Journal for General Philosophy of Science

, Volume 48, Issue 3, pp 341–369 | Cite as

Why Social Science is Biological Science

  • Alex Rosenberg


The social sciences need to take seriously their status as divisions of biology. As such they need to recognize the central role of Darwinian processes in all the phenomena they seek to explain. An argument for this claim is formulated in terms of a small number of relatively precise premises that focus on the nature of the kinds and taxonomies of all the social sciences. The analytical taxonomies of all the social sciences are shown to require a Darwinian approach to human affairs, though not a nativist or genetically driven theory by any means. Non-genetic Darwinian processes have the fundamental role on all human affairs. I expound a general account of how Darwinian processes operate in human affairs by selecting for strategies and sets of strategies individuals and groups employ. I conclude by showing how a great deal of social science can be organized in accordance with Tinbergen’s approach to biological inquiry, an approach required by the fact that the social sciences are all divisions of biology, and in particular the studies of one particular biological species.


Biology Darwinian theory Cultural evolution Strategies Packages of strategies Functions Selected effects 

1 Introduction

Homo sapiens is a biological species. That should be enough of a basis on which to argue that the social sciences are biological sciences, must proceed in accordance with the explanatory and evidential strictures of biology, and cannot be expected to transcend whatever limits biology faces as a domain of scientific inquiry. This is not just the claim that whatever social science turns out to be, it will be a compartment of biology because its domain is exhausted by Homo sapiens, a biological species. It is the claim that shape and strength of explanations, theories, models and regularities in the social science will be those we are already familiar with in biology, as will be the causal concepts invoked by them. This is not a reductionist thesis, in either of the two prevalent senses. It does not require that explanations and theories among the social sciences to be somehow derived from more fundamental ones in biology (though they will have to be so derived, if reductionism obtains in biology). Nor does it reduce the range of causal or explanatory variables to some subset of biological traits or properties, such as the genetic ones. The thesis is that social science requires the employment of experimental and field-study methods familiar from biology, and when successful results, in findings, theories, and models of human behavior and of human groups no different in kind from those biology is familiar with.

The first two sections of this paper offer another reason why the social sciences must be biological that may be more compelling and that provides more guidance about how they should proceed than the anodyne observation that human beings are biological creatures. In particular the argument underwrites a Darwinian approach to human affairs, but not a nativist, innatist, or genetically driven framework. In the third section we see how the argument establishes limitations on the explanatory aspirations of alternative theories, including especially rational choice theory, the currently most fashionable explanatory approach in several social and behavioral sciences. Then in Sect. 4 one equally fashionable objection to a biological approach to human affairs is disposed of. The fifth section develops a positive though still relatively general account of how Darwinian processes operate in human affairs. The last section explores how a research program about human affairs driven by Darwinian approach should proceed and how much of current, conventional social science can be expected to “recover.”

2 The Taxonomies and the Regularities of the Social Sciences Must Be Biological

There is a fairly direct argument for the claim that the taxonomies of the social sciences and the spatiotemporally restricted regularities, ceteris paribus laws, a priori mathematical models, and for that matter historical narratives they invoke to explain phenomena in their domains, identify biological categories and causal relations among biological categories.
  1. 1.

    Almost all significant features of human affairs—historical actions, events, processes, norms, organizations, institutions, etc.—have functions.

  2. 2.

    Functions are all adaptations,1 and the only source of adaptation in nature—including human affairs–are Darwinian processes of blind variation and environmental filtration.2


  3. 3.

    All regularities about the character and relationships among adaptations (or their direct results) are local evolutionary equilibria, which are eventually broken up by “arms races.”3 All biological regularities are “restricted,” limited in their spatiotemporal domains, and draw their explanatory power from underlying unrestricted Darwinian regularities.


  4. 4.

    All restricted regularities about human affairs are local equilibria (or their consequences) and are eventually broken up by arms races.

  5. 5.

    Regularities about human affairs have explanatory power because they are underwritten by unrestricted Darwinian regularities.

  6. 6.

    A social science in the business of explaining human behavior and human affairs should proceed by explicitly Darwinian research strategies—identifying adaptations and the processes that determine their emergence, proliferation, persistence and extinction.

The first two premises are controversial. The rest of the argument reflects fairly well established conclusions in the philosophy of biology about the nature of biological laws, models, theories, and explanations more generally. This section begins the process of justifying them. The task is not completed until the end of Sect. 2.

Premise 1 may seem dubious at first blush. How could almost everything in human affairs have a function and so be an adaptation? That sounds like an idea worthy of Pollyanna or Voltaire’s Dr. Plangloss. Even in biology, not everything turns out to be an adaptation. Much of evolution is a matter of drift—the play of chance on small and sometimes even large populations that leads to changes in the distribution of adaptations, and even to the persistence of non-adaptive and maladaptive traits. Moreover, important biological traits are themselves either the result of physical constraints or were acquired as adaptations early enough in evolutionary processes to remain fixed long after they ceased to be adaptations. Surely all the same must be said of the course of human affairs. Indeed, for obvious reasons, there may well be a greater role for drift and constrain in human affairs than biological processes.

Of course premise 1 needs to be understood as qualified by the reality of drift and constraint in human affairs. In fact the plausibility of the claim that premise 1 makes about the adaptedness of most features human affairs relies a great deal on the qualification ‘significant’. There will be many features of human affairs that are the result of drift, and yet little of what interests social scientists about human affairs is the result of random drift alone or even mainly. Similarly, social scientists will recognize constraints of many kinds as forcing subsequent features of human affairs to adapt to them. But few social scientists accord such constraints the fixed character that constraints—especially physical ones—have in biological evolution. In fact, the most revolutionary social changes break down the oldest, firmest, and most pervasive constraints. The real issue is whether such change is the result of blind variation and environmental filtration—cultural selection.

A little reoriented reflection on human affairs does suggest that, even more than in biology, significant features of social life are largely or even wholly functionally beneficial–for someone, or some group, or some practice.

The notion that all, most, many or at least some human social institutions have functions is not a new one. The idea goes back to Durkheim, and forward beyond Parsons. These 19th and 20th century functionalists were right about the functional character of almost all social institutions. But a serious oversight in their analysis condemned it to implausibility, and it went into eclipse long ago. The simple error functionalists made, which made their view sound so implausible, was to mis-identify the beneficiaries of the functions that institutions, practices, and organizations fulfilled.4 They assumed, quite myopically and wrongly, that the function of institutions, practices, organizations, was to fulfill the needs of people, of human beings. But it was obvious that many institutions, practices, organizations are in fact harmful to people, confer no net advantage or benefit on them, for instance most religions, or Chinese foot-binding, or tobacco smoking. This Panglossianism about all social institutions made functionalism a laughing stock when it was not pilloried as an invitation to complacence and conservatism: if almost all human institutions fulfilled functions for us, then it is tempting to reason that we should not change them lest we deprive ourselves of the benefits they confer on us. Whence the charge of complaisance and conservativism.

Only in the late 20th century did it become apparent that in these and other cases, a change in perspective—a Gestalt switch– would enable us to see what was not previously apparent: the relevant beneficiaries of those features of institutions, practices, organizations harmful to people were the institutions, practices, organizational structures themselves. Institutions, organizations, practices that, so to speak, prey on, parasitize people, treat people as niches, environments to be exploited in ways that enhance the persistence or replication of the trait with that feature. To vindicate functionalism we need to begin to think of people as the environment and think of types of institutions, practices and organizations as the things that survive, replicate, and spread or recede and become extinct owing to the degree their features exploit human characteristics. Institutions, practices, organizations emerge, persist, spread owing to how well their features function to encourage people to participate in them. Viewed this way the functionalist perspective becomes more difficult to resist: many socially significant institutions, practices, organizations, confer huge net benefits on people—money, the firm, the market price system. Many others confer huge net harms on people, but in so doing ensure their own persistence—think again of foot binding or tobacco smoking or heroin addiction. Other institutions confer benefits on some people, and harms on others—slavery for example. Most institutions–religions, for example– confer a mixture of harms and benefits on different mixtures of persons over time.

One way to effect the gestalt switch necessary to accept thoroughgoing functionalism about human affairs is to employ the game theorist’s notion of a ‘strategy.’ A ‘strategy’ is simply a rule, norm, procedure, of the form ‘Under condition X, do Y.’ Strategies may be reflexive or voluntary, moral, or ritual, matters of fashion or style, short-lived or not, obligatory or optional, complex or simple, consciously followed or not, beneficial to the agent employing them or harmful to him or her. People’s behaviors are determined by and express reveal, manifest the strategies they internalize. These strategies are traits, quasi-hard wired ones like left-handedness, or learned, like speaking French, or wearing miniskirts, that can come and go. They are acquired or suppressed (as in the case of left-handed), by social learning, by imitation or enforcement, by unconscious classical and operant conditioning, and transmitted from person to person. Individuals’ strategies interact with other strategies, cooperating with them, competing with them, subordinated to them, or subordinating them. Human social institutions, from a book club to Feudalism, are nested sets of coordinated strategies. Think of practices like patrilateral cross cousin marriage or purdah or the incest taboo. Think of organizations like the free masons or the parish council. Human affairs are a matter of nested institutions, organizations, practices, all composed of the strategies individuals employ. Then there are the strategies each individual employs to navigate through these institutions, organizations, practices—these packages of strategies. The institutions, organizations, practices have functions. They thrive or perish depending on how well the strategies they impose on people enable the institutions, organizations and practices to fulfill these functions for their beneficiaries—often themselves.

Focusing on the stategies instead of the humans whose behavior is described (and perhaps directed by them) enables us to embrace the thoroughgoing functionalism of theorists like Durkheim and Parsons without the Panglossian implausibility that daunted their approach. Think of strategies as symbionts, or parasites, or sometimes combinations of both, living on human life, and changing it for the better or for the worse, but always adapting to insure their own survival. (Much more on this approach below.)

