” Greek Democracy as a Major Evolutionary Transition: A Conversation with Josiah Ober “

The 1970s marked a conceptual breakthrough in evolutionary biology that was beyond Darwin’s imagination. Individual organisms can evolve, not only by small mutational steps from other individual organisms but also by social groups becoming so cooperative that they qualify as higher-level organisms in their own right. This is called a major evolutionary transition (MET).

The first example of a MET to become widely accepted was the symbiotic cell theory of Lynn Margulis, who proposed that nucleated cells evolved as symbiotic communities of bacterial cells.1 Then other METs were proposed in the 1990s by John Maynard Smith and Eors Szathmary, including the origin of life as groups of cooperating molecular interactions, the first cells, multicellular organisms, and eusocial insect colonies.2

The main requirement for a MET is a shift in the balance between levels of selection. Natural selection can operate among individuals within groups, which tends to result in socially disruptive behaviors, or between groups in a multi-group population, which results in internally cooperative behaviors.3 In most social species, within-group selection is sufficiently strong to prevent groups from acting with the unity of a single organism. In a MET, mechanisms evolve that shift the balance between levels of selection, suppressing disruptive within-group selection so that between-group selection becomes the dominant evolutionary force. The suppression of disruptive within-group selection is never complete. Even multicellular organisms, which have been evolving for billions of years, are still vulnerable to cancers.4

As if these conceptual developments in evolutionary biology weren’t revolutionary enough, the concept of METs makes great sense of human genetic and cultural evolution. Humans and chimpanzees might share 99% of their genes, but there is a night-and-day difference in their degree of cooperativity. Naked aggression is over 100 times more frequent in a chimp community compared to small-scale human communities.5 Between-group hostilities are another matter, but that is only to be expected from a multilevel perspective. Between-group selection results primarily in cooperation within groups, not necessarily cooperation between them.

To the best of our knowledge, the main factor in the evolution of within-group cooperation in humans is social control.6 Our distant ancestors found ways to suppress bullying and other forms of self-serving behaviors within their groups so that working as a group became the primary adaptive strategy. While the “reverse dominance” hypothesis of Christopher Boehm was advanced independently of the MET literature, the two concepts are convergent, since a MET is nothing more or less than the suppression of disruptive competition among individuals within groups. Recently, this is being described under the rubric of self-domestication.7

The human genetic MET resulted in myriad forms of cooperation, both physical and mental. Physical forms of cooperation include hunting, gathering, childcare, modification of the physical environment, defense against predators, and offense and defense against other human groups. Mental forms of cooperation include perception, memory, decision making, and the capacity for symbolic thought.

Symbolic thought is the maintenance of an inventory of symbols with shared meaning that informs action and is transmitted across generations. It is inherently a cooperative activity and thinking of it as part of a package of other cooperative activities, all of them made possible by a single MET, is a synthetic conceptual breakthrough.8

The capacity for symbolic thought resulted in a cultural stream of inheritance that first evolved by genetic evolution and then has been co-evolving with it ever since.9 To state this in evolutionary terms, every person is a collection of genes (their genotype) that partially influences their measurable properties (their phenotype). Every person is also a collection of symbols (call it their symbotype) that influences the very same phenotype. Selection operates on genotypes and symbotypes simultaneously, with symbotypic evolution by far the more rapid process.10

Cultural evolution is a multilevel process, no less than genetic evolution. In other words, a culturally acquired trait can spread by benefitting individuals, compared to other individuals within the same social group, or by benefitting the group, compared to other groups in a multi-group population. What I have described as two-level selection can be expanded to multiple levels, since humans often live in a nested series of groups (e.g., in modern life, towns, counties, states, nations…). The general rule is: Adaptation at any given level requires a process of selection at that level and tends to be undermined by selection at lower levels.

Cultural MLS theory provides a new theoretical lens for viewing the increase in the scale of human society over the last 10,000 years as a series of METs. Peter Turchin, who is trained in evolutionary biology and population dynamics, is the main proponent of this approach in books such as War and Peace and War and Ultrasociality: How Ten Thousand Years of War Made Humans the Greatest Cooperators on Earth. In addition to christening the field of Cliodynamics (the quantitative study of historical dynamics), Turchin has coordinated the assembly of a worldwide historical databank called Seshat for testing hypotheses about human cultural MLS. In a recent volume titled Seshat History of the Axial Age, what historians call the Axial Age is examined as a number of loosely coupled METs in which the major religions of the world provided a social glue enabling cooperation to take place at a larger scale than ever before—but always in the context of between-group economic and military competition at still larger scales.

Assembling a quantitative worldwide historical database requires collaborations with historians trained in more traditional social science perspectives and qualitative methods. Also, quantitative analysis requires coding qualitative information and must be complemented with the original “thick descriptions” that provide essential context. In this fashion, an explicitly evolutionary perspective is becoming integrated with other social science perspectives, especially the “new institutional economics” pioneered by Douglass North and represented by books such as Daron Acemoglu and James Robinson’s Why Nations Failwhich is strongly convergent with cultural MLS theory. This includes my own collaboration with Elinor Ostrom,11 another member of that tradition.

Against this background, a 2015 book titled The Rise and Fall of Classical Greece, by Josiah Ober, professor of political science classics at Stanford University acquires special significance. When I read it, cultural MLS and METs seemed to jump out of every page. It helped that Ober himself describes the Greek city-state (polis; plural poleis) as like an ant colony and cites the work of his Stanford colleague, the ant biologist Deborah Gordon. Also, Ober comes from the same new institutional economics tradition as Acemoglu, Robinson, and Ostrom.

One benefit of understanding classical Greece explicitly as a MET is that it can place the very concept of democracy on a stronger theoretical foundation. Many discussions of democracy offer purely cultural explanations, as something that emerged in certain times and places in human history, such as classical Greece or America and Europe in the 18th and 19th centuries. The concept of a MET helps us to see democracy in a broader and deeper evolutionary context, as something that is required for a society to function as an adaptive unit, in nonhumans in addition to humans, all the way back to the origin of life.


