File 015501
Game Theory and Morality by Moshe Hoffman, Erez Yoeli, and Carlos David Navarrete (File 015501)
Academic chapter exploring the intersection of game theory and moral intuitions, examining puzzles in human morality through evolutionary game theory frameworks.
Summary
This academic paper applies game theory to explain various moral puzzles and inconsistencies in human morality. The authors use Nash equilibrium concepts and evolutionary dynamics to argue that moral behaviors and intuitions converge toward game-theoretic solutions. The chapter examines phenomena including violations of human dignity (dwarf tossing), inefficient charitable giving practices, and culturally universal prohibitions on murder, proposing that these moral intuitions evolved to solve recurring social cooperation problems.
Game Theory and MoralityMoshe Hoffman , Erez Yoeli , and Carlos David NavarreteIntroductionConsider the following puzzling aspects of our morality:1. Many of us share the view that one should not use people, even if it benefits themto be used, as Kant intoned in his second formulation of the categorical imperative:“Act in such a way that you treat humanity, whether in your own person orin the person of any other, never merely as a means to an end, but always at thesame time as an end” (Kant, 1997 ). Consider dwarf tossing, where dwarfs wearingprotective padding are thrown for amusement, usually at a party or pub. It isviewed as a violation of dwarfs’ basic dignity to use them as a means for amusement,even though dwarves willingly engage in the activity for economic gain.Many jurisdictions ban dwarf tossing on the grounds that the activity violatesdwarfs’ basic human rights, and these laws have withstood lawsuits raised bydwarfs suing over the loss of employment (!).2. Charitable giving is considered virtuous, but little attention is paid to how justthe cause or efficient the charity. For example, Jewish and Christian traditionsadvocate giving 10 % of one’s income to charity, but make no mention of theimportance of evaluating the cause or avoiding wasteful charities. The intuitionthat giving to charity is a moral good regardless of efficacy results in the persistenceof numerous inefficient and corrupt charities. For example, the WishingWell Foundation has, for nearly a decade, ranked as one of CharityNavigator.M. Hoffman (*) • E. YoeliProgram for Evolutionary Dynamics , Harvard University ,One Brattle Square, Suite 6 , Cambridge , MA 02138 , USAe-mail: moshehoffman@fas.harvard.eduC.D. NavarreteDepartment of Psychology, and, the Ecology, Evolutionary Biology and Behavior Program ,Michigan State University , East Lansing , MI , USA© Springer International Publishing Switzerland 2016T.K. Shackelford, R.D. Hansen (eds.), The Evolution of Morality,Evolutionary Psychology, DOI 10.1007/978-3-319-19671-8_14289290M. Hoffman et al.com’s most inefficient charities. Yet its mission of fulfilling wishes by childrenwith terminal illnesses is identical to that of the more efficient Make-A-WishFoundation. Worse yet, scams masquerading as charities persist. One man operatingas The US Navy Veteran’s Association collected over 100 million dollars—over 7 years!—before anyone bothered to investigate the charity.3. In every culture and age, injunctions against murder have existed. If there is onething much of humanity seems to agree on, it’s that ending the life of anotherwithout just cause which is among the worst of moral violations. Yet culturesdon’t consider the loss of useful life years in their definition, even though it isrelevant to the measure of harm done by the murder. Why is our morality somuch more sensitive to whether a life was lost than to how much life was lost?There are numerous other examples of how our moral intuitions appear to be rifewith logical inconsistencies. In this chapter, we use game theory to provide insighton a range of moral puzzles similar to the puzzles described above.What Is Game Theory and Why Is It Relevant?In this section , we review the defi nition of a game , and of a Nash equilibrium , thendiscuss how evolution and learning processes would yield moral intuitions consistentwith Nash equilibria .Game theory is a tool for the analysis of social interactions. In a game, the payoffto each player depends on their actions, as well as the actions of others. Considerthe Prisoner’s Dilemma (Chammah & Rapoport, 1965 ; see Fig. 1 ), a model thatcaptures the paradox of cooperation. Each of two players chooses whether to cooperateor to defect. Cooperating reduces a player’s payoff by c > 0 while increasingthe other’s payoffs by b> c. Players could be vampire bats with the option ofsharing blood, or firms with the option of letting each other use their databases, orpremed students deciding whether to take the time to help one another to study. Thepayoffs, b and c , may represent likelihood of surviving and leaving offspring, profits,or chance of getting into a good medical school.Solutions to such games are analyzed using the concept of a Nash equilibrium 1 —a specification of each player’s action such that no player can increase his payoff bydeviating unilaterally. In the Prisoner’s Dilemma, the only Nash equilibrium is forneither player to cooperate, since regardless of what the other player does, cooperationreduces one’s own payoff.1Note that we focus on the concept of Nash equilibrium in this chapter and not evolutionary stablestrategy (ESS), a refinement of Nash that might be more familiar to an evolutionary audience. ESSare the Nash equilibria that are most relevant in evolutionary contexts. However, ESS is not welldefined in many of our games, so we will focus on the insights garnered from Nash and directlydiscuss evolutionary dynamics when appropriate.Morality Games291Fig. 1 The Prisoner’sDilemma. Player 1’s availablestrategies (C and D, whichstand for cooperate anddefect, respectively) arerepresented as rows . Player2’s available strategies (alsoC and D) are represented ascolumns . Player 1’s payoffsare represented at theintersection of each row andcolumn. For example, ifplayer 1 plays D and player 2plays C, player 1’s payoff isb. The Nash equilibrium ofthe game is (D, D). It isindicated with a circleGame theory has traditionally been applied in situations where players are rationaldecision makers who deliberately maximize their payoffs, such as pricingdecisions of firms (Tirole, 1988 ) or bidding in auctions (Milgrom & Weber, 1982 ).In these contexts, behavior is expected to be consistent with a Nash equilibrium,otherwise one of the agents—who are actively deliberating about what to do—would realize she could benefit from deviating from the prescribed strategy.However, game theory also applies to evolutionary and learning processes, whereagents do not deliberately choose their behavior in the game, but play according tostrategies with which they are born, imitate, or otherwise learn. Agents play a gameand then “reproduce” based on their payoffs, where reproduction represents offspring,imitation, or learning. The new generation then play the game, and so on. Insuch settings, if a mutant does better (mutation can be genetic or can happen whenagents experiment), then she is more likely to reproduce or her behavior imitated orreinforced, causing the behavior to spread. This intuition is formalized using modelsof evolutionary dynamics (e.g., Nowak, 2006 ).The key result for evolutionary dynamic models is that, except under extremeconditions, behavior converges to Nash equilibria. This result rests on one simple,noncontroversial assumption shared by all evolutionary dynamics: Behaviors thatare relatively successful will increase in frequency. Based on this logic, game theorymodels have been fruitfully applied in biological contexts to explain phenomenasuch as animal sex ratios (Fisher, 1958 ), territoriality (Smith & Price, 1973 ), cooperation(Trivers, 1971 ), sexual displays (Zahavi, 1975 ), and parent–offspring conflict(Trivers, 1974 ). More recently, evolutionary dynamic models have been appliedin human contexts where conscious deliberation is believed to not play an importantrole, such as in the adoption of religious rituals (Sosis & Alcorta, 2003 ), in theexpression and experience of emotion (Frank, 1988 ; Winter, 2014 ), and in the useof indirect speech (Pinker, Nowak, & Lee, 2008 ).292M. Hoffman et al.Crucially for this chapter, because our behaviors are mediated by moral intuitionsand ideologies, if our moral behaviors converge to Nash, so must the intuitions andideologies that motivate them. The resulting intuitions and ideologies will bearthe signature of their game theoretic origins, and this signature will lend clarity onthe puzzling, counterintuitive, and otherwise hard-to-explain features of our moralintuitions, as exemplified by our motivating examples.In order for game theory to be relevant to understanding our moral intuitions andideologies, we need only the following simple assumption: Moral intuitions andideologies that lead to higher payoffs become more frequent . This assumption canbe met if moral intuitions that yield higher payoffs are held more tenaciously, aremore likely to be imitated, or are genetically encoded. For example, if every timeyou transgress by commission you are punished, but every time you transgress byomission you are not, you will start to intuit that commission is worse thanomission.Rights and the Hawk–Dove GameIn this section we will argue that just as the Hawk –Dove model explains animal territoriality(Maynard Smith & Price, 1973 , to be reviewed shortly ), the Hawk –Dovemodel sheds light onto our sense of rights (Descioli & Karpoff, 2014 ; Gintis, 2007 ;Myerson, 2004 ).Let us begin by asking the following question (Myerson, 2004 ): “Why [does] apassenger pay a taxi driver after getting out of the cab in a city where she is visitingfor one day, not expecting to return?” If the cabby complains to the authorities, thepassenger could plausibly claim that she had paid in cash. The answer, of course, isthat the cabby would feel that the money the passenger withheld was his—that hehad a right to be paid for his service—and get angry, perhaps making a scene oreven starting a fight. Likewise, if the passenger did in fact pay, but the cabbydemanded money a second time, the passenger would similarly be infuriated. Thisexample illustrates that people have powerful intuitions regarding rightful ownership.In this section, we explore what the Hawk – Dove game can teach us about oursense of property rights.The reader is likely familiar with the Hawk – Dove game, a model of disputesover contested resources. In the Hawk – Dove game, each player decides whether tofight over a resource or to acquiesce (i.e. play Hawk or Dove). If one fights and theother does not, the fighter gets the resource, worth v . If both fight, each pays a costc and split the resource. That is, each gets v/2- c. If neither fights, they split