1Department of Ecology and Evolutionary Biology, University of California, 621 Young Drive South, Los Angeles, CA 90095-1606, USA. 2CNRS, Institut Jean Nicod-Ecole Normale Supérieure, 29 rue d’Ulm, 75005 Paris, France. 3National Security Affairs Department, Naval Postgraduate School, 1411 Cunningham Rd., Monterey, CA 93943, USA. 4Institut des Sciences de l’Evolution, Université Montpellier II, CNRS, Place Eugéne Bataillon, 34095 Montpellier, France. 5Politics and International Relations, University of Edinburgh, 15a George Square, Edinburgh, EH6 6LG, UK. 6Institute of the Environment, University of Arizona, Tucson, Arizona 85716, USA. 7Department of Anthropology, University of Connecticut, Storrs, Connecticut 06269-2176, USA. 8Department of Political Science, Baylor University, One Bear Place #97276, Waco, Texas 76798, USA. In addition to their primary affiliations, the authors are members of the Natural Security Working Group (www.naturalsecurity.arizona.edu) that has been supported by the National Center for Ecological Analysis and Synthesis, and the Office of Naval Research-Global.

Address correspondence to: D.T. Blumstein, email: marmots@ucla.edu

Knowing how to send and interpret signals is an essential part of both diplomacy and war. Political scientists have recognized that costly signals – gestures and actions that involve significant cost or risk – are central to politics and diplomacy since modeling doyen James Fearon built his Ph.D. thesis around the concept in the 1990s. Because these signaling systems are pervasive in nature (many of these strategies arise independently and repeatedly to solve common problems suggesting evolutionary pressure to select strategies offering the most success at the least cost), their underlying strategic logic has important implications to foreign policy challenges we face today. By capitalizing on solutions derived by evolution over 3.5 billion years of life on Earth, we may identify ideas that otherwise might not have been explored in a policy context potentially offering quick, novel, and effective options to increase strategic and combat effectiveness. Here we present 8 lessons from evolution for political science.

Lesson 1. Honest signals will be costly
The power of costly signaling in the animal world is captured in the famous example of the peacock’s tail. A series of studies have demonstrated that females select males with the longest and most elaborate tails. The benefits of such a selection criterion are clear: males that are able to allocate sufficient resources to grow this long tail, make it colorful, and keep it clean and healthy looking, survive despite their handicap. Males that can do this have energy to burn; they radiate their quality. The tail is thus an honest advertisement of the skills of acquiring food, avoiding predators, and controlling parasites and pathogens. Females choosing these showy males will ensure high quality genes being transmitted to their offspring.
Indeed, from this and many other studies of signaling and communication in a diversity of mammals, birds, fishes, and insects, we can conclude that a common feature of animal communication is that costly signaling is valued by receivers, not just the sender, and in a wide range of settings. This biological rule has important implications for several aspects of foreign policy including making positive gestures during negotiations.

An effective negotiator must communicate honesty to the receiver, thereby inducing trust. Negotiations proceed with repeated positive feedback: the receiver needs to honestly signal trust back to the negotiator. Complicating this interaction is the cross-cultural nature of diplomacy that may result in the misinterpretation of honest signals. Positive gestures are one way of building trust. To be effective in the long term, positive gestures must be costly, yet many positive gestures in foreign policy are ritualized into protocol whereby there are formal and invariant rules by which states, and their emissaries, interact. If they are expected as routine, they are unlikely to be valued—seen as a result of the situation rather than any cooperative disposition of the actor.#

Lesson 2: Ritualized signals may have little value, but may be used strategically

In animal systems, ritualized signals contain little information and do not vary much from individual to individual; consider the first, stereotypical behaviors in a courtship sequence. To avoid confusion, such displays are often ritualized. This lack of variation means that there is less of an opportunity for a receiver to associate a display with its underlying cost. And, while ritualized gestures might be commonplace ways of building trust – consider shaking hands, smiling, engaging in small talk – these displays may not be as effective as a genuinely costly display. This is not to entirely discount the importance of these displays, because effective cooperation between individuals is costly in that it takes time to develop and requires individuals to evaluate each other’s reputations that are built over time. However, for now, let’s consider isolated responses.

Systematically reducing the value of a signal to a receiver may be used strategically. The Egyptians capitalized on what became to Israelis a ritualized signal—maneuvers on the Israeli border in the months preceding the Yom Kippur War. Before attacking Israel, Egypt ran 40 military exercises on Israel’s borders#. This led Israel to discount the threat of a troop buildup on their border and enhanced its vulnerability.

However, there are also costs to ritualization. The recently scrapped Homeland Security Threat Level remained unchanged for years before being eliminated. What message did this invariant ritualized message send to travelers—or to prospective terrorists? Indeed, one lesson from nature is that one must consider the nature of the recipient in order to properly design a signal. And, in some cases, such ritualized signals may become meaningless and counterproductive.

Lesson 3: Unexpected signals may be more effective

Following a natural disaster, nations offer assistance to other countries. An offer of one million dollars in aid from a poor country is a much more meaningful contribution than the same offer from a wealthy one. And, individual citizens lining up to spontaneously help others (as often occurs following natural disasters) are truly meaningful gestures. For instance, following devastating 1999 and 2011 earthquakes in Turkey, members of the Israeli public spontaneously and immediately organized to collect food, clothing and other emergency necessities for Turkish citizens. While many governments formally responded, including the Israeli government, such responses are difficult to interpret since they are routine and likely to be strategically motivated. The spontaneity of the Israeli public response however appeared to be a sincere offer of help. The key insight is that it is not the absolute value, but rather the value relative to ability, and the sincerity of the donation that is likely to define a trustworthy display. Humans suffer from the so-called “correspondence bias”, which makes us more likely to assume the behavior of other actors is a result of their fundamental character, whereas our own behavior is a result of reacting to the situation—especially if the act impacts on us negatively. However, when the act impacts on us positively, we are more likely to assume the actor was motivated by situational constraints.

Lesson 4: Threats should be costly

Conversely, natural systems show us that negative gestures such as threats may also have to be costly to be effective. A striking lesson from evolution is that adversaries should organize their threats into a gradually escalating sequence, resorting to all-out fighting only if the less costly, earlier signals fail to induce their opponent to back down. Red deer competing for mates strut threateningly side-by-side, then bellow at each other, and only then lock antlers if one individual does not back away. The logic is impeccable: if the adversaries are badly mismatched, they will realize it during the first phase and back down quickly, saving both from wasting further time and energy. If the payoff for winning is not great, individuals should not escalate. But, if the payoff is great, more subtle differences will be detected at the second stage, again a mutually beneficial outcome. Truly dangerous fights will only occur when both have proved themselves to be so evenly matched that signaling alone cannot distinguish between them. Over millions of years, natural selection has crafted a finely-tuned “playbook” of signaling and escalation for species’ to work from as they attempt to resolve their conflicts of interest without getting killed in the process.

Humans face similar problems, and bargaining theories of war have investigated similar problems of incremental signals. However, evolution offers a useful new perspective on this because we face types of conflict that our ancestors never encountered, and thus to which our responses have not been molded by selection. In a world of cyber-warfare, weapons that kill at a distance, and remote command and control far from the battlefield, our evolved signaling mechanisms neither convey messages to the enemy nor bring us direct feedback (e.g., drone pilots fight thousands of miles from the battlefield, the sources of computer worms like Stuxnet are opaque and difficult to trace). This means we may expect an evolutionary “mismatch” between our behavior and our environment and may lead to unintended and un-checked escalation. Rival states may exchange costly signals prior to launching into war, for example military parades that display cutting edge technology, power projection through naval port calls, elevating level of alert status for forces as well as funding levels for security and defensive activities. However, different strategies and their deployment between adversaries in a crisis may lead to confusion and potentially catastrophic outcomes. Considering the stakes in incidents such as the nuclear alerts ordered by President Kennedy in the Cuban Missile Crisis, by Nixon during the Yom Kippur War of 1973, or North Korea’s response to accusations that it torpedoed and sank a South Korean warship in March 2010 – the question becomes a very urgent one.

Evolutionary thinking provides guidelines for formulating an appropriate policy response, particularly when it is unclear whether adversaries are following the same rules. Just as in conflicts in natural systems, honest signaling of intention in a series of reciprocal steps is the best way to obtain a peaceful resolution. And, developing a reputation for honest signaling is a powerful force in obtaining trust. In the absence of a reputation for honest signaling, it is difficult to know whether to escalate up or down in negotiations.

Lesson 5: State apologies should be costly

To be successful apologies must be costly. The cost of an apology is the political risk to the person or administration apologizing, and broader costs to pride and well-being within an apologizing society. This may explain why meaningful apologies are not very common, and most apologies happen long after the incident that stimulated it. Apologizing, after all, is a political calculation. Waiting until there is no political cost because opponents have died or moved on to some other issue lessens the value of an apology. In 1998, President Clinton apologized for his administration’s inaction during the Rwandan genocide four years earlier—an unusual and costly act for a President still in office. Israel and Germany have good diplomatic relations and there is also friendship among the populations (many Germans travel to Israel) and this is remarkable given the devastation in the living memory of so many Jews and Israelis. At least three factors were probably important: 1) public apologies which were followed up by real costs such as 2) reparation payments, and 3) a schooling system that makes German children more educated about the Holocaust than probably any other children outside Israel in the world. Nelson Mandela probably recognized the huge value of costly apologies when he set up the reconciliation commissions in South Africa after the fall of the apartheid. Rather than sending all whites involved in the apartheid state to jail, he created a forum for them to apologize face to face with their victims and wipe the slate clean.

