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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">plos</journal-id>
<journal-id journal-id-type="nlm-ta">PLoS Comput Biol</journal-id>
<journal-id journal-id-type="pmc">ploscomp</journal-id><journal-title-group>
<journal-title>PLoS Computational Biology</journal-title></journal-title-group>
<issn pub-type="ppub">1553-734X</issn>
<issn pub-type="epub">1553-7358</issn>
<publisher>
<publisher-name>Public Library of Science</publisher-name>
<publisher-loc>San Francisco, USA</publisher-loc></publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">PCOMPBIOL-D-14-00159</article-id>
<article-id pub-id-type="doi">10.1371/journal.pcbi.1003618</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Biology and life sciences</subject><subj-group><subject>Computational biology</subject><subj-group><subject>Evolutionary modeling</subject></subj-group></subj-group><subj-group><subject>Evolutionary biology</subject><subj-group><subject>Evolutionary theory</subject></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Social sciences</subject><subj-group><subject>Sociology</subject><subj-group><subject>Computational sociology</subject><subject>Social research</subject><subject>Social systems</subject></subj-group></subj-group></subj-group></article-categories>
<title-group>
<article-title>Barriers to Cooperation Aid Ideological Rigidity and Threaten Societal Collapse</article-title>
<alt-title alt-title-type="running-head">Cooperation, Ideological Rigidity, and the Societal Collapse</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Jusup</surname><given-names>Marko</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="corresp" rid="cor1"><sup>*</sup></xref></contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Matsuo</surname><given-names>Tadasu</given-names></name><xref ref-type="aff" rid="aff2"><sup>2</sup></xref></contrib>
<contrib contrib-type="author" xlink:type="simple"><name name-style="western"><surname>Iwasa</surname><given-names>Yoh</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref></contrib>
</contrib-group>
<aff id="aff1"><label>1</label><addr-line>Department of Biology, Kyushu University, Fukuoka, Japan</addr-line></aff>
<aff id="aff2"><label>2</label><addr-line>Faculty of Economics, Ritsumeikan University, Kusatsu, Japan</addr-line></aff>
<contrib-group>
<contrib contrib-type="editor" xlink:type="simple"><name name-style="western"><surname>Bonhoeffer</surname><given-names>Sebastian</given-names></name>
<role>Editor</role>
<xref ref-type="aff" rid="edit1"/></contrib>
</contrib-group>
<aff id="edit1"><addr-line>ETH Zürich, Switzerland</addr-line></aff>
<author-notes>
<corresp id="cor1">* E-mail: <email xlink:type="simple">mjusup@gmail.com</email></corresp>
<fn fn-type="conflict"><p>The authors have declared that no competing interests exist.</p></fn>
<fn fn-type="con"><p>Wrote the paper: MJ YI. Developed the basic model: TM YI. Generalized the model: MJ. Ran numerical simulations: MJ. Discussed the results and implications: MJ TM YI.</p></fn>
</author-notes>
<pub-date pub-type="collection"><month>5</month><year>2014</year></pub-date>
<pub-date pub-type="epub"><day>8</day><month>5</month><year>2014</year></pub-date>
<volume>10</volume>
<issue>5</issue>
<elocation-id>e1003618</elocation-id>
<history>
<date date-type="received"><day>26</day><month>1</month><year>2014</year></date>
<date date-type="accepted"><day>28</day><month>3</month><year>2014</year></date>
</history>
<permissions>
<copyright-year>2014</copyright-year>
<copyright-holder>Jusup et al</copyright-holder><license xlink:type="simple"><license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p></license></permissions>
<abstract>
<p>Understanding the factors that promote, disrupt, or shape the nature of cooperation is one of the main tasks of evolutionary biology. Here, we focus on attitudes and beliefs supportive of in-group favoritism and strict adherence to moral consensus, collectively known as ideological rigidity, that have been linked with both ends of the political spectrum. The presence among the political right and the left is likely to make ideological rigidity a major determinant of the political discourse with an important social function. To better understand this function, we equip the indirect reciprocity framework – widely used to explain evaluation-mediated social cooperation – with multiple stylized value systems, each corresponding to the different degree of ideological rigidity. By running game theoretical simulations, we observe the competitive evolution of these systems, map conditions that lead to more ideologically rigid societies, and identify potentially disastrous outcomes. In particular, we uncover that barriers to cooperation aid ideological rigidity. The society may even polarize to the extent where social parasites overrun the population and cause the complete collapse of the social structure. These results have implications for lawmakers globally, warning against restrictive or protectionist policies.</p>
</abstract>
<abstract abstract-type="summary"><title>Author Summary</title>
<p>Attitudes, beliefs, and resulting value systems may represent important motivational and decision-making factors that have strong impact on cooperation in a society. Accordingly, understanding the social function of value systems is a topic of great interest in evolutionary biology, but one where progress is made difficult by the sheer complexity of values-inspired behaviors. Here, we argue that considerable theoretical progress can be made within the indirect reciprocity framework. We show in the context of indirect reciprocity how to construct stylized value systems from a mathematically formalized notion of ideological rigidity. Our simulations indicate that politically imposed restrictions and protectionism favor the evolution of ideologically rigid value systems. The complete collapse of cooperation also arises as a possible evolutionary outcome.</p>
</abstract>
<funding-group><funding-statement>This study was partly supported by the Global Center Of Excellence (GCOE) Program “Asian Conservation Ecology”, Environmental Agency Fund (S9), and Grant-in-Aid for Basic Research (B) to YI. Japan Society for the Promotion of Science (JSPS) Postdoctoral Fellowship Program for Foreign Researchers (P13380) and the accompanying Grant-in-Aid for Scientific Research allowed MJ to conduct research in Japan. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.</funding-statement></funding-group><counts><page-count count="8"/></counts></article-meta>
</front>
<body><sec id="s1">
<title>Introduction</title>
