On the Evolution of Human Morality

By Herbert Gintis June 27, 2012 38 Comments

Steven Pinker’s thoughtful remarks concerning group selection present a useful occasion for clearing some misconceptions surrounding recent developments in the behavioral sciences concerning our understanding of moral vs. self-interested behavior. Initiated in 1966 by George C. Willams’ Adaptation and Natural Selection  and followed a decade later by Richard Dawkins’ The Selfish Gene, evolutionary biologists in the last quarter of the Twentieth century came to view humans as fundamentally selfish, contributing to society only when socially-imposed rewards and punishment render it in their self-interest to do so. Dawkins, for instance, opines in the opening pages of The Selfish Gene, “We are survival machines—robot vehicles blindly programmed to preserve the selfish molecules known as genes…. a predominant quality to be expected in a successful gene is ruthless selfishness. This gene selfishness will usually give rise to selfishness in individual behavior…. Anything that has evolved by natural selection should be selfish.”

Of course, it does not appear in our daily life that everyone is selfish, and if we introspect, most of us will agree that we try to behave, however successfully or unsuccessfully, as moral beings willing to sacrifice personal amenities in the pursuit of truth, justice, loyalty and compassion. Dawkins’ explanation is that human morality is a cultural facade laid upon our basically selfish human nature. “Be warned,” he states, “that if you wish, as I do, to build a society in which individuals cooperate generously and unselfishly towards a common good, you can expect little help from biological nature. Let us try to teach generosity and altruism, because we are born selfish.”

But why do fundamentally selfish beings, which is what humans are according to the selfish gene theory, accept cultural norms that contradict their natural strivings? Richard Alexander answered this question in 1987 in his The Biology of Moral Systems with his concept of indirect reciprocity, according to which we all continually evaluate others for possible future gainful interactions, and we reject individuals who violate norms of reciprocity. The somewhat more general answer offered by Pinker is that is that each of us conforms to social norms out of fear of losing our good reputation. What appears to be self-sacrifice is thus simply a superficial veneer covering our selfish natures. “Scratch an altruist,” biologist Michael Ghislin eloquently wrote in 1974, “and watch a hypocrite bleed.”

Pinker frames the issue in terms of sacrificing personal interests on behalf of the group. “What we don’t expect to see,” he writes, “is the evolution of an innate tendency among individuals to predictably sacrifice their expected interests for the interests of the group.” This is not the correct way to frame the issue. People do not generally “sacrifice on behalf of the group.” Rather, people have moral principles that the strive to uphold, and that compete with their material interests. When I behave honestly in a transaction, I may have no intention whatsoever of sacrificing on behalf of my transaction partners, much less on behalf of my society. I just do what I think is the morally correct thing to do. When I bravely participate in a collective action against a despotic regime, I am upholding my moral principles, not sacrificing on behalf of the group. Indeed, it is no sacrifice at all to behave morally, because we humans care about our moral worth in much the same way as we care about our material circumstances.

The past few decades have seen the massive accumulation of evidence in favor of the view that human beings are inherently moral creatures, and that morality is not a simple cultural veneer. Humans are born with a moral sense as well with a predisposition to accept and internalize moral norms their society, and often to act on these moral precepts at personal cost. In our book, A Cooperative Species, Samuel Bowles and I summarize a plausible model of human nature in which “people think that cooperating is the right thing to do and enjoy doing it, and that they dislike unfair treatment and enjoy punishing those who violate norms of fairness.” Most individuals include moral as well as material goals in their personal decision-making, and they willingly sacrifice material interests towards attaining moral goals. It is this view that I will defend in my remarks.

Pinker does not present, and indeed makes light of the body of research supporting the existence of a basic human moral sense, suggesting that there is only one piece of evidence supporting the view that people behave morally when their reputations are not at stake: “It seems hard to believe,” he says, “that a small effect in one condition of a somewhat contrived psychology experiment would be sufficient reason to revise the modern theory of evolution, and indeed there is no reason to believe it. Subsequent experiments have shown that most of the behavior in these and similar games can be explained by an expectation of reciprocity or a concern with reputation.” Because expectation of reciprocity and concern for reputation are basically selfish and do not involve a fundamental respect for moral values, Pinker is simply reiterating Dawkins’ message of a half-century ago that we are the selfish product of selfish genes.

1. Morality and Human Nature
Today the economics and psychology journals, including the most influential natural science journals, Science and Nature, are full of accounts of human moral and prosocial behavior. Pinker dismisses this evidence by asserting that “Any residue of pure altruism” beyond self-interested reciprocity and reputation building “can be explained by the assumption that people’s cooperative intuitions have been shaped in a world in which neither anonymity nor one-shot encounters can be guaranteed.” In other words what looks like moral behavior is just a form of mental error due to imperfections of the human brain.

The empirical evidence on cooperation in humans does not support Pinker’s view. The social interactions studied in the laboratory and field always involve anonymity, so subjects cannot help or harm their reputations, and they usually are one-shot, meaning that subjects cannot expect to be rewarded in the future for sacrifices they make at a given point in time.

