David Sloan Wilson has been perhaps the strongest advocate for group selection for several decades now. The article under consideration here is an attempt to show that human cultures have been created by and evolve by a form of group selection, presumably cultural group selection. I am afraid that I don’t find anything in this article convincing at all. It mostly consists of a series of assertions for which no evidence is provided. Since space is short, I shall take up just a few of Wilson’s arguments. In my discussion I shall think of group selection as behavior that has been selected “for the good of the group.”
- Widespread human cooperation is to be explained primarily by group selection. Certainly humans are a highly cooperative species and in every society there is an enormous amount of cooperation. But I see no reason why group selection is necessary to explain this. Groups are aggregates of individuals who are pursuing their own interests, which are both somatic and reproductive. But to achieve these interests they have to cooperate in numerous ways. Wilson gives the example of the difficulties of survival in the Arctic, pointing out the many forms of technology that are required, much of which can only be produced by cooperative efforts. No problem there. But the cooperation is easily explained by standard gene-level (or individual-level) evolutionary theory. What individual living in such an environment would be so foolish to think that he or she could somehow survive on his own? This would be well-nigh impossible. Of course members of Inuit groups cooperate, otherwise they would be driven to extinction as individuals. Cooperation doesn’t require selection at the level of the group. Ordinary individual selection will do just fine. (Sometimes I think that the group selectionists believe they are the first scholars to discover that there is such a thing as human cooperation.)
- Religion fascinates social scientists because it seems not to be utilitarian. I have not read many works on the sociology or anthropology of religion that take such a perspective. Indeed, I can’t think of a single example. Quite the opposite is the case. Nearly all social-scientific theories of religion stress its benefits, with different theorists proposing different benefits. Wilson is keenly interested in religion, indeed wrote a whole book on it (Darwin’s Cathedral) in which he tried to explain it by group selection. Many years earlier Durkheim proposed one of the most famous theories of religion ever developed. He argued that religion is really the worship of society and thus binds individuals together. Its function is to provide social cohesion. This is a type of group selection argument. Wilson believes that Durkheim got it basically right. I would claim, however, that he got it basically wrong. I have formulated a list of ten major criticisms that I will be developing in a book on religion that I am currently writing. Be that as it may, the really shocking thing Wilson says in his discussion of religion is that Rodney Stark, a leading sociologist of religion, “portrays religion as primarily dysfunctional.” Stark is rolling over in his grave at this very moment. The last person on earth who would make such a claim is Stark. He uses the well-known sociological theory known as exchange theory to explain how people use gods in order to gain rewards that are otherwise unavailable in daily life. Religion is not dysfunctional, but highly functional, and it is individuals that religion is highly functional for. Exchange theory is a highly individualist theory and Stark is a strong anti-Durkheimian. He is also, in my view, the best sociologist of religion since Max Weber.
- Cultural evolution is a product of group selection. Group selectionists talk a lot about cultural evolution, but in a very constricted way. Usually they have in mind just very small bits of it over very short periods of time. But let’s think about cultural evolution – I prefer to call it social evolution – on a long-time scale, the 10,000 years since the Neolithic Revolution. Social evolutionists are generally agreed that over this period of time there have been remarkably similar evolutionary trajectories all over the world. In terms of subsistence technology, we see a shift from hunting and gathering to horticulture (shifting cultivation), from horticulture to intensive agriculture, and from intensive agriculture to modern industrial capitalism. In terms of political life, we see an evolution from bands, to tribes, to chiefdoms, to states. Societies have grown much larger with much denser populations and greater complexity. It’s pretty much the same story wherever you look. We know that it is those societies smallest in scale that exhibit the highest levels of cooperation. But what happens when societies evolve? A big thing is increasing status differentiation and social stratification along with increasing concentration of power in the hands of a few. Hunter-gatherer societies are highly egalitarian and status competition is frowned upon and strongly policed. In horticultural societies we see the development of status competition that is lightly policed, if at all. In some of these societies “big men” hold large feasts to show how skilled they are. Some of them grow big inedible yams that they present to status rivals, who must respond with yams of equal or greater size or else lose status. Among the Kwakiutl of the Northwest Coast, chiefs held potlatches in which they gave their goods away, daring other chiefs to give their own goods away or lose status. Once we get to full-fledged chiefdoms, a major line of division opens up between a powerful chiefly class and the rest of the population. This class is capable of exercising a lot of coercion over everyone else. Once states are achieved, the level of coercion becomes much greater still. Chiefdoms and states can extract tribute from most of the population, and coerce a lot of young able-bodied males to fight in wars of conquest against other societies. Most of these men are not interested in fighting. They are given no choice. What kind of cooperation is that?