<p>Factors affecting cooperation in a society, such as attitudes, beliefs, and resulting value systems, are a subject of major interest in evolutionary biology. Some examples of considerable importance are in-group favoritism, nationalism, ethnocentrism, intolerance for dissent, submission to strong leadership, and support for tight control all of which were originally linked with the political right <xref ref-type="bibr" rid="pcbi.1003618-Jost1">[1]</xref>. However, evidence was presented to dispel such a link and argue that the same set of attitudes, by serving both the right and the left, is indicative of ideological rigidity rather than a position on the political spectrum <xref ref-type="bibr" rid="pcbi.1003618-Greenberg1">[2]</xref>. The alleged presence on both ends of the spectrum is likely to make ideological rigidity a potent force in directing the political discourse and ultimately shaping societies. Here, we set to investigate the social function of ideological rigidity, starting from a motivational premise that indirect reciprocity – a cooperation maintaining mechanism based on the evaluation of the reputation <xref ref-type="bibr" rid="pcbi.1003618-Nowak1">[3]</xref> – provides a proper framework for our investigation. To establish this premise, we emphasize the dual nature of the aforementioned attitudes and beliefs. On the one hand, ideologically rigid believe in the supremacy of one's group or, at least, distrust anyone who is not a member of this group. Such a belief, broadly termed in-group favoritism, represents an attractive phenomenon for the studies on indirect reciprocity <xref ref-type="bibr" rid="pcbi.1003618-Masuda1">[4]</xref>–<xref ref-type="bibr" rid="pcbi.1003618-Matsuo1">[6]</xref>. On the other hand, ideological rigidity is linked with attitudes that demand strict adherence to moral consensus. Corresponding ideas are again found in the indirect reciprocity framework, where social norms subjected to the evolutionary competition <xref ref-type="bibr" rid="pcbi.1003618-Pacheco1">[7]</xref>, <xref ref-type="bibr" rid="pcbi.1003618-Uchida1">[8]</xref> handle dissent from moral consensus in different ways. Our aim is to unify these ideas by incorporating the dual nature of ideological rigidity into stylized value systems and then examine the consequent evolutionary dynamics.</p>
<p>As the first step forward, we formalize the notion of ideological rigidity within the indirect reciprocity framework. In an indirect reciprocity game, members of a society, or players, encounter each other randomly, whereupon one player takes the role of a donor, while the other acts as a recipient. The donor can choose between two actions contingent on the recipient's reputation. By cooperating with the recipient, the donor incurs a cost <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e001" xlink:type="simple"/></inline-formula>, but the recipient benefits from a payoff <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e002" xlink:type="simple"/></inline-formula> for a net gain of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e003" xlink:type="simple"/></inline-formula> for the society. By defecting, the donor avoids the cost <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e004" xlink:type="simple"/></inline-formula>, the recipient gains nothing, and the society is exactly where it was before the encounter. Every action is scrutinized by observers who assign the donor an appropriate reputation for the next round of the game. Maintaining a favorable reputation improves the prospects of receiving the payoff <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e005" xlink:type="simple"/></inline-formula> afterwards, thus justifying the willingness to incur the cost <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e006" xlink:type="simple"/></inline-formula> in the first place. The payoff <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e007" xlink:type="simple"/></inline-formula> is potentially received in the next round of the game from a third player – hence the name indirect reciprocity – who then serves as a donor, while the current donor takes the role of a recipient. The detailed rules governing which action should be taken and how the reputation should be assessed are called action-assessment strategies and represent a stylized version of the donor's value system. Action-assessment strategies are a part of the central process called the reputation dynamics (see <xref ref-type="sec" rid="s3">Methods</xref>). For now, it is critical that the reputation dynamics can incorporate several action-assessment strategies, allowing us to distinguish between player types and place them appropriately on a scale of ideological rigidity.</p>
<p>The first key aspect of any action-assessment strategy, as the name suggests, is the action rule. We focus on the situation in which all players are discriminators, meaning that donors cooperate only with recipients who have a favorable reputation <xref ref-type="bibr" rid="pcbi.1003618-Nowak2">[9]</xref>. Because the action rule is the same for all players, making a distinction between player types requires other key aspects of action-assessment strategies to be more elaborate. One such aspect is the reputation assessment rule. Assessment rules are theoretical representations of social norms that govern the decision-making process of observers while assigning the reputation to donors for the next round of the game. We assume that information spreads from observers to other players rapidly (e.g. through gossip). Two assessment rules are considered. The first of the two rules is called simple-standing or the Sugden rule <xref ref-type="bibr" rid="pcbi.1003618-Leimar1">[10]</xref>–<xref ref-type="bibr" rid="pcbi.1003618-Ohtsuki1">[12]</xref>. It stipulates that a favorable reputation is assigned to a donor who cooperates with a recipient of favorable reputation or defects from a recipient of unfavorable reputation. An unfavorable reputation is assigned to a donor who defects from a recipient of favorable reputation. Importantly, a favorable reputation is assigned to a donor who cooperates with a recipient of unfavorable reputation, indicating that the Sugden rule liberally follows moral consensus. By contrast, the second of the two rules, called stern-judging or the Kandori rule <xref ref-type="bibr" rid="pcbi.1003618-Ohtsuki1">[12]</xref>, <xref ref-type="bibr" rid="pcbi.1003618-Kandori1">[13]</xref> strictly enforces moral consensus. Cooperation with a recipient of unfavorable reputation leads to an unfavorable reputation assignment for the donor. For an easy comparison, both assessment rules are summarized in <xref ref-type="table" rid="pcbi-1003618-t001">Table 1</xref>. These concepts are defined in a strict mathematical manner in the section on the reputation dynamics (see <xref ref-type="sec" rid="s3">Methods</xref>).</p>
<table-wrap id="pcbi-1003618-t001" position="float"><object-id pub-id-type="doi">10.1371/journal.pcbi.1003618.t001</object-id><label>Table 1</label><caption>
<title>The assessment rule.</title>
</caption><alternatives><graphic id="pcbi-1003618-t001-1" position="float" mimetype="image" xlink:href="info:doi/10.1371/journal.pcbi.1003618.t001" xlink:type="simple"/>
<table><colgroup span="1"><col align="left" span="1"/><col align="center" span="1"/><col align="center" span="1"/><col align="center" span="1"/><col align="center" span="1"/></colgroup>
<thead>
<tr>
<td align="left" rowspan="1" colspan="1">Observer's type</td>
<td colspan="2" align="left" rowspan="1">Sugden</td>
<td colspan="2" align="left" rowspan="1">Kandori</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Recipient's reputation</td>
<td align="left" rowspan="1" colspan="1">Favorable</td>
<td align="left" rowspan="1" colspan="1">Unfavorable</td>
<td align="left" rowspan="1" colspan="1">Favorable</td>
<td align="left" rowspan="1" colspan="1">Unfavorable</td>
</tr>
</thead>
<tbody>
<tr>
<td align="left" rowspan="1" colspan="1">Cooperation</td>
<td align="left" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">0</td>
</tr>
<tr>
<td align="left" rowspan="1" colspan="1">Defection</td>
<td align="left" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">0</td>
<td align="left" rowspan="1" colspan="1">1</td>
</tr>
</tbody>
</table>
</alternatives><table-wrap-foot><fn id="nt101"><label/><p>Values represent the probabilities that the donor is evaluated favorably conditional on all relevant circumstances (the observer's type, the recipient's reputation, and the donor's action).</p></fn></table-wrap-foot></table-wrap>