Pinker does cite a few studies that support his position. “Subsequent experiments have shown that most of the behavior in these and similar games can be explained by an expectation of reciprocity or a concern with reputation.” Let us see what these studies in fact say. Reciprocity, says Pinker, “is driven not by infallible knowledge but by probabilistic cues. This means that people may extend favors to other people with whom they will never in fact interact with again, as long as the situation is representative of ones in which they may interact with them again.” The only published paper he cites is by Andrew W. Delton, Max M. Krasnow, Leda Cosmides and John Tooby, “Evolution of Direct Reciprocity Under Uncertainty can Explain Human Generosity in One-shot Encounters.”[1] This paper (and several related papers coming out of the Center for Evolutionary Psychology at Santa Barbara, California) show that, in the authors’ words “generosity is the necessary byproduct of selection on decision systems for regulating dyadic reciprocity under conditions of uncertainty. In deciding whether to engage in dyadic reciprocity, these systems must balance (i) the costs of mistaking a one-shot interaction for a repeated interaction (hence, risking a single chance of being exploited) with (ii) the far greater costs of mistaking a repeated interaction for a one-shot interaction (thereby precluding benefits from multiple future cooperative interactions). This asymmetry builds organisms naturally selected to cooperate even when exposed to cues that they are in oneshot interactions.”

This statement is of course not only true, but completely obvious, and does not require sophisticated academic papers to validate its truth. However it does not explains human generosity. It is elementary logic that to say that P explains Q does not mean that if P is true then Q is true, but rather the converse: whenever Q is true, then P is true as well. In the current context, this means that whenever subject A sacrifices on behalf of stranger B in an experiment, it must be true that A is sufficiently uncertain concerning the probability of meeting B again, and A would incur a sufficiently large cost should A meet B again in the future, that it pays A to sacrifice now. The authors have not even attempted to show that this is the case. Nor is it plausible. The experiments under discussion assume subject anonymity,  subjects will never knowingly meet again. Pinker’s supposed counter-evidence is thus invalid. To my knowledge, there is simply no credible counter-evidence.

2. Human Morality in Everyday Life
Many readers will doubtless wonder if our view of human moral behavior, which is based on controlled laboratory and field studies, extends to real life.  Consider, for one example among many, political activity in modern societies.

In large democratic elections, the selfish individual will not vote because the costs of voting are positive and significant, but the probability that one vote will alter the outcome of the election is vanishingly small. Thus the personal gain from voting is vanishingly small. The cost, however, is a significant amount of time and energy that could have been devoted to other, materially rewarding, purposes.  It follows also that a selfish individual will generally not bother to form opinions on political issues, because these opinions cannot affect the outcome of elections.

Yet people do vote, and many do expend time and energy in forming political opinions. This behavior does not conform to the selfish gene model. Of course it could be argued that we only vote to enhance our reputation as a good citizen, but since who votes is normally not public information, and one’s voting history of little interest to employers and other social intimates, this is not a very plausible explanation.

It is a short step from the irrefutable logic of selfish political behavior that selfish individuals will not participate in the sort of collective actions that are responsible for the growth of representative and democratic governance, the respect for civil liberties, the rights of minorities and women in public life, and the like, that are characteristic of many modern societies. In the selfish gene model, only small groups of individuals who seek social dominance will act politically. Yet modern egalitarian political institutions are the result of such collective actions. This behavior cannot be explained by the selfish gene model.

Except for professional politicians and socially influential individuals, electoral politics is a vast morality play to which models of the selfish actor are a very poor fit.

Defenders of the selfish gene theory may respond that voters believe their votes make a difference, however untenable this belief might be under logical scrutiny. Indeed, when asked why they vote, voters’ common response is that they are trying to help get one or another party elected to office. When reminded that one vote cannot make a difference, the common reply is that there are in fact close elections, where the balance is tipped in one direction or another by only a few hundred votes. When reminded that one vote will not affect even such close elections, the common repost is that “Well, if everyone thought like that, we couldn’t run a democracy.” Agreed. But this is just the Kantian categorical imperative, an eminently moral value. People vote because it is simply the right thing to do.

Politically active and informed citizens appear to operate on the principle that voting is both a duty and prerogative of citizenship, an altruistic act that is justified by the categorical imperative: act in conformance with the morally correct behavior for individuals in one’s position, without regard to personal costs and benefits. Such mental reasoning, which has been called “shared intentionality,” is implicated in many uniquely human cognitive characteristics, including cumulative culture and language. Shared intentionality rests on a fundamentally prosocial disposition.

Human beings acting in the public sphere are, then, neither avid reputation mongers nor personal gain maximizers. Rather, they are in general what Aristotle called zoon politikon—political beings. And political beings are moral beings.

