David Sloan Wilson Interview

| TBS Staff

Are you ready to discover your college program?

David Sloan Wilson is SUNY Distinguished Professor in the departments of Anthropology and Biology at Binghamton University. Co-founder of the Evolution Institute, the Binghamton Neighborhood Project, and the combined Evolutionary Studies (EvoS) program at Binghamton (which has been imitated by several other universities), Professor Wilson is the author of numerous scholarly and popular books on evolutionary theory and its implications for human society. His research has been focused on the notion of group selection as a modification to mainstream neo-Darwinian evolutionary theory and on social and cultural evolution among human beings.

Among Professor Wilson’s many books are Unto Others: The Evolution and Psychology of Unselfish Behavior, with Elliott Sober (Harvard UP, 1998), Darwin’s Cathedral: Evolution, Religion, and the Nature of Society (University of Chicago Press, 2002), Evolution for Everyone: How Darwin’s Theory Can Change the Way We Think About Our Lives (Delacorte Press, 2007), The Neighborhood Project: Using Evolution to Improve My City, One Block at a Time (Little, Brown and Co., 2011), and, most recently, Does Altruism Exist?: Culture, Genes, and the Welfare of Others (Yale UP, 2015).


Thank you very much for agreeing to participate in this interview. This is just the first phase of a Focused Civil Dialogue between you and Professor Denis Noble on the question: “Is Neo-Darwinism Enough?” Accordingly, during most of this interview we will be focusing on your work in evolutionary theory. First, however, we would like you to tell our readers a little bit about yourself: when and where you were born; your family background; your education; your career—that sort of thing.

David Sloan Wilson

I was born in Norwalk, Connecticut, in 1949. My father was the novelist Sloan Wilson, who wrote two blockbusters: The Man in the Gray Flannel Suit (1956) and A Summer Place (1958). He achieved his fame in 1956, when I was seven years old. Being the son of a famous author was formative for me. I admired him tremendously, but I also worried that I couldn’t measure up to him. My boyish solution was to choose interests that he could admire but couldn’t do. He was a novelist, so I would become a scientist.

[Book Image]My family moved around a lot and I can scarcely remember my homes and hometowns. However, we spent every summer on the north end of Lake George, New York, one of the most beautiful lakes in the world. My father’s family owned a hotel on the site that was torn down after World War II, leaving 600 acres of property, about a mile of lakefront, and about two dozen cottages that were maintained as a private club called the Rogers Rock Club. Roger’s Rock is a cliff that rises 600 feet out of Lake George and it was on our property. Can you imagine a more glorious way to spend every summer? I spent most of my time outdoors and became a passionate fisherman, a love that has stayed with me.

My parents sent me to boarding school when I was 11. Their marriage was deteriorating and according to my mother, I was following my father around saying “I’m sorry! I’m sorry!” Knowing how much I loved my summers on Lake George, they sent me to a progressive boarding school called the North Country School near Lake Placid, New York, which is still going strong. It featured outdoor activities, including a working farm that the students helped to operate. I thrived there and it cemented my lifelong attachment to nature. I wear a denim barn jacket, not a gray flannel suit!

After graduating from the North Country School in the eighth grade, I tried to pick another boarding school that was just like it: the Woodstock Country School in South Woodstock, Vermont. It turned out to be a rather cruel social environment in which the students formed into cliques and the rules were too lax to keep them under control. There wasn’t physical violence, but I observed a lot of psychological stress. Also, this was the 1960s and drugs were making the scene. I wasn’t unduly affected, but it caused me to become somewhat of a loner, which suited my temperament anyway. I did not mourn when the school eventually went belly up.

When it came time to apply to college, I aimed high—Harvard, Yale and Princeton—only to be rejected by all three. I ended up going to my “safety” college, the University of Rochester, which provided a fine education. I earned my Ph.D. at Michigan State University in 1975, followed by a succession of postdoctoral and tenured faculty positions, ending up at Binghamton University [formerly, SUNY Binghamton—Eds.] in 1989.


Looking back, what would you say most influenced your decision to become an evolutionary biologist?