<p>Before introducing another key aspect of an action-assessment strategy, we make the assumption that the society consists of two separate parts. Namely, an inner circle (e.g. a nation state) is embedded into a much larger outer world (e.g. the international community), where the cooperation between the two parts of the society, though allowed, is made difficult (e.g. by the national border control). Members of the inner circle (i.e. insiders) thus have a high probability, denoted <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e008" xlink:type="simple"/></inline-formula>, of encountering other insiders, but only a small probability, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e009" xlink:type="simple"/></inline-formula>, of meeting a player from the outside world (i.e. an outsider). For an insider, cooperation with the outside world also carries an additional cost, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e010" xlink:type="simple"/></inline-formula> (e.g. a tariff). Herein, we are primarily interested in the evolutionary dynamics (see <xref ref-type="sec" rid="s3">Methods</xref>) of value systems inside the inner circle and the subsequent implications for ideological rigidity of insiders.</p>
<p>The division of the society into two separate parts leads us naturally to another key aspect of an action-assessment strategy. Within the inner circle, because encounters with the outside world are rare, it is fairly reasonable to presume that distrust towards outsiders can take root among a fraction of the insiders. These insiders exhibit strong in-group favoritism in the sense that all cooperation with outsiders is suspended and no benefits from the outside world are accepted. The remaining insiders, by contrast, reject in-group favoritism, maintain cooperation with the outside world, and receive the accompanying benefits. The setting we describe here is not without a historical precedent. A resemblance can be found in pre-modern Japan <xref ref-type="bibr" rid="pcbi.1003618-Matsuo1">[6]</xref>, where the two dominant value systems, one called bushido (the way of warriors) and the other called shonindo (the way of merchants), held opposing positions on in-group favoritism. In Western culture, many parallels can be drawn by examining the differences between the Maghribi and the Genoese <xref ref-type="bibr" rid="pcbi.1003618-Greif1">[14]</xref>. However, being primarily motivated by the bushido-shonindo dichotomy, we name the fraction of the insiders that embrace in-group favoritism “bushi”. The remaining insiders that reject in-group favoritism are named “shonin”.</p>
<p>The two introduced aspects of action-assessment strategies (Sugden vs. Kandori and shonin vs. bushi) allow us to distinguish four types of insiders. Sugden-shonin (hereafter Ss) liberally follow moral consensus, reject in-group favoritism, and hence are considered ideologically non-rigid. A step up on the scale of ideological rigidity are Kandori-shonin (Ks), who strictly enforce moral consensus, yet reject in-group favoritism. Sugden-bushi (Sb), while liberal towards moral consensus, endorse in-group favoritism. We consider the stance of Sb players to be more ideologically rigid than the stance of Ks players because in-group favoritism as defined herein limits the scope of cooperation far more strongly than the strict enforcement of moral consensus. The most ideologically rigid are Kandori-bushi (Kb), who enforce moral consensus and embrace in-group favoritism. Besides these four types of players we entertain the notion of social parasites in the form of unconditional defectors (often denoted AllD in the literature, hereafter simply Ad). For the outside world, which is much bigger than the inner circle, interactions with insiders are inconsequential. Modeling the evolutionary dynamics (see <xref ref-type="sec" rid="s3">Methods</xref>) of value systems in the outside world is possible using the same mathematical framework as for the inner circle, but with the probability of an outsider meeting another outsider set to unity. The inner circle is, therefore, a set of measure zero. Because we are interested in the evolutionary dynamics of value systems in the inner circle, the outside world is assumed to be in a stable equilibrium populated only by Ss or Ks players. Such a simple structure of the outside world can be justified by the fact that any other more complex structure would only diminish the benefits from cross-border encounters which is qualitatively captured by increasing the value of the parameter <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e011" xlink:type="simple"/></inline-formula>.</p>
</sec><sec id="s2">
<title>Results and Discussion</title>
<p>We explore the dual nature of ideological rigidity and its social function by means of indirect reciprocity games. Differences in adherence to moral consensus are reflected in the performance of the more liberal Sugden against the stricter Kandori rule (S and K in shorthand notation, respectively). Similarly, opposite attitudes towards in-group favoritism are reflected in the performance of open-minded shonin against distrustful bushi players (s and b, respectively). The focus is placed on the most illustrative cases, meaning a relatively closed inner circle <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e012" xlink:type="simple"/></inline-formula> in which the set of possible player types is either <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e013" xlink:type="simple"/></inline-formula> or <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e014" xlink:type="simple"/></inline-formula>. Though the model can handle any number of player types, having three types per simulation permits effective visualization and comparison of the results. We start with a technical description of two opposing situations, one where barriers to cooperation are low and the other where barriers are high (<xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1</xref>). To achieve this, we set both cost-benefit ratios, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e015" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e016" xlink:type="simple"/></inline-formula>, close to zero and subsequently increase either one towards unity. The results are then generalized by continuous mapping of the parameter space (<xref ref-type="fig" rid="pcbi-1003618-g002">Fig. 2</xref>) and finally by inclusion of social parasites (Ad) into simulations (<xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3</xref>).</p>
<fig id="pcbi-1003618-g001" position="float"><object-id pub-id-type="doi">10.1371/journal.pcbi.1003618.g001</object-id><label>Figure 1</label><caption>
<title>Performance of action-assessment strategies.</title>
<p>(a) Low barriers to cooperation make the Ss strategy successful against Kb and Sb strategies. (b) Under the same conditions as in (a), the Ks strategy performs worse than the Ss strategy. (c) High barriers to cooperation are detrimental for the Ss strategy. (d) Under the same conditions as in (c) the Ks strategy is successful against Kb and Sb strategies. (e) The success of the Ss strategy in (a) diminishes when the cost of cross-border cooperation is high. (f) Under the same conditions as in (e) the Ks strategy is affected less than the Ss strategy.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pcbi.1003618.g001" position="float" xlink:type="simple"/></fig><fig id="pcbi-1003618-g002" position="float"><object-id pub-id-type="doi">10.1371/journal.pcbi.1003618.g002</object-id><label>Figure 2</label><caption>
<title>A continuous map of the parameter space.</title>
<p>Relative performance of Ss and Ks strategies, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e017" xlink:type="simple"/></inline-formula>, is shown for all reasonable cost-benefit ratios. The black curve approximates the border at which both strategies are equally effective, i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e018" xlink:type="simple"/></inline-formula>.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pcbi.1003618.g002" position="float" xlink:type="simple"/></fig><fig id="pcbi-1003618-g003" position="float"><object-id pub-id-type="doi">10.1371/journal.pcbi.1003618.g003</object-id><label>Figure 3</label><caption>