It is difficult to think of tobacco smoking, or heroin addiction as having functions, because they are harmful. Their functions are not functions for us, but for themselves. To see this we need first to see how smoking or foot binding function in an environment composed of humans. They are sometimes (net-)beneficial and at other times (net-)harmful to humans, but they are practices with features that function to ensure their persistence and spread through human history at least until their environments change and their effects start to be selected against. Chinese foot binding is a clear example of how this works. Foot binding persisted for about 1000 years in China. It got started because women with bound feet were more attractive as wives. Bound feet were a signal of wealth, since only rich families could afford the luxury of preventing daughters from working. Girls with bound feet were easier to keep track of and so likelier to be virgins, etc. So, at first, when the practice arose, foot-bound girls had more suitors. Pretty soon every family that could afford it was binding daughters’ feet to assure they would get married. Result: when every girl’s feet were bound, foot binding no longer provided an advantage in the marriage market, and all foot-bound girls were worse off because they could not walk, suffered other health effects, etc. Foot binding starts out as an adaptation for some girls, and for some families, but by the time it becomes really widespread and fixed, it is actually a physical maladaptation that lowered every foot-bound girl’s fitness. But once every one was doing it, no one could get off the foot-binding merry-go round. Any one who stopped binding daughters’ feet condemns them to spinsterhood. Here we have a tradition, a norm—Bind daughters’ feet!–that by the time it was widely adopted ceased to convey any benefit on the people whose behavior it governs. Why did it persist despite its maladaptive effects on foot-bound girls? For whom or for what were its features adaptations? For itself, for the practice, norm, institution of foot binding! The practice persists, like a parasite, because of those of its features that functioned to exploit the “weaknesses” of humans and their institutions—marriage, the desire for virgin-brides and large dowries, the desire to control women before and after marriage.5

Once we widen the range of the possible beneficiaries of a function, the claim that almost everything of interest to social scientists in human affairs has functions becomes far less Panglossian. But can it be correct? Here is another deep reason to suppose that the taxonomies of all the social sciences individuate functionally: The vocabularies of all ordinary languages do so predominately. That is, almost every common noun in all languages is defined in terms of the causes and effects, especially their effects. Few things are defined in ordinary language even partly in terms of their material composition. The same goes for all the sciences, especially at their inceptions. There is not really any alternative, given that the goal of many sciences is to identify the structures that produce effects of interest. That is why they start with taxonomy of kinds characterized by effects. When it comes to biology, the effects we notice first and hit upon to define them are the one which appear to be functions—to confer benefits, advantages, to meet needs, etc. In fact as biologists have been reminded (Gould and Lewontin 1979), the tendency to taxonomies this way needs to be tempered by careful study of traits of interest to establish whether they really are functions, and if so what their functions are. Mere observation is often misleading about what functions are served and for which beneficiaries. This caution however does nothing to undermine the indispensability in biology of starting off with hypotheses about functions.

The same considerations enforce functional taxonomies in the social sciences, even among social scientists innocent of a biological agenda. It is hard to begin a research program in any other way than by describing the phenomena of interest in terms of effects of interest to the human scientist.

Moreover, the predictive and ameliorative goals that the human sciences impose upon their research programs assume that most of the significant features of human affairs have functions for some individuals and groups, and are dysfunctional for other groups. Though each of the social sciences may be neutral on the functional significance of the actions, events, norms, practices, and institutions in the domains of the other social and behavioral sciences, it will not be agnostic about those within its proprietary domain. This will be true at least so long as a discipline has ameliorative ambitions for social processes in its domain. The remodeling and redesign of political, legal, economic, social, or cultural institutions, rules, norms, and practices would be impossible if these items did not function to benefit or harm individuals, and groups of various sizes and compositions.

So much for premise 1 in the argument above. What of the second premise?

Premise 2 asserts that all functions are Darwinian adaptations, brought into existence, maintained and shaped by Darwinian processes, not biological but social.6 The reasons have only to be stated to be seen as obvious. Once premise 1 is accepted a serious problem confronts the social scientist. It cannot be a mere coincidence that almost everything of interest to social science has functions. Science does not tolerate coincidences on this scale. Either having functions is part of the causal process that brings socially significant features into existence, or their existence and their functions have a common cause. Of course the explanation could be selection bias: the social scientist’s interests are myopically focused only those features of human affairs with functions. Even if selection bias does explain the fact and even if, implausibly, much that is significant in shaping human affairs has no function, it is going to turn out that the only plausible causal process that brought into existence all those features of human affairs that do have functions is a Darwinian one.

Once we rule out accident or drift, there are only a limited number of possible causal processes that can bring about something that has a function: conscious design and fabrication, purposive future causation, immanent teleology, a benevolent deity or Darwinian natural selection. It requires no more than a sentence or two to rule out three of these alternatives. Everything we know about the physical and biological sciences prohibits the existence of and causal role of future purposes. It is not just that events in the future cannot bring events that precede them, the special theory of relativity even rules out simultaneous causation. Aristotelian immanent causation (“entelechies”) is equally untenable on evidential considerations if not also grounds of intelligibility, and inferences from the functionality of human institutions to the existence of a divine deity who planned and implemented them are excluded on similar grounds. This leaves only human intentional design and human contrivance to explain the character of social institutions, practices and organizations, or some process of blind variation and cultural selection.

The grip of human intentionality on the explanatory strategies of the social sciences has always been strong, and has strengthened in recent decades owing to the intellectual imperialism of economics, and the attendant prestige of rational choice theory. There is a widely held but mistaken view that economics has achieved some sort of “take off” as a science more completely than other disciplines. Combined with the mathematical tractability of aggregating individual constrained preference maximization as an account of human affairs, the mistake has encouraged the notion that social institutions, practices and organizations have “microfoundations” in rational choice theory’s formalization of intentional human choices. The unsatisfactory character of such approaches to explaining the functions of social institutions needs to be clearly established. Once this has been accomplished, the inevitability of a biological—a Darwinian—approach to human affairs will be evident.

3 The Limits of Intentional Design and Rational Choice in Human Affairs

In general, social scientists overestimate the role of intentional human design in human affairs. This is especially true among economists, and political scientists who employ the rational choice models economist have developed over the last century or so.7

To see most clearly the limits on intentional human design and rational choice approaches to the nature of human affairs it is best to focus on a number of important examples from the home base of such approaches: economics. So, let us consider how to explain the emergence of three central economic facts: the ubiquity of markets, the emergence of money, and the existence of firms. Each of these institutions fulfills an important need individuals have. None emerged from a rational choice process. It was the Nobel Prize winning economist Friedrich Hayek who earliest and most clearly recognized the problem facing economic theory of explaining the emergence of economic institutions. He called it the problem of ‘spontaneous order.’ But that is just to label the problem, as we will see.

In the case of the firm the human need is to solve a transaction cost problem, as Ronald Coase first noticed only in 1937. Without a solution to this problem, the division of labor must come to a stand still and with it almost all the productivity increases humans have contrived since the middle ages. No rational agent recognized what the problem was that every one faced, no one decided to invent the firm in order to solve this problem. It emerged “spontaneously” to “order” exchanges between individuals in ways that solved a transaction-cost problem. The firm is an example of “spontaneous order.” If the firm was not a conscious contrivance, nor the gift of a benevolent deity, its emergence demands an explanation.

Money solves the biggest problem of barter: what the economists call the double coincidence of wants. Without money if I want oranges and have only banana, I need to find someone who wants bananas and has oranges. What is more, since we cannot divide and store bananas and oranges, I will need to find someone who wants to trade in exact whole numbers of bananas and oranges that match up with the amounts I am prepared to trade. This is a problem that becomes intractable very early in human exchange. How does it get solved? Several times in distant cultures the same solution was hit upon: the emergence of a commodity with common features: portability, divisibility, durability, utility or widespread desirability, and short-term limits on its quantity. When money emerged no one around consciously recognized that it would have to have these features. No one intentionally, rationally chose to adopt some commodity owing to its having the features that solved the problem of the double coincidence of wants.

The emergence of money requires that agents solve another problem, one of coordination. Sooner or later they must all converge on the same commodity to serve as money. People must solve a “common knowledge” problem. Somehow each agent must be willing to adopt a certain commodity as money and must come to believe that every one else will adopt the same commodity, and must believe that every one else will be confident that every other agent has adopted the same commodity. You can see that this is a set of problems that can’t be solved by individual rational choice, that were not solved by some explicit social contract. The institution of money is another example of order emerging without any one intending it or taking steps to bring it about. Of course to say money emerged spontaneously is simply to label the problem. To model the acceptance among large numbers of rational individuals of a commodity as the numeraire, as the optimal solution to a cooperative game is not to solve this problem but to structure it in a way that cries out for a Darwinian approach to its solution: once the optimal outcome is retrospectively recognized, we can set about identifying forces that will select for variations in human behavior that approach and eventually realize it. It’s clear that this process will not involve conscious calculation by individuals of the sort that monetary economists recognize.

The third example, Hayek’s example, the system of market prices, is the most important but the most difficult to understand of these problems of spontaneous order. The unsolvable problem of socialist central planning is informational. Central planning faces the mathematical problem of converting a list of available inputs and a list of desired outputs into a list of production orders, and then continually updating this list as input availability changes and desired outputs change. Central planning faces the further problem of sending information about each of the changes in inputs and outputs only to those who need to have this information in order to change their production plans. The central planner cannot send the changes to everyone: we’d have to spend the better part of every day just trying to find the information we need from a daily massive data dump. But the central planner can no more figure out to whom exactly to send the updated information than it can figure out the initial production order. Updating this order as circumstances, change is still another challenge beyond the powers of rational, intentional choice and planning implementation. These are all what mathematicians call NP-hard problems (“Nondeterministic polynomial-time hard problems”). There is no known algorithmic, computerizable solution to such problems, and a good chance than none exists. This is the fundamental reason the soviet economies collapsed. Even a society composed exclusively of exemplars of New Socialist Man would not be able to solve the relevant NP-hard problem.