Insincere apologies may be particularly costly. Then Secretary of Defense Donald Rumsfeld’s apology about the Abu Ghraib prisoner abuse backfired when he tried to also explain that there was Arab mis-understanding of American culture which could not condone such behavior. This apology was widely regarded as a flop. Japan’s apologies for atrocities in China during World War II remain unvalued by the Chinese because of Japanese dignitaries’ visiting the Yasukuni Shrine to Japanese war dead, which includes the remains of convicted war criminals.

Lesson 6: Humans (and animals) are not necessarily rational: symbolic concessions may be useful

Studies of animals (and humans) also show us that in certain circumstances, individuals do not make economically rational decisions. There is healthy debate about whether these are indeed costly mistakes or evolved strategies. However, there are some lessons from this observation for security and policy.

One of these is that low cost actions may sometimes have a high value to recipients of these actions and, as Jeremy Ginges and colleagues found, the economically rational offers are not necessarily what people will accept#. Indeed, people’s core values must be recognized, often by symbolic concessions. In their research asking both Palestinians and Israelis about what sort of incentives might help move a peace process forward, they found that financial incentives were viewed very negatively, but symbolic concessions, such as apologies for past actions and removing anti-Semitic material from textbooks, went far both with leaders and citizens. Indeed, offers of reparation without the necessary symbolic concessions were viewed negatively. Here again, Nelson Mandela recognized the importance of symbolic concessions when he publicly supported the Springbok rugby team, even though, initially, his black supporters wanted to disband this symbol of apartheid. Understanding that economically irrational behavior is common and that symbolic concessions have great value may be essential in negotiating through seemingly intractable political quagmires (e.g., such as those in North Korea, Iran, and, as Atran and Ginges# write—Israel/Palestine).

Lesson 7: Weaker parties will advertise the strength of their conviction

Obviously not all diplomatic interactions and conflicts are between state actors. Are there unique insights for interactions with non-state actors? A characteristic of a state interacting with a non-state actor is power asymmetry.
Consider the rapid spread of suicide terrorism among weak and disenfranchised organizations fighting a stronger state. With the exception of Kamikaze pilots in WWII, we have not seen two strong opponents in the 20th century use suicide attacks as a military strategy. Indeed, one could argue that the Japanese only resorted to Kamikaze pilots once they realized they were losing because of a loss of pilots, fuel, and weapons. One interpretation of this is that being able to marshal legions of self-sacrificial volunteers creates an honest indicator of the amount of displeasure and the intention to continue fighting that the weaker party faces. When non-state actors fight against democratic state actors, the public opinion is as much a target as the military and the weaker party often wins by turning public opinion against ongoing conflict. Thus, in such asymmetrical combat, such as Chechens against the Russians, Hamas against Israel, or the LTTE against the Sri Lankan government, we should expect the weaker party to work hard to impress upon the stronger party the strength of their conviction. Suicide terrorism may be such a potent signal because it is an unbluffable signal of commitment.

Political scientists have recently recognized this, and have characterized five distinct ways in which terrorists use costly signals to advance their agenda: attrition (wearing down the enemy), intimidation (of opponents in their own population), provocation (of the enemy to violence and collateral damage that consolidates resistance against it), spoiling (of a peace process), and outbidding (of domestic rival parties)#. Moreover, natural systems show us that self-sacrifice is often associated with high relatedness. Social insect workers will die to help their highly-related group persist and the same logic might explain situations in which tightly knit groups, bound by kinship and religion, are willing to make the ultimate sacrifice. Holding this lens over the problem of terrorism clarifies the strategic logic that should guide a policy response.

Lesson 8: Signals are often species specific: it’s essential to know your audience

Species differ in their signals that are often directed to conspecifics. Indeed, divergent communication signals are used to ‘isolate’ species from each other and prevent costly, but mis-guided fights, and potentially wasteful reproductive attempts with the wrong species. The final insight comes from the widespread evolution of species-specific signals: it’s essential to know your audience in order to communicate effectively to it and the same signal may mean very different things to different audiences.

Consider how two populations may have difficulties communicating. The United States botched the messaging to the Muslim world after killing Osama bin Laden. To the audience in the United States there was a low cost propaganda release of video of bin Laden sitting on the floor watching a small TV aimed at demonstrating what a miserable and pathetic fellow the man was. But to would-be jihadis, the actual effect was the opposite. They saw a leader living in humble conditions—a positive image under Muslim law. It was a low cost, culturally blind signal sent by the United States that was filtered and amplified though Islamic culture and tradition into a demonstration of the ultimate high-cost sacrifice of the man for his people.

The lesson is clear: be sure you know who your audience is before signaling and realize that the same message can be interpreted quite differently by different audiences. A low-cost signal to one audience could illustrate high-cost behavior to another audience.

Conclusions

While humans, like every other organism, have a unique evolutionary history, the rules of evolution and natural selection act on us all. More work is needed to disentangle the logic of individual versus group selected mechanisms on policy, but identifying these rules will inevitably create more insights about successful strategic behavior. Importantly, nature’s rules are all around us just waiting to be discovered and explored to see whether they have modern-day applications.

References
Atran, Scott, and Jeremy Ginges. 2009. How words could end war. New York Times 24 January:WK12 (New York Edition).
Fearon, James D. 1995. Rationalist explanations for war. International Organization 49(3):379-414.
Ginges, Jeremy, Atran, Scott, Medin, Douglas, and Khalil Shikaki. 2007. Sacred bounds on rational resolution of violent political conflict. Proceedings of the National Academy of Sciences 104(18):7357-7360.
Husain, Ed. 2011. Did U.S. botch message with bin Laden videos? http://www.cfr.org/terrorism/did-us-botch-message-bin-laden-videos/p24939
Kydd, Andrew H., and Barbara F. Walter. 2006. The strategies of terrorism. International Security 31(1):49–80.
Rabinovich Abraham. 2004. The Yom Kippur War: The Epic Encounter That Transformed the Middle East. New York: Schocken Books.
Sagarin, Raphael D., and Terrence Taylor (eds.) 2008. Natural Security: A Darwinian Approach to a Dangerous World. Berkeley: University of California Press.
Schmitt, Eric. 2010. U.S. offers aid to rescue Pakistanis and reclaim image. New York Times, 15 August:A6 (New York Edition).

Commentaries 

Why cheap and ambiguous signals may serve diplomacy better than credible commitments.
Olivier Morin

In this interesting paper, Blumstein et al. argue for a view of diplomacy based on credible commitments, honest signals, and the prevention of misunderstandings caused by cultural distance. They hope to enlighten politicians with lessons drawn from evolutionary biology (though one might note that credible commitment was explored by Clausewitz before Darwin, and costly signalling is found in Thomas Schelling before Amos Zahavi). This comment challenges two points made by their article.

First, I argue that misdirection, ambiguity and provocation are worthy tools in a politician’s toolbox. Yes, such strategies may lead to military escalation — but sometimes that is precisely the point. I shall illustrate this with the story of the Ems dispatch, one of the most famous episodes of diplomatic signalling. Second, I will argue that Blumstein et al. misuse the notion of costly signalling, as used in economics and evolutionary biology (Zahavi, 1977). A costly signal in the strict sense is so costly to produce that the fact of producing it provides a credible information. Peacock’s tails and nuclear tests are costly signals sensu stricto. Threats, promises, symbolic gestures, and most other diplomatic moves are not costly in that sense. As a result, their informational value is different.

The Ems dispatch

In the summer of 1870, Chancellor Bismarck made a diplomatic move that would eventually make him the unifier of Germany and the victor of the Franco-Prussian war (Howard 2001). The move began when he encouraged a cousin of the King of Prussia to become candidate for the throne of Spain. A Hohenzollern in Madrid — that would mean an encirclement of France by friends of Prussia, a notoriously hostile power. Bismarck knew this. He knew this may cause violent reactions from the French, even a war in which Paris would play the part of the aggressor. Such a war would fit well into Bismarck’s plans for national unification.

Yet Napoleon III deftly defused the crisis. He uncovered Bismarck’s plan and managed to have the Hohenzollern claim to the Spanish throne withdrawn, with the agreement of the King of Prussia. A French agent went to Ems-Baden to seal the deal with the King. Bismarck, seeing his plan unravelling under his eyes, refused to be present at the meeting, and threatened to resign if the meeting took place. All in vain.

On the thirteenth of July, on the promenade of Ems, the King had a tense but courteous talk with the French ambassador. He confirmed the withdrawal of the Hohenzollern candidacy. Unbeknownst to Napoleon III, the French secretary of State had ordered the ambassador to take a hard line. The ambassador tried to push his advantage and obtain a commitment for the future. The King politely declined, and telegraphed Bismarck that the incident was over. The Chancellor did not react. He was nowhere to be found.