3. Cultural Evolution Theory is not Just History
“Most of the groupwide traits that group selectionists try to explain,” says Pinker, “are cultural rather than genetic…. Instead, they are traits that are propagated culturally… group selection … is not a precise implementation of the theory of natural selection… Instead it is a loose metaphor, more like the struggle among kinds of tires or telephones. For this reason the term `group selection’ adds little to what we have always called `history’.” There are two misconceptions in this statement. The first is that group selectionists are for the most part uninterested in genetic evolution. The second is that the concept of cultural evolution is a simply a metaphor.

In fact, the general framework within which we work is called gene-culture coevolution. This framework is quite clearly delineated in scientific terms (see, for instance, my overview, “Gene-culture Coevolution and the Nature of Human Sociality”[2] which I can summarize as follows.

First, as elucidated by Richard Lewontin in 1970, natural selection applies to any entity that follows certain rules. “The principle of natural selection as the motive force for evolution… embodies three principles.” These principles are first, phenotypic variation: “Different individuals in a popUlation have different morphologies, physiologies, and behaviors.” Second, differential fitness: “Different phenotypes have different rates of survival and reproduction in different environments.” And finally, fitness must be heritable: “There is a correlation between parents and offspring in the contribution of each to future generations.” Note that there is nothing about genes in this account of natural selection. Indeed, Darwin himself never heard of genes when he wrote The Origin of Species by Means of Natural Selection. “It is important to note,” writes Lewontin, “a certain generality in the principles. No particular mechanism of inheritance is specified… The population would evolve whether the correlation between parent and offspring arose from Mendelian, cytoplasmic, or cultural inheritance.”

Second, many forms of culture do indeed follow these three principles. Different societies have different cultural norms and technologies (phenotypic variation), some cultural forms contribute more to the fitness of individuals in the societies that embrace these forms, and therefore the cultural forms themselves have varying rates of having copies in succeeding generations (differential fitness), and finally, there is a degree of faithfulness in copying cultural objects from one generation to the next (heritability).

Third, there is a common underlying unity to genetic and cultural evolution. Genes transmit information from one generation to the next, encoded in DNA, that is used by new individuals for ontological development and adaptation to the environment. Cultural forms transmit information from one generation to the next, encoded in human brains, artifacts, and documents, used by new individuals for similar purposes.

Finally, when humans develop new cultural forms, such as language, tools, lethal weapons, control of fire and cooking of edibles, the long-term effect is the transformation of the human genome itself. In other words, cultural evolution leads to genetic evolution. Examples are cooking, which led to a vast reduction in the size of the human gut, language, which led to radical changes in the human larynx and tongue as well as a considerable increase in brain size, and lethal projectile weapons, which led to changes in the morphology of hand and shoulders, as well as a reorganization of the upper torso musculature. Consider, for instance, that our closest relative, the chimpanzee, spends five hours a day digesting comparing to one for humans, lacks the ability to produce complex vocalizations, and cannot throw a stone with more than minimal accuracy or force.

Gene-culture coevolution provides a plausible scenario for the development a moral sense in humans, a quality that appears to be absent or extremely rudimentary in other species. Of course, we may never know with certainty because the paleoanthropological record is extremely scanty. However, it is clear that the development of tools, weapons, and cultural objects required cultural norms promoting cooperation. Individuals who violated these norms were doubtless punished and shunned, and hence less likely to pass their genes on to the next generation. A genetic predisposition to conform to social norms was thus likely to be biologically fit, and hence to evolve through natural selection. Insofar as social norms contribute to the fitness of the groups that embrace these norms, so will the genetic predisposition to follow these norms. We call such a predisposition a moral sense.

4. The Group Selection Pseudo-controversy
Pinker begins his discussion of the group selection issue with the following question. “Human beings live in groups, are affected by the fortunes of their groups, and sometimes make sacrifices that benefit their groups. Does this mean that the human brain has been shaped by natural selection to promote the welfare of the group in competition with other groups, even when it damages the welfare of the person and his or her kin? If so, does the theory of natural selection have to be revamped to designate `groups’ as units of selection, analogous to the role played in the theory by genes?” There are two misconceptions in the very posing of this question.

If an altruistic behavior reduces the net fitness of the altruist and his kin, it cannot evolve. The “group selectionists” argue that while the altruist may be less fit that the selfish individuals in his group, groups with many altruists will expand at the expense of groups with few or no altruists, and this expansion can more than offset the fitness loss of the altruist. Because the altruists kin are more likely to be altruists and also are more likely to be in the altruist’s (this is called `limited dispersal’), the net effect of the altruistic act may be to increase the average fitness of the altruist’s kin.

The first misconception here is the view that group selection is incompatible with kin selection. It is not. Kin selection says that the fitness of an individual depends on the genes of his kin and not just his own genes. Group selection says the fitness of an individual depends on the characteristics of the group he is in, not just his own genes. The second misconception is that group selection means that the group is a “unit of selection.” This is not true. Group selection occurs when the fitness of individuals may be higher in one group rather than other, depending on the social structure of the group and its and distribution of genomes.