David Sloan Wilson (DSW): Greetings Josh!  Thanks for agreeing to have this conversation with me. I apologize for the long-winded introduction, but there was a lot of context to provide.

Josiah Ober (JO): I am delighted to begin the conversation. I have been fascinated with evolutionary biology and animal behavior since childhood, and likewise with the problem of social cooperation in large and small groups. It was more or less by accident that I ended up working in history and social science rather than biological science. And, as you mentioned, I have benefited immensely from talking with my good friend Deborah Gordon, whose work on ant behavior I find inspiring.

DSW: Let’s begin with the basic dynamic of cooperation and competition, within and between groups, which is the hallmark of MLS theory and also appreciated by new institutional economists. The thesis of Why Nations Fail, for example, is that nations vary along a continuum from inclusive to extractive.  The benefits of inclusive societies are widely shared by their members, whereas the benefits of extractive societies are confined to a small group of elites. Inclusive societies are always in danger of becoming more extractive, either by subversion from within or hostile takeovers from without. However, inclusive societies have an advantage over extractive societies in economic or military competition. Why? Because they function better as societies and their members are more willing to cooperate on their behalf. Is there anything that you would like to add to my description of Why Nations Fail, or more generally the tradition of new institutional economics?

JO: The new approach to institutional economics was a joint project of several scholars who had become disenchanted with the reigning neoclassical orthodoxy that reduced all behavior to individual-level utility maximization – that is to competition in markets. They saw that there was simply too much that was left unexplained, including strong forms of cooperation in large-scale democratic societies. They remained committed to the core concept of rationality and sought to explain why rational agents would choose not to defect from a cooperative order when it was in their short-term interest to do so. This led to new theories of social equilibrium, which accounted for the emergence of firms (Ronald Coase), states (Douglass North), and interstate order (Robert Keohane). As you noted, other scholars, notably Elinor Ostrom, sought solutions to the endemic problem of free-riding on the cooperative efforts of others, and the “tragedy of the commons” in which individually rational behavior degrades common pool resources. The real insights emerged when theories of rationality were tested against empirical evidence of actual behavior – most notably, I think, actual historical behavior. Once the long span of recorded human history is recognized as a source of evidence for theory testing, and choice theory was recognized as a tool for historical inquiry, the way was open to some really exciting new work. Why Nations Fail is one good example of that; Violence and Social Orders by North, John Wallis, and my colleague Barry Weingast is another; likewise, David Stasavage’s recent Decline and Rise of Democracy. In each case, the goal is to explain variation and similarity among historical societies, and differential levels of productivity (and thus adaptive success) by reference to different forms of social cooperation.

DSW: That’s very helpful. Now I’d like your own view of how your own academic background in political science is becoming integrated with the more explicitly evolutionary perspective represented by Peter Turchin and others, including the insights that you get from conversing with an ant biologist such as Deborah Gordon. What is the added value of this integration, beyond what new institutional economists have already discovered on their own? While we’re at it, please describe how your classics background enters the mix!

JO: I began my academic work in the field of ancient history, due largely to the accident of taking an undergraduate class at the University of Minnesota from an inspiring Greek historian, Thomas Kelly. My early work was on ancient Greek military history and the archaeology of borderlands; that got me interested in the deeper question of the similarity and differences among Greek city-states (there were about 1000 of them in the age of Aristotle), how democracy worked (or failed), and how the city-state world functioned as an extended ecology in which fierce competition (war) coincided with a range of innovative forms of cooperation (trade, interstate alliances, federalism). When (modern) Greek politics suddenly made field archaeology difficult, I moved on to the question of how distinctive forms of public communication (your symbotypes) in democratic Athens enabled cooperation across the lines of class, education, and status – and ultimately enabled democracy to avoid falling prey to elite capture. That is, in Acemoglu and Robinson’s terminology, sustaining inclusivity and fending off the tendency to exclusivity.

The resulting book, Mass and Elite in Democratic Athens, was well received and took me from Montana State, where I had begun my teaching career, to a professorship at Princeton, and from a department of history to classics. The great thing about classics as a field is that it is “pre-disciplinary” – so rather than being captive to a set of established theories and methods, I was free to wander into political theory, sociology… and ultimately biology. While I could now explain why democracy could escape the trap of exclusivity, I was still at a loss to explain why democratic Athens did so well, in terms of comparative performance, for so long.  I began to read more systematically in a range of fields, from organizational theory to primatology. And, by luck, I met Barry Weingast and some of the other institutional economists at Stanford during a fellowship year at CASBS (Center for Advanced Study in the Behavioral Sciences).

The upshot was Democracy and Knowledge – a book on how democratic institutions that incentivize cooperative behavior – especially “sharing what one knows” – can enable a state to out-perform more autocratic rivals. That project led to an offer from the political science department at Stanford, where I now split my appointment between Political Science and Classics. I got to know Deborah Gordon in 2006, soon after I joined the Stanford faculty. We have collaborated on a few small projects focusing on cooperation, and we have regular conversations and email exchanges about our shared interests. I know Peter Turchin through his visits to Stanford – his project is very impressive. Like some of my other Stanford colleagues, Ian Morris and Walter Scheidel, Peter does “big history” – very long time spans, highly comparative. My own approach has a narrower gauge: like a biologist who specializes in single species, I have kept my primary focus on the ancient Greek world, where there is a lot of data and where I feel I can drill down into the specifics of how organizations and institutions function.