- Inter–societal selection. The point about conquest is especially important because it is about one group defeating another. On the surface that looks like group selection. But is it? I would say no. It isn’t a matter of a whole society against another, but of the most powerful members of a society conquering another in order to gain the spoils. Do they want to share these spoils with the rest of the society? Not really. They may share some to keep the population relatively quiescent, but mostly they keep the spoils for themselves. That’s largely what preindustrial states and preindustrial warfare are all about. And what allows one group to defeat another is such things as bigger armies, superior military technology, more effective military strategies, and so on. It isn’t really about cooperation. But none of this is for “the good of the group.” It is for the good of those who are running the group. The sociologist Gerhard Lenski called this sort of thing inter-societal selection. I can’t see that it is any sort of group selection in Wilson’s sense.
- Consensus. Wilson stresses that the 45 participants in the Strungmann Forum all agreed that group selection is a vitally necessary concept. He seems to be implying that, because of this consensus, and because they agree with him, they must therefore be right. But who are these 45 people and what are their backgrounds? It would be quite easy to organize another forum and invite 45 different people ( or even 1,045 people) all of whom agreed that group selection is a wobbly concept that explains nothing. To be fair to Wilson, consensus in science is necessary, as Thomas Kuhn pointed out years ago. Research can march forward more efficiently this way. But the entire history of science is a story of consensus formed, consensus lost. Unfortunately, Wilson seems to want a permanent consensus, since he used that phrase in the title of an earlier article. The idea of a permanent consensus is dangerous because such a consensus can easily descend into dogma, and dogma doesn’t advance the cause of science.
In conclusion, I would just say I think Steven Pinker got it almost exactly right in his essay “The False Allure of Group Selection.” Of course there are cultural traits that may benefit the group as a whole. But all this can mean is that these traits benefit all of the individual members of a group. And of course there are individuals in societies who may display altruism (e.g., suicide bombers, men who volunteer for the military and are only too happy to fight for their country in faraway lands). But if one looks around I don’t think genuine altruism is the sort of thing one finds at a high level of frequency in human in any human society. And Pinker tells us why.
Stephen K. Sanderson. University of California, Riverside
Human cooperation is NOT easily explained by standard gene-level (or individual-level) evolutionary theory – *unless* you count memes as being a type of gene. Culture makes the difference between a modern city dweller and a cave man. The cooperation produced by culture is what you fail to explain if you only consider DNA genes.
Group selection is indeed unnecessary to explain cooperation – but only because it is equivalent to the widely-accepted theory of kin selection. Group selection is another way of looking at the same kind of process. As such, it is sometimes helpful – and sometimes confusing and redundant. I don’t think invoking Pinker helps with credibility here – Pinker seems unfamiliar with the literature on cultural evolution or group selection.
I tend not to think of cultural group selection as behavior that has been selected “for the good of the group.” I think of it in terms of behavior that brings advantages to one group compared to other groups.
So, agriculture was probably (a) bad for individuals (poorer grain-based diet) but (b) good for their genes (bigger population) and (c) often good for their groups (as with the Dravidian, or Austronesian, or Bantu expansions).
It is far too extreme to say that all cultural adaptations are a consequence of group level selection, whether cultural, biological or both. It seems as if we were losing the multilevel selection perspective. Adaptations are the result of complex interactions between cultural and biological evolution at all levels of selection (replicator, individuals, groups, etc).
When we are talking about the adaptations that are beneficial for the groups we must not speak of every group trait or alleged group adaptation. There are only some of them that are relevant to this group-level analysis.
For example: the artic knowledge about kayac building and surviving. Is it a group-level adaptation or is it an individual-level adaptation? Well, in my opinion there is a very simple criterium to find this out. We must ask ourselves: Does it require specialization and division of labor? If the answer is no, it is very likely that the dymanics of cultural evolution have been mainly at the individual level and/or the replicator level (memes). But if the answer is yes, it is very likely that the evolution of these cultural traits have also been developed under group-level selection pressures. We should remember that besides kin altruism this is one of the main features of eusocial animals: division of labor.
This is why sometimes small primitive hunter-gatherer societies are not the best sources to study group selection (but perfect for kin selection). More complex societies with strong social stratification, specialitation and division of labor seem to me much more likely to evolve under real multilevel selection pressures, since complex organizational configurations, that require confidence in a certain degree of general collaboration, are less likely to be aknowledged and adopted by rival groups than specific techniques or pieces of individual culture.
Specialization and division of labor seem quite compatible with the Allee effect to me.
The traditional way of looking for signs of group selection is to look for cases where individuals take fitness hit for the benefit of their group. The problem with that is that – if you do the accounting properly using inclusive fitness – all you find is organisms who are malfunctioning, manipulated, confused, or in unfamiliar environments.