A third misconception is that if genes are the only true replicators in evolutionary biology, and if genes are in some sense purely selfish replicators, then all biological species must ultimately sacrifice only for their close genealogical relatives. We can thus admit the complex division of labor and altruism in such eusocial species as termites and honeybees, but we must deny altruism in the case of humans, who cooperate widely with non-relatives. In fact, a careful development of gene-level fitness dynamics in a recent paper by Andy Gardner and J. J. Welsh, “A Formal Theory of the Selfish Genootnote Journal of Evolutionary Biology, 24, 2011. shows that even an inclusive fitness maximizing selfish gene can support altruistic behavior in its owner.

5. The Group Selection Issue is Scientific, not Political
The group selectionists, says Pinker “have drawn normative moral and political conclusions from these scientific beliefs, such as that we should recognize the wisdom behind conservative values, like religiosity, patriotism, and puritanism, and that we should valorize a communitarian loyalty and sacrifice for the good of the group over an every-man-for-himself individualism.” If this were true, it would, in my eyes, be a violation of scientific principles. Even if every known society has followed a certain practice (for instance, eating meat), and even if we can find evolutionary roots for this practice, it would not follow that this practice is morally defensible or practically desirable in our society. Indeed,, there is not a single paper or book that I authored or coauthored that drew any such political conclusions, or indeed any political conclusions at all. Nor am I aware of others who work in this tradition who have drawn such conclusions. Sociobiology is not liberal or conservative, or even middle of the road. It is just good science.

6. Conclusion
Some academic fields, including sociology, anthropology, and psychology, have no problem dealing with the fact that human beings are moral creatures, and that this morality is an important element in our success as a species. Others, including evolutionary and population biology, have had a harder time with this fact, because they have over the years developed theories that appear to show that  evolutionarily fit behavior is necessarily selfish behavior. However, the moral nature of human society, and the key role of morality in our success as a species, can be accommodated without requiring evolutionary biology to abandon its cherished accomplishments. The selfish gene versus group selection issue, when properly formulated, has little to do with the nature of human sociality.

[1] Proceedings of the National Academy of Sciences 108,32 2011.

[2] Proceedings of the Royal Society B 366, 2011.

Published On: June 27, 2012

Herbert Gintis

Herbert Gintis

Herbert Gintis (Ph.D. in Economics, Harvard University, 1969) is External Professor, Santa Fe Institute, and Professor of Economics, Central European University. He and Professor Robert Boyd (Anthropology, UCLA) head a multidisciplinary research project that models such behaviors as empathy, reciprocity, insider/outsider behavior, vengefulness, and other observed human behaviors not well handled by the traditional model of the self-regarding agent. His web site, www-unix.oit.umass.edu/~gintis, contains pertinent information. Professor Gintis published Game Theory Evolving (Princeton: Princeton University Press, 2000), and is coeditor, with Joe Henrich, Robert Boyd, Samuel Bowles, Colin Camerer, and Ernst Fehr, of Foundations of Human Sociality: Economic Experiments and Ethnographic Evidence from Fifteen Small-scale Societies (Oxford: Oxford University Press, 2004), and with Samuel Bowles, Robert Boyd and Ernst Fehr, Moral Sentiments and Material Interests: On the Foundations of Cooperation in Economic Life (Cambridge: MIT Press, 2005). He is currently completing a book with Professor Bowles entitled A Cooperative Species: Human Reciprocity and its Evolution.


  • Peter Turchin says:

    Herb, I agree 95% with what you wrote. The only place where I think you are conceding too much ground is here:

    The second misconception is that group selection means that the group is a “unit of selection.” This is not true. Group selection occurs when the fitness of individuals may be higher in one group rather than other, depending on the social structure of the group and its and distribution of genomes.

    This is correct as far as it goes, but I think that we also need models in which groups are units of selection. Samir Okasha makes a useful distinction between models of MLS1 and MLS2.

    Once a group becomes so well integrated and selfishness by its constituent elements so well suppressed, what’s to prevent us from using it as an organism? After all multicellular organisms are groups.

    I think this point needs to be stated explicitly. In this paragraph, for example:

    Second, many forms of culture do indeed follow these three principles. Different societies have different cultural norms and technologies (phenotypic variation), some cultural forms contribute more to the fitness of individuals in the societies that embrace these forms, and therefore the cultural forms themselves have varying rates of having copies in succeeding generations (differential fitness), and finally, there is a degree of faithfulness in copying cultural objects from one generation to the next (heritability).

    You are actually mixing MLS1 and MLS2 models. Norms are properties of a society as a group, but then you switch to the fitness of individuals. You implicitly define fitness of cultural forms in how many individuals they encompass. Yet a more natural way of defining fitness of such group-level characteristics is by fitness of the group itself. For example, the Romance-speaking ethnolinguistic group (which was an entity during the Roman Empire period) left around 40 descendants, some now extinct. The British Empire left several descendant polities (UK, Ireland, USA, Australia, Canada, etc). These are all group-level dynamics.