DSW:  I love it! And I’m so happy to introduce your work to an evolutionary audience through this conversation. Now I’d like to summarize the ecology and population structure of Greece. However you came by it, your account sounds like what an ecologist might write. Greek culture was adapted for a certain kind of subsistence based on grains, olive oil, and wine, that confined them to a certain climatic zone, fittingly called Mediterranean. Within that zone, there were literally hundreds of poleis interconnected by sea and land travel. People who identified as Greek by sharing the same language and customs (with ample variation) coexisted with other cultures large and small, including major empires such as Egypt, Persia, and Macedonia. Trade and warfare were thoroughly intertwined and warfare was chronic. This is the environment in which multilevel cultural evolution took place. It puts flesh on the bones of what I wrote earlier—that a given cultural trait might spread within a polis, where it might well be disruptive, or by causing a given polis to outcompete other poleis in economic and military competition. Have I accurately represented what I learned from your book? Do you have anything to add to my description?

JO:  Great description. Among the key points here are that from the time of Homer (ca. 8th c BCE) to the Roman takeover (2nd c BCE) hundreds of independent or semi-independent Greek city-states competed and cooperated in ways that generated a lot of economic growth and, famously, a highly influential culture. Growth and culture fed on each other, just as you say, at multiple levels of cooperation and competition: within each polis (competition for status and honors, typically mediated through a pretty thick civic culture), between poleis – with relationships ranging from hot war to federalism (shared citizenship and policy). Cooperative exchange (trade, culture) was facilitated by a shared language (many distinct dialects of Greek, but Greeks communicated readily), and other shared cultural attributes (religion, foodways, etc.) Competition between states in the context of ready communication drove institutional and technical advances, rewarding states (and individuals) capable of adapting to a quickly evolving environment. At the next level up, Greeks were engaged in trade (and war) with neighboring cultures – and over time the Greek language, institutions, technology, and even the organizational structure of the polis were adopted by some of their neighbors. Once again, this looks a lot like evolutionary adaptation as one species proves more capable of expanding its niche through the development of more effective forms of cooperation.

DSW: This is why I was so dazzled by reading your book! Now we can get to the main event. Please describe how democratic norms, customs, and institutions evolved in this metapopulation.

JO: I begin with the idea of citizenship as a generalized phenomenon in the Greek world, and one that takes a historically distinctive form: the basic notion of citizenship is that relatively many members of a given bounded community should have a role in making and implementing the decisions that affect the community’s fate. This is, of course, common in small foraging groups, but, famously, becomes difficult at scale, making more centralized forms of authority more efficient at producing the relevant public goods of security and welfare. The Greek world looked pretty “normal” in the Bronze Age of the 2nd millennium BCE – that is, palace-centered organizations with no evidence of what we think of as classical forms of citizenship. But in Greece, a “perfect storm” of social collapse knocked out the palaces in the late second millennium.

DSW: Please say more about the structure of palace-centered organizations and why they collapsed.

JO: Palace-centered social systems were the Bronze-Age norm, from western Asia and northeastern Africa (Egypt), through the eastern Mediterranean. The king was both a political ruler and a central figure in the maintenance of divine order. The palace (and associated temples) collected taxes on agricultural enterprise and returned security – repelling attacks by rival states, creating state granaries against famine, and, importantly, sustaining the cosmos through the performance of the correct rituals. Palace systems put a premium on the centralization of power in the palace and the person of the king; social order flows from the center to the periphery; tax revenues from the periphery to the center. This was a remarkably stable equilibrium (analogs are known world-wide, per Turchin), and it takes a lot to destabilize it. Erich Cline, in 1177: The Year Civilization Collapsed describes the “perfect storm” of disease, climate change, volcanic activity, and invasion that brought down the palaces of the Greek world within a century or so, completely quickly and completely eliminating the structures of belief, authority, and bureaucracy that had sustained them. Writing, for example, a skill essential for keeping palace records, but useless once the palaces were gone, was lost to the Greek world for perhaps 400 years.

The ensuing “Greek Dark Age” was one of general poverty: a big drop in population and consumption. During that period, it appears that the Greeks developed the core cultural norms that would enable the kinds of social cooperation characteristic of the polis: a strong norm of equality among adult males, restraint on the accumulation of power or wealth by individuals, forms of warfare that required high levels of cooperation among similarly armed men. These developments were facilitated by the “right” combination of geography (small valleys divided from one another by mountains, but easy travel by sea), climate: regular but sparse rainfall: not great rivers, primary food crops are storable: wheat, barley, olives [oil], grapes [wine]). And a catastrophe of the right level at the right time to knock out the old equilibrium. In any event, as the Greeks emerged from the Dark Age in the 8th century BCE the “normal form” of centralized government by a monarch or tiny ruling elite (“exclusion”) did not become the inevitable organizational form for the coalescing city-states – although “tyranny,” as the Greeks characterized highly centralized government, remained as a threat to citizen regimes. Over the next several hundred years, there was a lot of experimentation with different civic forms. And there were a lot of interchanges, both cooperative and competitive, between and within communities.

Part of the story of the most successful of these emerging city-states concerns political entrepreneurs who successfully advocated positive-sum bargaining solutions to cooperation problems: at Athens, these entrepreneurs include Solon, Cleisthenes, and Pericles. There, the citizen regime developed into full democracy (that is, universal native adult male participation in government). Athens was a success – that is, highly competitive in the polis ecology – because democratic institutions allowed for the effective aggregation of useful knowledge, while also promoting high levels of civic solidarity. This meant, among other things, that Greek democracies were magnets for an emerging class of intellectuals.

Like traders, Greek intellectuals (aka sophists, philosophers, poets, historians) could and did move around, share information, and compete for markets (students, patrons, attention). Among the big questions they asked, in the texts that are preserved for us, was – how does cooperation across a population of self-interested persons actually work? This is to say: the Greeks discovered the problem of social cooperation and came up with some interesting theories to help explain how it can be overcome. That is what I am currently working on in a book that is nearing completion: The Greeks and the Rational: The Discovery of Practical Reason. The point here is that it is at least a plausible hypothesis that “the Greek MET” became self-conscious of itself as a MET. Of course, they did not know the biology that provoked the original insights of MET. But they grasped the core insight (shared with mid-twentieth-century rational choice theorists) that cooperation is a problem that must be solved at the level of individual motivation and action: That is, cooperation can be achieved “bottom-up” via aligned incentives for (e.g.) paying the costs of punishment of norm-violation, as well as “top-down” via command and control. And that solving the problem is the key to competitive advantage in an environment of nested levels of interaction.