theresource and get v /2. As long as v/2 < c, then in any stable Nash equilibrium, oneplayer fights and the other acquiesces. That is, if one player expects the other tofight, she is better off acquiescing, and vice versa (see Fig. 2 ).Crucially, it is not just a Nash equilibrium for one player to always play Hawkand the other to always play Dove. It is also an equilibrium for both players to conditionwhether they play Hawk on an uncorrelated asymmetry —a cue or event thatMorality Games293Fig. 2 The Hawk–Dovegame. The Nash equilibria ofthe game are circleddoes not necessarily affect the payoffs, but does distinguish between the players,such as who arrived at the territory first or who built the object. If one conditions onthe event (say, plays Hawk when she arrives first), then it is optimal for the other tocondition on the event (to play Dove when the other arrives first).As our reader is likely aware, this was the logic provided by Maynard Smith toexplain animal territoriality—why animals behave aggressively to defend territorythat they have arrived at first, even if incumbency does not provide a defensiveadvantage and even when facing a more formidable intruder. Over the years, evidencehas amassed to support Maynard Smith’s explanation, such as experimentalmanipulation of which animal arrives first (Davies, 1978 ; Sigg & Falett, 1985 ).Like other animals, we condition how aggressively we defend a resource onwhether we arrive first. Because our behaviors are motivated by beliefs, we are alsomore likely to believe that the resource is “ours” when we arrive first. Studies haveshown these effects with children’s judgments of ownership, in ethnographies ofprelegal societies, and in computer games. In one such illustration, DeScioli andWilson ( 2011 ) had research subjects play a computer game in which they contesteda berry patch. Subjects who ended up keeping control of the patch usually arrivedfirst, and this determined the outcome more often than differences in fighting abilityin the game.This sense of ownership is codified in our legal systems, as illustrated by the quip“possession is 9/10ths of the law,” and in a study involving famous legal propertycases conducted by Descioli and Karpoff ( 2014 ). In a survey, these researchersasked participants to identify the rightful owner of a lost item, after reading vignettesbased on famous property rights legal cases. Participants consistently identified thepossessor of the found item as its rightful owner (as the judges had at the time of thecase). This sense of ownership is also codified in our philosophical tradition, e.g., inLocke ( 1988 ), who found property rights in initial possession. Note that, as has alsobeen found in animals, possession extends to objects on one’s land: In DeScioli and294M. Hoffman et al.Karpoff’s survey, another dictate of participants’ (and the judges’) property rightsintuitions was who owned the land on which the lost item was found.Also like animals, our sense of property rights is influenced by who created orinvested in the resource, another uncorrelated asymmetry. In locales that sometimesgrant property rights to squatters—individuals who occupy lands others have purchased—akey determinant of whether the squatters are granted the land is whetherthey have invested in it (Cone vs. West Virginia Pulp & Paper Co., 1947 ; Neuwirth,2005 ). Locke also intuited that investment in land is part of what makes it ours:In Second Treatise on Civil Government (1689), Locke wrote, “everyman has aproperty in his person; this nobody has a right to but himself. The labor of his bodyand the work of his hand, we may say, are properly his.”If the Hawk – Dove model underlies our sense of property rights, we would expectto see psychological mechanisms that motivate us to feel entitled to an object whenwe possess it or have invested in it. Here are three such mechanisms, which can beseen by reinterpreting some well-documented “biases” in the behavioral economicsliterature. The first such bias is the endowment effect : We value items more if we arein possession of them. The endowment effect has been documented in dozens ofexperiments, where subjects are randomly given an item (mug, pen, etc.) andsubsequently state that they are willing to sell the mug for much more than thosewho were not given the mug are willing to pay (Kahneman, Knetsch, & Thaler,1990 ). In the behavioral economics literature, the endowment effect has sometimesbeen explained by loss aversion, which is when we are harmed more by a loss thanwe benefit from an equivalent gain. However, the source of loss aversion is notquestioned or explained. When it is, loss aversion is also readily explained by theHawk – Dove game (Gintis, 2007 ).A second bias that also fits the Hawk – Dove model is the IKEA effect : Our valuationof an object is influenced by whether we have developed or built the resource.The IKEA effect has been documented by asking people how much they would payfor items like Lego structures or IKEA furniture after randomly being assigned tobuild them or receive them pre-built. Subjects are willing to pay more for items theybuild themselves.A third such bias that fits the Hawk – Dove model is the sunk cost fallacy (Mankiw,2007 ; Thaler, 1980 ), which leads us to “throw good money after bad” when weinvest in ventures simply because we have already put so much effort into them,arguably because our prior efforts lead us to value those ventures more.Possession and past investment are not the only uncorrelated asymmetries thatcan dictate rights. Rights can be dictated by a history of agreements, as happenswhen one party sells another deed to a house or car, or, as in our taxicab example,by whether a service was provided. There are also countless examples in whichrights were determined by perhaps unfair or arbitrary characteristics such as raceand sex: Black Americans were expected to give up their seat for Whites in the JimCrow South and women to hand over their earnings or property to their husbandsthroughout the ages.Hawk – Dove is not just a post hoc explanation for our sense of rights; it also leadsto the following novel insight: We can formally characterize the properties thatMorality Games295uncorrelated asymmetries must have. This requires a bit more game theory to illustrate;the logic is detailed in the section on categorical distinctions but the implicationsare straightforward: Uncorrelated asymmetries must be discrete (as in whoarrived first or whether someone has African ancestry) and cannot be continuous(who is stronger, whether someone has darker skin). Indeed, we challenge the readerto identify a case where our sense of rights depends on surpassing a threshold in acontinuous variable (stronger than? darker than?). More generally, an asymmetrymust have the characteristic that, when it occurs, every observer believes it occurredwith a sufficiently high probability, where the exact level of confidence is determinedby the payoffs of the game. This is true of public, explicit speech and handshakes,but not innuendos or rumors. (Formally, explicit speech and handshakesinduce what game theorists term common p-beliefs.)The Hawk – Dove explanation of our sense of rights also gives useful clarity onwhen there will be conflict. Conflict will arise if both players receive opposing signalsregarding the uncorrelated asymmetry, such as two individuals each believingthey arrived first, or when there are two uncorrelated asymmetries that point inconflicting directions, such as when one person invested more and the other arrivedfirst. The former source of conflict appears to be the case in the Israeli–Palestinianconflict. Indeed, both sides pour great resources into demonstrating their earlypossession, especially Israel, through investments in and public displays of archeologyand history. The latter source of conflict appears to be the case in many of thecontested legal disputes in the study by DeScioli and Karpoff ( 2014 ) mentionedabove. An example is one person finds an object on another’s land. Indeed, this turnsout to be a source of many legal conflicts over property rights, and a rich legal traditionhas developed to assign precedence to one uncorrelated asymmetry over another(Descioli & Karpoff, 2014 ). As usual, we see similar behavior in animals in studiesthat provide empirical support for Maynard Smith’s model for animal territoriality:When two animals are each given the impression they arrived first by, for example,clever use of mirrors, a fight ensues (Davies, 1978 ).Authentic Altruism, Motives, and the Envelope GameIn this section , we present a simple extension of the Repeated Prisoner’s Dilemmato explain why morality depends not just on what people do but also what they thinkor consider .In the Repeated Prisoner’s Dilemma and other models of cooperation, playersjudge others by their actions—whether they cooperate or defect. However, we notonly care about whether others cooperate but also about their decision-making process:We place more trust in cooperators who never even considered defecting. Toquote Kant, “In law a man is guilty when he violates the rights of others. In ethicshe is guilty if he only thinks of doing so.”The Envelope Game (Fig. 3 ) models why we care about thoughts and considerationsand not just actions (Hoffman, Yoeli, & Nowak, 2015 ). The Envelope Game296M. Hoffman et al.Fig. 3 A single stage of the Envelope Gameis a repeated game with two players. In each round, player 1 receives a sealed envelope,which contains a card stating the costs of cooperation (high temptation todefect vs. low temptation to defect). The temptation is assigned randomly and isusually low. Player 1 can choose to look inside the envelope and thus find out themagnitude of the temptation or choose not to look. Then player 1 decides to cooperateor to defect. Subsequently, player 2 can either continue to the next round or endthe game. As in the Repeated Prisoner’s Dilemma, the interaction repeats with agiven likelihood, and if it does, an envelope is stuffed with a new card and presentedto player 1, etc.In this model, as long as temptations are rare, large, and harmful to player 2, it isa Nash equilibrium for player 1 to “cooperate without looking” in the envelope andfor player 2 to continue if and only if player 1 has cooperated and not looked. Werefer to this as the cooperate without looking (CWOL) equilibrium. 2 This equilibriumemerges in agent-based simulations of evolution and learning processes. 