  • I have no problem with groups as objects of evolution provided they satisfy the usual requirements: replication, heredity, and selection. I am not sure this applies to your examples, because it is a small subset of cultural objects that are ‘inherited’ by the ‘offspring societies,’ so it might be better to focus on the particular innovations that are selected for. The fact that romance languages spread could be neutral diffusion, hitchhiking, etc.

    • Peter Turchin says:

      I also use language as a neutral marker for an ethnolinguistic group. Actually, historical successors of the Roman Empire arose by a process quite analogous to biological reproduction. For example, Francia, the polity established by the Merovingian Franks in Northern Gaul in the fifth century, inherited institutions from both the Roman empire and from the tribal confederation of the Franks. So it is possible for a group to inherit cultural variants from both (or more than two) ‘parents’, but also to get novel ones by recombination, and enitrely novel ones by ‘mutation’. Remember that institutions are group-level traits, it doesn’t make sense to speak of a person having an institution.

  • Group selection may occur when one group is more “fit” than another based on some kind of sum of individual traits, as long of course (as Herb points out) that these traits are passed on (vertically to daughter groups or horizontally via migration of people or communication of their behaviors/memes).

    Group selection may also happen when there is high or total integration of a norm, such that expressing the norm only carries a benefit to the average individual when it is expressed at a high frequency. So, singing a national anthem probably carries no advantage to someone at the beginning of a sporting event, but when expressed with high fidelity and at high frequency, it could forge resolve in inter-group competition (more aggressive on the job market resulting in higher overall group productivity, more likely to fight for one’s country and so forth).

  • tmtyler says:

    Re: “If an altruistic behavior reduces the net fitness of the altruist and his kin, it cannot evolve” – sure it can – think of cuckoos. The altruist can be being manipulated, by other agents, or by cultural elements. That’s the idea of “the extended phenotype”: the heritable information responsible for a trait may not be part of its owner’s genome.

    • hgintis says:

      The bird raising cuckoo young is no more an altruist than a rabbit eaten by a fox. Look up the definition.

      • Tim Tyler says:

        This sample definition (from Stanford Encyclopedia of Philosophy) apparently distinguishes between cuckoo and rabbit as follows:

        “In evolutionary biology, an organism is said to behave altruistically when its behaviour benefits other organisms, at a cost to itself. The costs and benefits are measured in terms of reproductive fitness, or expected number of offspring. So by behaving altruistically, an organism reduces the number of offspring it is likely to produce itself, but boosts the number that other organisms are likely to produce. This biological notion of altruism is not identical to the everyday concept. In everyday parlance, an action would only be called ‘altruistic’ if it was done with the conscious intention of helping another. But in the biological sense there is no such requirement.”

        The article goes on to discuss cuckoos in an example.

        We *could* argue definitions, but surely my usage – which classifies cuckoo foster parents as behaving altruistically towards unrelated cuckoo offspring – is the standard one.

        • hgintis says:

          Actually, you have identified a weakness in the Stanford Encyclopedia definition. It should say that altruism is only defined within species, or be much more specific about the mechanism involved. I have seen wider definitions than introspecific, in which the behavior must have a genetic basis and must have evolved through selection and adaptation. There certainly is no evolutionary problem understanding cuckoo parasitism, or any other form of parasitism. No one considers being parasitized as ‘altruistic.’

      • Peter Turchin says:

        Herb, so by excluding interspecific ‘altruism’, you also exclude the ‘great deception’ explanation, according to one version of which, it is malicious memes that fool people to behave seemingly altruistically? Because memes and genes are different ‘species’?

        • hgintis says:

          Genes and memes are not species. Species are biologically reproducing groups of organisms (see Mayr definition, and emmendations).
          I am not saying that the concept of altruism is not applicable between species. I am saying that in the population biology literature, only intraspecies altruism is defined and considered. I think the definition could be extended to interspecies interactions, but one would not end up including foxes catching slow rabbits or cukoos parasitizing other bird species.

      • tmtyler says:

        Defining altruism between arbitrary organisms is more general, more common, and IMO better than only defining it to refer to within species interactions. There’s no good reason to get into the tangled problem of when two individuals are part of the same species in this case. Anyhow, with humans manipulating other humans – or humans being manipulated by memes – both the donor and recipient of the altruism are typically human.

      • I think the question of whether interspecific interaction can be called “altruism” is interesting. But, to me, it is not the key distinction in the example, since I would have a hard time calling this behavior “altruistic” even if they were all conspecifics.

        Say there are sneaky cuckoos who sometimes sneak their eggs into naive cuckoos’ nests. What is the naive cuckoo’s options? It could abandon its eggs or eat them or something. If there is a sufficiently high probability the egg is its own, the better choice would be to hatch the egg and raise the offspring I would not consider the naive cuckoo’s raising of the offspring (which sometimes happens to not be its own) “altruism” because it is the best choice for maximizing individual fitness compared to the outside options.