Bismarck was busy exploiting a mistake in the ambassador’s strategy. The King had satisfied one French request, but not the other. Bismarck cut and rephrased the King’s telegram to showcase the King’s refusal. His version of the Ems telegram, sent to major French and German newspapers, depicts a harsh and bitter encounter. Nationalists on both sides further distorted the message. The dispatch humiliated Prussia, whose King had been harassed with an arrogant ultimatum. It incensed the French, who declared war on Prussia less than a week later. Bismarck had won his first move.

The Chancellor’s tactics may interest Daniel Blumstein and his coauthors, since it turns most of their advice on its head. Bismarck did not pay attention to Blumstein et al.’s lessons 1 and 4. He produced few signals, most of them deceptive. He did not intend to resign if the Ems meeting took place. The Ems dispatch itself was a deliberately distorted signal.

Oblivious to Blumstein et al.’s lessons 2 and 3, Bismarck used historical precedent in a predictable, almost boring way. Unifying German imperial powers with the Spanish throne had been a classic geostrategic move in European history since the time of Charles V. It had always been interpreted as an attempt to isolate France, and Bismarck used this common knowledge to provoke the French.

Against lesson 8, Bismarck knew that the Ems dispatch would give rise to divergent interpretations on both sides of the Rhine, with everyone distorting and amplifying the dispatch to suit their purposes (as he himself did). Yet he did not try to limit or control the disagreements, as he did not see them as obstacles. Quite the contrary.

“Costly” signals?

Yet, one might reply, Bismarck seems to follow at least one of Blumstein et al.’s lessons. In a way, he uses “costly” signals: every time he communicates something, he takes a risk. His foreign policy is like the peacock’s tail in this respect: it is a signal with a cost.

The analogy is misleading, though. The costly signals of modern costly signalling theory are not merely signals that have a cost to the producer. The point of costly signals is that certain signals are more reliable than others since their production is costly (Veblen 1899/1973, Schelling 1960, Zahavi 1977). For instance, a peacock’s tail is a reliable indicator of a peacock’s fitness because it would be hard to grow a beautiful tail without a good immune system. Or possessing two luxury cars is a reliable indicator of wealth because possessing such cars without being wealthy would be difficult. Those signals are especially reliable because they reveal a cost that has already been paid.

Compare those costly-to-produce signals with signals that are cheap to produce, though the emitters take risks when emitting them. Uttering a sentence, in itself, is easy, whether the sentence is true or not. Lying may have costly consequences, of course, but those costs lie in the future. However near that future might be, the costs are not paid by the lier when he pronounces his lie. Thus the production of a sentence, by itself, says nothing about its truth. In contrast, the mere fact of producing a beautiful tail is a good indicator of the reproductive potential that the tail indicates.

Blumstein et al. Are right to note that politicians sometimes use such signals, like military parades or nuclear tests. But most diplomatic threats and promises are not costly signals in the biological sense. They are not costly and honest in the way a peacock’s tail is costly and honest. Bismarck’s threat to resign if the Ems meeting took place was just cheap words. His utterance of the threat was not an honest signal of his intention to resign. Indeed, nothing Bismarck could have said could be a costly signal in the biological sense.

Technical or moral advice?

Bismarck’s warmongering was, of course, objectionable in many ways. A moral person would usually prefer diplomatic strategies based on credible commitment, and a careful avoidance of escalation and misunderstandings. Since the Cold War, this way of doing politics seems the least destructive on the global scale. Yet individual politicians do not necessarily have peace and security as their first objective. Credible commitment is a useful tool for them, but sometimes ambiguity and deception serve them better. Even for peace-making: Charles de Gaulle solved the Algerian crisis by appearing utterly uncommitted and unpredictable. Thus, while I share the moral outlook of Blumstein et al. and agree with them on ethical grounds, I do not think we should commend their strategic solutions to actual politicians — unless we make it clear that we are not giving technical advice to diplomats, but promoting one particular view of what international relations should look like.

References

Michael Howard, 2001. The Franco-Prussian War: The German Invasion of France 1870-1871. Oxford: Routledge.

Thomas Schelling, 1960. The Strategy of Conflict. Cambridge: Harvard University Press.

Thorstein Veblen, 1899/1973. The Theory of the Leisure Class. Boston: Houghton Mifflin.

Amos Zahavi, 1977. The cost of honesty (Further remarks on the handicap principle). Journal of Theoretical Biology, 67: 603-605.

Comments on “The peacock’s tale: Lessons from evolution for effective signaling in international politics,” by Daniel T Blumstein et al.

Blumstein et al. look to evolutionary theory regarding signaling in animal species to try to draw out some general rules of thumb that they think might be useful for diplomats and politicians in matters of foreign policy. The rationale for the exercise is that “By capitalizing on solutions derived by evolution over 3.5 billion year of life of Earth, we may identify ideas that otherwise might not have been explored in a policy context potentially offering quick, novel and effective options to increase strategic and combat effectiveness.”  

None of the eight rules of thumb, however, are novel propositions or observations, and some – like the eighth, “it’s essential to know your audience” – are mainstays of international relations literatures and diplomatic folklore or commonsense.  (Not to say such wisdom is always acted on!)  The core idea in several of them is that signals will not reliably convey their message unless they are actions (or words) that would be more costly for a potential bluffer to take.  This idea has been familiar in Economics and Political Science research for some time now, and in this specific formulation it derives not from evolutionary theory but from developments in information economics in the late 1960s and early 1970s.  

There are indeed some significant parallels between the strategic problems facing animals competing over territory or mates and that between two states, or a state and non-state group, at odds over territory or public policy.  In both cases violent conflict is typically costly and thus something both sides would like to avoid. But both sides would also prefer that the other side concede more of whatever resources or goods (which might be symbolic, in the human case) are at stake.  In both contexts, then, all can ideally be made better off if they can develop some kind of signaling system that allows the more motivated or strong types to credibly reveal this, so avoiding violent contests when a stronger or more motivated party faces a weaker or less motivated antagonist.  It is fascinating – though perhaps not so surprising, once we have seen the analogy and realized that evolution can sometimes select “best reply” strategies even without cognition – that animal contests (including sexual selection) often exhibit complex strategies of costly signaling.

What is less clear is whether there are specific findings from the study of signaling in (non-human) animal systems that have interesting or novel implications for diplomats and politicians engaged in international affairs.  As noted, none of these eight general rules of thumb are results or arguments that did not already emerge from social science studies of signaling in human contexts.  Perhaps, however, as the study of signaling in animal contexts proceeds it will produce findings that suggest novel ideas about signaling in human contexts.

For example, the canonical story about the peacock’s tail has, according to more recent research, gotten somewhat murkier.  There has always been the following theoretical objection:  If a bigger tail with more eyespots signals greater quality, then greater fitness should select out variation in tails so that residual variation in tail quality is not correlated with fitness.  Thus, in evolutionary equilibrium, expected fitness must be equal across larger and small tails, with the handicap of a larger train exactly compensating for a bird’s greater underlying quality.  

Empirically, the original study found a correlation between number of eyespots and mating success in a population in Britain.  However, more recent studies based on populations in France and Japan reached different conclusions.  In the French group, number of eyespots was unrelated to mating success, but the length of the train was.  In the Japanese birds, there was no correlation for either one.  In studies of three different North American populations there was also no correlation between number of eyespots and mating success (at least until almost the bird’s eyespots were clipped off).  

It could be that some other feature of the tail is a signal of mate quality.  According to Erol Akcay, an evolutionary biologist who is my source for this information, we just don’t know, and we also don’t have studies that directly test whether the peacock’s tail is a costly signal, since cost has not been measured and related to “quality.”  Akcay (personal communication) thinks it is likely that peacock’s tail is a signal of some kind, since it is hard to see it evolving without having some adaptive value.  But he thinks “what exactly it is a signal of and whether it is honest because it’s costly are open questions at this point.”

So perhaps things are not so straightforward. This is surely also the case for diplomacy.  For example, it is not uniformly true that “honest signals will be costly” (Lesson 1) or that (Lesson 4) “threats should be costly.”  At a minimum, these claims depend on a prior assumption that we are talking about a situation in which the signaler can have a strong incentive to misrepresent its preferences or type – to bluff – to the target. In some diplomatic contexts, “cheap talk” can be informative because the parties’ interests are sufficiently aligned that misrepresentation can be counterproductive. Or, it can be the case that in signaling between governments there are so many sources of publicly available information about what is going on that misrepresentation is not a big concern.  Finally, when states are signaling over multiple dimensions of policy, which is often the case, cheap talk can be informative even when there are incentives to misrepresent.#

A final comment is that to the extent that we do put stock in these costly signaling claims as rules of thumb, they may actually argue against the notion that evolutionary (or economic) theory can increase diplomats’ tactical success in contests.  The point of these arguments is that contests are like auctions in which the side that is genuinely more resolved or more capable is more likely to win.  The advice “Make your signals of resolve (or reassurance, if seeking peace) costly!” only makes sense if you actually are that resolved or you are that willing to risk a disadvantage in order to get to peace.  There is a parallel here with “Lesson 3:  Unexpected signals may be more effective,” where the authors suggest that “individual citizens lining up to help others (as often occurs after natural disasters) are truly meaningful gestures.”  The advice “deliberately make your gestures of reconciliation spontaneous in order to make them effective!” is a bit of contradiction in terms.

Evolutionary theory regarding signaling in animal contests may yet produce novel and interesting insights into signaling in roughly parallel human contexts.  Given the incredible complexity and diversity of animal signaling systems, I would expect that this could be the case.  But I’m not sure if these very high level rules of thumb optimally exploit the potential in this area.