That breakthrough in self-consciousness about the microfoundations of rationally cooperative behavior may be something that happened elsewhere in antiquity – but I do not know that it actually did. I think the discovery may have been unique and one of the primary (although little acknowledged) contributions to the Greeks to world culture.  I suppose that it was that very collective self-consciousness of “being a MET” (in your terms) that was one of the distinctive drivers of the growth of the Greek economy and culture. It is, perhaps, one of the ways that the Greeks are strikingly familiar (as well as in many ways strange) to us today.

DSW: Wow! Our main challenge in this conversation is to prevent it from growing into a book! In what follows, I’ll touch upon a number of major topic areas worth exploring from an explicitly evolutionary perspective. Let’s begin with what you have just emphasized; that the Greeks were conscious of their democracy-building efforts. It wasn’t entirely a blind process of cultural evolution. It was importantly directed, which doesn’t deny that there was also a large blind component. I am reminded of my own account of the Protestant Reformation in my book Darwin’s Cathedral, where dozens of reformers were consciously striving to create models of religious governance, but contingencies of history determined why some (such as Calvin in Geneva) succeeded and others (such as Zwingli in Zurich) failed. But I want to learn more about what preceded and coexisted with “self-consciousness about the micro-foundations of rationally cooperative behavior”. As you say, this is so familiar to us today that it is hard to imagine alternatives.

JO: It appears to me that various intuitions about individual and sub-group self-interest and the problems that it poses to higher levels of cooperation were available to the Greeks from the age of Homer – that is, already in our earliest post-Dark Age texts. But in the mid-fifth century BCE, with the growth of the wider Greek economy, and the rise of democratic Athens as a center for intellectual enterprise (and competition), the scene was set for those intuitions to be brought together into a theory of human behavior, predicated on ranked preferences, coherent beliefs, and strategic reasoning that involved calculating risk and taking into account the expected actions of self-interested others. It is that theory that, so I argue in my book in progress, that was the core of what the so-called Sophists taught their paying students. That theory of rational choice and action was taken over by Socrates, Plato, and Aristotle, and through them, became part of the western heritage of thought. And it is at least in part because it is part of our deep heritage from antiquity, that it seems so familiar today.

DSW: On a related point, I’d like to learn more about the range of variation in forms of governance that were not democratic. One of my favorite books is The Ritual Process, by Victor Turner, who makes a distinction between the “Structure” of a society, which is often hierarchical, and a spirit of “Communitas” in which everyone is regarded as a moral equal. According to Turner, many rituals serve to anchor Structure to Communitas, providing more bottom-up control of the elites than meets the eye. If nothing else, subjects of a chief or big man can argue with their feet if they have a choice of who rules over them, acting as a brake on extreme despotism.

Does this describe the social milieu from which democratic governance in Greece emerged?  Is an enlightened despot what Plato meant by philosopher-kings?

JO: David Stasavage’s book, mentioned above, gives a great sense of the range of mechanisms in which societies that were not democratic by a classical Greek standard, set limits on what Turner called “Structure” in hierarchical, central-authority systems. Stasavage distinguishes between societies in which the ruler is expected to consult regularly with a council that includes individuals he does not personally appoint and societies in which the ruler is not so limited. Certainly, the earliest Greek literary texts, by Homer and Hesiod, suggest that as they were emerging from the Dark Age, the residents of early Greek states expected rulers to consult with those who were in some ways their peers. The rough equality among a fairly extended group of elites was the precondition for the extension of the notion of equality across the adult male population of an extensive territory.

DSW: Continuing with the theme of “arguing with one’s feet”, In MLS theory there is a class of models called “Walkaway”, in which dispersal between groups is not random but contingent on the proportion of cooperators in the group.12 Both cooperators and defectors are free to come and go as they please, but the net result of free movement is an above-random segregation of cooperators and defectors into separate groups, which favors the evolution of cooperation by between-group selection. These models were developed with small ephemeral social groups in mind. Essentially, they are models of friendship that complement models of kinship, in which the segregation of cooperators and defectors into separate groups is due to genealogical relatedness.  But what you describe for the Greek Poleis—for example, Athens as “a magnet for an emerging class of intellectuals”, sounds similar. In general, to what extent did free movement contribute to cultural variation among the poleis?

JO: I think free movement was very important. Keep in mind that many states of the Greek ecology shared a language and other cultural features. So, the cost of “walking away” was relatively low. The historian Thucydides proposed that in very early times (he would be thinking, in our terms, of the late Dark Age), Athens was already a magnet for those who had lost out in power struggles elsewhere in the Greek world. Whatever the truth of that, it is certainly the case that there is a lot of movement, short and long-term, of people around the Greek world in the classical era. Some of that movement was structured: Greek colonies were established in, for example, Sicily and south Italy, and later on the coasts of the Black Sea. Greek colonization stories often concern internal conflicts in the “home country” that led to the expulsion or voluntary departure of those who left on colonizing expeditions. Some of those colonies in turn became major states and sent out colonies of their own. Overall, there was a good deal of “population churn” due to political conflict, and to individuals seeking their fortunes where they believed they would have the best chance. This is all very different from the conditions that many historians assumed pertained in static agricultural societies – what the French Annales School called the “longue durée.” An assumption of minimal change, with people staying close to where there were born, was standard in medieval historiography and influential in scholarship on the Greek world until recently.