<title>Ideological rigidity and the collapse of social structure.</title>
<p>(a) Social parasites cannot invade the inner circle dominated by any combination of ideologically non-rigid Ss and rigid Kb players. (b) Replacing ideologically non-rigid Ss with more rigid Ks players may polarize the inner circle to the extent where social parasites uncontrollably spread out and eventually suppress cooperation.</p>
</caption><graphic mimetype="image" xlink:href="info:doi/10.1371/journal.pcbi.1003618.g003" position="float" xlink:type="simple"/></fig>
<p>Low barriers to cooperation favor ideologically non-rigid Ss strategy (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1a, b</xref>). From ternary plots it is apparent that the vertices Ss and Ks share the property of being locally stable monomorphic attractors. By contrast, neither Kb nor Sb vertices have this property, but rather the whole segment connecting them is a locally stable dimorphic attractor (the Kb-Sb attractor). Comparing the sizes of the corresponding domains of attraction reveals the evolutionarily advantageous action-assessment strategy. When cost-benefit ratios are close to zero (<xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1a</xref>), the Ss attractor not only overshadows the Kb-Sb attractor in terms of the size of the domain of attraction (73% vs. 27%), but a rare occurrence of Ss players in an inner circle dominated by Sb players leads to a successful invasion. Under the same conditions (<xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1b</xref>), the Ks attractor fares less well, commanding a smaller domain of attraction than the Kb-Sb attractor (46% vs. 54%) and failing to successfully invade the inner circle dominated by any combination of Kb and Sb players.</p>
<p>Increasing barriers to cooperation make the ideologically non-rigid Ss strategy evolutionarily disadvantageous (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1c, d</xref>). For a society to maintain feasible cross-border interactions, ideological non-rigidity needs to be abandoned in favor of a more ideologically rigid Ks strategy. Namely, when the cost-benefit ratios <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e019" xlink:type="simple"/></inline-formula> is set close to unity, the domain of attraction of the Ss attractor (<xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1c</xref>) is greatly reduced in favor of the Kb-Sb attractor (&lt;1% vs. &gt;99%) despite Ss players still being able to invade an Sb-dominated inner circle. By contrast, the Ks attractor fares much better than originally (<xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1d</xref>). Its domain of attraction is now larger than that of the alternative (locally stable monomorphic) Kb attractor (62% vs. 38%) and a rare occurrence of Ks players in an Sb-dominated inner circle leads to a successful invasion. It is worth emphasizing that vulnerability to invasion by both Ss and Ks strategies makes the Sb strategy a weak candidate for the ideologically rigid.</p>
<p>The increasing cost of cross-border interactions aids ideological rigidity (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1e, f</xref>). The effect is twofold because the Sb strategy turns evolutionarily viable and the Ks strategy gains an (albeit marginal) evolutionary advantage over the Ss strategy. When the ratio <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e020" xlink:type="simple"/></inline-formula> is set close to unity and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e021" xlink:type="simple"/></inline-formula> is kept near zero, the benefit of encountering outsiders is reduced and, therefore, Ss and Ks strategies are negatively impacted. Accordingly, the inner circle dominated by Sb players can no longer be invaded by either Ss or Ks players. The domains of attraction of Ss and Ks attractors become smaller than that of the Kb-Sb attractor (36% vs. 64% and 38% vs. 62%, respectively). A notable difference between Ss and Ks strategies, however, is that the negative impact of high <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e022" xlink:type="simple"/></inline-formula> is worse for the former (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1a, e</xref>) than the latter (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1b, f</xref>) strategy. When <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e023" xlink:type="simple"/></inline-formula> is close to unity the domain of attraction of the Ss attractor is reduced to half its original size (from 73% to 36%), whereas the reduction for the Ks attractor is minimal (from 46% to 38%).</p>
<p>The three extreme cases examined heretofore are illustrative, but not exhaustive because any combination of cost-benefit ratios that satisfies <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e024" xlink:type="simple"/></inline-formula> is reasonable. Consequently, we map the parameter space in a continuous manner, focusing in particular on the relative performance of Ss and Ks strategies. We emphasize the relative performance because the evolutionary advantage of one strategy over the other changes with the location in the parameter space. By comparison, vulnerability to invasion suggests that for the ideologically more rigid the Sb strategy is a weak alternative. To summarize the relative performance of the two strategies over a wide range of cost-benefit ratios, we denote respectively by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e025" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e026" xlink:type="simple"/></inline-formula> areas of the domains of attraction corresponding to Ss and Ks attractors and introduce a performance indicator <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e027" xlink:type="simple"/></inline-formula>. Note that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e028" xlink:type="simple"/></inline-formula>, where positive (negative) values indicate the evolutionary advantage of Ss (Ks) players. The simulation results (<xref ref-type="fig" rid="pcbi-1003618-g002">Fig. 2</xref>) confirm the notion that Ss players have an evolutionary advantage when barriers to cooperation are low; that is, when the sum of the two cost-benefit ratios is close to zero. As barriers become higher, the Ks strategy turns out to be advantageous. Particularly detrimental for Ss players is the increase in the cost-benefit ratio <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e029" xlink:type="simple"/></inline-formula> because as it approaches unity the area <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e030" xlink:type="simple"/></inline-formula> tends to zero. Increasing <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e031" xlink:type="simple"/></inline-formula> affects both Ss and Ks players negatively, but the area <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e032" xlink:type="simple"/></inline-formula> is much more sensitive to the change in <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e033" xlink:type="simple"/></inline-formula> than <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e034" xlink:type="simple"/></inline-formula>.</p>