        Similarly, I would not consider getting my credit card information stolen an act of altruism, no matter how much it helps the identity thieves.

  • I’ve really appreciated reading the critical views of the Pinker essay in the Social Evolution Forum. I especially appreciated the above summary of Pinker’s misunderstanding of the empirical evidence of altruism and morality.

    For what it’s worth, I posted my initial response on Biased Transmission Monday. http://biasedtransmission.blogspot.com/2012/06/pinker-fundamentally-misunderstands.html

    I am looking forward to reading the official Edge responses.

  • My understanding of the work from the Center for Evolutionary Psychology in Santa Barbara that you cite is that currently observed altruism arises, in part, from ‘evolutionary-lag’. Because genomic change lags far behind changing environmental conditions, our current altruism exists because it was adaptive behaviour in the Paleolithic for the reasons they explain. Prof. Gintis appears to side-step this aspect of the Santa Barbara argument and thus, inadvertently, he creates a ‘straw-man’.

    • Except that a large amount of the variation in one-shot play can be explained by variation in culture. (See Henrich et al “In Search of Homo Economicus”) If genetic lag was so strong, you would not expect it to vary so much by culture.

      Also, the model does not have genetic lag in it anywhere. They simply show that if there is sufficient uncertainty about whether you are in a repeated or one-shot game, and making a mistake in one direction vs another is sufficiently costly, and then you should go with what gives the higher payoff. You can get there by rational choice, trial-and-error learning, cultural evolution, genetic evolution, gene-culture coevolution or whatever. Their model does not help distinguish between these mechanisms.

      As an aside – my guess is that if they allowed neutral invasion by cooperative strategies (ALLC), allowed other noncooperative strategies (e.g., suspicious TFT), and/or had a more reasonable mutation rate – they would have a harder time getting cooperation to work in their model.

    • hgintis says:

      Please do not tell me I created a straw man. I did no such thing. I commented on the articles cited in Pinker’s paper. They are faulty for the reasons I suggested.

      I know the position that contemporary altruism is now maladaptive but was adaptive in the early stages of human evolution. I reject that argument, which is completely different from the argument in the papers cited by Pinker.

      Human prosociality goes way beyond behavior in prisoner’s dilemma, public goods, and trust games. Rather it is connected to food sharing and collective child-rearing in early hominin evolution.

      At the risk of boring the reader, here is what Samuel Bowles and I said in A Cooperative Species (Princeton, 2011) on the matter:

      “Leda Cosmides and John Tooby (1992) … hold that in modern settings [altruistic cooperating and punishing] are fitness-reducing. Though the cooperation that was extended to family and reciprocating fellow group members enhanced the fitness of cooperators among our ancestors, according to Cosmides and Tooby, in the modern world of more ephemeral social contacts its expression is a maladaptive legacy of the distant evolutionary origins of human motivation. Because our ancestors rarely encountered strangers we just do not sufficiently distinguish, either in lab experiments or in real life, between one-shots and long lasting interactions, treating strangers much as if they were intimates. Dawkins explains it this way: “the lust to be generous and compassionate. . . is the misfired consequence of ancestral village life” (p. 222).

      There is little doubt that reputation-building in repeated interactions and a tendency of close family members to interact frequently contributed to the evolution of cooperation. But we think it unlikely that these mechanisms provide a sufficient explanation. First, modern humans are perfectly capable of distinguishing between situations in which reputation building and retaliation against freeriding are possible and situations in which they ar not. It is not likely that prosocial behavior in the latter situation is just a mistake. Second, the view that early humans lived in worlds with little contact outside one’s family—Dawkins’ ideal conditions for self-interested cooperation to flourish—is difficult to square with what is known about the Late Pleistocene and early Holocene. Like Jean-Jacques Rousseau’s philosophers, Dawkins, Huxley, and other biologists seem to have jumped on a faulty time machine, and have journeyed to an imaginary ancestral world.

      The evidence for a quite different picture of our ancestral condition is necessarily indirect, but convincing. Prehistoric foragers left few archaeological traces and the historical record contains few precontact histories extending over more than half a century. The best we can do is to make inferences from the available data including huntergatherer demographics, Late Pleistocene climate records, archaeological evidence on causes of deaths during the Pleistocene, and ethnographic and historical reports as well as genetic evidence from recent foragers.

      We will see that neither the likely size of groups, nor the degree of genetic relatedness within groups, nor the typical demography of foraging bands is favorable to the view that Late Pleistocene human cooperation can be adequately explained by kin-based altruism or reciprocal altruism. What is known or can reasonably be inferred about the Late Pleistocene and early Holocene suggests that ancestral humans did not live in small closed groups in which family and self-interest with a long time horizon alone were the cement of society. Rather our ancestors were cosmopolitan, civic-minded, and warlike. They almost certainly benefited from far-flung coinsurance, trading, mating and other social networks, as well as from coalitions and, if successful, warfare with other groups.”