3 Noticethat if player 1 could not avoid looking inside the envelope, or player 2 could notobserve whether player 1 looked, there would not be a cooperative equilibriumsince player 1 would benefit by deviating to defection in the face of large temptations.Not looking permits cooperative equilibria in the face of large temptations.The Envelope Game is meant to capture the essential features of many interestingaspects of our morality, as described next.Authentic Altruism . Many have asked whether “[doing good is] always and exclusivelymotivated by the prospect of some benefit for ourselves, however subtle”(Batson, 2014 ), for example, the conscious anticipation of feeling good (Andreoni,2Technically, the conditions under which we expect players to avoid looking and attend to lookingare ch > a /(1 − w ) > c lp + ch (1 − p ) and bp + d (1 − p ) < 0), where ch and cl are the magnitudes of thehigh and low temptations, respectively; p is the likelihood of the low temptation; a /(1 − w ) is thevalue of a repeated, cooperative interaction to player 1; and bp + d (1 − p ) is the expected payoff toplayer 2 if player 1 only cooperates when the temptation is low.3The simulations employ numerical estimation of the replicator dynamics for a limited strategyspace: cooperate without looking, cooperate with looking, look and cooperate only when the temptationis low, and always defect for player 1, and end if player 1 looks, end if player 1 defects, andalways end for player 2.Morality Games2971990 ), avoidance of guilt (Cain, Dana, & Newman, 2014 ; Dana, Cain, & Dawes,2006 ; DellaVigna, List, & Malmendier, 2012 ), anticipation of reputational benefitsor reciprocity (as Plato’s Glaucon suggests, when he proffers that even a pious manwould do evil if given a ring that makes him invisible; Trivers, 1971 ). At the extreme,this amounts to asking if saintly individuals such as Gandhi or Mother Teresa weremotivated thus, or if they were “authentic” altruists who did good without anticipatingany reward and would be altruistic even in the absence of such rewards.Certainly, religions advocate doing good for the “right” reasons. In the Gospel ofMatthew, Chapter 6, Jesus advocates, “Be careful not to practice your righteousnessin front of others to be seen by them. If you do, you will have no reward from yourFather in heaven,” after which he adds, “But when you give to the needy, do not letyour left hand know what your right hand is doing, so that your giving may be insecret. Then your Father, who sees what is done in secret, will reward you.”The Envelope Game suggests authentic altruism is indeed possible: By focusingentirely on the benefits to others and ignoring the benefits to themselves, authenticaltruists are trusted more, and the benefits from this trust outweigh the risk of, forexample, dying a martyr’s death. Moreover, this model helps explain why we thinkso highly of authentic altruists, as compared to others who do good, but with anulterior motive (consider, as an example, the mockery Sean Penn has faced forshowing up at disaster sites such as Haiti and Katrina with a photographer in tow).Principles . Why do we like people who are “principled” and not those who are“strategic”? For example, we trust candidates for political office whose policies arethe result of their convictions and are consistent over time and distrust those whosepolicies are carefully constructed in consultation with their pollsters and who “flipflop”in response to public opinion (as caricatured by the infamous 2004 Republicanpresidential campaign television ad showing John Kerry windsurfing and tackingfrom one direction to another). CWOL offers the following potential explanation.Someone who is strategic considers the costs and benefits to themselves of everydecision and will defect when faced with a large temptation, whereas someone whois guided by principles is less sensitive to the costs and benefits are to themselvesand thus less likely to defect. Imagine our flip-flopping politician was once againstgay marriage but supports it now that it is popular. This indicates the politician isunlikely to fight for the cause if it later becomes unpopular with constituents or riskslosing a big donor. Moreover, this model may help explain why ideologues that arewholly devoted to a cause (e.g., Hitler, Martin Luther King, and Gandhi) are able toattract so many followers.Don ’t Use People . Recall Kant’s second formulation of the categorical imperative:“Act in such a way that you always treat humanity, whether in your own person orin the person of any other, never simply as a means but always at the same time asan end.” In thinking this through, let’s again consider dwarf tossing. Many see it asa violation of dwarfs’ basic dignity to use them as a means for amusement, eventhough they willingly engage in the activity for economic gain. Our aversion tousing people may explain many important aspects of our moral intuitions, such as298M. Hoffman et al.why we judge torture as worse than imprisonment or punishment (torture is harmingsomeone as a means to obtaining information) and perhaps one of the (many) reasonswe oppose prostitution (prostitution is having sex with someone as a means toobtaining money). The Envelope Game clarifies the function of adhering to thismaxim. Whereas those who treat someone well as means to an end would alsomistreat them if expedient, those who treat someone well as an end can be trustednot to mistreat them when expedient.Attention to Motives . The previous two applications are examples of a more generalphenomenon: that we judge the moral worth of an action based on the motivationof the actor, as argued by deontological ethicists, but contested byconsequentialists. The deontological argument is famously invoked by Kant:“Action from duty has its moral worth not in the purpose to be attained by it but inthe maxim in accordance with which it is decided upon, and therefore does notdepend upon the realization of the object of the action but merely upon the principleof volition in accordance with which the action is done without regard for any objectof the faculty of desire” (Kant, 1997 ). These applications illustrate that we attend tomotives because they provide valuable information on whether the actor can betrusted to treat others well even when it is not in her interest.Altruism Without Prospect of Reciprocation . CWOL also helps explain why peoplecooperate in contexts where there is no possibility of reciprocation, such as inone-shot anonymous laboratory experiments like the dictator game (Fehr &Fischbacher, 2003 ), as well as when performing heroic and dangerous acts. Considersoldiers who throw themselves on a grenade to save their compatriots or stories likethat of Liviu Librescu, a professor at the University of Virginia and a Holocaust survivor,who saved his students during a school shooting. When he heard the shootercoming toward his classroom, Librescu stood behind the door to his classroom,expecting that when the shooter tried to shoot through the door, it would kill him andhis dead body would block the door. Mr. Librescu, clearly, did not expect this act tobe reciprocated. Such examples have been used as evidence for group selection(Wilson, 2006 ), but can be explained by individuals “not looking” at the chance offuture reciprocation. Consistent with this interpretation, cooperation during extremeacts of altruism is more likely to be intuitive than deliberative (Rand & Epstein,2014 ), and those who cooperate without considering the prospect of reciprocationare more trusted (Critcher, Inbar, & Pizarro, 2013 ). We also predict that people aremore likely to cooperate intuitively when they know they are being observed.The Omission–Commission Distinctionand Higher-Order BeliefsWe explain the omission –commission distinction and the means–by-product distinctionby arguing that these moral intuitions evolved in contexts where punishment iscoordinated. Then , even when intentions are clear to one witness for omissions andby-products , a witness will think intentions are less clear to the other witnesses .Morality Games299Why don’t we consider it murder to let someone die that we could have easilysaved? For example, we sometimes treat ourselves to a nice meal at a fancy restaurantrather than donating the cost of that meal to a charity that fights deadly diseases.This extreme example illustrates a general phenomenon: that people have a tendencyto assess harmful commissions (actions such as killing someone) as worse, ormore morally reprehensible, than equally harmful omissions (inactions such as lettingsomeone die). Examples of this distinction abound, in ethics (we assess withholdingthe truth as less wrong than lying (Spranca, Minsk, & Baron, 1991 )), in law(it is legal to turn off a patient’s life support and let the patient die, as long as onehas the consent of the patient’s family; however, it is illegal to assist the patient incommitting suicide even with the family’s consent), and in international relations.For example, consider the Struma, a boat carrying Jewish refugees fleeing Nazipersecution in 1942. En route to Palestine, the ship’s engine failed, and it was towedto a nearby port in Turkey. At the behest of the British authorities then in control ofPalestine, passengers were denied permission to disembark and find their way toPalestine by land. For weeks, the ship sat at port. Passengers were brought onlyminimal supplies, and their requests for safe haven were repeatedly denied by theBritish and others. Finally, the ship was towed to known hostile waters in the BlackSea, where it was torpedoed by a Russian submarine almost immediately, killing791 of 792 passengers. Crucially, though, the British did not torpedo the ship themselvesor otherwise execute passengers—an act of commission that they and theirsuperiors would undoubtedly have found morally reprehensible.Why do we distinguish between transgressions of omission and commission? Toaddress this question, we present a simple game theory model based on the insightby DeScioli, Bruening, and Kurzban ( 2011 ). The intuition can be summarized infour steps:1. We note that moral condemnation motivates us to punish transgressors. Suchpunishment is potentially costly, e.g., due to the risk of retaliation. We expectpeople to learn or evolve to morally condemn only when such costs are worthpaying.2. Moral condemnation can be less costly when others also condemn, perhapsbecause the risk of retaliation is diffused, because some sanctions do not workunless universally enforced or, worse, because others may sanction individualsthey believe wrongly sanctioned. This can be modeled using any game withmultiple Nash equilibria, including the Repeated Prisoner’s Dilemma and theSide- Taking Game. The Coordination Game is the simplest game with multipleequilibria, so we present this game to convey the basic intuition. In theCoordination Game, there are two players who each simultaneously choosebetween two actions, say punish and don’t punish. The key assumption is thateach player prefers to do what she expects the other to do, which can be capturedby assuming each receives a if they both punish, d if neither punish, b < d if onepunishes and the other does not, and c < a if one does not punish while the otherdoes (Fig. 4 ).3. Transgressions of omission that are intended are difficult to distinguish