The study of human cooperation today is the current state of a continuous line of intellectual inheritance from Adam Smith and David Hume, through Thomas Malthus, Charles Darwin, and Emile Durkheim, and more recently the biologists William Hamilton and Robert Trivers. But Adam Smith led in another direction, through David Ricardo, Francis Edgeworth, and Léon Walras to contemporary neo­classical economics, that at least until recently recognizes only self-regarding be­havior.

The twentieth century was an era in which economists and policy makers in the market economies paid heed only to the second Adam Smith, seeing social policy as the goal of improving social welfare by devising material incentives that induce agents who care only for their personal welfare to contribute to the public good. Ethics, in this paradigm, plays no role in motivating human behavior.

Contemporary research on human cooperation yields several insights. First, interdisciplinary research currently yields results that obeyed traditional disciplinary research goals. While the twentieth century was an era of increased disciplinary specialization, the twenty-first may well turn out to be an era of trans-disciplinary synthesis. Its motto might be: when different disciplines focus on the same ob­ject of knowledge, their models must be consistent where they overlap. Second, by combining economic theory (game theory in particular) with the experimental techniques of social psychologists, economists, and other behavioral scientists, we can empirically test sophisticated models of human behavior in novel ways. The data derived from this unification allows us to deduce explicit principles of human behavior that cannot be unambiguously derived using more traditional sources of empirical data.

The power of the experimental approach is obvious: it allows deliberate experimental variation of parameters thought to affect behavior while holding constant other parameters. Using such techniques, for example, experimental economists have been able to estimate the effects of prices and costs on altruistic behaviors, giving precise empirical content to a common intuition that the greater the cost of generosity to the giver and the less the benefit to the recipient, the less generous is the typical experimental subject (Andreoni and Miller 2002). The resulting “supply function of generosity” and other estimates made possible by experiments, are important in underlining the point that other-regarding behaviors do not contradict the fundamental ideas of rationality. They also are valuable in providing interdisciplinary bridges allowing the analytical power of economic and biological models, where other-regarding behavior is big news, to be enriched by the empirical knowledge of the other social sciences, where it is not.

Biological approaches have been misled by the apparent explanatory power of two theories: inclusive fitness and reciprocal altruism (Hamilton 1964, Williams 1966, Trivers 1971). These theories convinced a generation of researchers that, except for sacrifice on behalf of kin, what appears to be altruism—personal sacrifice on behalf of others—is really just long-run material self-regard. Ironically, human biology has settled in the same place as economic theory, though from a quite different starting point, and using a quite contrasting logic.

The experimental evidence supporting the ubiquity of non-self-regarding mo­tives, however, casts doubt on both the economist’s and the biologist’s model of the self-regarding human actor. Many of these experiments have in common a nexus of behaviors that we term strong reciprocity. Strong reciprocity is a predisposition to cooperate with others, and to punish those who violate the norms of cooperation, at personal cost, even when it is implausible to expect that these costs will be recovered at a later date.

Strong reciprocity contributes not only to the analytical modeling of human behavior, but also to the larger task of creating a cogent political philosophy for the twenty-first century. While the writings of the great political philosophers of the past were usually both penetrating and nuanced on the subject, they have come to be interpreted simply as having either assumed that human beings are essentially self-regarding (e.g., Hobbes and Locke) or, at least under the right social order, entirely altruistic (e.g., Rousseau, Karl Marx). In fact, people are often neither. Strong reciprocators are conditional cooperators, behaving altruistically as long as others are doing so as well, and altruistic punishers, applying sanctions to those who behave unfairly according to the prevalent norms of cooperation.

People cooperate not only for self-regarding reasons but also because they are genuinely concerned about the well-being of others, try to uphold social norms, and value behaving ethically for its own sake. People punish those who free-ride on the cooperative behavior of others for the same reasons. Contributing to the success of a joint project for the benefit of one’s group, even at a personal cost, evokes feelings of satisfaction and pride. Failing to do so is often a source of shame or guilt.

We came to have these “moral sentiments” because our ancestors lived in environments, both natural and socially constructed, in which groups of individuals who were predisposed to cooperate and uphold ethical norms tended to survive and expand relative to other groups, thereby allowing these prosocial motivations to proliferate. The first proposition concerns proximate motivations for prosocial behavior, the second addresses the distant evolutionary origins and ongoing per­petuation of these cooperative dispositions.

The Roots of Human Cooperation

Our Late Pleistocene ancestors inhabited the large-mammal-rich African savannah and other environments in which cooperation in acquiring and sharing food yielded substantial benefits at relatively low cost. The slow human life-history with prolonged periods of dependency of the young also made the cooperation of non-kin in child rearing and provisioning beneficial. As a result, members of groups that sustained cooperative strategies for provisioning, child-rearing, sanctioning non-cooperators, defending against hostile neighbors, and truthfully sharing information had significant advantages over members of non-cooperative groups.

In the course of our subsequent history we created novel social and physical environments exhibiting similar, or even greater, benefits of cooperation, among them the division of labor coordinated by market exchange and respect of rights of property, systems of production characterized by increasing returns to scale (irrigated agriculture, modern industry, information systems with network externalities), and warfare. The impressive scope of these modern forms of cooperation was facilitated by the emergence in the last seven millennia of governments capable of enforcing property rights and providing incentives for the self-interested to contribute to common projects.

But prior to the emergence of governments and since, cooperation has been sustained also by motives that led some people to bear costs on behalf of others, contributing to common projects, punishing transgressors, and excluding outsiders. In the pages that follow we will advance three reasons why these altruistic social preferences supporting cooperation outcompeted unmitigated and amoral self-interest.

First, human groups have devised ways to protect their altruistic members from exploitation by the self-interested. Prominent among these is the public-spirited shunning, ostracism, and even execution of free-riders and others who violate cooperative norms. Other group activities protecting altruists from exploitation are leveling practices that limit hierarchy and inequality, including sharing food and information.

Second, humans adopted prolonged and elaborate systems of socialization that led individuals to internalize the norms that induce cooperation, so that contributing to common projects and punishing defectors became objectives in their own right rather than constraints on behavior. Together, the internalization of norms and the protection of the altruists from exploitation served to offset, at least partially, the competitive handicaps born by those who were motivated to bear personal costs to benefit others.

Third, between-group competition for resources and survival was and remains a decisive force in human evolutionary dynamics. Groups with many cooperative members tended to survive these challenges and to encroach upon the territory of the less cooperative groups, thereby both gaining reproductive advantages and proliferating cooperative behaviors through cultural transmission. The extraordinarily high stakes of intergroup competition and the contribution of altruistic cooperators to success in these contests meant that sacrifice on behalf of others, extending beyond the immediate family and even to virtual strangers, could proliferate. Modern-day nationalism is an example.

This is part of the reason why humans became extraordinarily group-minded, favoring cooperation with insiders and often expressing hostility toward outsiders. Boundary-maintenance supported within-group cooperation and exchange by limiting group size and within-group linguistic, normative and other forms of heterogeneity. Insider favoritism also sustained the between-group conflicts and differences in behavior that made group competition a powerful evolutionary force.

Why did humans, rather than chimps, lions, or meerkats, develop such exceptional forms of cooperation? The answer lies in the human cognitive, linguistic and what physical capacities that made us especially good at all of the above, and more. These capacities allow us to formulate general norms of social conduct, to erect social institutions regulating this conduct, to communicate these rules and they entail in particular situations, to alert others to their violation and to organize coalitions to punish the violators. No less important is the psychological capacity to internalize norms, to experience such social emotions as shame and moral outrage, and to base group membership on such non-kin characteristics as ethnicity and language, which in turn facilitates costly conflicts among groups. Equally essential was the developmental plasticity of humans and our long period of maturation, the latter initially a result of the particular feeding niche that early humans occupied. Also important is the unique human capacity to use projectile weapons, a consequence of which is to lower the cost of coordinated punishment of norm violators within a group, to reduce the costs of hunting large animals, with concomitant benefits accruing to groups with widely endorsed sharing norms, and to render intergroup conflicts more lethal. A result was to elevate group-level competition to a more powerful evolutionary force. These exceptional aspects of human livelihoods and social interactions, we will show, have favored the evolution of an individual predisposition to cooperate with others and to punish those who exploit the cooperation of others. But more than individual-level motivation is involved. The regulation of social interactions by group-level institutions plays no less a role than altruistic individual motives in understanding how this cooperative species came to be. Institutions affect the rewards and penalties associated with particular behaviors, often favoring the adoption of cooperative actions over others, so that even the self-regarding are often induced to act in the interest of the group. Of course it will not do to posit these institutions a priori. Rather, the historical evidence indicates that they could have coevolved with other human traits in the relevant ancestral ecologies and social environments.

Cooperation and Competition

The tension between the relentless logic of self-interest and the ubiquity of collective action in real-world settings was eventually resolved by a series of experiments by psychologists and economists, most notably by Ernst Fehr and his colleagues (Fehr and Gachter 2000, Herrmann et al. 2008). The experiments confirmed that self-interest is indeed a powerful motive, but also that other motives are no less important. Even when substantial sums of money are at stake, many, perhaps most, experimental subjects were found to be fair-minded, generous toward those similarly inclined, and nasty toward those who violate these prosocial precepts. In light of these results, the evidence that the tragedy of the commons is sometimes averted and that collective action is a motor of human history is considerably less puzzling. The puzzle, instead, is how humans came to be like this.