David Sloan Wilson

When I decided to become a scientist at an early age, I had the white-coated lab scientist in mind. Gradually, I became aware that it was possible to become an ecologist and study organisms in their natural environment in addition to the laboratory. As you can imagine, given my love of nature, it was a very easy decision to become an ecologist. One of the great things about Rochester was the ability to work in the labs of the professors, more like a graduate student than an undergraduate. I started to function in this mode as a sophomore, thanks to an ecologist named Conrad Istock, who became my first mentor.

Book Image
Ernst Mayr’s Animal Species and Evolution (1963)

I was lucky to receive my higher education at a time when the biological sciences were becoming conceptually unified by evolutionary theory. I learned from Conrad that you can’t be an ecologist without also being an evolutionary biologist, because all species are a product of natural selection. I was a poor student when it came to the reductionistic branches of biology, but “natural selection thinking” came very easily to me. I remember reading Ernst Mayr’s 800-page Animal Species and Evolution (Harvard UP/Belknap Press, 1963) in a week, as if it were a novel.

Here is how I experienced conceptual unification as a student. Conrad was studying life history evolution in insects in his laboratory, but I became interested in vertical migration in zooplankton. No problem. In both cases, it was a matter of reasoning about what traits might evolve in a given species occupying a given environment. The same reasoning made it easy to switch from studying vertical migration in zooplankton to selective feeding in zooplankton—same organism, different set of traits. In graduate school I had the opportunity to study selective feeding in ant lions—different organism, same set of traits. Then I learned how to construct mathematical models of selective feeding—what would evolve in any species, given certain selective pressures. Natural selection thinking was the opposite of the old joke about experts learning more and more about less and less until they knew everything about nothing. Natural selection thinking was a passport to the study of all subjects. That’s the kind of higher education in biology that I and my peers received in the 1970s: the decade that witnessed Theodosius Dobzhansky’s famous dictum “Nothing in biology makes sense except in the light of evolution” in 1973 [1] and the publication of E.O. Wilson’s Sociobiology in 1975.[2]

The thesis of Sociobiology was that a single theoretical framework can explain social behaviors in all species, from microbes to humans. It was celebrated as a triumph—except for the final chapter on humans, which created a storm of controversy. I was at Harvard on my first postdoctoral position in 1975, working with the ecologist Tom Schoener on the adequacy of body size as a niche difference. I read a manuscript copy of Sociobiology that Ed Wilson had placed in the library of the Museum of Comparative Zoology before it was published. I regarded the final chapter as uncontroversial and was surprised when it created such a fuss.

Studying humanity from an evolutionary perspective had a special attraction for me because it is similar to the novelistic enterprise of trying to understand the human condition. My father did this by seeing through the lens of his personal experience. I could do it by seeing through the lens of evolutionary theory. I sometimes describe myself as a novelist trapped inside the body of a scientist!


You are best known for two things: (1) for your work on the conceptual foundations of the theory of natural selection, especially on the so-called “hierarchical model” of the selection (which we will ask you to explain shortly); and (2) for your public advocacy of evolutionary theory as a way of understanding human nature and human society, and even as a means of practical problem-solving in the social and political spheres.

In connection with the second interest, you have been heavily engaged in institution-building. You are best-known for three initiatives, in particular: The EvoS curriculum, the Evolution Institute, and the Neighborhood Project. Could you please tell us a little bit about the nature and aims of each of these remarkable undertakings?

David Sloan Wilson

Environment of Evolutionary Adaptedness

As background, it is important to establish that the study of study of evolution in relation to human affairs has a very different history than the study of evolution in the biological sciences. The latter developed more or less continuously since Darwin, while the former became politically incorrect early in the twentieth century. This is why the final chapter of Sociobiology created such a fuss. Terms such as “Evolutionary Psychology” and “Evolutionary Anthropology” weren’t coined until the 1980s, signifying a renewed effort to rethink the human-related academic disciplines from an evolutionary perspective, and even these had an air of scandal about them. As a result, evolutionary training in higher education is very largely restricted to the biological sciences. If you’re not a biology or a physical anthropology major, you won’t learn much about evolution, and that’s the way it has been since before the professors in the human-related departments were born. This problem exists around the world, unlike religious creationism, which is largely (although not entirely) confined to the United States.