fromunintended transgression, as is the case when perpetrators are simply not paying300M. Hoffman et al.Fig. 4 The CoordinationGame. In our applications, Astands for punish, and Bstands for don’t punishattention or do not have enough time to react with better judgment (DeScioliet al., 2011 ). Relative to the example of the tennis player with the allergydescribed above, it is usually hard to distinguish between a competitor who doesnot notice his opponent orders the dish with the allergen versus one who noticesbut does not care. In contrast, transgressions of commission must be intendedalmost by definition.4. Suppose the witness knows an omission was intentional: In the above example,the tennis player’s opponent’s allergy is widely known, and the witness saw theplayer watch his opponent order the offending dish, had time to react, thoughtabout it, but did not to say anything. The witness suspects that others do notknow the competitor was aware his opponent ordered the dish, but believes thetennis player should be condemned for purposely withholding information fromhis competitor. However, since the witness does not wish to be the sole condemner,she is unlikely to condemn. In contrast, when a witness observes a transgressionof commission (e.g., the player recommends the dish), the witness isrelatively confident that others present interpret the transgression as purposelyharmful, since his recommendation reveals that the player was obviously payingattention and therefore intended to harm his opponent. So, if all other individualspresent condemn the tennis player when they observe the commission, each doesnot anticipate being the sole condemner.For the above result to hold, all that is needed is the following: (1) The more the costsof punishment decrease, the more others punish and (2) omissions are usually unintended(Dalkiran, Hoffman, Paturi, Ricketts, & Vattani, 2012 ; Hoffman et al., 2015 ). 44In fact, even if one knows that others know that the transgression was intended, omission will stillbe judged as less wrong, since the transgression still won’t create what game theorists call commonp-belief, which is required for an event to influence behavior in a game with multiple equilibria.Morality Games301This explanation for the omission–commission distinction leads to two novelpredictions: First, for judgments and emotions not evolved to motivate witnesses topunishment but to, say, motivate witnesses to avoid dangerous partners (such as theemotion of fear; in contrast to anger or moral disgust), the omission–commissiondistinction is expected to be weaker or disappear altogether. Second, for transgressionsof omission that, without any private information, can be presumed intentional(such as a mother who allows her child to go hungry or a person who does not giveto a charity after being explicitly asked), we would not expect much of an omission–commissiondistinction in moral condemnation.As with the all models discussed in this chapter, the game theoretic explanationfor the omission–commission distinction does not rest on rational, conscious, strategiccalculation. In fact, in this particular case, all reasonable evolutionary dynamicmodels lead away from punishing omissions. The fact that the above results do notrest on rational, strategic thinking is particularly important in this setting since thereis evidence that the distinction between omissions and commissions is not determineddeliberately but rather intuitively (Cushman, Young, & Hauser, 2006 ) andappears to be evolved (DeScioli et al., 2011 ) and that consciously considering whatothers believe is an onerous process (Camerer, 2003 ; Epley, Keysar, Van Boven, &Gilovich, 2004 ; Hedden & Zhang, 2002 ).This same model can explain several other puzzling aspects of our morality. Thefirst is the means –by-product distinction. This distinction has been documented instudies that ask respondents to judge the following variants of the classic “trolley”problem. In the standard trolley “switch” case (Foot, 1967), a runaway trolley ishurtling toward a group of five people. To prevent their deaths, the trolley must beswitched onto a side track where it will kill an innocent bystander. In studies usingthis case, the vast majority of subjects choose the utilitarian option, judging it permissibleto cause the death of one to save five (e.g., Cushman et al., 2006 ; Mikhail,2007). In the “footbridge” variant (Thomson, 1976 ), the trolley is hurtling towardthe group of five people, but the switch to divert it is inoperable. The only way tosave the five is to push a man who is wearing a heavy backpack off a bridge onto thetrack, thereby slowing the trolley enough so the five can escape, but killing the man.In contrast to the standard switch version, where causing the death of one person isbut a by-product of the action necessary to save five, most subjects in the footbridgecase find it morally impermissible to force the man with the backpack onto thetracks (Cushman et al., 2006 ; DeScioli, Gilbert, & Kurzban, 2012 )—that is, whenthe man is used as a means to saving the five—even though the consequences are thesame, and the decision to act was made knowingly and deliberately in both cases.Such effects are found in less contrived situations, as well. Consider the real-lifedistinction between terrorism, in which civilian casualties are used a means to apolitical goal, and anticipated collateral damage, which is a by-product of war, evenwhen the same number of civilians are knowingly killed and the same political endsare desired (say increased bargaining power in a subsequent negotiation).The explanation again uses “higher-order beliefs” and is based on the key insightin DeScioli et al. ( 2011 ) and formalized in Dalkiran et al. ( 2012 ) and Hoffman et al.( 2015 ): When the harm is done as a by-product, the harm is not usually anticipated.302M. Hoffman et al.So even when a witness knows that the perpetrator anticipated the harm, the witnessbelieves other witnesses will not be aware of this and will presume the harm was notanticipated by the perpetrators. For instance, suppose we observe Israel killing civiliansas a by-product of a strategic raid on Hamas militants. Even if we knew Israelhad intelligence that confirmed the presence of civilians, we might not be sure otherswere privy to this information. On the other hand, when the harm is done as ameans, the harm must be anticipated, since otherwise the perpetrator would have nomotive to commit the act. Why would Hamas fire rockets at civilian towns with nomilitary presence if Hamas does not anticipate a chance of civilian casualties?Consequently, it is Nash equilibrium to punish harm done as a means but not harmdone as a by-product.Similar arguments can be made for why we find direct physical transgressionsworse than indirect ones, a moral distinction relevant to, for instance, the UnitedStates’ current drone policy. Cushman et al. ( 2006 ) found that subjects condemnpushing a man off a bridge (to stop a train heading toward five others) more harshlythan flipping a switch that leads the man to fall through a trap door. Pushing thevictim with a stick is viewed as intermediate in terms of moral wrongness. Suchmoral wrongness judgments are consistent with considerations of higher-orderbeliefs: When a man is physically pushed, any witness knows the pushing wasintended, but when a man is pushed with a stick some might not realize this, andeven those who realize it might suspect others will not. Even more so when a buttonis pressed that releases a trap door.It is worth noting that the above argument does not depend on a specific modelof punishment, as in DeScioli and Kurzban’s ( 2009 ) Side-Taking Game. The abovemodel also makes the two novel predictions enumerated above, but neverthelesscaptures the same basic insight. It is also worth noting the contrast between theabove argument and that of Cushman et al. ( 2006 ) and Greene et al. ( 2009 ), whosemodels rest on ease of learning or ease of mentally simulating a situation. It is notobvious to us how those models would explain that the omission–commission andmeans–by-product distinctions seem to depend on priors or be unique to settings ofcoordinated punishment.Why Morality Depends on Categorical DistinctionsWe explain why our moral intuitions depends so much more strongly on whether atransgression occurred than on how much damage was caused. Our argument againuses coordinated punishment and higher-order beliefs : When a categorical distinctionis violated, you know others know it was violated , but this is not always true forcontinuous variables .Consider the longstanding norm against the use of chemical weapons. This normrecently made headlines when Bashar al-Assad was alleged to have used chemicalweapons to kill about a thousand Syrian civilians, outraging world leaders who hadMorality Games303been silent over his use of conventional weapons to kill over 100,000 Syriancivilians. A Reuters/Ipsos poll at the time found that only 9 % of Americans favoredintervention in Syria, but 25 % supported intervention if the Syrian governmentforces used chemical weapons against civilians (Wroughton, 2013 ). In the past, theUnited States has abided by the norm against the use of chemical weapons even atthe expense of American lives: In WWII, Franklin D. Roosevelt chose to eschewchemical weapons in Iwo Jima even though, as his advisors argued at the time, theiruse would have saved thousands of American lives. It might even have been morehumane than the flame-throwers that were ultimately used against the Japanese(“History of Chemical Weapons,” 2013). We say that the norm against chemicalweapons is a categorical norm because those who abide by it consider whether atransgression was committed (did Assad use chemical weapons?), rather than focusingentirely on how much harm was done (how many civilians did Assad kill?).Other norms are similarly categorical. For instance, in the introduction to this chapter,we noted that across cultures and throughout history, the norm against murderhas always been categorical: We consider whether a life was terminated, not the lossof useful life years. Likewise, discrimination (e.g., during Jim Crow) is typicallybased on categorical definitions of race (the “one drop rule”) and not, say, the darknessof skin tone. Human rights are also categorical. A human rights violationoccurs if someone is tortured or imprisoned without trial, regardless of whether itwas done once or many times and regardless of whether the violation was helpful in,say, gaining crucial information about a dangerous enemy or an upcoming terrorattack. We even assign rights in a categorical way to all Homo sapiens and not basedon intelligence, sentience, ability to feel pain, etc.Why is it that we attend to such categorical distinctions instead of paying moreattention to the underlying continuous variable? We use game theory to explain thisphenomenon as