Social Preferences and Social Dilemmas

Social preferences are a concern for the well-being of others and a desire to uphold ethical norms. By contrast with self-regarding preferences, which are based on states concerning oneself alone, we stress other-regarding and ethical preferences, the former defined as valuations based at least in part on states that occur to others. Social preferences include not only generosity toward others and a preference for “fair” outcomes, but also what Thomas Hobbes called the desire for “eminence,” Thorstein Veblen’s “pecuniary emulation” exemplified by a desire to “keep up with the Joneses” (Veblen 1899), Charles Horton Cooley’s “looking-glass self” according to which our self-esteem is dependent in part upon what others think of us, so we attempt to favorably impress others as a means of raising our subjective self-esteem (Cooley 1902, Brennan and Pettit 2004), and Aristotle’s character virtues, such as honesty and courage, which are personal values that promote prosocial behavior(Aristotle 2002[350BC]).

Social preferences assume special importance in interactions termed social dilemmas, that is, interactions in which the uncoordinated actions of individuals result in an outcome that is Pareto inefficient, meaning that there exists some other feasible outcome such that at least one member could be better off while no member would be worse off. Examples of social dilemmas modeled by game theorists are the prisoner’s dilemma, the public goods game, sometimes termed an n-person prisoner’s dilemma, the so-called war of attrition and other so-called arms race interactions, the tragedy of the commons and the common pool resource game in which contributing to the common project takes the form of forgoing the overexploitation of a jointly utilized resource such as a fishery, water supply, or forest. We say a person free rides if he benefits from the contributions of other group members while himself contributing less or nothing at all.

Another-regarding player cares about not only his own payoff, but that of his partner as well. Such a player might reason as follows. “I feel sufficiently positive toward a partner who cooperates that I would rather cooperate even if by doing so I forgo the larger payoff ($15) I could have had by defecting. If my partner defects, I of course prefer to defect as well, both to increase my earnings, and to decrease the earnings of a person who has behaved uncharitably toward me.” If Bob and Alice reason in this manner, and if each believes the other is sufficiently likely to cooperate, both will cooperate. Thus, both mutual cooperate and mutual defect are equilibria in this new game, transformed from the old by augmenting the material payoffs with the players’ concerns about one another.

Genes, Culture, Groups, and Institutions

We define culture as the ensemble of preferences and beliefs that are acquired by means other than genetic transmission. Culture is an evolutionary force in its own right, not simply an effect of the interaction of genes and natural environments.

An alternative but we think incorrect approach holds that while preferences and beliefs that are transmitted culturally may constitute the proximate causes of behavior, they in turn are entirely explained by the interaction of our genetic makeup and the natural environment. It is of course true that natural environments and genes affect the evolution of culture. But it is also true that culture affects the relative fitness of genetically transmitted behavioral traits. C. J. Lumsden and Edward O.Wilson (1981), Luigi Luca Cavalli-Sforza and Marcus Feldman (1981), Robert Boyd and Peter Richerson (1985), William Durham (1991), Richerson and Boyd (2004) and others have provided compelling instances of these cultural effects on genetic evolution.

Recognizing the intimate interactions between genes and culture in humans, Edward Wilson, Charles Lumsden, Robert Boyd, Peter Richerson, Luigi Luca Cavalli-Sforza and Marcus Feldman began workinginthe1970’s on the parallels between genetic and cultural evolution and their interactions, their work initiating the modeling of gene-culture coevolution, the second concept underpinning a plausible explanation of the origins and nature of distinctive cooperation among humans. According to gene-culture coevolution, human preferences and beliefs are the product of a dynamic whereby genes affect cultural evolution and culture affects genetic evolution, the two being tightly intertwined in the evolution of our species.

In our gene-culture coevolution model of group-structured populations, the process of differential replication affects the frequency of both individual traits, generosity toward fellow group members, say, and group traits, a system of consensus decision making or property rights. Though inspired by biological approaches, especially those of Cavalli-Sforza and Feldman(1981), Boyd and Richerson (1985), and Durham(1991),like these authors, we do not privilege biological explanation. This approach may be summarized as follows.

First, while genetic transmission of information plays a central role in our account, the genetics of non-pathological social behavior is for the most part unknown. Knowledge of the genetic basis of the human cognitive and linguistic capacities that make cooperation on a human scale possible has expanded greatly in recent years, but virtually nothing is known about genes that may be expressed in cooperative behavior, should these exist. No “gene for cooperation” has been discovered. Nor is it likely that one will ever be found, for the idea of a one-to-one mapping between genes and behavior is unlikely given what is now known about gene expression, and is implausible in light of the complexity and cultural variation of cooperative behaviors. Thus, when we introduce genetic transmission in our models, our reasoning operates at the phenotypic level.

Second, as is conventional in all models of selection, relative payoffs, whether in terms of fitness, material reward, social standing or some other metric, influence the evolution of the population shares of various behavioral types, higher payoff behaviors tending to increase their frequency in a population. The resulting so called payoff monotonic dynamic is often implemented using “as if” optimization algorithms, though in doing this we do not attribute conscious optimization to individuals. Nor do we conclude that the resulting outcomes are in any sense optimal. In general they are not. The aggregation of individually optimal choices is universally suboptimal, except under highly unrealistic conditions.

Individuals with higher payoffs may produce more copies of their behaviors in subsequent periods either through the contribution of their greater resources to differential reproductive success or because individuals disproportionately adopt the behaviors of the more successful members of their group. The latter may occur voluntarily, as when youngsters copy stars, or coercively, as when dominant ethnic groups, classes, or nations impose their cultures on subjugated peoples. Of course, cultural transmission may also favor lower payoff behaviors (think of smoking or fast food).

Third, because positive feedbacks are common in the processes of behavioral and institutional change we study, otherwise identical populations may exhibit quite different trajectories, reflecting the multiplicity of equilibria that is typical of models with positive feedbacks. The outcome that occurs need not be that with the higher average payoff. The process of selection among equilibria may be on such a long time scale that two populations described by exactly the same model may exhibit dramatically different distributions of behaviors for thousands of generations. The process of determining which of many possible equilibria will occur, termed equilibrium selection, thus assumes major importance.

Finally, the emergence, proliferation and biological or cultural extinction of collections of individuals such as foraging bands, ethno-linguistic units, and nations, and the consequent evolutionary success and failure of distinct group-level institutions such as systems of property rights, marital practices, and socialization of the young, is an essential, sometimes the preeminent, influence on human evolutionary processes. The maintenance of group boundaries (through hostility toward “outsiders”, for example) and lethal conflict among groups are essential aspects of this process. Within-group non-random pairing of individuals for mating, learning and other activities also plays an important part.

Herbert Gintis
Santa Fe Institute and Central European University

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Accordingly, although most political scientists are committed to a general view that the interests and beliefs of human beings and human groups are the primary drivers of political behavior and political change, a good deal of attention by “institutionalists” is directed to relishing the ironies or bemoaning the tragedies of rationality ignored and interests contradicted. Indeed you do not need a political scientist to point out numerous examples of institutional forms or collective beliefs or norms that are severely suboptimal for precisely those populations and communities that uphold and honor them. Political scientists, as well as pundits, are well aware of the obstacles sclerotic institutions pose to good policy, progress, and a general sense that our political communities work for us rather than against us. References abound to “institutional inertia,” “the stickiness of institutions,” or the institutionalization of answers to questions that current circumstances no longer pose.

However, if it is well understood that institutions cannot change fluidly with changing needs and changing insights, it is also known that institutions do change, and sometimes they adapt. What is not well understood are the limits to the effectiveness and pace of institutional change and, specifically, why some institutions are exceedingly resistant to change, even when the deficiencies of the practices, policies, and predicaments associated with them are fully appreciated by influentials as well as ordinary people. In this essay I want to suggest the contribution that evolutionary theory, as a tool in the hands of trained social scientists, can make to finding answers to these questions.

In part because political scientists are so aware of how bad things are compared to what they theoretically could be, they commonly reject evolutionary theory and thinking as inappropriate for application to the worlds they seek to explain. This rejection is usually based on a fundamentally incorrect understanding of evolution as “survival of the fittest”–short-hand for the (incorrect) idea that Darwinian evolution produces the “best” version of what could be out of a ferocious competition among versions that can be. Let us not linger over the reasons for this error. There are many candidates for explanations including past abuses of evolutionary theory by Social Darwinists and some sociobiologists, cultural or psychological fears, and religious commitments. One oft-neglected explanation is perhaps the easy identification of such a Panglossian understanding of evolution with the principle of neoclassical economics that free, unconstrained competition in a market place can yield a Pareto optimal set of prices for guiding the most efficient use of resources possible.

In any case, this notion—that “history is efficient”—is certainly not an implication of evolutionary theory, not in the natural world and not in the social world either. Indeed it is the very effectiveness of evolutionary theory in accounting for suboptimality that offers political scientists and social scientists more generally, an approach to explaining the prevalence, not only of institutional suboptimality, but to the combination of adaptation and extreme resistance to change that institutions display.