As you might guess from my life story, I am among those who have been fearlessly and joyously expanding evolutionary theory beyond the biological sciences to include all things human. I regard humans as just another species that my evolutionary passport qualifies me to study. Over the years I have studied human-related topics as diverse as altruism, epistemology, personality, language, Machiavellianism, decision-making, gossip, physical attractiveness, emotion, and religion. These studies took place in parallel with my research on insects, fish, birds, parasites, and microbes.

The institution-building phase of my career began in 2003. I could see that historians would look back upon the twenty-first century as a period of synthesis of knowledge about humanity, comparable to the synthesis of biological knowledge during the twentieth Century (which continues). Given the conservatism of academic culture, however, I could also see that without a concerted effort, decades would be required for the twenty-first century synthesis to be reflected in higher education, on my campus or any other campus. I therefore worked to create a campus-wide program for teaching evolution across the curriculum that we called EvoS (for Evolutionary Studies and pronounced as one word).

We built EvoS from elements that exist at most colleges and universities, including a scattering of faculty who are already employing an evolutionary perspective across human-related disciplines, but who are isolated within their respective departments. My book Evolution for Everyone (Delacourt, 2007) was written on the basis of EvoS and numerous academic articles describe the program in detail.[3] A sister program was started at SUNY New Paltz by the evolutionary psychologist Glenn Geher. Together we wrote and received a NSF grant to develop our programs and create a multi-institution consortium. There is now a consortium website and online journal, about six full-fledged campus-wide programs, more nascent programs, and many more single courses that use Evolution for Everyone as a text. That’s pretty good, but there is still a long way to go.

The Binghamton Neighborhood Project uses the community surrounding Binghamton University as a “field site,” as an evolutionary ecologist would use the term. You can’t get started studying a species from an evolutionary perspective unless you know something about the species in relation to its environment. That’s why field research is the starting point of evolutionary inquiry and all laboratory research needs to be informed by field research. Yet, most research in the human behavioral sciences is not like this and the most field-oriented disciplines, such as sociology and cultural anthropology, have also historically been the most phobic about evolution. Thus, doing what comes naturally as an evolutionary ecologist turns out to be a new model for basic and applied community-based research on humans, as I describe in my book The Neighborhood Project (Little, Brown, 2011).

A year after I started to do community-based research, I was approached by a retired political scientist and lifelong humanist named Jerry Lieberman with the idea of creating a think tank. Jerry wanted it to be science-based and was persuaded by my book Evolution for Everyone that it should be informed by evolutionary theory. I was already hooked on the idea of applying evolutionary theory to the solution of real-world problems, so I eagerly accepted Jerry’s invitation and the Evolution Institute (EI) was born.

I’ll have the opportunity to describe EI projects later in this interview, but here I will mention its communication outlets: This View of Life (for a general audience), the Social Evolution Forum (for a professional audience), Evonomics.com (for an economics audience), and PROSOCIAL Magazine (for the community of groups using PROSOCIAL, which I will describe later). Collectively, they reach hundreds of thousands of readers every month, spreading the word about the twenty-first-century synthesis in the same way as EvoS.


Now, let us move on to your work on the foundations of evolutionary theory.

T.H. Morgan's Fly Room
T.H. Morgan in the “Fly Room” at Columbia University

From the publication of the Origin of Species in 1859, to the rediscovery of Mendel’s work on inheritance by Hugo de Vries, William Bateson, and others around the turn of the twentieth century, to the further elucidation of genetic principles by T.H. Morgan in his “fly room” at Columbia in the 1920s, to the “modern synthesis” of natural selection with population genetics by R.A. Fisher, Sewall Wright, J.B.S. Haldane, and others in the 1940s, the theory of evolution had always revolved primarily (with some exceptions) around individual organisms.

Darwin’s key idea—supplemented and reinforced, but not essentially changed by the discoveries in genetics—was that random genetic variation at the genetic level leads to differential fitness at the phenotypic level of individual organisms, which in turn drives differential reproduction and thus change in the distribution of genes within the population to which the individuals belong.

Against this background, the idea for which you are most famous—that natural selection takes place at more than one level (is “hierarchical”), including that of the social group (“group selection”)—assumes a radical aspect.