Yet the problem is solved all day, everyday, instantaneously by the system of market prices. The market price system is an information storage, retrieval and calculation system—a vast virtual computer—that provides the closest approximation to mathematically correct solutions to the central planners’ calculation problems and at no cost whatever.

The market price system performs a function indispensible not just to modern life, but to all human life beyond the Pleistocene. It meets a need that cannot have been foreseen by humans, no matter how rational. It is a solution to that need that no human or coalition of humans could have filled by intentional design and contrivance. Indeed it is a solution that rational choice would have led individuals to try to undermine or subvert in their own interests. But the solution to the problem people face is so ingenious it automatically and successfully responds to such subversion attempts. Even the strongest exponents of rational choice theory have recognized this feature of markets: their prescription for the elimination of monopoly, externalities, insider trading, and other “market failures” is to leave them alone. The excess “rents” these failure that produce send price-signals to the rest of the rational agents in the economy that will change their behavior and compete away the rents and the market failures.

The market price system operates continually to meet a need that no human or set of humans could fulfill by intentional and deliberate action. The function served by the market price system cannot be met by people, no matter how rational they are, and no matter how powerful and inexpensive their information storage, retrieval and computational resources are. And the market price system emerged, like money, spontaneously, independently, repeatedly and without malice of human forethought throughout human affairs, across the globe.

These three examples of spontaneous order highlight the economist’s version of a problem facing all social sciences. If rational choice theory is incapable of dealing with the problem in its home base, it has few prospects of dealing with it elsewhere in human affairs. But the problem of spontaneous order is pervasive.

Of course rational choice theory is but a formalization of the common sense folk psychology. This theory is at work, especially in narrative history, to explain how many human institutions, organizations and practices were intentionally designed and implemented in order to function in foreseen ways. And it is undeniable that such institutions exist. But institutions, practices and organizations that result from conscious human contrivance—the US constitution, the UN Charter, various diplomatic treaties, are a minority, probably a small minority of human norms, practices and institutions.8 They do not last as long, do not spring up repeatedly and independently, and have far more limited impact on the character of everyday human affairs.

Once we have excluded divine benevolence, future teleology, Aristotelian entelechies, and random chance, the only well established, available mechanism that explains the emergence, persistence, spread and recession and extinction of things with functions is a Darwinian process of blind variation and environmental filtration. The “only” problem that faces the social scientist is to identify the details of this process. Let us see how far we can get in doing so.

4 Regularities in Human Affairs are All Local Equilibria

Step 3 in the argument tells us that all “laws,” regularities, generalizations, explanatory models and theories about human affairs will be at most descriptions of “local equilibria” waiting to be broken up by arms races among the adapted institutions, practices and organizations they describe. Why is this?

If human affairs are mostly the emergence, persistence and improvement of adaptations by Darwinian processes, then we can expect only a limited number of different kinds of regularities about the relationships between them, regularities of the sort already familiar in biology and exemplified in the original argument set out above. To begin with, in human affairs individual adaptations will emerge, then there will be pairs or larger sets of adaptations each of which constitute selective components of one another’s environment, they will either accommodate to one another, or cooperate with one another, over long periods, or will complete with one another over such periods. In either of the first two cases, any variations in either that can exploit to its selective advantage the accommodation or the cooperation will do so. The result will eventually be the emergence of exploitation or a competition somewhere. The exploitation or competition may persist in a local equilibrium. But it must eventually break down into an extinction or an arms race as further variants of each adaptation emerge. The results of the emergence of adaptations and of the persistence of these alternative relations between adaptations are the regularities we recognize in the social sciences.

First, there are regularities about the (real, often latent) function of a behavior or norm or practice or institution: e.g. “Firms (function to) solve the transaction cost problem.” Then there will be regularities about the co-occurrence of adaptations in the same individuals or lineages of them, regularities of co-occurrence of adaptations in two or more distinct individuals or lineages—cases of co-evolution or mutualism, or parasitism (the most famous example of a regularity of co-evolution or mutualism in political science is given below). These regularities will in effect record local equilibria among adaptations, ones that last as long as the historically contingent circumstances that brought them about to obtain.

But, as in biology, each individual’s or group’s adaptation sets a design problem for the individuals and groups with which it finds itself in local equilibrium. The existence of these mutual design problems together with the persistent but blind variation among adaptations means that the prospects for arms races are ever present. Beneath every local equilibrium there is a seething rumble of blind variations continually being tested by and testing the local equilibrium. The latter must always eventually be broken up by one of these variants that precipitate an arms race. Whence the restricted character of every explanatory regularity and all the models in social science.

Examples of these restricted regularities and models are easy to identify. Consider perhaps the most robust regularity in international relations, perhaps even in the whole of political science: Arguably, no two democracies have ever gone to war with one another. There is apparently not a single exception to this regularity since democracies emerged at the end of the 18th century, even though the number of pairs of countries that could have gone to war with each other since 1776 is literally in the several thousands.9 Apparently, the trait of being a democracy and the trait of not going to war with a democracy are for the moment at least co-adapted to each other, and this probably helps explain several things, like why democracies do better economically than even market-economy dictatorships (they engage in fewer wars), and why the number of democracies seems to be increasing. It has also guided foreign policy—the US and European encouragement of new democracies to ensure stability and peace.

But, nothing is forever. We can be confident that somewhere or some when, some democracy is going to find a way to exploit this regularity by attacking some completely unsuspecting fellow-democracy, lulled into a false sense of the permanence of peace among democracies. How can we be so confident? Just as Mother Nature searches through biological design space for variations on that trait that can take advantage of the environment of other adaptations the variation faces, so to it will search for variations in the human social design space. And the rate of variation will be vastly accelerated in comparison to biological evolution. For human evolution does not have to wait a 20-year generation for a genetic variation to change a trait the way that biological evolution must.

One more example, this time of a mathematical model that is explanatory over a restricted domain, but whose explanatory power is destroyed by an arms race breaking up a local equilibrium. Consider again what was for a long time the most influential model in economics, and perhaps even all of social science, the LM/IS graphs and equations of Keynesian macroeconomics. This set of graphs and equations enabled economists of the third quarter of the 20th century successfully to model the stable relationship between sets of macroeconomic variables, including investment and savings, consumption and gross national income, the interest rate and the money supply, and with one another.

The stagflation of the 70’s put an end to the model’s general acceptance, and resulted in its replacement by newer ones, including the rational expectations model. This model explained why the superseded model was no longer a basis for effective intervention. The analysis of why the Keynesian model ceased to work was roughly that, if it ever worked at all, then the co-adaptations it identified, were broken up by an arms race. The model’s widespread dissemination, or at least the fact that economic agents had become acquainted with the governmental interventions it guided, resulted in a change in their choices, one which rendered Keynesian fine-tuning ineffective.

The beautiful and temporarily powerful model of the capitalist economy that John Maynard Keynes inspired ceased to work because the relationships it described broke down once some of the institutions, groups and individuals the model included, began to exploit the fact that other institutions were guided by it, to frustrate the policies the model guided. Result: 10 or 15 years after it became widely known, the Keynesian model became the victim of an arms race.

Examples are not arguments even though they can be proliferated ad infinitum. But there is a simple argument that they buttress and which explains them. It shows exactly why restricted regularities and mathematical models are explanatory and why they are the best we can hope for in the explanation of human affairs.

It is widely recognized among philosophers of biology that there are no completely invariant regularities, no laws, in biology except for those reported in the Darwinian theory of natural selection. A variety of arguments have been offered for this consensus view, of which perhaps the most influential is John Beatty’s (1995) “evolutionary contingency thesis”: all other regularities in biology—from the most invariant to the least—obtain only as a result of the operation of natural selection on initial conditions that have obtained in the history of the Earth, and are subject to abrogation by the operation of natural selection on later conditions.

The general argument is obvious, and it has immediate implications for biological arms races. Since nature builds adaptations by a process of environmental filtration of random variations, when environments change adaptations can become maladaptations and vice versa; variations neutral in fitness in one environment can become adaptive or maladaptive in another one. But nothing is forever: even the most stable environmental conditions will sometimes change, and sometimes even quickly. Consider how the asteroid impact at the Cretaceous-Tertiary boundary 65 million years ago changed the environment and killed off all dinosaurs within a few years. Thus no regularity thrown up by the process of natural selection is immune to breakdown as a result of environmental change.

Once the evolutionary environment comes to include creatures and their effects on one another, the life-times of regularities about creatures’ adapted traits are reduced from the scale of billions of years (archebacteria—whose environment has not changed for 3 billion years) to multiple geological epochs (oxygen-respirators) to hundreds of millions of years (vertebrates) to weeks and months in the case of others (the AIDS-virus). Owing to the role of environmental change, even the most established and long lasting regularities in biology are not as invariant as any well-established regularity of physical science.10

Given the slowness of most environmental changes, regularities about individual species can remain invariant over geologically long periods. Changes such as the shift to an oxygen-rich atmosphere, or continental drift or the on-set of ice ages will break up some invariances and create new ones. Other, more rapidly occurring species-making or species-changing processes such as earthquakes, major droughts, will have similar results. But the invariances produced will be hard to break down for the same reason that massive and long lasting environmental change was required to put them in place.

However, matters begin to change quite radically once members of any species become part of the selective environment of members of another species. In these circumstances regularities about the latter species become comparatively shorter lived, more temporary and more spatially restricted as well. The reason of course is that species become part of one another’s selective environments when they compete, or one predates upon or parasitizes the other, or both are predated by a third species, or both predate a third species, or both cooperate for that matter. Under all these circumstances, nature is persistently searching through design space seeking variations in both species that will provide them with a selective advantage over the other. Until it finds one, the two species are locked into a local equilibrium, one that may be noticed by the naturalists, and be reported in a temporarily invariant regularity. The theory of natural selection however assures us that, if it can, natural selection will break up these local equilibria along with the regularities that describe them and their consequences.11

When traits are genetically coded, the arms race process will be relatively slow, though much faster than non-arms race adaptational change. Since favorable mutations are rare, biological invariances between genetically encoded traits will often be locked in as the result of some relatively long term stable equilibrium in the arms race. Besides interspecific competition, there is also a great deal of intraspecific competition, arms races between lineages within a single species, which also makes and breaks invariances at an even faster rate than intraspecific competition does.