A common trope among historically minded political scientists is that severely suboptimal outcomes are the product of the inheritance from the past of an institutional form or policy implemented to serve a particular purpose under particular circumstances.[i] The simplest form of this explanation is that the outcome is fully explained by “path dependence” and “inertia;” that is to say by the contingency of what happened in the past and by the difficulty of changing the status quo, even if the status quo no longer reflects the purposes or circumstances that resulted in the contingent outcome that was deposited in the present by the past.

Of course many political scientists offer very nuanced and learned “process-tracings” of these outcomes and the ironies and tragedies associated with them. But for more systematic and generalizable explanation for the regularity of this type of outcome, we can profitably turn to evolutionary theory. To appreciate how, we must understand the standard concept in evolutionary theory of a “fitness landscape.”

From an evolutionary point of view, the particular history of a polity or society is one path through the “state-space” of possible ways that society could have changed over time, from one combination of characteristics to another. Since each of these distinctive combinations of characteristics is a separate point, or “state” in the space of possible states, the trajectory of the society over time is a path through the state space achieved by movement from one state to another “accessible” to it. The idea of “accessibility” here reflects whatever laws are governing the behavior of relevant entities so that once relevant elements are configured in a particular way, a subset of possible successor states exists that includes the state that actually materializes. As a social or political institution or practice changes over time, slowly or rapidly, it can be imagined to be exploring a particular, and, in all likelihood, relatively small, portion of the state space it inhabits. Evolutionary mechanisms are ways of explaining patterns of movement through the state space that do not rely on calculated strategic choices at the level of the entity moving through the space. Instead, these mechanisms rely on the outcome of competition for replicative success among large populations of variants at levels of analysis below the ontological level of the entity moving through the state space. Another way of saying this, indicating the links between evolutionary and complexity theories, is that the path through the state space is an emergent property of unguided evolutionary processes at a lower level of analysis.

Thus, in evolutionary biology, the unit of selection reflects a particular level of analysis. The effects of evolution are traced on levels higher than that of the unit of selection. Sometimes the unit of selection is the codon, sometimes the gene, sometimes the trait which might be expressed by a combination of genes; sometimes a species and sometimes varieties within a species. Whenever evolutionary, or at least natural selection, questions are asked at one of these levels, we identify variation, a selection criteria arising from circumstances/competition, and retention ability across generations of replication. The result of processes of variation, selection, and retention at any particular level is a pattern of outcomes at a higher (emergent) level. For example, giant tortoises in the Galapagos Islands vary, island by island, by the shape of their shells near their necks and by the length of their necks. This pattern of differences at the level of tortoise anatomy resulted from competition, at a lower level of analysis, among different rates of reproductive success for individuals on different islands with traits (slightly longer necks and slightly notched shells) that marginally advantaged the individuals possessing those traits wherever food was relatively higher. The result was that the state space for the configuration of neck and shell on giant tortoises was explored along different routes on different islands.

A “fitness landscape” is a heuristic device for analyzing the opportunities and challenges of changing for the “better” (by whatever metric is imagined as selecting or pressuring behavior). For a simple example, picture a three dimensional grid comprised of columns rising from a checkerboard along the Y (vertical) axis. (See Figure 1) The squares on the checkerboard represent all the different ways the entity could behave or be constituted so as to combine every available value on the X (horizontal) axis (Attribute 1, e.g. neck length) with everyone on the Z (depth) axis (Attribute 2 e.g. shell shape). In other words, the checkerboard is the space of possible states the entity can “be.” Since evolution seeks explanations for patterns of change in response to circumstances without endowing the units of selection with the capacity to look far ahead for their success, the only states that can be achieved by an institution, policy, or practice are those adjacent to it. The columns rising from the squares on the grid register their relative “fitness” by their different heights. If change is incremental and myopically driven toward whichever adjacent (i.e. only very marginally different) column is higher, then “hill-climbing” will be normal. The trajectory of an entity through the state space will be upward.

Figure 1.

However, and this is the key to understanding why history is “inefficient” from an evolutionary point of view, ravines and chasms may exist in the landscape, or may come into existence as changes in circumstances degrade the effectiveness of strategies (ways of being; traits; call them what you will) that worked in the past while enhancing strategies that performed (or would have performed) poorly in the past. In such a rugged landscape, many better ways of being are not “accessible” without going “downhill,” or becoming less “fit” in the process of becoming more fit. But if the processes producing change are truly myopic, or only responsive to direct local stimuli and information, then they themselves—i.e. evolution, per se, will never produce downhill trajectories. Absent non-evolutionary, or at least non-natural selection, processes of change, a stabilized set of practices (i.e. an institution) can evolve to, or become stuck on, one of many suboptimal “local maxima” that may exist in the state space.

In our checkerboard illustration, this occurs if an entity is surrounded by columns that are lower than the column it occupies. Without look-ahead powers that evolution does not, per se, assume, that entity will not be able to improve its “fitness” by relocating to a higher, but distant and therefore non-accessible location. To do so would require some extraordinary event that exogenously relocates the entity to a position from which it could hill-climb to the highest peak available in the state space. This is of course not impossible, but it would run against the grain of normal interactions. In any event, the predicament outlined here proves the point that history’s inefficiencies can be modeled evolutionarily with no contradiction whatsoever to the fundamentals of evolutionary theory, i.e. to the claim that unguided change in the deployment or appearance of strategies drawn from a repertoire of those available arises from the immediate successes and failures that determine rates of replication of alternatives. In the world of institutions this means that memos identifying pathologies and offering plans for institutional change that entail high short-term costs in favor of long-term gains will tend to be out-replicated, and effectively suppressed, by memos warning of immediate pain, discounting the future heavily, and distorting the benefits of staying the course.

Let us look more closely at how these ideas could be used to do some work in the political world. We can do this by recognizing that political institutions, political practices, or the campaign strategies of politicians are entities that can be moved through state spaces. The trajectories can be understood as the product of evolution to the extent that they result from competition at lower levels among varieties of organizational forms, rhetorical appeals, political positioning, slogans, and formulas–these are the equivalent of the genes, codons, traits, or varieties that compete to drive the trajectory of evolution in biological contexts, depending on the level of analysis of the entity whose position in the state space is under examination.

Students of the political economy of advanced industrial societies are familiar with the “Japanese model.” This refers to the distinctive and enormously successful combination of institutional forms and attendant policies that produced the stupendous success of the Japanese economy in the 1980s. We need not go into detail here, but the Japanese economic miracle is widely understood to be due to effective planning and implementation by a developmentalist state intent on harnessing its resources for efficient production and prevailing against international competitors.[ii]  Key ingredients in the “Japanese model,” imitated to one extent or another by a number of East Asian and Southeast Asian countries in the 1970s and 1980s, included a deferential political culture; well-trained, well-coordinated, and political protected bureaucracy; a dense patron-client network with massive corporations working closely with banks and the national bureaucratic apparatus toward goals of growth of the economy; job security for the middle class; a disciplined working class; massive subsidies for agriculture; and the effective exclusion of women from the upper levels of the work force. Trusting in the wisdom of the state, and willingly following its directives, Japanese firms fully cooperated in tax, trade, and monetary policies to override market forces and endow Japan with tremendous competitive advantages in the global market place. Between 1960 and 1980 annual per capita growth grew from 36% of the American rate to 72%.

But in the early 1990s the real estate market tanked. Banks incurred immense losses on their books in order to prevent large firms from failing. This was imperative based on the corporatist organization of the economy and the concomitant absence of a welfare state to cushion unemployment. The political system churned away, producing government after government, but no reforms that could help Japan confront the huge challenges from its own ageing work force and from increasingly severe global competition. In her closely argued and extensively researched study of what happened to the Japanese economic miracle, Jennifer Amyx identifies the gradual crystallization of the Japanese model, but asks why it seemed incapable of grappling with the challenges posed in the late 1980s and early 1990s. Amyx credits Japan with a gradualist, evolutionary path to the economic model to which it owed its immense success, but then notes the sudden reversal of Japanese economic fortunes and the seeming inability of Japan to adjust in a timely manner to patently new circumstances. The result was a national debt more than twice as large as the OECD average by 2008 and an economy, once the envy of the world, now entering its third decade of stagnation. In her 2004 study, Amyx focuses her questions on the crucial finance sector of the Japanese economy, noting the difficulty existing theoretical approaches to institutional adaptation had in explaining Japan’s predicament.

Existing theories cannot explain the length of delayed government response to banking problems, the magnitude of breakdown seen in Japanese finance, or why Japanese authorities are unable to restore the financial sector to health even thirteen years after the onset of the crisis.[iii]

As Amyx makes clear, the problem was not that Japanese experts did not, and do not, understand the problem or were/are not committed to change or reversing Japan’s economic performance. The problem is that the kind of transformation required threatened and has threatened even more pain for the Japanese people and for the government, in the short run, than would be experienced (in the short run) by continued stagnation. Evolution toward optimal policies arising from the kind of gradualism that has characterized Japanese institutional adaptation in the past has not occurred, and will not occur. In essence, Japan has been stuck on the “local maximum” it had achieved in the 1980s in a fitness landscape that changed to reduce the relative fitness of its historically evolved strategy, but whose ruggedness has prevented normal processes of institutional adaptation to replace the “less fit” strategy with a “more fit,” but not immediately accessible alternative. While it is likely, if not certain, that Japan will eventually change its policies, when and how that happens will likely have little to do with myopic, evolutionary processes of natural selection per se, but will depend heavily on shifts in circumstances that remove ruggedness from the fitness landscape, on far-sighted leadership prepared to whether high political costs, and/or the force majeure associated with devastating shocks.