follows: Suppose that two players (say, the United States andFrance) are playing a Coordination Game in which they decide whether to punishSyria, and each wants to sanction only if the other sanctions. We assume the UnitedStates does not want to levy sanctions unless it is confident France will as well,which corresponds to an assumption on the payoffs of the game (if we reverse thisassumption, it changes one line in the proof, but not the result).We model the underlying measure of harm as a continuous variable (in ourexample, it is the number of civilians killed). For simplicity, we assume this variableis uniformly distributed, which means Assad is equally likely to kill any number ofpeople. This assumption is, again, not crucial, and we will point out the line in theproof that it affects. Importantly, we assume that players do not directly observe thecontinuous variable, but instead receive some imperfect signal (e.g., the UnitedStates observes the body count by its surveyors).Imagine a norm that dictates that witnesses punish if their estimate of the harmfrom a transgression is above some threshold (e.g., levy sanctions against Syria ifthe number of civilians killed is estimated to be greater than 100,000). As it turnsout, this is not a Nash equilibrium. To see why, consider what happens when theUnited States gets a signal right at the threshold. The United States thinks there is a304M. Hoffman et al.50 % chance that France’s estimates are lower than its own 5 and, thus, that there isa 50 % chance that France’s estimates are lower than the threshold. This furtherimplies that the United States assesses only a 50 % chance that France levies sanctions,so the United States is not sufficiently confident that France will sanction, tomake it in the United States’s interest to sanction.What we have shown so far is that for a threshold of 100,000, it is in the interestof the United States to deviate from the strategy dictated by the threshold normwhen it gets a signal at the threshold. This means that 100,000 is not a viable threshold,and (since 100,000 was chosen arbitrarily) there is no Nash equilibrium inwhich witnesses punish if their estimate of the harm from a transgression is abovesome arbitrary threshold.It should be noted that this result only requires that there are sufficiently manypossibilities, not that there is in fact a continuum. Neither does it require that thedistribution is uniform nor that the Coordination Game is not affected by the behaviorof Assad. The only crucial assumptions are that the distribution is not too skewedand that the payoffs are not too dependent on the behavior of Assad (for details, seeDalkiran et al., 2012 ; Hoffman, Yoeli, & Dalkiran, 2015 ).What happens if such norms are learned or evolved and subject to selection?Suppose there is a norm to attack whenever more than 100,000 civilians are killed.Players will soon realize that they should not attack unless, say, 100,100 civiliansare killed. Then, players will learn not to attack when they estimate 100,200 civiliansare killed and so on , indefinitely. Thus, every threshold will eventually“unravel,” and no one will ever attack. 6Now let’s consider a categorical norm, for example, the use of chemical weapons.We again model this as a random variable, though this time, the random variablecan only take on two values (0 and 1), each with some probability. Again,players do not know with certainty whether the transgression occurred, but insteadget a noisy signal. In our example, the signal represents France or the United States’sassessment of whether Assad used chemical weapons, and there is some likelihoodthe assessors make mistakes: They might not detect chemical weapons when theyhad been used or might think they have detected chemical weapons when none hadbeen used.Unlike with the threshold norm, provided the likelihood of a mistaken signal isnot too high, there is a Nash equilibrium where both players punish when theyreceive a signal that the transgression occurred. That is, the United States and Franceeach levy sanctions if their assessors detect chemical weapons. This is becausewhen the United States detects chemical weapons, the United States believes France5This is where the assumption of a uniform distribution comes in. Had we instead assumed, forinstance, that the continuous variable is normally distributed, then it would not be exactly 50–50but would deviate slightly depending on the standard deviation and the location of the threshold.Nevertheless, the upcoming logic will still go through for most Coordination Games, i.e. anyCoordination Game with risk dominance not too close to .5.6As with omission, this follows from iterative elimination of strictly dominated strategies (seeHoffman et al., 2015 , for details).Morality Games305likely detected them and will likely levy sanctions. So the United States’s bestresponse is to levy sanctions. Similarly, if the United States does not detect chemicalweapons, it expects France did not and will not levy sanctions, so the UnitedStates is better off not levying them.This result is useful for evaluating whether it is worthwhile to uphold a norm.The Obama administration was harshly criticized for threatening to go to war afterthe Assad regime used chemical weapons but not earlier, although the regime hadalready killed tens of thousands of civilians. The model clarifies that Obama’s positionwas not as inconsistent as his critics had charged: The norm against chemicalweapons may be worth enforcing since it is sustainable, whereas norms againstcivilian casualties are harder to sustain and hence might not be worthwhile toenforce.Let’s return to some more of our motivating examples. Our model can explainwhy we define murder categorically: It is not possible to punish differently for differentamount of quality life years taken, but it is possible to punish differentially fora life taken. As with omission–commission, however, we do expect sadness or griefto depend greatly on life years lost, even if the punishment or moralistic outrage willbe less sensitive. This is a prediction of the model that, as far as we know, has yet tobe tested.Similarly, the “one-drop” rule is a categorical norm, so it can be socially enforcedin an apartheid society. In contrast, consider a rule that advocates giving up one’sseat for someone with lighter skin. Since this is based on a threshold in a continuousvariable, while it might be enforceable by a unilateral authority, it cannot be enforcedby “mob rule.” Other forms of discrimination, such as discriminating against theless attractive, or the less tall, or the elderly, all being continuous variables, cannotbe socially enforced via coordinated punishment, and hence, we expect such discriminationto be of a different form. In particular, it will not be based not on punishingviolators. For example, male CEOs might still prefer young attractive femalesecretaries, and taller men are more likely to be hired as CEOs, not because ofcoordinated rewards or punishment but because those who hire the CEOs or secretariesare likely to be satisfying their own preferences or doing what they expect willlead to higher profits.Likewise, the number of victims tortured by a regime or the number of livessaved by torturing is continuous. Thus, a regime cannot be punished by a coordinatedattack by other countries or by a coordinated rebellion by its citizens based onthe number of people tortured or the paucity of reasons for such torture. But, aregime can be attacked or overthrown depending on whether a physical harm wasinflicted on a citizen by the state. Hence, human rights are treated as inalienable,even in the absence of an a priori justification for this nonutilitarian norm. And whyare human rights ascribed to all living Homo sapiens ? Perhaps not because of agood logical a priori argument, but simply because violations of human rights areenforceable by coordinated punishment, but no regime can be punished for harmingany “person” of less than a certain degree of consciousness.Finally, here is one last application. The model might also explain why revolutionsare often caused by categorical events, such as a new tea tax or a single, widely306M. Hoffman et al.publicized self-immolation, and not a breach of a threshold in, say, the quality of lifeof citizens or the level of corruption. This explanation requires simply that we recognizerevolutions as a coordination problem (as argued in Morris & Shin, 2002 ;Chwe, 2013 ), where each revolutionary chooses whether to revolt, and each is betteroff revolting only if sufficiently many others revolt.Quirks of Altruism and the Repeated Prisoner’s Dilemmawith Incomplete InformationThe Repeated Prisoner ’s Dilemma has famously been used as an explanation forthe evolution of cooperation among non-kin (Axelrod & Hamilton, 1981 ; Dawkins,2006 ; Pinker, 2003 ; Trivers, 1971 ). In this section , we show how the same basicmodel can be used to explain many of the quirky features of our pro-social preferencesand ideologies .Recall that in the Prisoner’s Dilemma, each of two players simultaneouslychooses whether to cooperate. Cooperation reduces a player’s own payoffs by c > 0while increasing the other’s payoffs by b > c . The only Nash equilibrium is for neitherplayer to cooperate. In the Repeated Prisoner’s Dilemma, the players play astring of Prisoner’s Dilemmas. That is, after the players play a Prisoner’s Dilemma,they learn what their opponent did and play another Prisoner’s Dilemma against thesame opponent with probability δ (and the game ends with probability 1 − δ ). As iswell known in the evolutionary literature, there are equilibria in which players endup cooperating, provided δ > c /b . In all such equilibria, cooperation is sustainedbecause any defection by one player causes the other player to defect. This is calledreciprocity. As the reader is surely familiar, there is ample evidence for the RepeatedPrisoner’s Dilemma as a basis for cooperation from computer simulations (e.g.,Axelrod, 1984 ) and animal behavior (e.g., Wilkinson, 1984 ). The model can beextended to explain contributions to public goods if, after deciding whether to contributeto a public good, players play a Repeated Prisoner’s Dilemma (see, e.g.,Panchanathan & Boyd, 2004 ) (Fig. 5 ).The key to understanding these quirks is that players often have incompleteinformation. For example:1. Players do not always observe contributions. It is intuitive that, for cooperationto occur in equilibrium, contributions need to be observed with sufficiently highprobability.2. Others cannot always tell whether a player had an opportunity to contribute. Fordefection to be penalized, it must be the case that others can tell that a player hadthe opportunity to cooperate and did not (i.e. the player should not be able to hidethe fact that there was an opportunity to cooperate).3. Sometimes, there are two ways to cooperate, and one has a higher