  • Thanks for that response Prof.Gintis. My post cited “the work from CEP” rather than the articles cited in Pinker’s paper. The work of CEP does indeed refer to the lag phenomenon and you cite one example of this in your response (Cosmides and Tooby, 1992). Indeed, you did set out your case against any significant impact of this phenomenon on contemporary altruism in ‘A Cooperative Species’ and you have kindly given an extract here. I suggest that this point needed to be raised to give a well-rounded picture of the CEP position and your counter-argument.

  • tmtyler says:

    I think the “Evolution of Direct Reciprocity Under Uncertainty can Explain Human Generosity in One-shot Encounters” paper is a nice one that correctly identifies and exposes some nonsense that was being used to support group selection arguments. The Santa Barbara folk are wrong about a number of things, but this time doesn’t seem to be one of them.

    • hgintis says:

      Altruism in humans is not “being used to support group selection arguments.” Most researchers working on this problem have no interest in biology disputes over levels of selection. The Santa Barabara article you mention does not explain human altruistic punishment or cooperation at all, for reasons I mention in my reply to Pinker. In fact, their argument is rather trivial and obvious, not needing any extensive analytical elaboration.

      • tmtyler says:

        To clarify, the paper says that evidence for human altruism in one-shot interactions has been used to advance “reconceptualizations of economic rationality and proposals of generalized other-regarding preferences to accounts of altruism produced variously through genetic group selection, cultural group selection, or gene–culture coevolution”. It cites the authors involved: Boyd, R., Gintis, H., Bowles, S., Richerson, P.J., Fehr, E., Fischbacher, U., Van Vugt, M., Van Lange, P.A.M., Henrich, J., Haidt, J., Wilson, D.S. and Sober, E.

        • hgintis says:

          That sentence makes no sense to me. Whatever it means, the fact is that cultural group selection and gene-culture coevolution can be supported easily without reference to human other-regarding preferences. Indeed, we tend to use gene-culture coevolution as a possible explanation of some aspects of human prosociality, rather than the reverse.

      • tmtyler says:

        It wasn’t what I was originally talking about, but one example of people using academic work on human cultural group selection to promote group selection may be found the “Beyond Demonic Memes” article. It says: “One of the sleights of hand performed by Dawkins in The God Delusion, which takes a practiced eye to detect, is to first dismiss group selection and then to respectfully cite the work of Richerson and Boyd without mentioning that their theory of cultural evolution is all about group selection.” The “Survival of the selfless” article has sections about the work of Christopher Boehm and Peter Turchin. I don’t think it can be denied that the group selection enthusiasm associated with much evolutionary anthropology is feeding back into the larger group selection debate in the form of evidence that maybe group selection has something going for it after all.

        • hgintis says:

          I have grown to have an intellectual interest in group selection, as an intriguing part of evolutionary and population biology. But I came to this from using it to try to explain the results we got on other-regarding preferences. I would not attempt to use human morality as the central way to “prove” or “justify” group selection. The latter is a much larger issue and involves all social species.
          Of course, others might have had different intentions.
          Intentions don’t matter much to me. Only results.

  • tmtyler says:

    Morality is surely largely a “cultural veneer”. We know that our distant, uncultured ancestors bashed each other’s skills in with rocks, ate their brains, and generally regularly engaged in rape and pillage. Humans are more moral than chimpanzees largey because of their memetic symbionts – in conjunction with the fact that memes spread by social contact, and therefore like to inhabit sociable hosts. Sure, there will have been some coevolution – and I’m sure humans are genetically more sociable than chimps – but let’s not underestimate the continued significance culture in modern times. Without culture, humans are still pretty bestial – “Lord of the Flies” style. Humans have a tendency to romanticise the past. However, let’s not forget how nasty and savage things were only a few thousand years ago.

    • hgintis says:

      You just haven’t read the literature on early hominin social organization., There is no evidence in favor of your view. See my paper “The Roots of Human Hyper-Cognition” on my web site, or Chris Boehm’s “Hierarchy in the Forest” and his new book, Christopher Boehm, Moral Origins: The Evolution of Virtue, Altruism, and Shame (New York: Basic Books, 2011).

      • tmtyler says:

        No evidence in favor of a powerful association between cultural development and moral behaviour? You are kidding, right? Pinker just wrote a popular book on this very topic, for example. The shifting moral zeitgeist is a well-known and dramatic phenomenon. In your defense, you cited a paper of yours which claims that: “Darwin’s theory of sexual selection was a serious error” and “there are no convincing examples of runaway sexual selection in the biological literature”.

        • hgintis says:

          By “your view” I mean “Morality is surely largely a “cultural veneer”. We know that our distant, uncultured ancestors bashed each other’s skills in with rocks, ate their brains, and generally regularly engaged in rape and pillage.”
          Hominins have always bashed each other’s heads in, and still do. This does not fully characterize human morality. Please read the material I cited before going on about this.