I think that hardly ever individuals really take a hit for the benefit of the group. That notion is too narrow to address cooperation: a lose-win business. In my opinion it is more logical to expect a a winwin-like business. Indeed we can expect that sometimes culture will manipulate the instincts of kin altruism as you propose, but there is also the possibility that the individuals only apparently take a hit, because there are indirect ways to get a benefit from a given behaviour. Namely social and sexual benefits. I go with Geoffrey Miller that sexual selection by mate choice can account for the indirect but extremely important benefits that certain kinds of cooperation bring.
Suicide bombers are a classic example of taking a fitness hit for the benefit of unrelated others. It isn’t plausible that they are helping their blood relatives – rather they are victims of manipulation. The memes inside the suicide bombers aren’t sacrificing themselves for nothing, though – they are dying so that copies of themselves inside other human hosts might live. Such things aren’t common – but they do happen often enough to need explaining.
I agree with almost all of this–the one thing I’d add is that small-scale local wars (i.e. all wars for the first 2 million years of humanity) do often involve everyone pitching in and everyone gaining (mostly gaining the territory of the conquered, and often their women). That’s still individual benefits, but it leads to groups succeeding and other groups failing. Think of the Yanomamo–who were largely fighting over land, not for some nutty desire just to fight (as Harris and Chagnon and so on imply) The wars over major loot, with the leaders of the winners hogging all the loot, come in only with civilization or very late chiefdoms.
all best, Gene Anderson
Modern warfare sometimes features solders who are manipulated into helping apparent kin by apparent father figures using tools such as patriotism. However, even in the modern world, being a soldier is often job, and many get paid for doing it. Probably both features of warfare are ancient – though perhaps modern “propaganda machines” have increased the potential of the first mechanism.
Speaking of the Inuit, they took over the Arctic from the Dorset people (around 1000-1400 AD). I suspect this is because as a group they had some cultural advantages in that environment. (Maybe they had some genetic advantages too.)
Steve, your point 1 on cooperation ignores the insights of Mancur Olson and Garrett Hardin. Sure everybody wants to live in a cooperative society, but it is even better to live in such a society and not contribute to cooperation – to free-ride. If all, or even a majority of individuals are rational actors, who are only interested in maximizing their evolutionary fitness, then they will choose to free-ride and cooperation will unravel. The big question is how such collective-action problems were solved by evolution. A number of theories have been proposed and all, except one, founder on either logical or empirical grounds. The only theory that works is multilevel selection.
On the other hand, I agree that David toots the horn of group selection too much. Clearly, not all, and perhaps even a minority, of cultural traits were selected “for the good of the group.” And some cultural groups go extinct because they consist of individuals better adapted to the environment, not because they are more cooperative (think of early modern Europeans dying like flies in tropical Africa).
Still, the main question of social evolution (which is also the core issue of this Forum) is how huge and (somewhat) cooperative societies could evolve. Neither you nor Pinker offer a good explanation of this major evolutionary transition.
… then they will choose to free-ride and cooperation will unravel.
That is what actually happens periodically and what happens now in most developed countries and what causes those secular cycles you described in Secular Cycles. Free-riders or social parasites evolve at every social level among the elites, among those who represent the state, among those who develop into criminals, and even among those who live on charity.
Right. We made a lot of progress understanding how changing social, demographic, and economic factors create conditions conducive either to cooperation, or to its unraveling. And the multilevel selection theory provides an over-arching theoretical framework for understanding these effects.
Use *inclusive* fitness and this “problem” goes away. Ants build fantastic cooperative edifices, and their behaviour is completely consistent with – and is well explained by – inclusive fitness theory. If you think some kind of separate theory of group selection is required to explain such behaviour, your theory has led you off the rails.
Tim, how does kin selection account for unrelated genes cooperating in the genome?
That is only to do with kin selection (or group selection) in the case where genes exist in multiple copies in the same genome (e.g. LINEs and SINEs). Unrelated genes collaborate to build bodies through the Allee effect.
Peter,
I agree but would accentuate that as long as we agree that group/multilevel selection occurs in cultural evolution, that we no longer depend upon group selection within genetic evolution. All we need to show is that culture is possible in humans, then the selection mechanism for groups is introduced. I think the burden of proof for genetic group selection should be very demanding. Certainly it seems unnecessary.
Group selection is widely agreed by modern evolutionists to be kin selection. As such, it is ubiquitous in both the organic and cultural realms. The argument that group selection was of minor significance in the organic realm was based on a conception of group selection formulated by its critics. That formulation now appears to be archaic and unimportant.
Thanks Tim,
Could you do me a favor and direct me to further information on how group selection is now agreed to be kin selection? I am obviously not following.
Perhaps try my “Group selection and kin selection: formally equivalent” article – and its references. There’s also a nearby post on the positions of some of the naysayers.
Thanks Tim,
Very interesting.