The upshot is that all invariances among genetically encoded traits are restricted. During the periods that they obtain, they are vulnerable to being undermined by random variations that break up co-adaptational equilibria. As the rate of variation increases, the life span of an invariant regularity will decrease as will the spatial range over which it obtains. In the case of competition between very fast breeding species—say, between parasites and their host-targets—for example phage and bacteria or bacteria and humans, regularities may remain invariant only for a few years, months or even weeks. (Consider the lifetimes of antibiotic effectiveness).

In biology, restricted regularities are explanatory and its mathematical models applicable, because they reflect local equilibria underwritten by unrestricted regularities—laws of nature—that Darwin discovered.12 These regularities are restricted owing to the inevitability of arms races ordained by the unrestricted regularities Darwin discovered. Insofar as the social sciences individuate their descriptive taxonomies functionally, and seek regularities that obtain between instances of their taxonomic categories, all of their regularities and models will have to be similarly restricted, as open to exceptions, ceteris paribus hedges, and as predictively imprecise as those of biology. After all, social science is a branch of biology.

We now have a well grounded explanation for why no social science has succeeded in uncovering real laws in its domain, why at best the quantitative models advanced in the most self-consciously mathematical (parts of each of the) disciplines have limited application, unimprovable precision, and are overtaken by events. But the pay-off is not purely negative. We now have the theoretical framework for a research program in the social sciences that expect the kinds of successes in explanation and application that evolutionary biology has secured: identify the functions that manifest themselves in individual and aggregated behavior, and account for them as adaptations by identifying the selective forces that bring them about, shape their changes, and determine their trajectories of persistence, spread and extinction. Before sketching the shape of this research program, one widely discussed objection to the approach must be disposed of.

5 Disposing of the ‘No-Memes’ Objection to Darwinism About Human Affairs

The argument given at the outset of this paper will be accused of invoking Darwinian processes without showing that their necessary conditions obtain, and it will be argued that a crucial one among these necessary conditions cannot obtain in human affairs.

The objection is simple: Darwinian processes require replicators and interactors. In particular it requires high fidelity replicators to store and transmit traits faithfully enough and long enough for environmental filtration to shape them into adaptations and to maintain them as adaptations. In the biological domain these replicators are the genes. There are no equivalent replicators in human affairs, or at least there are not enough of them for Darwinian processes to explain human affairs. Darwinism of the sort evinced in the argument given at the outset requires cultural replicators–memes. But there are no memes.13 No memes, ergo no Darwinian processes. (Cf. Sperber 2000 for the origin of this widely mooted argument).

The issue is serious for Darwinian social science. That natural selection is an attractive metaphor in the description of human affairs is both unexceptional and uninteresting. The issue is whether it is more than a metaphor. Are all social processes literally, actually, really matters of blind variation and environmental filtration? That is the issue. If a real Darwinian process requires actual replicators, then the no-memes critique must be addressed. It can be.

Darwinian social scientists should respond to this objection in several ways. To begin with, it is obvious that the literal application of Darwinian theory to a domain does not require that the domain contains gene-like replicators. Biological replicators—genes—did not predate Darwinian processes. Natural selection presumably got its start prior to the emergence of these replicators, and for that matter, prior to the emergence of any recognizable very high fidelity informationally rich replicators. Moreover, natural selection is likely to produce replicators of the sort that the genes constitute only when environments change slowly, when evolution is extremely gradual, cumulative, and atomistic in its shaping of individual traits, one by one, for adaptations. When one or more of these conditions do not obtain, adaptive evolution may employ replicators and processes of replication quite unlike genes. (Cf. footnote 9 above).

Following Richerson and Boyd (2006), defenders of Darwinian approaches to culture have also argued that cultural replicators need not have the high fidelity features of genes, since there are a variety practices, norms, and institutions in human culture which have emerged as adaptations precisely because they preserve the adaptive informational content of replicators even under conditions of low copy fidelity. (Cf. Driscoll 2008, for a useful recent discussion.)

It may be granted that Darwinism about human affairs does require cultural replicators, probably a variety of quite different kinds of replicators, and some of them may be gene-like. But the memes Darwinism requires about human affairs will be huge in number, short in life spans, and extremely difficult to individuate, for the very same reason regularities in the human sciences are restricted: because of the ubiquity of arms races. So, it will be no surprise that few obvious examples of memes can be provided now, or perhaps ever.

To see the real problem with the argument from no memes to no Darwinian processes in human affairs consider its first premise; memes have to be like genes, because natural selection only works in culture when there are gene-like replicators in culture. But, the argument continues, there are no such gene-like replicators in human affairs. Ergo no Darwinian processes in human affairs. The trouble with this argument is that it rests on an idea of what genes are and how they work that was obsolete about a hundred years ago. This is the one trait-one gene idea, the notion that most or many significant observed inherited traits are controlled by a single gene.

There are only a small number of such traits in any mammal, and in humans only about 7 such traits are known. For example, tongue rolling or the widow’s peak. All other inherited traits in humans, like eye- and skin color, even sexual characteristics, are the each result of the inheritance of many and in some cases a huge number of genes. In fact, genes do not really transmit or control the appearance of any of the biological traits common sense and folk biology think they do. Each gene controls the production of a protein or other large molecule. There are 25,000 of these genes, switched on and off in every cell of our bodies. It is the protein molecules they code for, and the order in which the proteins are produced by the genes that build and operate biological machinery. Many different traits that do not have anything to do with each other are built or controlled by the same gene; many traits that look to us absolutely the same to us across individuals—say eye color—are the result of different sets of different genes in different people. And when we actually locate the genes inside the nucleus of our somatic cells, and in the sperm and egg that develop into our bodies when they combine, these genes can differ from one another substantially without that difference making any difference for the proteins they produce.

The moral for memes is obvious: if memes are like genes then any single meme will by itself almost never either control the appearance of a behavior or action or anything else that common sense or even sophisticated social science is interested in. It will take many, many memes working together to produce anything of interest to the human sciences, and we will never be able to detect or identify memes by doing anything like garden-variety sociology, economics, anthropology, or even psychology. Whatever memes are, they are going to be as complicated and hard to identify as genes are, or even more so!

If memes are anything like genes, it is going to be very hard to identify them, isolate or individuate them, and learn the details of how they work. Doing any of these things for “memetics” will be orders of magnitude more difficult than what a century of molecular biology has done for genetics. And the reason for all three of these difficulties will in large measure be the ubiquity of arm races cutting short the life time of any regularity about a particular package of memes. The shorter this life time, the more difficult it will be for any social or behavioral scientist to design a method for identifying these package, still less identifying their component memes. To figure out the mechanism of transmission of some package of memes that controls a socially interesting adaptated behavior, will require, first, a long enough lived regularity about that adapted behavior, second, another long enough lived regularity about how the particular package of memes that codes for the behavior and transmits it does each of these things. Since in a social environment of accelerating change, these packages of memes and the memes themselves will be transmitted faster and faster, and their modes of transmission and control interfered with more and more effectively, as arms races accelerate, the chances of identifying and locating memes become harder and harder as culture’s change accelerates.

This is not an argument for the existence memes. At most it is a well-grounded explanation for why social and behavioral scientists are unlikely to find them, an excuse erected on the criticism of a bad argument against the very possibility of memes. If there are memes, regularities about their transmission, mode of action, and realizations in the brain, will be complex, short lived, and completely beyond the reach of any hypothesis testing in the social sciences. So, if we can’t identify them, why suppose that there are any memes?

The real argument for memes is the two premises of the original argument. If these premises are reasonable and support the conclusion that human affairs must be Darwinian in nature, then that’s the argument for memes, or whatever replicators are required by Darwinian processes in human affairs.

Should the human sciences cease work until neuroscience or some even more molecular sub-discipline has established the existence or nonexistence of memes? Did the rest of biology stop working while geneticists did the century of research that was required first to establish the existence of genes and then to identify their location, composition and model of action? Of course not. The same goes for the human sciences.

6 The Smallest Units of Darwinian Cultural Selection

The task of identifying the memes that transmit and encode matters of interest to the social scientist can be left, for the foreseeable future, to cognitive neuroscience. Meanwhile, the Darwinian social scientist needs to get on with the tasks of identifying the regularities reflecting local equilibria in human affairs, tracking the incipient and inevitable arms races that break them up, and employing this knowledge to ameliorate human life and perhaps even design human institutions that move us to preferred equilibria and protect us from the harms so often precipitated by arms races. How should the Darwinian social scientist proceed to do this? Here the obvious source of guidance is that part of Darwinian biology which has succeed while remaining agnostic about the underlying molecular genetics that drives the emergence and breakup of local equilibria: Darwinian population biology as it figures in agriculture, some aspects of medicine, and increasingly in human behavioral ecology and evolutionary anthropology. This approach begins with biologically significant traits—hypothesized adaptations, tests hypotheses about whether they are and what they are adaptations for—i.e. what their functions are or have been in the past. Then it goes on to the rest of four questions Tinbergen identified as on the research agenda of an evolutionary discipline: questions about ontogenic development, evolutionary origin, and mechanism of operation. But where to start? What are the traits to be subjected to evolutionary analysis? We need to identify some broad class, category, kind of aspect, unit, feature of human affairs whose instances recur regularly enough and share enough in common so that some interesting generalizations can be framed about them, or so that we can apply to them some models already effective in systematizing biological phenomena. The rest of this paper is devoted to arguing for the adoption of a single such ‘unit’ among all subdivisions of a Darwinian approach to human affairs: a reconfiguration of the concept of a ‘strategy’ introduced in Sect. 1 above.