DSW: Now I’d like to consider the cultural evolution of four kinds of structure that enable cooperation and functional organization to take place at larger scales: federalism, institutions, trade, and interstate alliances. Beginning with federalism, I was fascinated to learn that Athens developed a multi-tier governance structure. The lowest level, called the deme, consisted of 150-250 free adult males, which fits nicely with Robin Dunbar’s well-known work.13 The next level, which you call a “tribe” fascinates me because it appears to be entirely made up in a way that is designed to integrate coastal, inland rural, and urban populations, which would not otherwise consider themselves part of the same group. Is this an example of the conscious democracy-building efforts that we discussed previously? Please comment more generally about how Federalism emerged and why it was important.

JO: The Athenian system of demes (politically recognized villages) grouped into completely artificial “tribes” (with membership drawn from demes from 3 distinct parts of the territory controlled by Athens) was indeed a “federal” system, in that it enabled local identities and interests and knowledge to be integrated into a “national” system of government. It was a brilliant idea, attributed to the legislator Cleisthenes, and institutionalized in the aftermath of the revolution that established democracy in Athens. Meanwhile, in other parts of Greece, federal systems were worked out between city-states. Shared citizenship facilitated both trade and military security, allowing groups of small states to compete effectively with much larger rivals. The study of Greek federalism was put on a new and sound footing by Emily Mackil’s fine book, Creating a Common Polity: Religion, Economy, and Politics in the Making of the Greek Koinon

DSW: Proceeding to institutions, you have already commented on how they incentivize democratic governance. Please tell us more about how they arose and the roles that they play. Is this another example of conscious cultural evolution?

JO: The Greeks realized that cooperation at scale was difficult, especially when really dangerous undertakings (like war) required voluntary buy-in by people with very different positions in a social-economic order: by rich and poor, aristocrat and commoner, farmer and trader, and so on. The development of democracy at Athens after the “big bang” of the democratic revolution and the deme/tribe reforms of Cleisthenes, show a lot of awareness of the need to keep ordinary and elite Athenians “on the same page.” Those institutional reforms included pay for political service (making participation practical for the poor) and the chance to win honors for the wealthy (allowing them the prestige that they desperately desired). Cooperative participation in the community also offered other incentives – notably the chance to expand valuable social networks. Citizens gained desirable roles in the system of collective self-government both by lotteries (thus enabling the relatively disadvantaged to take part) or election, allowing those with merit and ambition to compete for especially important positions.

DSW: Proceeding to trade, modern economists rightly point out that markets are remarkable for enabling cooperation and coordination among people who are total strangers to each other. Yet, the metaphor of the invisible hand notwithstanding, unregulated markets do not automatically function for the common good. The role of the state in shaping and regulating markets is a hot contemporary topic, so it will be fascinating to learn from you how they culturally evolved in ancient Greece and the degree to which they were part of conscious democracy-building efforts.

JO: The Greek interstate economy was highly market-driven, as Alain Bresson, The Making of the Ancient Greek Economy: Institutions, Markets, and Growth in the City-States has demonstrated in wonderful detail. But internal economies were to a degree regulated: market inspectors established weights and measures and penalized cheating. In Athens, state officials (indeed highly expert state-owned slaves) were available to those trading in the market to test and thereby guarantee the values of coins that were used in exchange. These officials (and other state businesses) were paid for by levying taxes on imports and exports (usually at a rate of 2%). The Greeks seem to have had a sense that free trade drove prosperity, but also that without regulation, trade can degenerate into piracy.

DSW: Finally, we get to interstate alliances, which is the emergence of a new tier of multilevel governance. How did they form—and just as important, how did they fall apart? Did the member states insist on the same democratic rights that existed on a smaller scale?

JO:  Alliances ranged from very strong and long-lasting (as in the federal leagues studied by Mackil) to short-term and relatively ephemeral mutual defense pacts. Powerful states, like Athens, Sparta, and Thebes, attempted to create long-lasting alliance systems that would benefit themselves. These alliances were established with greater or lesser degrees of coercion – and were seldom fully successful for very long periods of time. The Athenians had some sense that supporting democracies abroad tended to stabilize their alliance network, but they never pursued a systematic policy of democracy promotion. Member states in alliances increasingly demanded, and got, guarantees of fair treatment (e.g. no military garrisons on their home territories), but this was a matter of negotiation, not a recognition of inherent “rights.”

DSW: In my introduction, I identified the ability to suppress disruptive self-serving behaviors within a group as a key requirement for a MET. Presumably conscious democracy builders of ancient Greece were well are of this. What are some of the mechanisms that they invented, or which otherwise emerged, to suppress disruptive within-group selection?

JO: Greek systems of law (within a given polis) were all about suppressing self-serving behaviors. Ultimately the system of justice and punishment aimed at creating norms of self-moderation, whereby the residents of a territory would voluntarily choose to refrain from self-aggrandizing behavior. In the first instance, they refrained because of fear of detection and punishment. But over time, the result of the system of laws was the internalization of self-moderation as an ethical norm. This is made intentional and explicit in the philosophical works of Plato and Aristotle, but the process is quite clear in the workings of the well-documented legal system of Athens – and (although less well documented) other Greek states. Among the most interesting mechanisms, and perhaps most disturbing from a modern “rights” standpoint, was ostracism. In a given year, one Athenian could be expelled from Athens for a period of 10 years, by a plurality vote of his fellow citizens. Ostracism did not require a trial or even charges of criminal wrong-doing. It might be seen as an example of “majoritarian tyranny” over the individual. But notably, the Athenians were very moderate in its use: only about a dozen ostracisms were actually carried out in the course of the 180-year history of the full democracy.

DSW: Now let’s focus specifically on the powerful elites. Generally, they are part of the problem, wishing to run the poleis for their own benefit, just as in modern times. But some elites become patrons of democratic governance, such as Franklin Delano Roosevelt in modern times, who was called “a traitor to his class”. What role did elites play in Greek democratic governance, both positive and negative?