Re: “If all, or even a majority of individuals are rational actors, who are only interested in maximizing their evolutionary fitness, then they will choose to free-ride and cooperation will unravel.”
You don’t think rational actors can cooperate? Why do you think that? It’s contrary to practically all the game theory I’ve come across. Kin selection and reciprocity can produce cooperation among rational actors. This is surely an old and well-understood point.
Re: “The only theory that works is multilevel selection.”
After accounting for the kin-selection / group-selection equivalence that seems incorrect. It’s like saying that: “the only theory that works is kin selection”. In fact, kin selection accounts for some cooperation. Reciprocity accounts for some more cooperation. Reputations account for some cooperation. Manipulations by memes account for some cooperation. This just isn’t the type of problem where one solution does all the work.
If you count an organism and its own offspring to be a “group”, than I think it would have to be admitted that the majority of a typical organisms’ adaptations exist to benefit some kind of group. If you don’t count an organism and its offspring to be a “group” then one would have to ask at this stage what kind of theory of group selection we are talking about – and whether or not it really deserves to use the name.
Thanks to Sanderson for his commentary–my critical reply notwithstanding! Thanks also to the respondents.
All aspects of Sanderson’s commentary need to be subjected to the equivalence test. You can’t evaluate multilevel selection without comparing fitness differentials within and among groups. Along with Pagel, Sanderson appears unwilling or unable to do this–yet he provides sufficient information in his examples for the reader to make the appropriate comparisons. When this is done, Sanderson emerges as a group selectionist. In other words, many of the traits that he posits do not evolve by virtue of fitness differentials within groups and require fitness differentials between groups.
Sanderson writes as if terms such as “self-interest” are unitary and self-explanatory, when they are neither. Defining “self-interest” as “the trait that maximizes an individual’s absolute fitness” is not the same as “the trait that maximizes the relative fitness of the individual within its group.” Both definitions are easy to grasp, so it’s a matter of clearly stating one’s definition and being able to translate among frameworks that employ different definitions. Most of the confusion surrounding group selection at this point can be attributed to the needless refusal of some people to become “bilingual” as I elaborate in my previous SEF essay titled “Clash of Paradigms”.
As an empirical matter, in many contexts, people are more sensitive to their relative advantage within groups than their absolute advantage, where “advantage” is a rough surrogate for fitness. This is captured by a proverb in which a genie grants a man a wish, subject to the constraint that his neighbor will get double. “Put out one of my eyes”, the man says.
On religion, if you read the progression of ideas in a book such as Durkheim’s Elementary Forms of Religious Life, it begins with the puzzle of why religions, with their counterfactual beliefs and costly practices, exist. Their secular utility is not at all obvious. Either they lack secular utility–which provides one category of explanation–or they have secular utility after all, which provides another category of explanation that Durkheim favored.
On Rodney Stark, when I categorized his formal theory of religion as a byproduct theory, it was so clearly stated by Stark that I merely had to quote him at length. His formal theory is distinct from his empirical work on religion, which indeed demonstrates many secular benefits. Please consult Darwin’s Cathedral for more.
On cultural evolution over long time scales, this is Peter Turchin’s specialty, which he describes as a process of multi-level cultural evolution. In addition to Turchin’s books, I recommend his article titled “Warfare and the Evolution of Social Complexity: A Multi-level Selection Approach” ( http://cliodynamics.info/PDF/WarComplx.pdf ). I hope that Sanderson will be conscientious and appropriately bilingual when he relates his ideas to Turchin’s.
On inter-societal selection, there is a big problem conflating proximate and ultimate causation, as I describe in my general reply. How an individual thinks and the consequences of their actions on themselves and others must be defined and evaluated separately. All four combinations are possible: Selfishly motivated individuals who act selfishly, selfishly motivated individuals who act altruistically, altruistically motivated individuals who act altruistically, and altruistically motivated people who act selfishly. Why is Sanderson so sure about the motives of other people? Why is he so sure that wars aren’t really about cooperation, when “solidarity” is the key factor according to Turchin? I hope that more lengthy treatments by Sanderson are more judicious than his commentary.
On consensus, it’s silly for anyone to think that if X people believe something, it must be true, no matter what the value of X. On the other hand, science is thankfully not a frictionless pendulum of ideas. It is a fact that the earth is round, which has caused large numbers of people to accept the fact. Similarly, it is a fact that traits often evolve based on the differential contribution of groups to the total gene/meme pool, despite being selectively disadvantageous within groups. It is gratifying that large numbers of individuals are starting to accept this fact, putting the flat-earthers in the minority.
Re: “But let’s think about cultural evolution – I prefer to call it social evolution – on a long-time scale […]”
Note that “culture” and “society” are quite different things. Ants have society without culture. These terms refer to different things – and are hardly interchangeable.