Recall its suggestion above that we treat human institutions, groups, practices as packages of strategies employed by people. The features, characteristics, traits of institutions, organizations, practices, are composed of these packages of strategies. At the basement level of individual agents, the strategies they employ are their own individual adaptations—traits that have pay-offs for them or for someone else that result in these strategies persisting—being used over and over, and spreading—by imitation or instruction, reinforcement or coercion, or receding by operant punishment, or legal sanction, etc. Individual strategies are traits of individual people. Their cognitive equipment is what passes them on, modifies them.

The word ‘strategy’ is not quite right to describe the full range of patterns of individual behavior it has to include to do the work required by a Darwinian approach. A strategy will be everything from scratching itches, speaking with an accent, signing one’s name, walking around and not under ladders, crossing one’s self before take-off, using a compass, employing corporal punishment, polygamy, employing the 1662 Book of Common Prayer, saving for retirement, voting, all the way to choosing a spouse, or ordering in the Old Guard at Waterloo. It may well include every repeatable (but not necessarily repeated) bit of action or behavior that it is not genetically encoded (such as responding to operant reinforcement or classical conditioning). There is also an ambiguity in the word ‘strategy’ between describing the behavior and describing the ‘rule’ that ‘governs’ the behavior. ‘Governed’ in quotes because there should be no suggestion that behaviors reflecting strategies need be intentional or strategies the agent is all cases conscious of acting upon. In the sense of rule, ‘strategies’ have a normative flavor: ‘if (or when) in circumstance C, then do B!’ A strategy is often expressed in a norm—categorical or instrumental. But agents need not be conscious of their obligatory force. So, think about a strategy(-type) as a repeatable pattern or instance of behavior (a strategy token) people can repeatedly perform.

Now, as noted in Sect. 1, the key gestalt switch Darwinian social science requires us to make is to shift focus from the selection of agents who behave in accordance with strategies to thinking about the selection among behaviors or strategies that describe them. Darwinian evolutionary theory is a theory about trait fitness and trait-frequencies, not individual fitness. Counting individuals offspring-numbers is a matter of bookkeeping, not the causal process that results in their changes. Similarly, cultural evolution is not a matter of more adapted individuals having more offspring, but of more adapted strategies having more offspring-strategies, copies of themselves. It is a matter of completion and coadaptation of copies among strategies being played more frequently, by more people, over a longer time. [Some evolutionary game theorists have been self-consciously doing this for quite a while. See Skyrms (2010)]. If memetics were simple, we could dispense with the behaviors that token strategies, and just count the instances of a certain neural pattern that encodes a strategy spreading from brain to brain. Watching their demographic expansion and contraction over time, would enable us to measure the memes’ relative fitnesses via their effects on behavioral strategies. Alas, not even genetics is that simple. So, in both cases the scientist must track the traits that interact directly with environments—phenotypes in the biological case, behaviors in the social case, and eventually perhaps infer the character of the molecularly encoded genes and the neurally encoded memes. Long before they can do so, much important biological and social science will have built up.

The easiest way to expound the approach to Darwinian cultural evolution as carried by strategies is by a genealogical illustration, starting at the hunter/gatherer origin of social institutions. This route to recognizing that strategies are the basic explanatory unit in human affairs is not the only one. But it is a natural one.

Well before Homo sapiens, humans were employing strategies first to survive and then to thrive, moving up the food chain on the African savanna by cooperating with one another. In particular the domestic division of labor involves already having distributed roles in accordance with gender based strategies in child rearing, gathering, hunting, fire preservation, etc. these social strategies are formed by a familiar process of operant learning, and transmitted by a combination of learning (imitate the most successful strategy, the most frequent strategy, eventually the most prestigious strategy—once social institutions have become established). Even prior to the strategies that combine to constitute the domestic division of labor, the environment had been operating to select for packages of cooperative strategies that constitute symbolic communication—speech—out of gestures, eye movements, and the theory of mind our ancestors shared with primates. But we may take some Darwinian account of the emergence of language as given for present purposes. The initial emergence of distinct behavioral strategies between male and female in the domestic division of labor well before the appearance of homo sapiens, indeed prior to the appearance of homo, or primates for that matter, has significant productivity effects that increase longevity and population size, which in turn provides further opportunities for the division of labor, for the appearance and selection for still more distinct strategies, whether in the fabrication of tools, specialization in targets of predation, food preparation, even trade. Over time individual strategies get “packaged” together into rudimentary social practices, institutions and organizations. The time scale required even for the precipitation of the earliest of these packages will have been great, and the selection pressures that shaped them must have been varying in force or even intermittent until population densities increased much beyond nuclear families. Through out this and subsequent periods, new strategies are continually emerging through blind variation, and mostly being extinguished by selection against them, or drift. For example, the strategy of combining a hand ax and a shaft into a compound tool surely appeared more than once in the million years between the emergence of the former and the eventual proliferation of the latter. But the variant was not copied with sufficient frequency to withstand drift until populations grew large enough.

Strategies played by more than one individual may conflict or synergize. When they do the latter, there will be selection for the packages that combine them. When they synergize, enhancing one another’s fitness, the result is a local equilibrium. Such equilibria may be very short lasting or quite long lasting. The local equilibria among strategies may be harmful or beneficial to all, some or none of the individuals or groups that employ them. Long lasting packages of such strategies in local equilibrium will typically be recognizable as historically venerable institutions—feudalism, the Roman Catholic Church, chattel slavery. When strategies make individuals or groups compete, they may survive even as individuals or groups playing them become extinct. The result may be extinction of the strategies as well. Or the conflict between players may also eventuate in a local equilibrium in strategies. Packages of strategies in local equilibria will themselves become environmental settings, niches that structure the replication and transmission of lower level strategies that make up the packages among lineages playing the individual strategies. Since transmission of strategies is not genetic, but by learning, imitation, enforcement, and the lineages of these strategies will spread and shrink (when they become maladaptive) through populations of genetically unrelated individuals.

But as in other processes of natural selection, local equilibria are perpetually threatened by random variations that may be selected for owing to their ability to exploit and break up local equilibria. The history of evolution is a history of arms races breaking out, putting an end to local equilibria, and eventually settling into new local equilibria. Sometimes this process proceeds with glacial slowness (for example consider camouflage and camouflage detection between predator and prey); sometimes it moves with great speed (consider the response of the AIDS virus to antiretroviral). The rate of break-up of local equilibria in cultural selection will run the gamut from the time scale of cultural epochs to faster than arms races in the bacterial cases. The brevity and fragility of most local equilibria between strategies makes it difficult to identify them, and impossible to exploit them. But the few long lasting ones are usually of far greater significance for human life. Consider the local equilibrium that characterized church/state relations in Europe from 1500 to the Thirty Years’ War.

Packages of strategies, all the way from the smallest—those of the domestic division of labor—to the largest—the ultra social complex state—are built from this smallest package by a Darwinian process of selection operating to package together those smallest packages that synergies, and to select for variants in smaller packages that enhance synergies. As the process of blind variation and environmental filtration of packages proceeds, new environments are created that have selective effects on the smaller packages of strategies that operate within the niches created by larger ones, selecting for variations among the strategies played within these smaller packages.

The natural and the cultural environments that constitute the original packages of strategies are sufficiently similar across the locations where human social evolution proceeded to independents produce a good deal of convergent cultural evolution. Consider some of the major cultural universals: marriage rules and other sexual mores, religion, inequality and hierarchy precipitated by the transition from hunter/gather modes to agriculture. And of course, differences in local circumstances will select for quite different packages of strategies as well, thus producing the diversity and complexity of institutions, practices and organizations that characterize human culture.

It is crucial to emphasize that the unit of adaptation in this approach is not the individual person or group of people who adopt a strategy which confers some benefit on him, her or them. The unit of adaptation is the strategy itself, or its token-instance played by individuals or groups of them (consciously, unconsciously, willingly, under coercion, incentivized, etc.). People are the environments in which strategies are selected for, owing to whether their effects on people who play them and on whom they are played by others effect the probability or frequency with which they are copied or not copied. It is open to memeticists to hold that strategies are “represented” in memes in minds, and that a bit of strategic behavior is the meme’s “phenotypic” effect. But as we have seen, this is not a commitment Darwinian cultural evolution need embrace, or having embraced, need actually confirm at this stage of social scientific research. At this stage, a Darwinian social science need commit itself to nothing more than the evolution of strategies. Of course the hard work in each of the social sciences will be to identify the strategies significant to the emergence and persistence of the local equilibria each of the social sciences makes its own explanatory focus. The term ‘strategy’ is an explanatory one, not an observational one. Just as in genetics, where the challenge is to determine whether an observable trait is a phenotype (by making guesses regarding its genotype and testing them), similarly the empirical challenge in the social sciences is taxonomizing behaviors into strategies.

It will be objected that strategies, even adapted ones, do no proliferate, “reproduce” over time, but may be played by a roughly constant number of people at a roughly constant rate, and so cannot count as replicators whose fitness increase is a matter of off-spring size. No reproductive increase, no replicators, no replicators no Darwinian process. This critique reflects oversight about important evolutionary processes.

As a Darwinian process, cultural evolution repeatedly results in “major transitions”, as Maynard Smith and Szathmary called them: occasions in which individual strategies enter into packages that persist and in which the individual strategies cease to reproduce independently and competitively. In purely biological evolution there have been several such transitions: from independent replicators to chromosomes, from prokaryotes to eukaryotes, from asexual to sexual reproducers, from single cell creatures to metazoans, from individual organisms to colonies with non-reproductive castes and the division of labor—e.g. the social insects. In each case, individuals have surrendered their independent replication strategy (but not their other strategies) to the ensemble of which they are members. The mystery is why selection for reproductive fitness should result in the surrender of independent reproduction in favor of controlled reproduction as part of a larger individual. This is a crucial question on the research agenda of evolutionary biology and tentative answers are increasingly available. The answers will probably differ for each of the major transitions. But the important fact to bear in mind is that major transitions that shut down or control the replication of particular packages of strategies will occur in the case of Darwinian cultural evolution at least as frequently as they occur in purely biological evolution. This means that uncontrolled replication will be surrendered for controlled reproduction.