<p>Pursuing a technical description of the simulation results so far sheds new light on the evolution of value systems as implied by the indirect reciprocity framework, but remains silent on the underlying mechanisms. We are thus required to make an extra effort to access these mechanisms and in return gain an intuitive grasp of the mathematical formalism being applied. Starting from a comparison of bushi and shonin players, the former are at a fundamental disadvantage because of refusing to interact with the outsiders. Such a situation is exemplified by the success of Ss over Sb players in <xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1a</xref>. Bushi, in fact, may not represent an evolutionarily viable alternative at all without sufficiently closed borders (i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e035" xlink:type="simple"/></inline-formula> slightly below unity). If this criterion is met, the difference between Sugden and Kandori rules plays a major role. Players adopting the Sugden rule, due to their liberal stance, receive benefits and incur costs of cooperation more often than players adopting the Kandori rule – a clear advantage when <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e036" xlink:type="simple"/></inline-formula> is low. As <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e037" xlink:type="simple"/></inline-formula> approaches unity, the fortunes reverse. We can now understand why Sb players can resist Ks players in <xref ref-type="fig" rid="pcbi-1003618-g001">Fig. 1b</xref>, as well as the success of Kandori players in <xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1c and d</xref>. Protectionism (i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e038" xlink:type="simple"/></inline-formula> slightly below unity) naturally helps bushi agenda, which is best illustrated by how Ss players lose their advantage over Sb players as the cost of cross-border interactions increases (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1a, e</xref>). The same effect is visible by comparing <xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1b and f</xref>, although it is much weaker because the Sugden rule to a certain extent shields Sb against Kb players even when the cost of cross-border cooperation is low. The described mechanisms can be used to explain all intermediate outcomes in <xref ref-type="fig" rid="pcbi-1003618-g002">Fig. 2</xref>.</p>
<p>Having determined that the Kb strategy shields the ideologically rigid from invasion, while the ideologically less rigid should choose between Ss and Ks strategies contingent on how high barriers to cooperation are set, we consider the effect of social parasites on the society. Indirect reciprocity games with two sets of possible player types, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e039" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e040" xlink:type="simple"/></inline-formula>, reveal important qualitative differences (<xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3</xref>). With moderately low barriers to cooperation, the conflict between Ss and Kb strategies in the presence of social parasites results in three distinct domains of attraction accompanied with three locally stable monomorphic attractors, as well as three dimorphic and one trimorphic equilibria (<xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3a</xref>). Though the size of each domain of attraction is parameter-dependent, the remarkable outcome is that there are no openings for an invasion. By contrast, the conflict between Ks and Kb strategies in the presence of social parasites lacks a trimorphic equilibrium and leaves the dimorphic equilibrium of Ks and Kb players vulnerable to invasion in the case of a rare occurrence of Ad players (<xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3b</xref>). Therefore, by making the ideologically more rigid Ks strategy evolutionarily advantageous over the Ss strategy, rising barriers to cooperation not only aid ideological rigidity, they even threaten the collapse of the social structure.</p>
<p>Looking at the results in <xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3</xref>, what we truly observe are the negative consequences of the maxim “the enemy of my enemy is my friend” as well as the way to avoid these consequences. When the inner circle is populated with an ideologically more rigid combination of Ks and Kb (along with Ad) players, even if they initially treat each other favorably, after a while a Ks player will cooperate with an outsider and be assigned an unfavorable reputation by the Kb observer. Such a player is bound to be denied cooperation from a Kb donor, resulting in an unfavorable reputation assignment for this donor from the Ks observer. A rift between Ks and Kb players forms. Ad players may eventually take the advantage of such a rift (<xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3b</xref>) because when they defect from a Ks recipient they receive benefits from a Kb donor and vice versa. Note that with stern-judging the rift only widens after every interaction of either a Ks or a Kb donor with an Ad recipient. The reason is that the Ad recipient is treated favorably by one side and unfavorably by the other, resulting with certainty in an unfavorable reputation assignment for the donor. Replacing Ks players with ideologically non-rigid Ss players does not prevent the rift from opening. Ss players, however, mend the rift after a Kb donor cooperates with an Ad recipient by assigning a favorable reputation to this donor. Liberal attitude towards moral consensus thus makes it impossible for social parasites to invade the inner circle (<xref ref-type="fig" rid="pcbi-1003618-g003">Fig. 3a</xref>) and can be seen as a layer of stability for the social structure. To confirm the robustness of the described mechanisms, we performed simulations (results not shown) with two additional sets of possible player types: <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e041" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e042" xlink:type="simple"/></inline-formula>. It turned out that only the ideologically rigid combination of Ks and Kb players was vulnerable to an invasion by Ad, agreeing with the notion that liberal attitude towards moral consensus had a stabilizing effect on the society. In the context of the model robustness, we did not simulate unconditional cooperators nor the first-order scoring rule because it was shown that neither could maintain stable cooperation <xref ref-type="bibr" rid="pcbi.1003618-Uchida1">[8]</xref>; all else being equal, the former got eliminated in the presence of Sugden, Kandori, and Ad, whereas the latter, if not eliminated, became indistinguishable from Ad. The presence of unconditional cooperators, nonetheless, might have favored Kandori over Sugden to a certain extent because the Sugden rule would have encouraged more cooperativeness and hence higher costs in comparison with the Kandori rule <xref ref-type="bibr" rid="pcbi.1003618-Uchida1">[8]</xref>.</p>
<p>In the modern world, two omnipresent processes affecting barriers to cooperation are the technological development (lowering <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e043" xlink:type="simple"/></inline-formula>) and globalization (lowering <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e044" xlink:type="simple"/></inline-formula>). Because both of these processes make cooperation easier, our results imply (to the extent game theoretical representations are valid in a complex reality) that the modern world is conducive of ideologically non-rigid societies with presumably an increasing number of functioning democracies and more economic liberties <xref ref-type="bibr" rid="pcbi.1003618-Lawson1">[15]</xref>. Support can be found in indisputable growth of electoral democracy among the world's nations, especially over the past three decades, although the Democracy Index compiled by the Economist Intelligence Unit suggests that the overall quality of democracy is stagnating since the financial crisis of 2007–2008. As for economic liberties, the average Economic Freedom of the World Index reported by the Fraser Institute indicates steady increase from 1980 until 2006, but again a period of stagnation during the 2008–2012 global recession. Looking at our results from a different angle, a remarkable implication is that restrictive or protectionist policies aid the creation of ideologically rigid societies. Perhaps then it is not surprising that the above indices are stagnating in the midst of a five-years long recession. This is, after all, the second worst economic contraction since the Great Depression of the 1930s, which itself brought on a number of restrictive or protectionist policies, coinciding with the rise of multiple totalitarian regimes and ending only after the deadliest conflict in the human history.</p>