benefit, b .Then, the only way this more effective type of cooperation can be sustained inequilibrium is if others know which cooperative act is more effective.Morality Games307Fig. 5 The Repeated Prisoner’s Dilemma. Two players play a Prisoner’s Dilemma. They eachobserve the other’s action, then, with probability δ , play another Prisoner’s Dilemma against thesame opponent, etc.Technically, for the second and third point, what is needed is common knowledgethat a player had an opportunity to cooperate or of the more effective means ofcooperation. If observers were to know one purposely chose to defect or chose theless cooperative act, but they do not know that others know this, then observersthink others will think punishment is not warranted, and observers will not punish.The argument is analogous to the discussion of higher-order beliefs in the omission–commissionsubsection and formalized in Dalkiran et al. ( 2012 ) and Hoffmanet al. ( 2015 ).Interpreting the Quirks of AltruismBelow we discuss some of the quirky features of altruism identified by economistsand psychologists. In each case, we will argue that these features might be puzzling,but not when viewed through the lens of the above model:Insensitivity to Effectiveness . We are surprisingly insensitive to the impact of ourcharitable contributions. We vote because we “want to be a part of the democratic system,”or we “want to make a difference,” despite the fact that our likelihood of swingingan election (even in a swing state) is smaller than our likelihood of being struck bylightning (Gelman, Silver, & Edlin, 2012 ). Why is our desire to “make a difference” or“be a part of the system” immune to the actual difference we are making? Our charitablecontributions or volunteer efforts suffer from the same insensitivity. Why doesanyone give money or volunteer time to Habitat for Humanity? The agency flies highearners who have never held a hammer halfway across the world to build houses thatwould be substantially more cheaply built by local experts funded by the high earners.Experimental evidence demonstrates our insensitivity: Experimental subjects are willingto pay the same amount to save 2000, 20,000, or 200,000 birds (Desvousges et al.,2010 ). Likewise, when donors are told their donations will be matched, tripled, orquadrupled, they donated identical amounts (Karlan & List, 2006 ). Why do we give somuch, but do not ensure our gifts have a large impact?The explanation follows directly from the above model: It is often the case thatobservers do not know which acts are effective and which are not and, certainly, this308M. Hoffman et al.usually is not commonly known. Thus, they will not reward or punish based oneffectiveness, and we ourselves will not attend to effectiveness in equilibrium. Thisexplanation suggests that if we want to increase efficacy of giving, we ought tofocus on making sure donors’ friends and colleagues are aware of the efficacy ofdifferent options. In fact, this is perhaps more important than informing the donorof efficacy, since the donor will be motivated to uncover efficacy herself.Magnitude of the Problem . We are surprisingly unaware of and unaffected by themagnitudes of the problems we contribute to solving. How many of those who participatedin the recent ALS Ice Bucket Challenge have even the vaguest sense of thenumber of ALS victims? (Answer: about 1/100th the victims of heart disease.) Howmuch happier would these individuals have been if the number of ALS victims werecut in half? Multiplied by 100? The same questions could be asked about AIDS orcleft lips. If we were actually motivated by our desire to rid the world of such afflictionsas we often proclaim, then we would be happier if there were fewer afflictedindividuals and less happy if there were more. But we are not even aware of thesenumbers, let alone affected by them. This suggests an alternative motivation thanthe one we proclaim.On the other hand, if we give in order to gain social rewards, it does not matterwhether the problem is large or small, provided others recognize it as a problem andthe social norm is to give. If our learned or evolved preferences were drasticallyimpacted by the magnitude of the crises, we would be sensitive to whether the problemwas solved, perhaps motivating us to ensure that others solve it, which wewould not get credit for, or perhaps motivating us to devote too much of ourresources to solving it, beyond what we would actually get rewarded for.Observability . There is overwhelming evidence that people give more when theirgifts are observed. Much of this evidence comes from the lab, where it has beendemonstrated a myriad of ways (e.g., Andreoni & Petrie, 2004 ; Bolton, Katok, &Ockenfels, 2005 ; List, Berrens, Bohara, & Kerkvliet, 2004 ). For instance, whenparticipants play a public goods game in the laboratory for money, their contributionsare higher when they are warned that one subject will have to announce to theroom of other participants how much they contributed (List et al., 2004 ). However,evidence also comes from real-world settings, which find large effects in settings asdiverse as blood donation (Lacetera & Macis, 2010 ), blackout prevention (Yoeli,Hoffman, Rand, & Nowak, 2013 ), and support for national parks (Alpizar, Carlsson,& Johansson-Stenman, 2008 ). In Switzerland, voting rates fell in small communitieswhen voters were given the option to vote by mail (Funk, 2006 ), which makesit harder to tell who did not vote, even though it also makes it easier to vote. In fact,our willingness to give more when observed extends to subtle, subconscious cues ofbeing observed: People give twice as much in dictator games when there are markingson the computer screen that vaguely represent eyes (Haley & Fessler, 2005 ),and they are more likely to pay for bagels in their office when the payment box hasa picture of eyes above it (Bateson, Nettle, & Roberts, 2006 ).These results should not surprise anyone who believes our pro-social tendencies areinfluenced by reputational concerns (though the magnitudes are surprisingly large).Morality Games309The effectiveness of subconscious cues of observability points to a primary role forreputations in our learned or evolved proclivities toward pro-social behavior. Thelarge impact of subtle cues of observability, however, calls into question alternativeexplanations not based on reputations.Explicit Requests . When we are asked directly for donations, we give more than ifwe are not asked, even though no new information is conveyed by the request. In astudy of supermarket shoppers around Christmas time, researchers found that passersbywere more likely to give to the Salvation Army if volunteers not only rangtheir bell but explicitly asked for a donation (Andreoni, Rao, & Trachtman, 2011 ).If our motive is to actually do good, or perhaps proximally to feel good by the act ofgiving, we should not be impacted by an explicit request.However, if we evolved or learned to give in order to gain rewards or avoid punishmentas described above, then we ought to be more likely to give when, if we didnot give, it would be common knowledge that we had the option to give and chosenot to. The explicit request makes the denial common knowledge.It is worth emphasizing that our evolved intuition to respond to explicit asks maybe (mis)applied to individual settings that lack social rewards. Imagine you areapproached by a Salvation Army volunteer in front of a store in a city where you arevisiting for one day only. A literal reading of the model would suggest that youshould be no more likely to respond to an explicit request. But it is more realistic toexpect that if your pro-social preferences were learned or evolved in repeated interactionsthen applied to this new setting, you would respond in a way that is notoptimal for this particular setting and nonetheless give more when explicitly asked(just as our preferences for sweet and fatty foods, which evolved in an environmentwhere food was scarce, lead us to overeat now that food is abundant).Avoiding Situations in Which We Are Expected to Give . In the same supermarketstudy, researchers discovered that shoppers were going out of their way to exit thestore through a side door, to avoid being asked for a contribution by the SalvationArmy volunteers. In another field experiment, those who were warned in advancethat a solicitor would come to the door asking for charitable donations were morelikely to not be home. The researchers estimated that among those who gave, 50 %would have avoided being home if warned in advance of the solicitor’s time ofarrival (DellaVigna et al., 2012 ). In a laboratory analog, subjects who would haveotherwise given money in a $10 dictator game were willing to pay a dollar to keepthe remaining nine dollars and prevent the recipient from knowing that a dictatorgame could have been played (Dana et al. 2006 ). If our motive were to have animpact, we would not pay to avoid putting ourselves in a situation where we couldhave such an impact. Likewise, if our motive were to feel good by giving, we wouldnot pay to avoid this feeling.In contrast, if we evolved or learned to give in order to gain rewards or avoidpunishment, then we would pay to avoid situations where we are expected to give.Again, this would be true even if, in this particular setting, we were unlikely to actuallybe punished.310M. Hoffman et al.Norms . People are typically conditionally cooperative , meaning that they are willingto cooperate more when they believe others contribute more. For example, studentsasked to donate to a university charity gave 2.3 percentage points more whentold that others had given at a rate of 64 % than when they were told giving rateswere 46 % (Frey & Meier, 2004 ). Hotel patrons were 26 % more likely to reuse theirtowels when informed most others had done the same (Goldstein, Cialdini, &Griskevicius, 2008 ). Households have been shown to meaningfully reduce electricityconsumption when told neighbors are consuming less, both in the United States(Ayres, Raseman, & Shih, 2012 ) and in India (Sudarshan, 2014 ).Such conditional cooperation is easily explained by the game theory model:When others give, one can infer that one is expected to give and may be sociallysanctioned if one does not.Strategic Ignorance . Those at high risk of contracting a sexually transmitted disease(STD) often go untested, presumably because if they knew they had the STD,they would feel morally obliged to refrain from otherwise desirable activity thatrisks spreading the STD. Why is it more reproachable to knowingly put a sexualpartner at risk when one knows one has the STD than to knowingly put a sexualpartner at risk by not getting tested? There is evidence that we sometimes pursuestrategic ignorance and avoid information about the negative consequences of ourdecisions to others. When subjects are shown two options, one that is better