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      Group Cooperation without Group Selection: Modest Punishment Can Recruit Much Cooperation

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          Abstract

          Humans everywhere cooperate in groups to achieve benefits not attainable by individuals. Individual effort is often not automatically tied to a proportionate share of group benefits. This decoupling allows for free-riding, a strategy that (absent countermeasures) outcompetes cooperation. Empirically and formally, punishment potentially solves the evolutionary puzzle of group cooperation. Nevertheless, standard analyses appear to show that punishment alone is insufficient, because second-order free riders (those who cooperate but do not punish) can be shown to outcompete punishers. Consequently, many have concluded that other processes, such as cultural or genetic group selection, are required. Here, we present a series of agent-based simulations that show that group cooperation sustained by punishment easily evolves by individual selection when you introduce into standard models more biologically plausible assumptions about the social ecology and psychology of ancestral humans. We relax three unrealistic assumptions of past models. First, past models assume all punishers must punish every act of free riding in their group. We instead allow punishment to be probabilistic, meaning punishers can evolve to only punish some free riders some of the time. This drastically lowers the cost of punishment as group size increases. Second, most models unrealistically do not allow punishment to recruit labor; punishment merely reduces the punished agent’s fitness. We instead realistically allow punished free riders to cooperate in the future to avoid punishment. Third, past models usually restrict agents to interact in a single group their entire lives. We instead introduce realistic social ecologies in which agents participate in multiple, partially overlapping groups. Because of this, punitive tendencies are more expressed and therefore more exposed to natural selection. These three moves toward greater model realism reveal that punishment and cooperation easily evolve by direct selection—even in sizeable groups.

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          Via freedom to coercion: the emergence of costly punishment.

          In human societies, cooperative behavior in joint enterprises is often enforced through institutions that impose sanctions on defectors. Many experiments on so-called public goods games have shown that in the absence of such institutions, individuals are willing to punish defectors, even at a cost to themselves. Theoretical models confirm that social norms prescribing the punishment of uncooperative behavior are stable-once established, they prevent dissident minorities from spreading. But how can such costly punishing behavior gain a foothold in the population? A surprisingly simple model shows that if individuals have the option to stand aside and abstain from the joint endeavor, this paves the way for the emergence and establishment of cooperative behavior based on the punishment of defectors. Paradoxically, the freedom to withdraw from the common enterprise leads to enforcement of social norms. Joint enterprises that are compulsory rather than voluntary are less likely to lead to cooperation.
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            Strong reciprocity and human sociality.

            H Gintis (2000)
            Human groups maintain a high level of sociality despite a low level of relatedness among group members. This paper reviews the evidence for an empirically identifiable form of prosocial behavior in humans, which we call "strong reciprocity", that may in part explain human sociality. A strong reciprocator is predisposed to cooperate with others and punish non-cooperators, even when this behavior cannot be justified in terms of extended kinship or reciprocal altruism. We present a simple model, stylized but plausible, of the evolutionary emergence of strong reciprocity. Copyright 2000 Academic Press.
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              Formidability and the logic of human anger.

              Eleven predictions derived from the recalibrational theory of anger were tested. This theory proposes that anger is produced by a neurocognitive program engineered by natural selection to use bargaining tactics to resolve conflicts of interest in favor of the angry individual. The program is designed to orchestrate two interpersonal negotiating tactics (conditionally inflicting costs or conditionally withholding benefits) to incentivize the target of the anger to place greater weight on the welfare of the angry individual. Individuals with enhanced abilities to inflict costs (e.g., stronger individuals) or to confer benefits (e.g., attractive individuals) have a better bargaining position in conflicts; hence, it was predicted that such individuals will be more prone to anger, prevail more in conflicts of interest, and consider themselves entitled to better treatment. These predictions were confirmed. Consistent with an evolutionary analysis, the effect of strength on anger was greater for men and the effect of attractiveness on anger was greater for women. Also as predicted, stronger men had a greater history of fighting than weaker men, and more strongly endorsed the efficacy of force to resolve conflicts--both in interpersonal and international conflicts. The fact that stronger men favored greater use of military force in international conflicts provides evidence that the internal logic of the anger program reflects the ancestral payoffs characteristic of a small-scale social world rather than rational assessments of modern payoffs in large populations.
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                Author and article information

                Contributors
                Role: Academic Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                20 April 2015
                2015
                : 10
                : 4
                : e0124561
                Affiliations
                [1 ]Department of Psychology, Harvard University, Cambridge, Massachusetts, United States of America
                [2 ]Department of Political Science, College of Business, Center for Behavioral Political Economy, Stony Brook University, Stony Brook, New York, United States of America
                [3 ]Center for Evolutionary Psychology, University of California Santa Barbara, Santa Barbara, California, United States of America
                [4 ]Department of Psychological and Brain Sciences, University of California Santa Barbara, Santa Barbara, California, United States of America
                [5 ]Department of Anthropology, University of California Santa Barbara, Santa Barbara, California, United States of America
                Universidad Carlos III de Madrid, SPAIN
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MMK AWD LC JT. Performed the experiments: MMK AWD. Analyzed the data: MMK AWD. Wrote the paper: MMK AWD LC JWD.

                Article
                PONE-D-14-47166
                10.1371/journal.pone.0124561
                4404356
                25893241
                d6ae05ac-3a8a-4ac1-bba9-fc361b890605
                Copyright @ 2015

                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
                : 20 October 2014
                : 3 March 2015
                Page count
                Figures: 3, Tables: 2, Pages: 17
                Funding
                This research was made possible by a National Institutes of Health Director’s Pioneer Award to LC ( http://www.nih.gov/), National Science Foundation grant #0951597 to LC and JT ( http://www.nsf.gov/), and John Templeton Foundation grant #29468 ( http://www.templeton.org/). The opinions expressed in this publication are those of the authors and do not necessarily reflect the views of the John Templeton Foundation. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Custom metadata
                All relevant data are within the paper and its Supporting Information files.

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