<p>Analyzing the social role of ideological rigidity within the indirect reciprocity framework, we uncovered evolutionary outcomes that warn against restrictive or protectionist government policies. Yet to prevent from falling into the trap of naive rationalism or worse interventionism, a constant remainder is needed that these outcomes follow from a mathematically tractable representation of immensely complex human concepts such as value systems. We, therefore, perceive the present and similar studies as theoretical constructs that identify the potential, rather than the actual, drivers of social phenomena. Keeping this important distinction in mind does not diminish the multitude of opportunities for the field. Our understanding of the factors that promote (e.g. punishment), disrupt (e.g. corruption) or shape the nature (e.g. spontaneous in-group favoritism) of cooperation is still quite limited.</p>
</sec><sec id="s3" sec-type="methods">
<title>Methods</title>
<sec id="s3a">
<title>Reputation dynamics</title>
<p>The reputation dynamics control intra-generational partitioning of players according to their reputation. Intuitively, the outcomes of the reputation dynamics specify probabilities that the generation of players of a particular type will be assigned a particular reputation. More formally, we are concerned with a discrete probability measure defined on a sample set, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e045" xlink:type="simple"/></inline-formula>, where the sample set is built from two basic constituents, the set of player types, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e046" xlink:type="simple"/></inline-formula>, and the set of all possible reputations, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e047" xlink:type="simple"/></inline-formula>. Because five distinct player types are considered, the set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e048" xlink:type="simple"/></inline-formula> could generally be any combination of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e049" xlink:type="simple"/></inline-formula>. It is beneficial, however, to display the results of extensive numerical simulations on ternary plots by referring only to the most illustrative outcomes. The main reason is that handling three player types at once permits us to effectively visualize and compare the results. The set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e050" xlink:type="simple"/></inline-formula> is accordingly limited to 3-combinations with repetitions (or 3-multisets) of the set <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e051" xlink:type="simple"/></inline-formula>, where F and U denote a favorable and an unfavorable reputation, respectively. Hence, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e052" xlink:type="simple"/></inline-formula>. With these basic constituents, the sample set is given by the Cartesian product of the form <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e053" xlink:type="simple"/></inline-formula>, so that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e054" xlink:type="simple"/></inline-formula> implies <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e055" xlink:type="simple"/></inline-formula>. The discrete probability measure of concern, denoted by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e056" xlink:type="simple"/></inline-formula> because it is closely related to the so-called honor score <xref ref-type="bibr" rid="pcbi.1003618-Ohtsuki2">[16]</xref>, is fully defined by specifying how it operates on the elements of <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e057" xlink:type="simple"/></inline-formula>. To emphasize the dependence on the type and reputation of players, we introduce a short-hand notation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e058" xlink:type="simple"/></inline-formula>. The letters <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e059" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e060" xlink:type="simple"/></inline-formula> are used throughout the text to denote player types, primarily that of a donor and a recipient, respectively. The letters <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e061" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e062" xlink:type="simple"/></inline-formula> are reserved for the type of observers and outsiders, respectively. It is also useful to reserve the letters <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e063" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e064" xlink:type="simple"/></inline-formula> for the reputation of donors and recipients, respectively. Note that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e065" xlink:type="simple"/></inline-formula> and, therefore, stand for three different reputations. When interested only in the reputation from the viewpoint of type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e066" xlink:type="simple"/></inline-formula> observers, we can write <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e067" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e068" xlink:type="simple"/></inline-formula>. In addition, to keep formulas for the probabilities <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e069" xlink:type="simple"/></inline-formula> more tractable, it is helpful to introduce two auxiliary functions as shown henceforth.</p>
<p>The first of the two auxiliary functions, denoted <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e070" xlink:type="simple"/></inline-formula>, is called the action rule. Because each interaction in the game involves two players, a donor and a recipient, the action rule specifies the probability of an action <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e071" xlink:type="simple"/></inline-formula> being undertaken by the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e072" xlink:type="simple"/></inline-formula> donor towards the recipient with the reputation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e073" xlink:type="simple"/></inline-formula> from the donor's viewpoint. Only two actions are possible, so that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e074" xlink:type="simple"/></inline-formula>, where C and D stand for cooperation and defection, respectively. It follows, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e075" xlink:type="simple"/></inline-formula>. For <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e076" xlink:type="simple"/></inline-formula> the action rule is<disp-formula id="pcbi.1003618.e077"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pcbi.1003618.e077" xlink:type="simple"/></disp-formula>and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e078" xlink:type="simple"/></inline-formula>. Ad players never cooperate, thus <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e079" xlink:type="simple"/></inline-formula>.</p>