As noted, the problem of institutional rigidity is well-known and pervasive, and it would be wrong to characterize all instances as the result of being stranded on a local maximum. Determined, albeit mistaken, policies of powerful elites; the effects of vested and well-positioned interests or veto-players; the existence of determinative but latent or obscured functions; or the suppression of ideas about how change might be achieved are examples of alternative explanation for the failure of institutions to adapt. Nonetheless, as suggested above, the effects of being stranded on a local maximum is a powerful explanation and valuable tool for thinking about the predicaments faced by political systems and how they might be escaped.

In the 1950s it was clear to most Frenchmen that the Fourth Republic was a severely dysfunctional institution, but it was governed by parties and leaders that had evolved to perform well within its institutions and no matter how many opportunities they were given to adapt the regime to the requirements of the political system, they could not. It was the genius of de Gaulle to realize this, to withdraw from “le systeme,” and then use the regime-threatening problem of Algeria to replace it wholesale with the “Fifth Republic.” There was no gradual, evolutionary path from the Fourth to the Fifth Republic, but there was a revolutionary path. It is widely understood, in Israel and outside the country, not only that the country desperately needs a peace agreement with the Palestinians. But it is just as well understood that Israeli political institutions, however effective they are at maintaining democracy and producing opportunities for political office and patronage to those in power, have insured that every Israeli government has been coalition-based and reliant on small religious and highly ideological right-wing groups that prevent any realistic peace plan from being put forward. Gaullist solutions in Israel have been attempted—by Rabin and, to an extent, by Sharon–but so far Israeli elites have failed to weather the storm of political opposition associated with efforts to “deinstitutionalize” deeply embedded arrangements.[iv] In the United States, the Madisonian system described in Federalist 10, that prevents tyranny by dividing and balancing power among states, Houses of Congress, and branches of the Federal government, has also institutionalized a kind of gridlock in so many domains that the confidence of the American people in its government is falling to record lows. Just as Washington may well be understood as stuck on a local maximum—fit enough to allow incumbents to be re-elected, but not fit enough to solve the problems posed to it in the twenty-first century while enabling re-election–so may we understand the predicament of the Republican Party in this election cycle. As has been widely observed, any candidate wishing to win the Republican nomination may be forced to position himself or herself in such a way as to attract Tea Party support; thereby greatly complicating if not rendering impossible the rapid adaptation that will be necessary to achieve a position on the rugged “electability” fitness landscape near the position that wins by attracting independents and conservative Democrats.

Identifying a syndrome in politics with an evolutionary dynamic does not itself solve any problems. On the other hand, our understanding of evolution in fields as far removed from one another as psychology, botany, agronomy, pest control, pharmaceutical research, and cancer treatment, has helped enormously to explore state spaces for solutions and improvements that were not imagined beforehand. By understanding key predicaments in political life with the same conceptual, theoretical, and analytic equipment used to solve problems in evolutionary theory, we can begin to see the quandaries we face more clearly and imagine more systematically possible opportunities to escape or overcome them.

COMMENTARIES

Daniel Nettle, University of Newcastle, daniel.nettle@newcastle.ac.uk

The individual and the greater good: A comment on Lustick

Many of us biologically-minded folk have been appealing for years for social scientists to take evolution seriously (Nettle 2009). Thus, it is very gratifying to read Lustick’s thoughtful analysis of how institutional political scientists could employ evolutionary concepts. There are good and bad ways of bring evolution into social science (and you can find plenty of examples of both in recent literature). Bad ways have two characteristics: the evolution they appeal to is a highly simplified and sometimes wrongly characterized version of the nuanced edifice of evolutionary biology; and in their enthusiasm to embrace their Darwinian idea, they are less rigorous than they could be in the deployment of methods and knowledge base of the social science discipline which they purport to be expanding. The good ways, happily, are represented by Lustick’s work: it grows from a deep grounding in political science itself, and employs a sophisticated evolutionary metaphor in which both adaptation and history are important.

Lustick’s essay is based on the idea of institutional forms as replicating themselves through time, giving rise to a process of institutional descent with modification which happens more slowly than the change in the environment of people’s lives. Although institutional adaptation does occur, the fitness landscape has a complex shape, such that institutions can become trapped on local maxima. This analysis potentially has a lot to recommend it. However, I was surprised be did not invoke another important principle in evolutionary thinking, namely conflicts of interest between the individual and the collective as a source of suboptimality in design. The idea of the social dilemma – the tragedy of the commons – is already much discussed in political science, and indeed evolutionary biology got much of its thinking in this arena from social scientists (e.g. Ostrom 1990).

In evolutionary genetics, for example, there are many aspects of genomes which perpetuate themselves despite having no functionality for the organism as a whole (Burt and Trivers 2006). Transposable genetic elements make many copies of themselves despite not serving any function at the organismal level. Segregation distorters bias meiosis in their favour, and can spread despite being costly to the health of the animal carrying them. The replicatory interests of the segregation-distorting allele and the rest of the genome of the organism carrying it are partly, but not perfectly, aligned. The consequence is that things which are inefficient at the organismal level often persist.

How might we apply these ideas at the institutional level? The long-term efficiency of government might, hypothethically, be served by reducing the size of the political class. However, any individual leader introducing this reform reduces the size of his alliance base, and consequently risks personal loss of power. All politicians might agree that this is the reform needed, but it is not good for the career of any of them. Thus, it is a very hard reform to introduce. The same is true by definition of any policy which makes governments unpopular over the timescale of the electoral cycle, even if it would be good for society in the very long term. These social traps essential arise because the interests of society and of individuals, whether politicians or not, are partly but not perfectly aligned.

Something I found interesting in Lustick’s analysis is the potential role of rare massive upheaval in overcoming these traps. If society occasionally becomes completely destabilized, then no individual has any possibility of doing well personally by continuing with the status quo, then an adaptive radiation of new and better institutional forms is possible. I can only hope that, in this era of financial crisis and looming environmental problems, our political masters understand this.

References

Burt, A. and R. Trivers (2006). Genes in Conflict: The Biology of Selfish Genetic Elements.  Cambridge, MA: Belknap Press.  Nettle, D. (2009). Beyond nature versus culture: Cultural variation as an evolved characteristic. Journal of the Royal Anthropological Institute 15: 223-40.

Ostrom, E. (1990). Governing the Commons: The Evolution of Institutions for Collective Action. Cambridge: Cambridge University Press.

Comment on Ian Lustick’s  “Institutional Rigidity and Evolutionary Theory: Trapped on a Local Maximum”

David Sloan Wilson
President of Evolution Institute and SUNY Distinguished Professor of Biology and Anthropology, Binghamton University

Thanks to Ian Lustick for his stimulating essay. He correctly notes that stasis and change are two sides of the same evolutionary coin, for biological evolution no less than the cultural evolution of human political institutions. Sewall Wright, one of the fathers of population genetics theory, was the first to appreciate that when phenotypic traits have a complex genetic basis, natural selection can result in multiple stable local equilibria (the “peaks” of a multi-peak landscape), which are internally stable by definition but can differ in their absolute fitness (the “altitude” of each peak). His shifting balance theory was a complex scenario involving selection among populations occupying different adaptive peaks (Provine 1986). He originally developed the theory for individual traits with a complex genetic basis (such as coat color in guinea pigs), but it applies equally to social adaptations with a complex basis, where it is called “equilibrium selection” (Binmore and Samuleson 1997, Boyd and Richerson 1992, Samuelson 1997)

Applying these and other evolutionary ideas to stasis and change in political institutions is even more complex than Lustick suggests. I would like to introduce three additional factors and conclude with a reflection on how to manage the study of highly complex systems in both biology and the human-related sciences.

1) Multilevel selection theory needs to be distinguished from evolution on multi-peaked landscapes. The classic group selection model posits two traits, selfish and altruistic, in a multi-group population. The altruistic trait is selectively disadvantageous in all groups containing both types; there is no local equilibrium favoring altruism. Nevertheless, altruism can evolve in the total population if the differential fitness of groups containing the most altruists outweighs the selective disadvantage of altruism within groups.  A political institution can be dysfunctional, not because it is trapped on a small peak, but because individuals or subgroups are maximizing their relative advantage within the institution, at the expense of the institution as a whole and even their own long-term welfare (Wilson 2004).

2) The term “evolutionary mismatch” refers to adaptations to one environment that become dysfunctional in a changed environment. Our adaptations for evaluating and copying behaviors evolved in the context of small-scale social interactions and can easily malfunction in the context of large-scale political institutions, resulting in the paradox of practices that work but don’t spread and spread but don’t work. This class of institutional dysfunction needs to be distinguished from both multiple adaptive peaks and multilevel selection (Wilson et al. 2011)

3) Rational thought and intentional planning might seem to be furthest removed from evolution, especially with respect to escaping local maxima. However, these are better regarded as evolutionary processes in which both variation (efforts to imagine alternatives) and selection (explicitly stated goals) are highly organized. Moreover, our genetically evolved reasoning abilities might be better adapted to winning arguments than solving collective action problems (Mercier and Hugo 2011),  accounting for some of our reasoning inabilities in addition to our abilities.