forthemselves but worse for their partners and one that is worse for themselves but betterfor their partners, many choose the option that is better for their partners. But,when subjects must first press a button (at no cost) to reveal which option is betterfor their partners, they choose to remain ignorant and simply select the option thatis best for themselves (Dana, Weber, & Kuang, 2007 ).This quirk of our moral system is again easy to explain with the above model.Typically, information about how one’s actions affect others is hard to obtain, sopeople cannot be blamed for not having such information. When one can get suchinformation easily, others may not know that it is easy to obtain and will not punishanyone who does not have the information. For example, although it is trivially easyto look up charities’ financial ratings on websites like charitynavigator.org, fewpeople know this and could negatively judge those that donate without first checkingsuch websites. And even when others know that one can get this informationeasily, they might suspect that others do not know this, and so avoid punishing,since others won’t expect punishment. To summarize, strategic ignorance preventscommon knowledge of a violation and so is likely to go unpunished. We againemphasize that we will be lenient of strategic ignorance, even when punishment isnot literally an option.Norm of Reciprocity . We feel compelled to reciprocate favors, even if we knowthat the favors were done merely to elicit reciprocation and even if the favor askedin return is larger than the initial one granted (Cialdini 2001). For instance, membersof Hare Krishna successfully collect donations by handing out flowers to disembarkingpassengers at airports, even though passengers want nothing to do withthe flowers: They walk just a few feet before discarding them in the nearest bin.Morality Games311Psychologists and economists sometimes take this “norm” as given, without askingwhere it comes from, and a naive reading of Trivers would lead one to think that weshould be sensitive to the magnitude of the initial favor and whether it ismanipulative.However, according to the above model, reciprocity is the Nash equilibrium,even if the favors are not evenly matched or manipulative, since, in equilibrium, weare neither sensitive to such quantitative distinctions nor to whether the initial reciprocitywas manipulative, unless these facts are commonly known.Self-Image Concerns . People sometimes play mental tricks in order to appear tothemselves as pro-social. For example, in an experiment, subjects will voluntarily takeon a boring task to save another subject from doing it, but if given the option of privatelyflipping a coin to determine who gets the task, they often flip—and flip, and flipagain—until the “coin” assigns the task to the other subject (Batson, Kobrynowicz,Dinnerstein, Kampf, & Wilson, 1997 ). Why would we be able to fool ourselves andnot, say, recognize that we are gaming the coin flip? Why do we care what we thinkof ourselves at all? Are there any constraints on how we will deceive ourselves?Such self-image considerations can be explained by noting that our self-image canact as a simple proxy, albeit an imperfect one, for what others think of us, and also thatwe are more convincing to others when we believe something ourselves (Kurzban,2012 ; Trivers, 2011 ). This explanation suggests that the ways we deceive ourselvescorrespond to quirks described throughout this section—for example, we will absolveourselves of remaining strategically ignorant even when it is easy not to, or be convincedthat we have done good by voting, even if we cannot swing an election.Framing Effects . Whether we contribute is highly dependent on the details of theexperiment, such as the choice set (List, 2007 ) and the labels for the differentchoices (Ross & Ward, 1996 ; Roth, 1995 ). Such findings are often taken as evidencethat social preferences cannot be properly measured in the lab (Levitt & List, 2007 ).We believe a more fruitful interpretation is simply that the frame influenceswhether the laboratory experiment “turns on” our pro-social preferences, perhapsby simulating a situation where cooperation is expected (Levitt & List, 2007 ).One-Shot Anonymous Giving : We give in anonymous, one-shot settings, such asdictator games. We also sacrifice for others in the real world when there is no chanceof reciprocation: Heroes jump on grenades to save their fellow soldiers or block thedoor to a classroom with their bodies to prevent a school shooter from entering(Rand & Epstein, 2014 ). This is often seen as evidence for a role of group selection(Fehr & Fischbacher, 2003 ).However, an alternate explanation is that we do not consider the likelihood ofreciprocation (Hoffman et al., 2015 ), as described above. To explain the laboratoryevidence, there are two more possibilities. First, subjects may believe there is somechance their identity will be revealed and feel the costs of being revealed as selfishare greater than the gains from the experiment (Delton, Krasnow, Cosmides, &Tooby, 2011 ). Second, we again emphasize that learned or evolved preferences andideologies are expected to be applied even in novel settings to which they are notoptimized.312M. Hoffman et al.ConclusionIn this chapter we have showed that a single approach–game theory , with the helpof evolution and learning –can explain many of our moral intuitions and ideologies.We now discuss two implications .Group Selection . Our chapter relates to the debate on group selection, wherebygroup level competition and reproduction is supposed to occasionally cause individualsto evolve to sacrifice their own payoffs to benefit the group (e.g., Wilson,2006 ). One of the primary pieces of evidence cited in support of group selection isthe existence of human cooperation and morality (Fehr & Fischbacher, 2003 ; Fehr,Fischbacher, & Gächter, 2002 ; Gintis, Bowles, Boyd, & Fehr, 2003 ; Haidt, 2012 ;Wilson, 2010 , 2012 ), in particular: giving in one-shot anonymous laboratory experiments,intuitively sacrificing one’s life for the group (jumping on the grenade), andcontributions to public goods or charity. However, we have reviewed an alternativeexplanation for these phenomena that does not rest on group selection. It also yieldspredictions about these phenomena that group selection does not, such as that peopleare more likely to cooperate when they are being observed and there is variancein the cost of cooperation. The approach described here also explains other phenomena,such as categorical norms and ineffective altruism. These lead to social welfarelosses, which is suboptimal from the group’s perspective. The categorical normagainst murder, for example, leads to enormous waste when keeping alive, sometimesfor years, those who have virtually no chance of a future productive life.Admittedly, despite their inefficiencies, these moral intuitions do not rule outgroup selection, since group selection can be weak relative to individual selection.But it does provide a powerful argument that group selection is unnecessary forexplaining many interesting aspects of human morality. It also suggests that groupselection is, indeed, at most, weak. One example that makes this especially clear isdiscrete norms. Recall that we argued that continuous norms are not sustainablebecause individuals benefit by deviating around the threshold. Notice that this benefitis small, since the likelihood that signals are right around the threshold is low.Group selection could easily overwhelm the benefit one would get from deviatingfrom this Nash equilibrium, suggesting group selection is weak (i.e. there are fewgroup- level reproductive events, high migration rates, high rates of “mutation” inthe form of experimentation among individuals, etc.).Logical Justification of Moral Intuitions . In each of the applications above, weexplained moral intuitions without referring to existing a priori logical justificationsby philosophers or others. Our explanation for our sense of rights does not rely onLocke’s “state of nature.” No argument we gave rests on God as an orderly designer,on Platonic ideals, on Kant’s concepts of autonomy and humanity, etc. What doesthis mean for these a priori justifications? It suggests that they are not the source ofour morality and are, instead, post hoc justifications of our intuitions (Haidt, 2012 ).To see what we mean, consider the following analogy. One might wonder whywe find paintings and sculptures of voluptuous women beautiful. Before theMorality Games313development of sexual selection theory, one might have argued that perfect spheresare some kind of Platonic solid, and inherently desirable, or that curvy hips yieldgolden ratios. But with our current understanding of sexual selection, we recognizethat our sense of beauty has evolved and that there is no platonic sense of beautyoutside of that shaped by sexual selection. Any argument about perfect spheres isunparsimonious and likely flawed. Without the help of evolution and game theory,did philosophers conjure the moral equivalents of perfect spheres and golden ratios?The state of nature, the orderly designer, Platonic ideals, autonomy, and humanity,etc.—perhaps these arguments are also unfounded and unnecessary.ReferencesAlpizar, F., Carlsson, F., & Johansson-Stenman, O. (2008). Anonymity, reciprocity, and conformity:Evidence from voluntary contributions to a national park in Costa Rica. Journal of PublicEconomics, 92 (5), 1047–1060.Andreoni, J., & Petrie, R. (2004). Public goods experiments without confidentiality: A glimpseinto fund-raising. Journal of Public Economics, 88 (7), 1605–1623.Andreoni, J. (1990). Impure altruism and donations to public goods: A theory of warm-glow giving.The Economic Journal, 100 , 464–477.Andreoni, J., Rao, J. M., & Trachtman, H. (2011). Avoiding the ask: A fi eld experiment on altruism,empathy, and charitable giving . Technical report, National Bureau of Economic Research.Axelrod, R. M. (1984). The evolution of cooperation. New York: Basic Books.Axelrod, R., & Hamilton, W. D. (1981). The evolution of cooperation. Science, 211 (4489),1390–1396.Ayres, I., Raseman, S., & Shih, A. (2012). Evidence from two large field experiments that peercomparison feedback can reduce residential energy usage. Journal of Law, Economics, andOrganization , 2–20.Bateson, M., Nettle, D., & Roberts, G. (2006). Cues of being watched enhance cooperation in areal-world setting. Biology Letters, 2 (3), 412–414.Batson, C. D. (2014). The altruism question: Toward a social-psychological answer . Hillside, NJ:Psychology Press.Batson, C. D., Kobrynowicz, D., Dinnerstein, J. L., Kampf, H. C., & Wilson, A. D. (1997). In avery different voice: Unmasking moral hypocrisy. Journal of Personality and Social Psychology,72 (6), 1335.Bolton, G. E., Katok, E., & Ockenfels, A. (2005). Cooperation among strangers with limited informationabout