JO: The book I mentioned above, Mass and Elite in Democratic Athens, dealt with exactly that question. My conclusion was that the “symbotype” of Athens, in which mass audiences responded vocally and by voting to proposals (and legal conflicts) between elites served to control elite aspirations and reign in the tendency of elites to seek to capture the state.  The tension between mass and elite remained, but it was a dynamic one, in that highly ambitious elites were given the opportunity to take important leadership roles, if and only if they demonstrated fealty to democratic ideals. That demonstration was both in their public speeches, but also in the observable conduct of their lives. Elites who lived as if they scorned the many (think gold toilet seats) were despised and unlikely to gain the positions of influence they sought. Moreover, they were likely to be looked at askance by large juries of ordinary citizens when, as inevitably happened, they came into legal conflicts with their elite rivals. Once again, the incentives encouraged Athenians to “look down the game tree” – and to moderate their behavior accordingly. This did not mean “dumb down to the lowest common denominator,” far from it: those who sought responsible positions of leadership without also demonstrating competence were quickly rejected. But it did mean “public displays of anti-democratic arrogance had high costs and thus low expected utility.”

DSW: In my introduction, I emphasized two dimensions of moral systems: a compulsory dimension that demands conformance to norms, and a voluntary dimension of wanting to help others and one’s group as a whole, motivated by emotions such as sympathy, empathy, friendship, patriotism, and love. These must go together because the compulsory dimension makes it safe to express the voluntary dimension. We have just covered the compulsory dimension in our discussion of social control. Can you now comment on the voluntary dimension? One reason I ask is that your emphasis on “self-consciousness about the micro-foundations of rationally cooperative behavior” might be taken to imply that the Greeks framed democracy entirely in terms of “what’s in it for me?” self-interest. I’d be surprised if there were not also an ample dose of public-spirited behavior.

JO: Exactly as you say, in the Athenian democracy, for example (as opposed to the Spartan aristocratic oligarchy) the compulsory part made it safe for individuals to be empathetic, public-spirited, and to trust their fellows, without risking a “sucker’s payoff” – as they would in a classic Prisoner’s Dilemma game. The creation of a public culture of “civic friendship” (to use Aristotle’s term) was essential to the Athenian success story, and to the success of other Greek poleis. The ideal of public service, grounded in a rational recognition of the credible commitment of others to that same ideal, is strongly expressed in much Greek literature: for example the famous “funeral oration of Pericles” in Book 2 of Thucydides’ great history of the Peloponnesian War.

DSW: Now I’d like to explore the exclusion of women and slaves from democratic governance. Was gender equality and universal democracy among men ever even discussed? If I understand correctly, there was not a big ethnic divide between masters and slaves. It was more a matter of such things as who was victorious in warfare. In general, how did the Greeks justify the exclusionary nature of their democracies?

JO: Gender equality was indeed broached on a philosophical plane – notably in Plato’s Republic, where the warrior-ruler class of Guardians included men and women on equal terms. And in his comedy Assemblywomen, Aristophanes imagined an alternative Athens in which the women ran the government on egalitarian, even communistic lines. But notably, even in Aristophanes’ alternative world, the women-rulers maintained familiar conditions of slavery. Many slaves were non-Greeks (for example, Thracians from what is now Bulgaria and Scythians from the Black Sea region) – but Greeks also enslaved other Greeks, especially as prisoners of war or when a polis was utterly destroyed by a rival. Slavery was typically thought of as a matter of bad luck: while no one would choose the life of a slave; some had the misfortune of being enslaved. The Greeks had no fantasies about “happy slaves” and Aristotle’s notion that some persons were “slaves by nature” seems to have been his own idea, and not widely shared. Given that it was seen as misfortune that could befall anyone, rather than a moral wrong, the Greeks felt little need to justify slavery. They saw that all other peoples known to themselves employed coerced labor of some sort or another. Although some Greek thinkers regarded slavery as in some sense “unnatural,” we do not have evidence of anything like an abolition movement. On the other hand, manumission of individual slaves was fairly common (certainly not universal: mine slaves surely usually died as slaves), and sometimes more or less contractual: Skilled slaves could and did buy their own freedom with money they earned above and beyond that which they paid over to their masters.

DSW: In the same vein, it seems that warfare was a never-ending feature of Greek life, taken for granted as a way to acquire resources, along with farming and trade. Did anyone even imagine a world without war? Sparta is a particularly frightening example, to me at least. You describe it as a predatory culture pure and simple, not even extending rights to the farmers who provided food for them! In ecological and cultural evolutionary terms, because Spartans could become specialized as full-time warriors, they were victorious in battle and therefore had a niche in a multi-niche cultural ecosystem, like lions and hyenas on the African savannah. Morality had nothing to with it, other than to organize democratic governance within their own ranks. Please help me make sense of this, against the background of our modern sensibilities!

JO: Like slavery, war was generally taken as a given of human experience by the Greeks. There were some calls for an end to intra-Greek warfare (by, for example, Plato in the Republic and the orator Isocrates). And in the fourth century BCE, the age of Plato and Aristotle, there were several attempts to create a “common peace”- an agreement among leading Greek states to cease hostilities indefinitely. But those attempts invariably broke down due to the ambitions of one or another of the parties. Peace was regarded as a blessing (indeed as a kind of divinity), and it was associated with prosperity and welfare. But inter-state rivalry meant that war remained endemic until the Roman takeover. Since the Romans did not tolerate large-scale violence within their empire (except by themselves), that finally put an end to warfare in the Greek world – for a while. Sparta was a special case of military specialization – highly effective as a rentier-state, Spartan institutions were designed to enable a highly cohesive elite of violence specialists to extract resources from a much larger population of dominated agrarian laborers. In the end, the system proved unsustainable, as other states proved more innovative in devising new military tactics – and because the Spartans’ own practice of weeding out Spartans who had failed in any of their manifold public duties undermined the demographic basis of the Spartan army. By the time of Aristotle, there were only about a thousand rights-holding Spartan families left, down from a high of perhaps 9000. Highly adaptive at its outset, the Spartan system was doomed to demographic failure by its own severe logic.