Godfrey-Smith has aptly called this state of affairs “de-Darwinization.”14 The simplest and clearest cases of de-Darwinization occur when free-living single cell organisms combine into a single multicellular organism. Many of the strategies that characterize the individual cells persist because they synergies with other copies of themselves and with other quite different strategies into a package that enables the hitherto free-living cells to persist over longer periods. But the packaging requires the suppression of one strategy: in constrained independent reproduction. Multicellular creatures persist only when they successfully suppress individual unconstrained fitness maximization by their component cells, when they suppress a strategy that had been selected for when cells were individual free-living units. When multicellular individuals fail to do this the result is cancer—the unconstrained multiplication of individual cells at the expense of the rest of the component cells and the strategies they are “playing.” According to Maynard Smith and Szathmary major transitions have occurred some seven times in biological evolution, each time involving the suppression of reproductive strategies of the component individual units. Once larger packages of coordinated strategies are selected for, these component strategies are de-Darwinized in Godfrey-Smith’s useful term. That is, the component packages of strategies surrender their reproductive strategies while continuing to play their other ones to mutual advantage. Along with de-Darwinization, the transition to multicellularity will select for division of labor, including the specialization of some cells to engage in reproduction on behalf of the entire ensemble of cells composing the multicellular individual–the sequestration of the germ line, as Weismann put it. At this point survival of the fittest be one of survival of the fattest … the individual package of strategies played by the multicellular organism persists for a long time as it enforces controlled replication among the strategies played by component packages and individual strategies. Multicellular individuals do reproduce, though at much slower rates than the individual cells from which they are composed used to do before combing. This slows down rates of adaptive evolution to the generation times of the multicellular individuals. In some cases, reproduction ceases almost altogether. Consider the quaking aspen. A grove of these trees is a just one organism, all the trees being ramets of one genet.15

Of course multicellular individuals need to, and before the emergence of sexual reproduction, did not by and large reproduce via sequestration of germ line and genetic recombination. Mostly they reproduced by various sorts of fission, in which they either lost a few cells that eventually grew as a colony of the original multicellular individual, or disaggregated into individual component cells which later regrouped into new individuals after asexual reproduction.

As noted, there is a major mystery about the major transitions in which successive individuals surrender their selfish genetic strategy of maximizing their own representation in offspring generations in exchange for membership in larger individuals. Why did free-living mitochondria and nuclear cells join to provide the eukaryotic cell? Why did eukaryotic cells join to make multicellular organisms, why did multicellular organisms such as the social insects surrender their reproductive strategies to a single queen or to classes of reproducers? These are important questions on the agenda of biological Darwinian evolution. They are the problems of the major transitions, in which lower level packages of strategies give up their reproductive autonomy.

The packaging of individual strategies into sets and their de-Darwinization is a crucial process in the emergence, persistence, adaptation and succession of human institutions, practices and organizations. Individual strategies will coadapt to one another tightly enough to ensure the package’s integrity even under circumstances that would challenge the individual strategies severely. In many cases there will be selection for strategies of enforcement of the co adaptations, and strategies that encourage or even control the deployment and reproduction of other strategies. If they are relatively weak they may be expressed in preferences of participants. If stronger these strategies will look to participants and to us like norms. In fact they will constitute norms. Norming strategies will enforce the conformity of new instances of strategies to the form of their predecessors, thus ensuring a copy-fidelity that preserves the package of strategies over time. But de-Darwinization ensures that few strategies will spread like wildfire in a culture, or certainly not in a traditional one, narrowly constrained by natural circumstances and larger packages of strategies that include strong enforcement strategies among their repertoires. This means that in Darwinian cultural evolution we should not expect to find a large number of Individual strategies engaged in a great deal of replication, recombination or variation. They will too often be constrained by the iterated hierarchy of packages of packages of packages of strategies in which they figure, to behave like the strategies of free-living individuals engaged in unconstrained rapid asexual reproduction of the sort that genes engage in. Insofar as individual strategies are memes or behaviors under the control of memes, then memes will turn out to be very different from germ line genes, and much more like somatic genes in de-Darwinized populations of memes.

7 Tinbergen’s 4 Questions and the Future of Social Science

Niko Tinbergen (1963) famously organized the agenda of behavioral biology, and all of biology for that matter, around four questions: what is the trait’s current function, by what particular Darwinian trajectory did it emerge, how does it develop in the individual, and what is the mechanism whereby it delivers its function.

It was long supposed that in biology answers to all four of these questions required appeal to the operation of genes—germline and somatic. We now recognize that in their home base of biology for many species the answers to all four questions increasingly include Darwinian cultural processes. This is just as we would expect given the continuities between Homo sapiens and other species. Darwinian cultural evolution is not confined to humans. Human cultural evolution is a subcategory, not an extension of, still less a metaphorical application of Darwinian evolution. The analysis of human affairs fits neatly into preestablished components of biology.

Tinbergen’s first question, “What is it for, what is its function?” has preoccupied this paper. Biologists have come to see answers to this question nuanced in at least three respects we should also expect in human affairs. First, identifying a trait’s function is a highly fallible matter requiring clever experimentation and observation. Some traits of some organisms are actually imposed on them by other organisms: they are adaptations for the imposing organisms. What is more, functions change over time. As environments change traits with one function can be shaped, pruned, combined by natural selection to take on quite different functions. Gould and Vrba (1982) called such traits exaptations. Feathers are a prime example: evolved for warmth, then selected for flight. Exaptations will be common among human practices. Second, selection operates at many different levels—the individual lineage, the family and local interbreeding population, indeed perhaps even the species. Strategies are traits. In the biological domain, selection for packages of strategies—e.g. among the caste-organized social insects—is not uncommon. Third, as noted above, cultural selection operates in the biological domain as well as genetic selection (we return to this important matter below). Other species besides ours have culture and their cultures evolve by Darwinian processes. Human Darwinian cultural evolution is of a piece with that of other organisms.

The second question on the agenda of the biologist is clearly inseparable from the first: what was the particular sequence of events that resulted in the emergence of the functional trait. Since identifying current function is largely based on identifying what current and past “design problems” the trait’s environment posed, answering the functional question requires us to address the phylogenetic question. Of course the role of drift, small population numbers, physical constraint, operate to channel Darwinian natural selection. But as the previous paragraph noted, increasingly biologists are appreciating the role of developmental plasticity, social learning, long-term niche construction, historical contingencies (founder effects), and other apparently cultural factors in filtering variations to generate the actual trajectory whereby a biological trait emerged and so identify its function. The details of natural history, insofar as they can be recovered, answer this question. Human history needs to be examined to identify the “design problems” and the Darwinian processes that solved them. The traditional explanations offered in human history and even data about their explanantia however may not be of much use in this inquiry.

Tinbergen’s third question is how does the trait work, how does it accomplish the function which the process of adaptive evolution conferred up on it. Lessons from the study of behaviors such as bird song reveal that a variety of top-down and bottom up strategies can elucidate mechanisms at several different finenesses of grain, including somatic gene expression, neural processing, sensory stimulation and deprivation, hormonal changes and seasonal factors, for that matter (Bateson and Laland 2013). There are obvious and severe limitations on the application of experimental methods to learn about equivalent processes that enable traits to accomplish their functions for humans and on humans, but these are not in principle obstacles to biological social science. In the case of individual behavioral strategies, these are tasks for cognitive neuroscientists. In the case of packages of strategies selected for their adaptational functions, answering the “how does it work” question, the role of sociologists, political scientists, economists, is obvious. There may even be room for some “thick description” of the sort cultural anthropologists are eager to provide. But in most cases the answers are unlikely to involve much appeal to human design, and intentional maintenance.

The fourth question is the developmental or ontogenetic one: how does the trait emerge, develop, appear among the organisms that bear it? Tinbergen was particularly interested in non-genetic determinants of the development of animal behaviors, dispositions and capacities, including social learning in birds, rodents, cetaceans, and insects, that operate on genetic inheritance to shape predation, feeding, nesting, communication, etc. This is just the sort of cultural learning long supposed to be limited to humans. Behavioral biologists have shown that these learned traits are faithfully transmitted over many generations by niche construction and other epigenetic processes. The nature/nurture distinction has pretty much lost whatever role it had in biology.

In areas of biology that have developed greatly since Tinbergen proposed the quadripartite division of biological inquiry, answers to each of his four questions have had important spillover implications for answers to others (Bateson and Laland 2013). There are now entire subdisciplines that intersect two or more of his questions: ‘evo-devo”—the study of how developmental patterns are selected or, and how these patterns channel or constrain evolutionary trajectories. But such combinations built on separate first draft answers to each of the questions, and especially to the first and second of them: what is it for? By what process of variation and selection did it emerge? Testing hypotheses about answers to each of these two questions are the only way to proceed in answering the other, and both need at least a well-supported answer before we go on to address questions 3 and 4.

The proprietary taxonomy of biology was functional long before Darwin. No one had to argue that it had to be, and once Darwin’s theory emerged, evidence based answers to the first two questions organized the rest of the discipline. The same assumption that traits are individuated in terms of their function is equally strong in the human sciences. It is this assumption that forces social science to be Darwinian. But this step in turn raises three questions at least: what will a properly biological division of the social sciences into manageable research programs look like, how much of social science will Darwinian cultural evolutionary theory “recover” or “preserve,” and what will the pay-off to such research programs be.

One obvious way of assigning tasks to distinct research programs starts by making distinctions between traits, that is, between strategies. The current division of the social sciences can be construed as implausibly dividing these into psychological, sociological, political, economic, or some such a taxonomy of kinds. This taxonomy is probably mistaken. Two reasons for this are evident. It has not eventuated in much significant explanatory agreement and cumulative predictive improvement. It reflects common sense taxonomies we have excellent reasons to suspect from the wrongness of such taxonomies in physics, chemistry, and biology.