      • Peter Turchin says:

        Folks, please keep the debate within the civilized bounds.

      • tmtyler says:

        Yes, hominins still do bash each other’s heads in – the issue is how much they do it, and to what extent the frequency has to do with how cultured they are. We can see, roughly, the impact of culture on moral behaviour by looking at modern cultural variation, hunter-gatherer tribes, natural experiments in cultural isolation and by looking at history.

        The traditional way of considering such issue is to ask how much “morality” (measuread as a scalar somehow) is explained by genetic variation in the population, how much by cultural variation in the population, and how much is explained by other environmental factors.

        Looking at some of Pinker’s statistics, we have:

        “Tribal warfare was nine times as deadly as war and genocide in the 20th century. The murder rate of Medieval Europe was more than thirty times what it is today.”

        Such figures may suffer from being a little cherry-picked – but the effect they describe is a real one – and they do illustrate the dominance of cultural factors over genetic ones – in that particular moral area. Culture has large and important effects on moral behaviour – just as it does on practically every other category of human behaviour. So: describing the effect of culture on morality as a “veneer” is using the wrong word, really.

    • Peter Turchin says:

      Actually, I was referring to Tim’s comment and I slightly edited it to make it less confrontational. I just wanted to catch this before it escalates and I would have to start deleting comments.

  • tmtyler says:

    The article “A Formal Theory of the Selfish Gene” rather takes for granted that selfish genes can produce altruistic organisms, saying: “selfish gene theory represents an attempt to reduce organismal altruism to the self-interested behaviour of a more fundamental agent (Dawkins, 1976, 1978, 1982)”. AFAICS, the paper makes no futher mention of altruistic organisms. Instead it argues that genes themselves may exhibit altruism towards other genes. The argument given in the paper says that this is because they are maximising their inclusive fitness, not their personal fitness. However, if you simply define altruistic behavior – following Trivers (1971) – as being:

    “behavior that benefits another organism, not closely related, while being apparently detrimental to the organism performing the behavior, benefit and detriment being defined in terms of contribution to inclusive fitness.”

    …this suppopsed problem with selfish gene theory goes away, and genes become “selfish” once again.

    I think using “fitness” to mean “personal fitness” is liable to mislead. These days, most understand that “inclusive fitness” is biology’s optimand – and thus more deserving of the title. The term “altrusim” should probably be defined accordingly – making kin selection and group selection into non-explanations for altruism.

    • hgintis says:

      This sounds right to me. One point: inclusive fitness “people” love to say that the object of selection (gene,individual, or what have you) “behaves in a way that maximizes inclusive fitness.” However, when there is frequency dependent payoffs (the usual case in complex social organisms), this is certainly not proved, and there are many counterexamples.

      Natural selection does NOT select organisms that behave as though maximizing their inclusive fitness (or any other kind of fitness, as far as I can tell).

  • tmtyler says:

    Richard Dawkins retracted all that stuff about being “born selfish” in 1989 – as follows:

    ‘I do with hindsight notice lapses of my own on the very same subject. These are to be found especially in Chapter 1, epitomised by the sentence ‘Let us try to teach generosity and altruism because we are born selfish’. There is nothing wrong with teaching generosity and altruism, but ‘born selfish’ is misleading. In partial explanation, it was not until 1978 that I began to think clearly about the distinction between ‘vehicles’ (usually organisms) and the ‘replicators’ that ride inside them (in practice genes: the whole matter is explained in Chapter 13, which was added in the Second Edition). Please mentally delete that rogue sentence and others like it’.

    • hgintis says:

      Thanks for this. Do you have a citation I could use?
      In fact, however, most if not all of the “group selection” skeptics believe in some form of human behavior as either a “big mistake” (the Santa Barbara school, for Instance), or completely self-regarding, once reputation is included. What this has to do with “group selection” I do not know.

      • tmtyler says:

        It’s actually from “The Selfish Gene: 30th Anniversary edition” (2006), Introduction, page ix. I had the wrong version in my last post.

      • tmtyler says:

        Traditional explanations for altruism do include kin selection. Kin selection and group selection have turned out to be different ways of modelling the same phenomenon. Kin selection is not more “self-regarding” than group selection is (they are equally other-regarding). Wherever you would say “group selection” there’s a corresponding more orthodox explanation in terms of kin selection.

        I think it is correct to say that kin/group models are under-used. One problem is that the expansion of relatedness to include inheritance not based on DNA has not yet properly gone mainstream. That should bring with it a whole new wave of kin/group models.

  • Daan says:

    “Cultural forms transmit information from one generation to the next, encoded in human brains, artifacts, and documents..” But basically in behaviour; artifacts and documents are abstract forms of information-transmission, all that is essentially needed is social interaction such as parenting-behaviour.
    Note for instance that the first artifacts arose about 100000 years ago, while homo sapiens evolved 100000 years before that.

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