DSW: Now I’d like to focus on elements of Greek culture that appear non-economic. In modern life, economics seems to exist as a silo that is completely isolated from philosophy, culture, and the arts. But historically, all aspects of a culture are intimately entwined. One of my favorite examples is Venice: Lion City: The Religion of Empire, by Garry Wills, which shows how all aspects of Venetian life, including its own version of Christianity and the rich artistic culture for which it is famous, were entwined with the politics and economics of a mercantile empire. In the case of classical Greece, how was the efflorescence of culture for which it is so well known connected to economics and democratic governance?

JO: For a long time, ancient Greece was imagined, rather romantically, as a world in which general poverty was mysteriously conjoined with extraordinary cultural achievement. In fact, the astounding achievements of the Greeks in various cultural domains (literature, philosophy, science, medicine, architecture, visual arts, performing arts) coincided with a surprisingly high and sustained level of economic growth. As in Venice, or the Dutch republics of the 16th and 17th century, wealth promoted high culture both through private channels of patronage and through state channels. The latter is especially evident in democratic Athens, where the Athenian state-sponsored architectural and sculptural masterpieces (notably the Parthenon), drama (comedy and tragedy), and other visual and performing arts. Meanwhile, private wealth provided the leisure that was the precondition for the development of philosophy, historiography, and so on. Democracy was clearly a stimulus to culture in that it fostered a norm of innovation. Armand D’Angour, in his book The Greeks and the new: Novelty in Ancient Greek Imagination and Experience, has underlined the positive evaluation of “the new” in Greek, and especially democratic Athenian, culture. Competitors in various fields (including formal state-sponsored contests) therefore sought to stand out by their innovations, as well as by their proven excellence in established domains. We can track the Greek competitive drive to bold innovation in many fields, including vase-painting, drama, and science. The growth of wealth, along with the constraints on inequality (progressive taxation) contributed to democratic, trade-oriented Athens being at the center of those cultural developments.

DSW: In his book, Money and the Early Greek Mind: Homer, Philosophy, Tragedy, the classics scholar Richard Seaford specifically attributes the cultural evolution of Greek analytic thought to the invention of coinage. Do you have an opinion on this specific thesis?

JO: I tend to shy away from techno-sociological explanations of historical change. I think Seaford is right to claim that the quick adoption of coinage by a number of Greek states, and the move away from electrum (an alloy of gold and silver – meaning that the metal value of the coin was very hard to determine) to a standard, pure silver coinage, facilitated the development of Greek trade and culture alike. But I don’t think that it is possible to demonstrate a causal arrow from “coinage” to “culture.” The basic idea of money (denominating value in a given weight of precious metal) long antedated coinage and the Greek economic and cultural efflorescence. I’d tend to see the widespread adoption of the use of money in the form of coined silver as one of several innovations that arose in the context of the competitive/cooperative multi-state polis ecology.

DSW: Now let’s finish up with the decline of Greek democracy. Why did the Athenian empire fail in competition with other empires…

JO:  Athens built an empire, ultimately dominating hundreds of small states in the eastern Mediterranean, in the aftermath of the great war against Persia. The Athenian empire lasted for two generations, from, roughly, the 460s to 404 BCE, when Athens finally lost the 27-year Peloponnesian War. It failed, at least in part, because the Athenians never found an effective way to scale up their highly participatory system of citizenship: Tribute-paying imperial subjects gained in some ways from being under Athenian control (protection against Persia and pirates, facilitated trade through a central market and a standard coinage) but they were not even “semi-citizens” and eventually the cost-benefit equation turned against Athens, leading to widespread revolts. It is, however, a mistake (albeit a common one) to correlate Athenian democracy with empire. Athenian democracy emerged a long generation before the empire, and it lasted for another three generations after the end of the empire. As it turned out, Athens, as a Greek-style democracy, was not very effective as an imperial hegemon. The really long-lasting, geographically extensive, and ethnically diverse empires of the ancient western-Asian/Mediterranean world (Persia, Rome) were not democratic.

DSW: …and how did Greek culture continue to thrive after being subsumed by larger empires? What I infer from your book is that democratic culture had become so deeply entrenched and was so successful at the scale of the single polis or federation of poleis that their conquerors were content to extract rent from them and otherwise leave them alone. Is that roughly correct?

JO: Yes. As the historian John Ma has shown, in work in progress, the Greek city-state form, and the core cultural norms of polis citizenship, proved to be remarkably robust. Ma suggests that the real golden age of Greek democracy – at least in terms of the number of city-states with democratic internal government – may have been after the incorporation of many of the city-states into the imperial kingdoms established by the Macedonian successors of Alexander the Great. The Macedonian kings realized that crushing local self-governance would simply be too costly, so they allowed local self-government and set taxes at a moderate level. That had the effect of sustaining relative prosperity across the ecology of city-states. Even the Roman conquest did not eliminate polis culture, in Ma’s view – indeed, he argues that there are discernible traces of polis civic culture into the fourth century CE.

DSW: Finally, one major form of cultural evolution is based on imitation rather than conquest. Greek culture appears to have been highly contagious—Hellenization—even after it lost political and military dominance. What accounts for its contagious quality?