This degree of complexity might seem daunting, but it is not evolutionary theory that makes the topic complex.  It is inherent in the subject matter and must be faced by anyone who studies political institutions from any perspective.  The question is whether an explicitly evolutionary perspective adds value to other perspectives. Along with Lustick, I think that the answer is emphatically “yes”–for the cultural evolution of political institutions on rugged adapted landscapes and the additional factors that I have briefly identified in this commentary.

A manuscript titled “Evolution as a General Theoretical Framework for Economics and Public Policy” (Wilson and Gowdy 2011) makes some of these points at greater length and is equally relevant to the field of political science.

References

Binmore, K., & Samuelson, L. (1997). Muddling throught: Noisy equilibrium selection. Journal of Economic Theory, 74, 235-265.

Boyd, R., & Richerson, P. J. (1992). Punishment allows the evolution of cooperation (or anything else) in sizable groups. Ethology and Sociobiology, 13, 171-195.

Mercier, H., & Sperber, D. (2011). Why do humans reason? Arguments for an argumentative theory. Behavioral and Brain Sciences, 34, 57-111.

Provine, W. B. (1986). Sewall Wright and Evolutionary Biology. Chicago: University of Chicago Press.

Samuelson, L. (1997). Evolutionary games and equilibrium selection. Cambridge, MA: MIT Press.

Wilson, D. S. (2004). The New Fable of the Bees. In K. R. (Ed.), Advances in Austrian Economics (Vol. 9, pp. 201-220). Greenwich CN: JAI Press.

Wilson, D. S., & Gowdy, J. (2011). Evolution as a General Theoretical Framework for Economics and Public Policy. First article of special issue planned for the Journal of Economic Behavior and Organization.

Wilson, D. S., Hayes, S. C., Biglan, A., & Embry, D. (2011). Evolving the Future: Toward a Science of Intentional Change. Behavioral and Brain Sciences, submitted.

Evolution predicts suboptimality, but not only because of getting stuck on local peaks

Hanna Kokko – Evolution, Ecology & Genetics, Research School of Biology, ANU College of Medicine, Biology & Environment. Hanna.Kokko@anu.edu.au

That institutions can become trapped on local maxima (with respect to some measure of performance) does not come as a surprise to any evolutionary biologist. I am probably not alone in my field when welcoming a move towards interdisciplinary understanding of complex systems, political and social science included. In addition to ‘food for thought’ provided by essays such as that by I.S. Lustick, researchers working in this area would also greatly benefit from moving towards quantitative rigour in this area (e.g. Turchin 2006), a step that biologists took long ago (for a historical review see Kingsland 1995).

There is one aspect in Lustick’s otherwise thought-provoking essay that warrants comment: it remains silent on the fact that even if an evolutionary process found itself on a slope towards highest peak, subsequently reaching it, the solution thus found can be maladaptive compared with a hypothetically reachable peak that could be found if entities forming the group (e.g. individuals in a society) forgot about their selfish short-term interests and pulled together to reach a common goal. This problem is one of the levels of selection, and it provides a clear counterexample for those who believe in invisible hands that create the best possible society simply by letting competition run free.

Lustick mentions that the unit of selection can be a codon, or a gene, or perhaps an entire species. Biologists agree, but only when the statement comes with a reminder that processes acting on the lower end of this spectrum often override selection on the higher units. This is why reaching outcomes that are good for the group is difficult, except — arguably — in the special case of multicellular individuals, where specialized adaptations exist to make sure that individual cell lineages do not take over and start proliferating at the expense of how well the entire organism functions and survives.

After all, it’d be terrible if some cells in your liver, or brain, started dividing haphazardly, without the law and order that states they have to serve the greater good of the multicellular society (i.e. you). Unimaginable? In fact, I have just described cancer, which quite commonly causes death in senescing animals (though not in plants, which do not have animal-like circulatory systems that would allow rogue cells to spread around the entire body). Despite cancer-killing cells and other similar adaptations shown by tightly regulated multicellular creatures like you and me, the system may, to its detriment, sometimes fail to extinguish the selfish, short-term interests of individual cells (Lewis et al. 2008). Here, it is irrelevant that cells do not really have aspirations in any cognitive sense. Short-term proliferation of cells can be selected for in the simple sense that more cells of the cancerous type is by definition a short-term reproductive improvement in this lineage. Given a short enough timeframe, this remains true despite extinction looming in the near future: cancer often kills, and the death of the organism kills the cancer.

We tend to think of cancer as a medical rather than an evolutionary problem. The beauty of evolutionary theory, however, is that it provides a framework for thinking about the commonalities of problems occurring at every level of selection. Fishermen unable to resist the temptation to overfish this year, despite fish stocks depleting to levels that threatens the entire industry? A virulent pathogen spreading in the local population of schoolchildren, even though this means that soon everyone will be immune, and the virus has nowhere to go? These are all examples of the tragedy of the commons, which means that evolution often favours short-term success over more prudent behaviour, even if the latter meant better performance as a whole in the fitness landscape (Rankin et al. 2007).

In fact, human institutions could be viewed as adaptations that try to keep some level of order intact at a higher level of selection that is vulnerable to the invasion of short-term interests. Societies fund police forces that have been given the power to punish thieves, for essentially the same reason as our bodies produce cancer-killing cells: to prevent detrimental selfishness from spreading. It is also the same reason why we spend money on negotiations over quotas for dwindling stocks of cod in the sea — or over the right to dump CO₂ in the atmosphere. The struggle between interests of different entities, each of which takes a shorter term view than would be ideal, is the root of much what is problematic in the world. Add to that the types of myopia that Lustick mentions — the inability of an evolutionary process to be farsighted enough to jump to distant peaks — and the power of evolutionary theory to predict suboptimal design should be clear to anyone.

Kingsland, S.E. 1995. Modeling nature. The University of Chicago Press, Chicago.

Lewis, Z., Price, T.A.R. & Wedell, N. 2008. Sperm competition, immunity, selfish genes and cancer. Cellular and Molecular Sciences 65:3241-3254.

Lim, M., Metzler, R. & Bar-Yam, Y. 2007. Global pattern formation and ethnic/cultural violence. Scence 317:1540-1544.

Rankin, D.J., Bargum, K. & Kokko, H. 2007. The tragedy of the commons in evolutionary biology. Trends Ecol. Evol. 22:643-651.

Turchin P. 2006. War and Peace and War: The Life Cycles of Imperial Nations. Pi Press

Commentary on Ian Lustick’s “Institutional Rigidity and Evolutionary Theory:  Trapped on a Local Maximum”

Bradley A. Thayer
Professor Department of Political Science
Baylor University

Ian Lustick has produced an important argument by thoughtfully applying evolutionary ideas to a major problem in the study of institutions.  Lustick demonstrates how an evolutionary approach may explain why institutions resist change even when their faults and limitations are well understood, while at the same time remain adaptive.  To advance his argument, he draws on the central concept of evolutionary theory:  natural selection.  Evolution through natural selection operates through variation within a population, a selection criterion or criteria arising from competition, and replication.

From this foundation, Lustick artfully creates a “fitness landscape” for illuminating why institutional change is so difficult—in sum, entities are trapped on a local maximum, plans for change entail short term cost for long term gain, and will lose out to those discounting the future.  When these ideas are applied to political problems, they can explain major phenomenon like the rigidity of political systems in important and novel ways.

In this brief commentary, I evaluate Lustick’s use of evolutionary ideas and conclude that Lustick’s approach is a model of how evolutionary ideas and theory may be applied to social science.

Lustick’s use of evolutionary ideas is well done.  Here I offer two points to place the discussion in greater context.  A critical point in the operation of evolution through natural selection is the selection pressure on variation within a population.  Selection pressure is broader than Lustick’s treatment, and should be thought of as competition for resources among conspecifics, for example, institutions and states, but also with other species, predators, and changing immediate, seasonal, and long-term ecological conditions (the impact of the international system, if you will).  The last point underscores the consideration of time.  Benign ecological conditions, for example, abundance of resources or few predators reduce selection pressure and may do so over a considerable period of time.  The reverse is likely to accelerate change over a relatively short period.  Recognizing the broader scope of selection pressures will only assist the study of institutions.

Second, evolution lacks teleology.  With that in mind, it is important to stress a major point:  “better” and “best” are relative, and are so for given ecological conditions.  As conditions change, what was the “best” for a specific condition may be fatal for the species in new conditions.  Better to be “good enough” and adaptable to changing ecological conditions than the “best” and inflexible.  Here evolution agrees with Voltaire’s quip that the perfect is the enemy of the good enough.  As I read Lustick’s consideration of “fitness” in his presentation of his “fitness landscape,” he is implicitly sensitive to this point, which I would encourage to be drawn out in the future.

Finally, Lustick deserves great credit for advancing the goals of consilient social science, the use of insights from the life sciences to inform, improve, and augment our understanding of major social problems.  Sven Steinmo’s recent examination of political economies of Sweden, Japan, and the United States joins Lustick as another excellent example of the use of a consilient approach to aid our comprehension of institutional change.[1]  As other scholars join Lustick and Steinmo in a consilient approach, the knowledge of institutions, and politics and, more broadly, social science may advance.