<p>The second auxiliary function, denoted <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e080" xlink:type="simple"/></inline-formula>, is called the assessment rule. Because observers assign new reputations to donors after every interaction, the assessment rule represents the probability that the donor will be assigned the reputation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e081" xlink:type="simple"/></inline-formula> by the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e082" xlink:type="simple"/></inline-formula> observer if the recipient's reputation from the observer's viewpoint is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e083" xlink:type="simple"/></inline-formula> and an action <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e084" xlink:type="simple"/></inline-formula> is taken. Consequently, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e085" xlink:type="simple"/></inline-formula>. The case <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e086" xlink:type="simple"/></inline-formula> is presented in <xref ref-type="table" rid="pcbi-1003618-t001">Table 1</xref>, where the distinction between <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e087" xlink:type="simple"/></inline-formula> and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e088" xlink:type="simple"/></inline-formula> formally defines Sugden and Kandori observers. Similarly with the action rule, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e089" xlink:type="simple"/></inline-formula>. Ad players treat every donor unfavorably, thus <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e090" xlink:type="simple"/></inline-formula>.</p>
<p>By combining action and assessment rules, we can formally express how action-assessment strategies determine the probabilities that the donor will be assigned the reputation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e091" xlink:type="simple"/></inline-formula> conditional on all relevant circumstances. Such conditional probabilities are crucial for calculating the probabilities <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e092" xlink:type="simple"/></inline-formula>. If the recipient is an insider, the relevant circumstances are specified by the type of observer (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e093" xlink:type="simple"/></inline-formula>) and donor (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e094" xlink:type="simple"/></inline-formula>), as well as the recipient's reputation from observer's (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e095" xlink:type="simple"/></inline-formula>) and donor's (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e096" xlink:type="simple"/></inline-formula>) viewpoints. Accordingly, we introduce<disp-formula id="pcbi.1003618.e097"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pcbi.1003618.e097" xlink:type="simple"/><label>(1)</label></disp-formula></p>
<p>If the recipient is an outsider, both action and assessment differ between shonin and bushi players, thus providing a way to formalize the distinction between the two. The former players make an effort to learn the outsider's reputation, whereas the latter simply dismiss the outsider as a player with an unfavorable reputation. Such a situation implies that outside recipients, who are by assumption of type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e098" xlink:type="simple"/></inline-formula>, are perceived favorably by the observer and the donor, i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e099" xlink:type="simple"/></inline-formula>, if and only if <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e100" xlink:type="simple"/></inline-formula>. Consequently, we can introduce the probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e101" xlink:type="simple"/></inline-formula> as an analogue to <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e102" xlink:type="simple"/></inline-formula> conditional only on the type of observer (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e103" xlink:type="simple"/></inline-formula>), donor (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e104" xlink:type="simple"/></inline-formula>), and outsider (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e105" xlink:type="simple"/></inline-formula>) by<disp-formula id="pcbi.1003618.e106"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pcbi.1003618.e106" xlink:type="simple"/><label>(2)</label></disp-formula></p>
<p>We mentioned that the reputation dynamics controlled partitioning of players according to their reputation within a generation. Therefore, each generation plays many rounds of the game, whereby every player serves both as a donor and as a recipient once per round. When serving as a donor, the player encounters either an insider recipient with probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e107" xlink:type="simple"/></inline-formula> or an outsider recipient with probability <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e108" xlink:type="simple"/></inline-formula>. If the recipient is an insider, the probability of it being a player of type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e109" xlink:type="simple"/></inline-formula> is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e110" xlink:type="simple"/></inline-formula> and the probability of its reputation being <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e111" xlink:type="simple"/></inline-formula> is <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e112" xlink:type="simple"/></inline-formula>, where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e113" xlink:type="simple"/></inline-formula> denotes the current round of the game. Using <xref ref-type="disp-formula" rid="pcbi.1003618.e097">Eqs. (1)</xref> and <xref ref-type="disp-formula" rid="pcbi.1003618.e106">(2)</xref>, the probability that the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e114" xlink:type="simple"/></inline-formula> donor is assigned the reputation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e115" xlink:type="simple"/></inline-formula> for the next round, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e116" xlink:type="simple"/></inline-formula>, becomes<disp-formula id="pcbi.1003618.e117"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pcbi.1003618.e117" xlink:type="simple"/><label>(3)</label></disp-formula>After (infinitely) many rounds of the game, the probabilities <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e118" xlink:type="simple"/></inline-formula> converge to the equilibrium values defined by <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e119" xlink:type="simple"/></inline-formula>. These equilibrium values are then used to simulate the evolutionary dynamics of value systems inside the inner circle.</p>
</sec><sec id="s3b">
<title>Evolutionary dynamics</title>