reputation. Journal of Public Economics, 89 (8), 1457–1468.Cain, D., Dana, J., & Newman, G. (2014). Giving vs. giving in. Yale University Working Paper.Camerer, C. (2003). Behavioral game theory: Experiments in strategic interaction . Princeton, NJ:Princeton University Press.Chammah, A. M., & Rapoport, A. (1965). Prisoner’s dilemma; a study in confl ict and cooperation. Ann Arbor, MI: University of Michigan.Chwe, M. (2013). Rational ritual: Culture, coordination, and common knowledge. Princeton, NJ:Princeton University Press.Cone v. West Virginia Pulp & Paper Co. , 330 U.S. 212, 67 S. Ct. 752, 91 L. Ed. 849 (1947).Critcher, C. R., Inbar, Y., & Pizarro, D. A. (2013). How quick decisions illuminate moral character.Social Psychological and Personality Science, 4 (3), 308–315.Cushman, F., Young, L., & Hauser, M. (2006). The role of conscious reasoning and intuition inmoral judgment testing three principles of harm. Psychological Science, 17 (12), 1082–1089.314M. Hoffman et al.Dalkiran, N. A., Hoffman, M., Paturi, R., Ricketts, D., & Vattani, A. (2012). Common knowledgeand state-dependent equilibria. In Algorithmic game theory (pp. 84–95). New York: Springer.Dana, J., Cain, D. M., & Dawes, R. M. (2006). What you don’t know won’t hurt me: Costly (butquiet) exit in dictator games. Organizational Behavior and Human Decision Processes, 100 (2),193–201.Dana, J., Weber, R. A., & Kuang, J. X. (2007). Exploiting moral wiggle room: Experimentsdemonstrating an illusory preference for fairness. Economic Theory, 33 (1), 67–80.Davies, N. B. (1978). Territorial defense in the speckled wood butterfly (pararge aegeria): The residentalways wins. Animal Behaviour, 26 , 138–147.Dawkins, R. (2006). The selfi sh gene . Oxford, UK: Oxford University Press.DellaVigna, S., List, J. A., & Malmendier, U. (2012). Testing for altruism and social pressure incharitable giving. The Quarterly Journal of Economics, 127 (1), 1–56.Delton, A. W., Krasnow, M. M., Cosmides, L., & Tooby, J. (2011). Evolution of direct reciprocityunder uncertainty can explain human generosity in one-shot encounters. Proceedings of theNational Academy of Sciences, 108 (32), 13335–13340.DeScioli, P., Bruening, R., & Kurzban, R. (2011). The omission effect in moral cognition: Towarda functional explanation. Evolution and Human Behavior, 32 (3), 204–215.DeScioli, P., Gilbert, S. S., & Kurzban, R. (2012). Indelible victims and persistent punishers inmoral cognition. Psychological Inquiry, 23 (2), 143–149.DeScioli, P., & Wilson, B. J. (2011). The territorial foundations of human property. Evolution andHuman Behavior, 32 (5), 297–304.DeScioli, P., & Kurzban, R. (2009). Mysteries of morality. Cognition, 112 (2), 281–299.Descioli, P., & Karpoff, R. (2014). People’s judgments about classic property law cases . BrandeisUniversity Working Paper.Desvousges, W. H., Johnson, F. R., Dunford, R. W., Boyle, K. J., Hudson, S. P., Wilson, K. N.,et al. (2010). Measuring nonuse damages using contingent valuation: An experimental evaluationof accuracy . Research Triangle Park, NC: RTI Press.Epley, N., Keysar, B., Van Boven, L., & Gilovich, T. (2004). Perspective taking as egocentricanchoring and adjustment. Journal of Personality and Social Psychology, 87 (3), 327.Fehr, E., & Fischbacher, U. (2003). The nature of human altruism. Nature, 425 (6960), 785–791.Fehr, E., Fischbacher, U., & Gächter, S. (2002). Strong reciprocity, human cooperation, and theenforcement of social norms. Human Nature, 13 (1), 1–25.Fisher, R. A. (1958). The genetic theory of natural selection . Mineola, NY : Dover.Frank, R. H. (1988). Passions within reason: The strategic role of the emotions. New York: WWNorton & Co.Frey, B. S., & Meier, S. (2004). Social comparisons and pro-social behavior: Testing “conditionalcooperation” in a field experiment. American Economic Review, 94 , 1717–1722.Funk, P. (2006). Modern voting tools, social incentives and voter turnout: Theory and evidence.In Annual Meeting of the American Economic Association , Chicago, IL.Gelman, A., Silver, N., & Edlin, A. (2012). What is the probability your vote will make a difference?Economic Inquiry, 50 (2), 321–326.Gintis, H. (2007). The evolution of private property. Journal of Economic Behavior & Organization,64 (1), 1–16.Gintis, H., Bowles, S., Boyd, R., & Fehr, E. (2003). Explaining altruistic behavior in humans.Evolution and Human Behavior, 24 (3), 153–172.Goldstein, N. J., Cialdini, R. B., & Griskevicius, V. (2008). A room with a view-point: Using socialnorms to motivate environmental conservation in hotels. Journal of Consumer Research, 35 (3),472–482.Greene, J. D., Cushman, F. A., Stewart, L. E., Lowenberg, K., Nystrom, L. E., & Cohen, J. D.(2009). Pushing moral buttons: The interaction between personal force and intention in moraljudgment. Cognition, 111 (3), 364–371.Haidt, J. (2012). The righteous mind: Why good people are divided by politics and religion .New York: Vintage.Haley, K. J., & Fessler, D. M. (2005). Nobody’s watching? Subtle cues affect generosity in ananonymous economic game. Evolution and Human Behavior, 26 .Morality Games315Hedden, T., & Zhang, J. (2002). What do you think I think you think? Strategic reasoning in matrixgames. Cognition, 85 (1), 1–36.Hoffman, M., Yoeli, E., & Nowak, M. A. (2015). Cooperate without looking: Why we care whatpeople think and not just what they do. Proceedings of the National Academy of Sciences,112 (6), 1727–1732.Hoffman, M., Yoeli, E., & Dalkiran, A. (2015). Social applications of common knowledge . HarvardUniversity Working Paper.Kahneman, D., Knetsch, J. L., & Thaler, R. H. (1990). Experimental tests of the endowment effectand the Coase theorem. Journal of Political Economy, 98 , 1325–1348.Kant, I. (1997/1787). Groundwork of the metaphysics of morals (1785). Practical Philosophy,108 .Karlan, D., & List, J. A. (2006). Does price matter in charitable giving? Evidence from a largescalenatural fi eld experiment . Technical report, National Bureau of Economic Research.Kurzban, R. (2012). Why everyone (else) is a hypocrite: Evolution and the modular mind.Princeton, NJ: Princeton University Press.Lacetera, N., & Macis, M. (2010). Social image concerns and prosocial behavior: Field evidencefrom a nonlinear incentive scheme. Journal of Economic Behavior & Organization, 76 (2),225–237.Levitt, S. D., & List, J. A. (2007). What do laboratory experiments measuring social preferencesreveal about the real world? The Journal of Economic Perspectives, 21 , 153–174.List, J. A. (2007). On the interpretation of giving in dictator games. Journal of Political Economy,115 (3), 482–493.List, J. A., Berrens, R. P., Bohara, A. K., & Kerkvliet, J. (2004). Examining the role of socialisolation on stated preferences. American Economic Review, 94 , 741–752.Locke, J. (1988/1688). Locke: Two treatises of government student edition . Cambridge, UK:Cambridge University Press.Mankiw, N. G. (2007). Principles of economics . Thomson Learning.Milgrom, P., & Weber, R. J. (1982). The value of information in a sealed-bid auction. Journal ofMathematical Economics, 10 (1), 105–114.Morris, S., & Shin, H. S. (2002). Social value of public information. The American EconomicReview, 92 (5), 1521–1534.Myerson, R. B. (2004). Justice, institutions, and multiple equilibria. Chicago Journal ofInternational Law, 5 , 91.Neuwirth, R. (2005). Shadow cities: A billion squatters, a new urban world . New York: Routledge.Nowak, M. A. (2006). Evolutionary dynamics: Exploring the equations of life . Cambridge, MA:Harvard University Press.Panchanathan, K., & Boyd, R. (2004). Indirect reciprocity can stabilize cooperation without thesecond-order free rider problem. Nature, 432 (7016), 499–502.Pinker, S. (2003). The blank slate: The modern denial of human nature . UK: Penguin.Pinker, S., Nowak, M. A., & Lee, J. J. (2008). The logic of indirect speech. Proceedings of theNational Academy of Sciences, 105 (3), 833–838.Rand, D. G., & Epstein, Z. G. (2014). Risking your life without a second thought: Intuitivedecision- making and extreme altruism . Available at SSRN 2424036.Ross, L., & Ward, A. (1996). Naive realism: Implications for social conflict and misunderstanding.In T. Brown, E. Reed, & E. Turiel (Eds.), Values and knowledge (pp. 103–135).Roth, A. E. (1995). Bargaining experiments. In J. H. Kagel & E. R. Alvin (Eds.), The handbook ofexperimental economics (pp. 253–342). Princeton, NJ: Princeton University Press.Sigg, H., & Falett, J. (1985). Experiments on respect of possession and property in hamadryasbaboons ( Papio hamadryas ). Animal Behaviour, 33 (3), 978–984.Smith, J. M., & Price, G. (1973). The logic of animal conflict. Nature, 246 , 15.Sosis, R., & Alcorta, C. (2003). Signaling, solidarity, and the sacred: The evolution of religiousbehavior. Evolutionary Anthropology: Issues, News, and Reviews, 12 (6), 264–274.Spranca, M., Minsk, E., & Baron, J. (1991). Omission and commission in judgment and choice.Journal of Experimental Social Psychology, 27 (1), 76–105.316M. Hoffman et al.Sudarshan, A. (2014). Nudges in the marketplace: Using peer comparisons and incentives to reducehousehold electricity consumption . Technical report, Harvard University Working Paper.Thaler, R. (1980). Toward a positive theory of consumer choice. Journal of Economic Behavior &Organization, 1 (1), 39–60.Thomson, J. J. (1976). Killing, letting die, and the trolley problem. The Monist, 59 (2), 204–217.Tirole, J. (1988). The theory of industrial organization . Cambridge, MA: MIT Press.Trivers, R. (2011). The folly of fools: The logic of deceit and self-deception in human life.New York: Basic Books.Trivers, R. L. (1971). The evolution of reciprocal altruism. Quarterly Review of Biology, 46 ,35–57.Trivers, R. L. (1974). Parent-offspring conflict. American Zoologist, 14 (1), 249–264.Wilkinson, G. S. (1984). Reciprocal food sharing in the vampire bat. Nature, 308 (5955), 181–184.Wilson, D. S. (2010). Darwin’s cathedral: Evolution, religion, and the nature of society . Chicago:University of Chicago Press.Wilson, E. O. (2012). The social conquest of earth . New York: WW Norton & Company.Wilson, D. S. (2006). Human groups as adaptive units: Toward a permanent consensus. InP. Carruthers, S. Laurence, & S. Stich (Eds.), The innate mind: Culture and cognition . Oxford,UK: Oxford University Press.Winter, E. (2014). Feeling smart: Why our emotions are more rational than we think. Public Books.Wroughton, L. (2013). As Syria War Escalates, Americans Cool to U.S. Intervention: Reuter/IpsosPoll. Reuters , August 24, 2013. Accessed at http://www.reuters.com/article/2013/08/25/us-syria- crisis-usa-poll-idUSBRE97O00E20130825 on 12 March 2015.Yoeli, E., Hoffman, M., Rand, D. G., & Nowak, M. A. (2013). Powering up with indirect reciprocityin a large-scale field experiment. Proceedings of the National Academy of Sciences,110 (Suppl. 2), 10424–10429.Zahavi, A. (1975). Mate selection: A selection for a handicap. Journal of Theoretical Biology,53 (1), 205–214.