship). It is difficult to imagine how these separate theories can all be integrated scientifically and how these processes can be combined in the animal's mind. To solve this problem, we first surveyed the empirical patterns and then we developed an individual-based model (called GrooFiWorld) in which individuals group, compete and groom. The grooming rule is based on grooming out of fear of defeat and on the anxiety reducing effects of grooming. We show that in this context this rule alone can explain many of the patterns of affiliation as well as the differences between egalitarian and despotic species. Our model can be used as a null model to increase our understanding of affiliative patterns of primates.

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          Most cited references26

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          Modulation of aggressive behaviour by fighting experience: mechanisms and contest outcomes.

          Experience in aggressive contests often affects behaviour during, and the outcome of, later contests. This review discusses evidence for, variations in, and consequences of such effects. Generally, prior winning experiences increase, and prior losing experiences decrease, the probability of winning in later contests, reflecting modifications of expected fighting ability. We examine differences in the methodologies used to study experience effects, and the relative importance and persistence of winning and losing experiences within and across taxa. We review the voluminous, but somewhat disconnected, literature on the neuroendocrine mechanisms that mediate experience effects. Most studies focus on only one of a number of possible mechanisms without providing a comprehensive view of how these mechanisms are integrated into overt behaviour. More carefully controlled work on the mechanisms underlying experience effects is needed before firm conclusions can be drawn. Behavioural changes during contests that relate to prior experience fall into two general categories. Losing experiences decrease willingness to engage in a contest while winning experiences increase willingness to escalate a contest. As expected from the sequential assessment model of contest behaviour, experiences become less important to outcomes of contests that escalate to physical fighting.A limited number of studies indicate that integration of multiple experiences can influence current contest behaviour. Details of multiple experience integration for any species are virtually unknown. We propose a simple additive model for this integration of multiple experiences into an individual's expected fighting ability. The model accounts for different magnitudes of experience effects and the possible decline in experience effects over time. Predicting contest outcomes based on prior experiences requires an algorithm that translates experience differences into contest outcomes. We propose two general types of model, one based solely on individual differences in integrated multiple experiences and the other based on the probability contests reach the escalated phase. The difference models include four algorithms reflecting possible decision rules that convert the perceived fighting abilities of two rivals into their probabilities of winning. The second type of algorithm focuses on how experience influences the probability that a subsequent contest will escalate and the fact that escalated contests may not be influenced by prior experience. Neither type of algorithm has been systematically investigated.Finally, we review models for the formation of dominance hierarchies that assume that prior experience influences contest outcome. Numerous models have reached varied conclusions depending on which factors examined in this review are included. We know relatively little about the importance of and variation in experience effects in nature and how they influence the dynamics of aggressive interactions in social groups and random assemblages of individuals. Researchers should be very active in this area in the next decade. The role of experience must be integrated with other influences on contest outcome, such as prior residency, to arrive at a more complete picture of variations in contest outcomes. We expect that this integrated view will be important in understanding other types of interactions between individuals, such as mating and predator-prey interactions, that also are affected significantly by prior experiences.
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            The integration of dominance and social bonding in primates.

            Social dominance is usually viewed from the perspective of intragroup competition over access to limited resources. The present paper, while not denying the importance of such competition, discusses the dominance concept among monkeys and apes in the context of affiliative bonding, social tolerance, and the reconciliation of aggressive conflicts. Two basic proximate mechanisms are supposed to provide a link between dominance and interindividual affiliation, namely, formalization of the dominance relationship (i.e., unequivocal communication of status), and conditional reassurance (i.e., the linkage of friendly coexistence to formalization of the relationship). Ritualized submission is imposed upon losers of dominance struggles by winners; losers are offered a "choice" between continued hostility or a tolerant relationship with a clearly signalled difference in status. If these two social mechanisms are lacking, aggression is bound to have dispersive effects. In their presence, aggression becomes a well-integrated, even constructive component of social life. In some higher primates this process of integration has reached the stage where status differences are strongly attenuated. In these species, sharing and trading can take the place of overt competition. The views underlying this "reconciled hierarchy" model are only partly new, as is evident from a review of the ethological literature. Many points are illustrated with data on a large semi-captive colony of chimpanzees (Pan troglodytes), particularly data related to striving for status, reconciliation behavior, and general association patterns. These observations demonstrate that relationships among adult male chimpanzees cannot be described in terms of a dichotomy between affiliative and antagonistic tendencies. Male bonding in this species has not been achieved by an elimination of aggression, but by a set of powerful buffering mechanisms that mitigate its effects. Although female chimpanzees do exhibit a potential for bonding under noncompetitive conditions, they appear to lack the buffering mechanisms of the males.
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              The winner and loser effect: integrating multiple experiences.

              An important question in state-dependent behaviour is how multiple influences on state are integrated to determine current behaviour. Aggressive behaviour is known to be affected by a prior contest experience. Nevertheless, whether and how multiple prior fighting experiences are integrated into a fighting decision remain unexplored. In this study, individuals of Rivulus marmoratus (Cyprinodontidae), a hermaphroditic fish, were given different combinations of two prior fighting experiences to investigate: (1) the effect of penultimate experiences on the probability of winning a subsequent contest; (2) the relative effect of a recent win and loss; and (3) whether the effect of a winning experience was as short lived as observed in other species. Penultimate and recent fighting experiences were given to the test fish approximately 48 and 24 h prior to the dyadic contests, respectively. From the results of the five types of contests staged, we conclude that: (1) penultimate fighting experiences had a significant effect on the probability of winning a subsequent contest; (2) a more recent experience had a more pronounced effect than an earlier experience, which suggested that the effect of a fighting experience would decay and/or the effect of a recent experience would interfere with the effect of an earlier experience; (3) no asymmetric effect between a winning experience and a losing experience was detected; and (4) the effect of both a winning and a losing experience lasted for at least 48 h in R. marmoratus which was the maximum time tested in these experiments. The possible reasons for the differences in results among studies of experience effects on contest outcomes are discussed. Copyright 1999 The Association for the Study of Animal Behaviour.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS Comput Biol
                plos
                ploscomp
                PLoS Computational Biology
                Public Library of Science (San Francisco, USA )
                1553-734X
                1553-7358
                December 2009
                December 2009
                24 December 2009
                : 5
                : 12
                : e1000630
                Affiliations
                [1]Theoretical Biology, Centre for Ecological and Evolutionary Studies, University of Groningen, Groningen, The Netherlands
                University College London, United Kingdom
                Author notes

                Conceived and designed the experiments: IPG CKH. Performed the experiments: IPG. Analyzed the data: IPG CKH. Wrote the paper: IPG CKH. Reimplementation of the model in C++: HH.

                Article
                09-PLCB-RA-0591R3
                10.1371/journal.pcbi.1000630
                2792710
                20041175
                6191161d-7bea-43c2-ad08-e5f8386beb78
                Puga-Gonzalez et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
                History
                : 26 May 2009
                : 30 November 2009
                Page count
                Pages: 17
                Categories
                Research Article
                Computational Biology
                Computational Biology/Systems Biology
                Ecology/Behavioral Ecology
                Neuroscience/Animal Cognition
                Neuroscience/Behavioral Neuroscience
                Neuroscience/Cognitive Neuroscience

                Quantitative & Systems biology
                Quantitative & Systems biology

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