JO: Hellenic polis culture was indeed highly attractive to people on the periphery of the core Greek world – notably to the Etruscans and Romans in Italy, to the Macedonians to the north and east of core Greece, and to local dynasts on the fringes of the Persian empire in what is now western Turkey. Greek trade goods and artistic forms extended much further – into what is now France, Egypt, and the steppes of eastern Europe and western Asia. Most of this cannot be explained by imperialism or colonialism, in any ordinary sense of those terms. As you say, Greek culture was taken up by others long after – and geographically far beyond – when and where the Greeks were able to assert political or military power. I suppose that Greek culture was attractive because of its perceived quality – because Greek pottery and architecture were regarded as beautiful as well as functional, Greek philosophy as profound, Greek drama as moving, and so on. The decentralized competitive/cooperative ecology of Greek communities had, in brief, resulted in the creation of cultural products (arts and institutions alike) that were (and are) recognized as worthy of admiration and thus of emulation.

DSW: Thank you for taking this deep dive with me! To conclude, what are the most important lessons that Greek democracy teaches us about our current moment in world history?

JO: Perhaps just these three points. First, that a centralized system of spatial and economic organization (the “empire model”) with a top-down system of authority (“king or commissars”) is not the default, nor even necessarily the most adaptive form, of human social order. Nor is it the only way to produce welfare or cultural advances. An ecology of many relatively small communities, some of them highly democratic, engaged in both cooperative and competitive relations, was, and is, an alternative. Whether one form will ultimately drive out the other remains to be seen, but human history offers no reason to suppose it will. Next, as we apparently need to learn time and again, democracy (in practice rather than just in name) is not about to become the default form of human political organization. Democratic citizenship is an equilibrium that can be quite robust, once established, but it is not easy to get up and running.  Sustaining democracy requires compromises and a recognition that self-interest must be moderated by a sense of the longer-term value of voluntary, non-coerced, local cooperation. And finally, democracy ought not to be confused with ideal justice. Neither by our standards nor indeed by their own was democratic Athens an ideally just community. Nor, of course, is modern America. Perfectly just communities are indeed imaginable. Greek philosophers living in democratic Athens inaugurated utopian thinking; democracy encourages critical inquiry into the sources of injustice. But in the real world, democracy is a bargain among competing interests, leavened by a shared commitment to a common good. It gives many, although never all, persons residing in a bounded territory a chance to be active participants in determining their own collective fate. Adopting Aristotle’s terms, it enables us, as citizens, to exercise our natural capacities, to be most completely the kind of social, communicative, reason-using, political animals that we are. And that, for me and for at least some others, is of great value.


[1] Margulis, L. (1970). Origin of Eukaryotic Cells. New Haven: Yale University Press.

[2] Maynard Smith, J., & Szathmary, E. (1995). The Major Transitions in Evolution. New York: W.H. Freeman.

[3] Wilson, D. S. (2015). Does Altruism Exist? Culture, Genes, and the Welfare of Others. New Haven, CT: Yale University Press.

[4] Aktipis, A. (2020). The Cheating Cell: How Evolution Helps Us Understand and Treat Cancer. Princeton: Princeton University Press.

[5] Wrangham, R. (2019). The Goodness Paradox: The Strange Relationship Between Virtue and Violence in Human Evolution. New York: Pantheon.

[6] Boehm, C. (2011). Moral Origins: The Evolution of Virtue, Altruism, and Shame. New York: Basic Books.

[7] Wrangham, R. (2019). The Goodness Paradox: The Strange relationship Between Virtue and Violence in Human Evolution. New York: Pantheon; Hare, B., & Woods, V. (2020). Survival of the Friendliest: Understanding Our Origins and Rediscovering Our Common Humanity. New York: Random House.

[8] Deacon, T. W. (1998). The Symbolic Species. New York: Norton.; Jablonka, E., & Lamb, M. (2006). Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life. Cambridge, MA: MIT Press.

[9] Richerson, P. J., & Boyd, R. (2005). Not By Genes Alone: How Culture Transformed Human Evolution. Chicago: University of Chicago Press.

[10] Wilson, D. S., Hayes, S. C., Biglan, A., & Embry, D. (2014). Evolving the Future: Toward a Science of Intentional Change. Behavioral and Brain Sciences37, 395–460.

[11] Wilson, D. S., Ostrom, E., & Cox, M. E. (2013). Generalizing the core design principles for the efficacy of groups. Journal of Economic Behavior & Organization90, S21–S32. https://doi.org/10.1016/j.jebo.2012.12.010

[12] Aktipis, C. A. (2004). When to walk away: Contingent movement and the evolution of cooperation. Journal of Theoretical Biology231(2), 249–260.

[13] Dunbar, R. I. M. (1996). Grooming, Gossip and the Evolution of Language. Cambridge, Mass: Harvard University Press.


Josiah Ober

Josiah Ober, Mitsotakis Professor in the School of Humanities and Science, Stanford University, works on historical institutionalism and political theory, focusing on democratic theory and the contemporary relevance of the political thought and practice of the ancient Greek world. He is the author of Demopolis: Democracy before Liberalism (2017), The Rise and Fall of Classical Greece (2015), and other books, mostly published by Princeton University Press, including Mass and Elite in Democratic Athens (1989), Political Dissent in Democratic Athens (2008), and Democracy and Knowledge (2008). He has also published about 100 articles and chapters, including recent articles in American Political Science ReviewPhilosophical Studies, Polis, Public Choice, Critical Review, and Transactions of the American Philological Association. Work in progress includes books on instrumental rationality in classical Greek thought and the role of civic bargains in the emergence and persistence of democratic government.


David Sloan Wilson

David Sloan Wilson is SUNY Distinguished Professor of Biology and Anthropology at Binghamton University. He applies evolutionary theory to all aspects of humanity in addition to the rest of life, both in his own research and as director of EvoS, a unique campus-wide evolutionary studies program that recently received NSF funding to expand into a nationwide consortium. His books include Darwin’s Cathedral: Evolution, Religion, and the Nature of SocietyEvolution for Everyone: How Darwin’s Theory Can Change the Way We Think About Our Lives, and The Neighborhood Project: Using Evolution to Improve My City, One Block at a Time and Does Altruism Exist? Culture, Genes, and the Welfare of Others. .