<p>The evolutionary dynamics of value systems inside the inner circle is modeled using the replicator equations. If we denote the fraction of the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e120" xlink:type="simple"/></inline-formula> players at the generational time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e121" xlink:type="simple"/></inline-formula> with <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e122" xlink:type="simple"/></inline-formula>, so that <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e123" xlink:type="simple"/></inline-formula>, then the same fraction in the next generation, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e124" xlink:type="simple"/></inline-formula>, is given by<disp-formula id="pcbi.1003618.e125"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pcbi.1003618.e125" xlink:type="simple"/><label>(4)</label></disp-formula>where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e126" xlink:type="simple"/></inline-formula> is the fitness of the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e127" xlink:type="simple"/></inline-formula> players and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e128" xlink:type="simple"/></inline-formula> is the average fitness (both at time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e129" xlink:type="simple"/></inline-formula>). Fitness is a function of the equilibrium probabilities <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e130" xlink:type="simple"/></inline-formula> (and the parameters <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e131" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e132" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e133" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e134" xlink:type="simple"/></inline-formula>) because players of a given type receive the payoff <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e135" xlink:type="simple"/></inline-formula> to the extent they are perceived favorably by their respective donors and incur the cost <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e136" xlink:type="simple"/></inline-formula> to the extent they perceive their recipients favorably. Assuming that outsiders are of type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e137" xlink:type="simple"/></inline-formula>, i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e138" xlink:type="simple"/></inline-formula>, the above considerations can be written in general mathematical terms as<disp-formula id="pcbi.1003618.e139"><graphic position="anchor" xlink:href="info:doi/10.1371/journal.pcbi.1003618.e139" xlink:type="simple"/></disp-formula>where <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e140" xlink:type="simple"/></inline-formula> is an arbitrary basic level of fitness and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e141" xlink:type="simple"/></inline-formula> is the usual Kronecker delta symbol. Besides <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e142" xlink:type="simple"/></inline-formula>, two additional summands appear in the last equation. The first of the two summands represents the difference between the benefits received and the costs incurred from within-group encounters. Note that the benefits from type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e143" xlink:type="simple"/></inline-formula> players are received only if the reputation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e144" xlink:type="simple"/></inline-formula> is favorable, i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e145" xlink:type="simple"/></inline-formula>. Similarly, when encountering type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e146" xlink:type="simple"/></inline-formula> players the costs are incurred only if the reputation <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e147" xlink:type="simple"/></inline-formula> is favorable, i.e. <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e148" xlink:type="simple"/></inline-formula>. The second of the two summands also represents the difference between benefits and costs, but now as a result of cross-border encounters. Here, insiders who are of the same type as outsiders, i.e. when <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e149" xlink:type="simple"/></inline-formula>, receive the benefits if their reputation is favorable (<inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e150" xlink:type="simple"/></inline-formula>) and incur the cost <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e151" xlink:type="simple"/></inline-formula> with certainty because the outside world is assumed to be in a stable equilibrium populated by the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e152" xlink:type="simple"/></inline-formula> players. Social parasites behave opportunistically in the sense that they receive the benefits whenever their reputation is favorable from the viewpoint of the type <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e153" xlink:type="simple"/></inline-formula> observer.</p>
</sec><sec id="s3c">
<title>Implementation</title>
<p>We performed numerical simulations based on the described methodology to (i) visualize convergence of the model over the generational time scale, (ii) delineate the domains of attraction, and (iii) estimate their sizes. We achieved these goals in several steps. First, we defined a grid with 5050 points distributed uniformly over the ternary domain <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e154" xlink:type="simple"/></inline-formula>, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e155" xlink:type="simple"/></inline-formula>, and <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e156" xlink:type="simple"/></inline-formula>. Coordinates of each grid point served as the initial conditions for one model run. Every run consisted of many generational time steps, where in a single step multiple rounds of the game played out according to <xref ref-type="disp-formula" rid="pcbi.1003618.e117">Eq. (3)</xref>. Instead of presetting the number of rounds, we waited until the difference <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e157" xlink:type="simple"/></inline-formula> reached the desired accuracy. The resulting approximation of the equilibrium probabilities <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e158" xlink:type="simple"/></inline-formula> allowed us to calculate the fitness of all player types and advance their respective fractions, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e159" xlink:type="simple"/></inline-formula>, into the next generation, <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e160" xlink:type="simple"/></inline-formula>, using <xref ref-type="disp-formula" rid="pcbi.1003618.e125">Eq. (4)</xref>. Eventually, the model converged to one of the attractors, forming a link between the starting point and the attractor. In the second step, we chose a uniform subset (134 points) of the initial grid for which sample paths over the generational time <inline-formula><inline-graphic xlink:href="info:doi/10.1371/journal.pcbi.1003618.e161" xlink:type="simple"/></inline-formula> were stored and subsequently visualized in the ternary plots (<xref ref-type="fig" rid="pcbi-1003618-g001">Figs. 1</xref> and <xref ref-type="fig" rid="pcbi-1003618-g003">3</xref>). For the visualization, we used curved arrows to characterize the direction and the rate of convergence along sample paths; the longer an arrow, the faster the convergence rate along that particular path. The third step began after completing all 5050 runs for a fixed parameter set. Because every grid point had been linked with an attractor, we could isolate the neighboring points that belonged to two different domains of attraction. Segments connecting such neighboring points were further subdivided with five equidistant points to provide the initial conditions for extra runs in which an even closer pair belonging to two different domains of attraction could be determined. The process continued until the distance between the neighboring points reached the desired accuracy and thus revealed the location of the border between the adjacent domains of attraction. In the final step, we calculated the fraction of grid points linked to each attractor as an estimate of the size of the corresponding domain of attraction.</p>
</sec></sec></body>
<back>
<ack>
<p>We thank Joung-Hun Lee for constructive criticism on an earlier version of the manuscript.</p>
</ack>
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