These considerations suggest that the principle subdivisions of a Darwinian science of human affairs will look quite different from the current set of social sciences. There are several possibilities, some of which “recover” more of current social science than others, some of which proliferate subdisciplines, and some of which make separate sciences—even inexact ones—impossible, because they reflect the operation of multiple and inseparable causes for every explanandum of interest. Which alternative is right is a vast empirical question. But some Darwinian social scientists have already taken sides on various possibilities that are worth illustrating.

Runciman (2009) has argued that Darwinian processes operate among humans in three different ways: biological selection, social selection and cultural selection:

At three different levels of natural cultural and social selection, there are three different types of behavior: evoked behavior, where the agent is responding directly and instinctively to some feature of the environment; acquired behavior, where the agent is imitating or has learned from some other agent, whether directly or indirectly; and imposed behavior, where the agent is performing a social role underwritten by institutional inducements and sanctions. [p. 8]

Like a long tradition of sociologists, going back at least as far as Durkheim, Runciman insists that imposed behavior is not reducible to learned behavior, so that there are two distinct levels of Darwinian selection operating in human affairs:

The difference is not that social, as opposed to cultural, rules have to be followed…The difference is that information encoded in institutional practices make society roles what they are independently of how successive incumbents have come to learn to perform them or what their individual motives are for doing so…

The…differences between one society and another are just as much the outcome of a process of heritable variation and competitive selection irreducible to cultural evolution as the beliefs and attitudes which distinguish one culture from another are the outcomes of a process of heritable variation and competitive selection irreducible to biological evolution. [p. 39]

Runciman treats selection of individual strategies as historically prior to the selection of social institutions—what I have called packages of strategies: “…once material resources began to be accumulated and stored and families of households to settle in designated locations…, the necessary conditions for the transition from acquired [culturally selected, learned behavior] to [socially] imposed behavior are in place” [p. 41]. Runciman treats the shift as a major transition in the Maynard Smith/Szysmary sense: “the culture-to-society transition…was a revolutionary transformation of interpersonal relationships and consequential behavior patterns without precedent in the earlier history of the human species.” [p. 42]

If the shift from hunter/gatherers to sedentary groups produced a qualitatively different unit of variation/selection from the individual strategies that characterize the biological and domestic division of labor, then there will be scope for an autonomous Darwinian sociology. The difference between small packages that characterize biological/domestic divisions and larger ones may of course be only a matter of degree. Selection for group strategies may already have kicked in among the primates, enabling Homo to succeed on the African savanna. The difference between group selection and correlated individual selection is either semantic or also a matter of degree. Whether there is full blooded group selection (a Maynard Smith/Szysmary major transition) or only a complicated set of pairwise correlated individual strategies, there will still be scope for a scientific division of labor between the study of institutions, organizations, practices on the one hand, and individual behavioral ones on the other. In effect, a Darwinian approach “recovers” a recognizable distinction between social science—the study of packages of strategies and behavioral science—the study of individual ones (with social psychology somewhere in between).

Nelson and Winter (1982) long ago argued for an evolutionary theory of economic change to supplant the mainstream approach with its heavy reliance on (global) equilibrium thinking. In particular, they introduce the business “routine” as the unit of selection, and used it to offer an analysis of firms, industries, and markets. “Routines” are pretty obviously strategies as elaborated here.

Our general term for all regularities and predictable behavior patterns of firms is “routine.” …. In our evolutionary theory, these routines play the role that genes play in biological evolutionary theory. They are persistent features of the organism and determine its possible behavior…; they are heritable in the sense that tomorrow’s organisms generated from today’s (for example by building a new plant) have many of the same characteristics, and they are selectable in the sense that organisms with the same routines may do better than others, and if so, their relative importance in the population (the industry) is augmented over time. [p. 14]

Two notions that Nelson and Winter needed to more fully articulate the parallel between strategies and genes were the parallel between routines and somatic (non-germline) genes in controlling repeated behavior within the same individual (organism or firm), and the importance of de-Darwinization of replicators that are parts of larger individuals. De-Darwinized but highly adapted routines, like highly adapted genes, will increase the economic strength and size of the individual firms they figure in, but may not increase the number of such firms—this is selection of the fattest, not fittest, as in the growth of Aspen ramets.

Two questions arising from their proposal are obvious: first, why has it met with such resistance in a discipline which should be much more concerned with change and local equilibria rather than with stasis and general equilibria; second, why has the approach not generated a research program fruitful enough to displace the ruling paradigm in economics. Answers to the first question would take us far afield. They would largely reflect the assumptions economists since Walras have made to attain mathematical tractability of their theory at the expense of relevance—e.g. constant returns to scale, probabilistic risk instead of radical uncertainty, maximizing of measureable objective functions. Seeking answers to the second question are more important here, and will have a sobering impact on what we should demand of all the research programs of Darwinian social science.

The most significant problem facing a Darwinian approach to human affairs—whether in economics or elsewhere—is the mismatch between what a Darwinian theory can provide and what social scientists, policy makers and other consumers of social science, want from theories in the human sciences.

Darwinian biological science faces significant limits on application and prediction. On the one hand, generation time between replication and development is often so long that great patience, lots of resources, and a research protocol whose lifetime exceeds that of individual scientists, is required to conduct experiments. Moreover, there are more variables that matter in Darwinian evolution than we can identify or control and quasi-experiments exploiting experimental controls imposed by nature are few and far between. Add in the random character of mutation and recombination, the importance of statistical drift, and it becomes obvious why great predictive expectations of Darwinian theory in biology are unreasonable. Even in laboratory contexts, where many of these problems can be mitigated, precise quantitative predictions are impossible, owing to the fact that rates of random variation increase radically as generation time drops and population size increases. What is more, inferences from laboratory results to predictions about natural setting are fraught to say the least. This is the problem well known in social science of “external validity.” The best we can do in Darwinian biological science is the development of models—Hardy–Weinberg equilibrium, Lotka-Volterra predator/prey equations, Turing’s pattern formation models, Fisher sex-ratio result, game theory model such as iterated prisoner’s dilemma or Hawk/Dove, etc. These models explain post facto but no one can employ them in reliable prediction since we can’t establish the values of their independent variables reliably enough, nor control for exogenous factors, nor otherwise establish whether their application conditions obtain. These limitations have in part motivated the focus of molecular biologists on domains in which such problems do not arise, or can be mitigated, with resultant predictive pay-offs in medical and agricultural biotechnology.

The problems a Darwinian approach to human affairs faces must be far greater than those which Darwinian approaches face in the biological realm. In considering them it has to be rigorously borne in mind that there is no alternative approach that does not face these and other more serious problems. Darwinian social science is the only game in town. The first and most serious obstacle to predictive testing and technological application of Darwinian theories in any social domain is the very opposite of the first problem identified above for Darwinian biological science: instead of the length of experiments having to be vastly beyond the life times of even many generations of investigators, in the social realm, the pace of change is so fast that there is not enough time to construct and carry out experiments that will have much relevance to social phenomena even a few days later than the dates of the experiments. Consider the limitation on market research. The number and values of the independent variables in any Darwinian model we might seek to apply in the social domain will be too great and too difficult to measure, and they will change too rapidly for replication. Then there is arms race phemomenon that in the biological domain eventually unpredictably unravels all local equilibria, even ones that last geological epochs. In human affairs the rates of variation are so great that many local equilibria that policy-designers may wish to exploit last at most for weeks, days, or even just hours.

It is fair to say therefore that Darwinian approaches to domains treated by more traditional theories will not do much better than they have done by way of explanations that can be confirmed via their predictive consequences. Consider the widespread disensus in political science regarding alternative theories about international relations, the behavior of states and transnational actors. There are at least a dozen competing theories (realism, neorealism, liberalism and post-liberalism, rational choice theory, constructivism, Marxian theory, and others) employed to explain and predict the behavior of nations and especially the likelihood of conflict and cooperation between them. In this domain retrospective explanations are plentiful but no theory has a track record of increasing (or even regular) predictive success. A Darwinian theory that focuses on variation and selection for strategies—ones played by states, or ministries, or individual political agents, etc. is not going to do any better than these theories. So why should it be preferred to them?

To begin with, we can already be confident that a Darwinian approach to human affairs is right and that any incompatibility with other theories must be resolved in its favor. That is because humans are biological creatures and we have pretty well conclusive evidence that such creatures must be shaped by Darwinian processes. Furthermore, Darwinian processes are the only way the functional kinds identified in all the social sciences could have arisen. This means that the Darwinian approach can provide a non-ad hoc explanation of its own predictive limitations and those of competing theories. We know from the original domain of application what the difficulties are and we can see how they arise in the human domain even more forcefully. A good deal of the explanation for particular explanatory failures, predictive disconfirmation and general fruitlessness of many research programs in the human sciences turns on the facts about their target subject matter that inhibit general theory—local equilibrium, arms races, frequency dependent and other reflexive forms of strategy selection. Thus, for example, the problems facing explanation and prediction in economics are well understood from a Darwinian perspective on human affairs.

But since the Darwinian theory of human cultural evolution is a derived special case of the more general Darwinian theory, it secures substantial indirect empirical confirmation, including predictive vindication, from the successes of the theory in the purely biological domain. This will of course be cold comfort to those who seek to apply the theory to predictions in the human domain, but it will mitigate criticism that explanations exploiting Darwinian cultural evolutionary theory cannot satisfy even modest demands for indirect predictive improvements. Every Darwinian prediction vindicated in domains such as agriculture, medicine, genomics or molecular biology provides such indirect support for Darwinian cultural evolutionary theory. Moreover, the relationship between models and data in the purely biological domain provide good guidance for what we should expect by way of successful application of the theory in the human domain. Fisher’s sex-ratio theory cannot explain why all sexually reproducing species have a 1 to 1 sex ratio, because they do not! But Darwinian theory can explain why many species have such a ratio, why Homo sapiens have a 1:1.05 ratio, why some paper wasps have a 7:1 ratio. Similarly, a Darwinian approach will provide guidance about developing/applying models in human affairs, and also guide our identification of factors that falsify the models in some domains, obstruct their improvement in others, and explain away their exceptions in non-ad hoc ways (A good example of this sort of guidance is how Darwinian theory helps us evaluate the significance of findings such as Henrich et al. 2004).

The upshot is that first of all, a Darwinian approach is unlikely to recover or vindicate a good deal of received social theory (as opposed to making some good use of at least some of the data it has mounted up); second, it may not enhance our ability to predict, control, or even fine tune human affairs. But this is not something a scientifically sophisticated approach to human affairs should find troubling. Third, what a sophisticated science of human affairs should do is obvious: identify the functions—almost certainly latent, un-noticed—of social events, states, processes, and then build models that explain (and even modestly to predict) their emergence, persistence, spread and recession as the result of multilevel processes of environmental filtration—a.k.a. natural selection. How good we can get at achieving these explanations and predictions will depend on the ratio of the rate at which we can acquire and organize data and the rate at which the local equilibria we seek to identify are overtaken by arms races of various sorts.


  1. 1.

    There is a vast literature on these two notions. To a first approximation, and for present purposes, a function can be defined as the behavior of a system’s component that meets a need, confers a benefit or provides an advantage to the system, and whose occurrence or presence is contingent on its doing so. An adaptation is a “selected effect,” a trait that has emerged through a process of Darwinian (but not necessarily genetic) selection owing to its enhancing the fitness of systems that bear it. The factual thesis that all functions are adaptations is uncontroversial in biology. That function should be defined in terms of selected effects is a somewhat controversial claim in the philosophy of biology. See Rosenberg and McShea (2008) for an introduction.

  2. 2.

    Variation is usually qualified as random, not blind. But its blindness to need or benefit is crucial. Darwin described the process he discovered as natural selection. A better label is ‘environmental filtration.’ It emphasizes the crucially passive character of the process Darwin misleadingly called ‘selection.’ I use ‘environmental filtration’ hereafter as a terminological variant on ‘natural selection’ to emphasize this point.

  3. 3.

    This notion is explained further below. Competing (and for that matter cooperating) adaptations will effect one another’s fitness, and eventually find themselves in a local equilibrium. Owing to the persistence of random variation among all traits, including adaptations, there is a constant threat that one or the other will change perhaps only slightly but enough to exploit the other adaptation. This breaks up the local equilibrium, and precipitates an “arms race” in which variations in the second adaptation that can respond to the first one’s new variant will be selected for. Thus is provoked an “arms race.”

  4. 4.

    In general, the beneficiary of a trait (such as a function) is the individual that benefits from the presence of the trait. One feature of an individual instance of a practice (or hereafter, a strategy’s feature) confers a benefit on that practice or some other practice, if it enhances the persistence of that practice or the replication of further particular instances of that practice (or strategy).

  5. 5.

    See Mackie (1996) for an introduction to Chinese foot binding in a game-theoretical framework. I am indebted here to Andres Luco.

  6. 6.

    As becomes clear below, “Darwinian processes” are construed somewhat broadly to include operant learning, which is only Darwinian selection operating ontogenetically, and other means of producing non-hard wired environmentally appropriate behavior put in place by Darwinian selection. The argument of this paper is certainly not that human institutions and behaviors are the result of selection operating on genes. Quite the contrary, Darwinian cultural selection is mainly social and only in a few early but important cases matters of dual genetic/cultural coevolutionary processes, e.g. lactose tolerance and pastoralism.

  7. 7.

    Recognizing the limitations on rational choice theory or intentional human design to be identified here does not however relegate mathematical models that employ it, or experimental and evolutionary economic results that employ it to explanatory and predictive irrelevance. Just as biological processes appear to approach optima of various sorts well modeled by attributing “design” to mother nature, the same kind of instrumental approach in the social sciences can sometimes make use of rational choice models. More on the role of models below.

  8. 8.

    There is further reason to suppose that the processes of human conscious intentional creation are themselves Darwinian ones carried out within the brain(s) of those who intend them. For the explanation of the apparently purposive creations of human intention faces the same problems as the explanation of other biological adaptations: once we have ruled out future causation, immanent teleologies or vital/spiritual forces and disembodied minds, there seems no alternative to treating brain processes that eventuate in individual actions as Darwinian ones as well. Pursuing this argument would take us into the intricacies of neuroscience. But see Dennett (1995), Campbell (1965).

  9. 9.

    See Maoz and Russett (1993). Rare dissenters from this view have invoked the US Civil War, and the belligerency of Kaiser Wilhelm’s Germany as counterexamples (Layne 1994), since both parties to the civil war considered themselves democracies, and Germany had an elected (if largely powerless) Reichstag. Suffice it to say that these counterexamples are controversial and have been rejected on a variety of counts. If accepted they would not undermine the argument advanced here.

  10. 10.

    Consider what was until recently thought to be the most invariant of biological regularities: all genes are composed of DNA. For a long time this regularity was subject to no exception. But because it remained invariant over a very long period, its operation provided an environment that would allow for the selection for any new biological system that could take advantage of the fact that all genes are composed of DNA. Such a system eventually came into existence—the RNA viruses, whose genes are made of RNA and which parasitize the machinery of DNA replication (the HI virus is the most notable example of these viruses). Thus, the regularity that all genes are made of DNA gives way to the regularity that they are all made of nucleic acids (either RNA or DNA). But we can be sure that the arms race of evolutionary competition will eventually undermine this new invariant regularity, by producing an alternative means of genetic transmission that exploits the regularity (unless it already has done so, by bringing about the prion protein that transmits Mad Cow disease). The same arms race between DNA and RNA and prions also disposes of another invariance of molecular biology, the so-called Central Dogma (in its strong form) that the flow of genetic information is always from DNA to RNA to proteins.

  11. 11.

    Natural selection even produces locally invariant regularities between traits that are not adapted at all, but are correlated as the by - products of traits mutually selected for. For example, consider a remarkable discovery of Darwin’s: In all mammalian species subject to domestication at least some examples are “piebald”— i.e. have spots, usually white on dark—and this trait is heritable. Darwin’s observation has since been widely confirmed, even in “natural experiments,” in domestication of hitherto wild and non-piebald species such as the mink have produced this trait. Presumably, being piebald is not an adaptation, and in general animal breeders do not select for it. The relationship between being domesticated and being piebald is nevertheless invariant, or has been hitherto. However, we pretty much know why. Domestication has always proceeded by allowing the tamest, least aggressive young to reproduce with one another. Tameness is a hereditary trait. At least some of the genes involved in tameness behavior are probably located close together on the same chromosomes as recessive genes that control for variegated coat color. Repeated interbreeding always brings out the recessive trait of piebald coat in at least some descendants. So long as those chromosomes are not broken in meiosis at points between the herd genes and the piebald genes, the regularity that domestic species have some piebald members will be invariant. But of course this generalization is evolutionarily contingent. There are several obvious circumstances—human and natural interventions—that can and some day probably will break it down. Besides a suite of mutations, a founder effect recombination that breaks the chromosomal link, or (equivalently) a persistent program of artificial selection to breed non-piebald domestic animals, there is the possibility of a new move in some arms race we have not noticed breaking the invariance. Being piebald may become an adaptational disadvantage owing to the conspicuousness or other fitness lowering effect of such marks in an evolutionary arms race with predators or parasites.

  12. 12.

    R. A. Fisher’s model of the sex ratio is a particularly powerful illustration of the restricted character of generalizations and mathematical models in biology. It is a regularity that in almost all vertebrate species, indeed in almost all sexually reproducing species, the sex ratio is 1:1—50% males, 50% females. That there is almost always, almost exactly the same number of men as women, was long treated as strong evidence of the benevolence of God. Fisher developed a mathematical model to show that the 1:1 sex ratio generalization is an adaptationally advantageous stable equilibrium that results from a Darwinian process of blind variation and passive environmental filtration. Women have varying hereditary predispositions to give birth to males or to females. Whenever the sex ratio departs from 1:1 in favor of more females, those mothers who disproportionately bare male children will have more and fitter grand children, since their sons are scarcer relative to females and can be choosier. More grand children carrying genes that favor having boys results in more boys and so moves the sex ratio back to 1:1. When ratio begins to favor males over females the same process in reverse shifts it back to 50% of each. Whence the stable sex ratio equilibrium. But of course the model gives false results for a number of species, including us. In humans the long run equilibrium sex ratio at birth is 1.05 to 1, slightly favoring male births. Why? Because boys’ mortality rates are higher than girls’, or at least were higher in the environment that selected for homo sapiens. Darwinian natural selection had to fine tune the sex ratio to make it 1:1 at sexual maturity. Doing that required more boys at birth than girls. So, the model needs to be revised: what is equalized is not the sex ratio, but the amount of parental investment in males and females. Additionally, there are several species of insects in which the sex ratio is heavily biased towards females.

  13. 13.

    I employ the term ‘meme’ here to identify whatever it is that is the replicator in Darwinian cultural evolution. At about the same time Dawkins coined the term Wilson and Lumsden (1981) introduced the word ‘culturgen’ to name whatever fills the replicator role in culture. Had things turned out differently, Wilson’s term would have become the meme for ‘memes’ instead.

  14. 14.

    In biological cases, “De-Darwinization” frequently occasions and in fact is required for reproduction and thus selection at higher levels. This will not be invariable in cases of social and cultural de-Darwinization.

  15. 15.

    The aspen trees in a grove are all parts of a single organism—ramets of a genet. They are genetic clones, the offshoots of a spreading root system, which persists and grows new buds, even as its older buds, the trees, grow up and die off. The single individual may survive thousands of years. But it remains the unit exposed to the vicissitudes of a Darwinian process. Many human institutions—packages of strategies—will be like the aspen.


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© Springer Science+Business Media Dordrecht 2017

Authors and Affiliations

  1. 1.Department of PhilosophyDuke UniversityDurhamUSA

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