One of the main phenomena that the theory of group selection was developed to explain is what biologists call “altruism,” a term of art meaning a phenotypic trait “that contributes to group advantage at the expense of disadvantage to itself.”[4] Here, “advantage/disadvantage” is understood to mean Darwinian reproductive success/failure (increase/decrease in the proportional representation of an organism’s genes in subsequent generations). An example would be sterile castes among social insects, such as soldier ants, worker bees, and the like.

This phenomenon—organisms which seem to act for the good of the group to which they belong, rather than their own good—seemed very difficult to account for along standard Darwinian lines which had always emphasized relative fitnesses and competition among individuals. This is the problem (and the historical context), as we understand it, of the notion of “group selection” that you have advanced in your work.

Assuming little more knowledge of evolution on the part of our readers than has been sketched above, could you please explain to us in simple terms how you view the “hierarchical” nature of natural selection? That is, in a nutshell, how would you characterize your own main theoretical contribution to evolutionary theory?

David Sloan Wilson

Let me revise your historical account a little bit. According to the Christian worldview, a universe created by an all-powerful and beneficent God must be harmonious from top to bottom, from the smallest insect to the stars in heaven. The first Enlightenment thinkers, such as Isaac Newton, thought that science and reason would affirm scripture in this respect. Darwin’s theory led to a very different conclusion: that the kind of functional organization that we associate with a human implement such as a watch or a single organism such as an insect, might cease to exist at larger scales, such as an animal society or a multi-species ecosystem.

Sterile Worker Bees

Darwin was led to this conclusion by considering the evolution of traits that are “for the good of the group” and therefore morally praiseworthy in human terms. He could see that acting for the benefit of others or one’s group as a whole typically placed the “altruist” at a selective disadvantage, compared to more selfish individuals within the same group. This was a problem of the first rank.

Not only did Darwin clearly see the problem, but he also saw the outline of a solution. Groups of altruists have a selective advantage over groups of selfish individuals, as surely as selfish individuals have an advantage over altruistic individuals within groups. As Edward O. Wilson and I put it in our 2007 review article titled “Rethinking the Theoretical Foundation of Sociobiology,”[5] selfishness beats altruism within groups, altruistic groups beat selfish groups, and everything else is commentary. Thus, the basic concept of multilevel selection theory was present at the start of evolutionary theory and was required to address a fundamental problem concerning levels of functional organization.

Notice that in the basic scenario imagined by Darwin, the evolving traits (altruistic and selfish behaviors) can be measured in individuals. In a genetic model, they are both coded by genes. Multilevel selection theory is concerned with fitness differences in a nested hierarchy of scales—favoring selfish traits (and genes) at the scale of individuals within groups and favoring altruistic traits (and genes) at the scale of groups within a multi-group population. This will become important when we consider the tortuous history of the group selection controversy.


The idea of group selection was first brought into the mainstream of modern evolutionary discourse by V.C. Wynne-Edwards in his classic work, Animal Dispersion in Relation to Social Behaviour (Edinburgh: Oliver & Boyd, 1962). After much rancorous debate, Wynne-Edwards’s work was considered to have been thoroughly “debunked,” and was cast into the outer darkness by the neo-Darwinian mainstream. Your work has had a crucial impact on the grudging tolerance now accorded to group selection today.

Could you please tell us (1) how you became convinced that Wynne-Edwards was at least partly right; and (2) without going into too much technical detail, how your own view of group selection differs from his?

David Sloan Wilson

I wish it was otherwise, but scientists are as prone to constructing simplified patriotic histories as other people. Good scholarship is required to set the record strait. Fortunately, this period in the history of evolutionary thought is beginning to get the attention it deserves. The brief account that Elliott Sober and I gave in Unto Others has withstood the test of time, and has been supplemented by books such as Evolutionary Restraints by Mark Borrello and The Price of Altruism by Oren Harman.[6] Here is a thumbnail history.

As important as the altruism question was, it was eclipsed by even more important questions such as the nature of heredity during the first half of the twentieth century. R.A. Fisher, J.B.S. Haldane, and Sewall Wright, the three main architects of population genetics theory, each considered the problem briefly, sketching simple mathematical models along the lines of Darwin’s verbal scenario. In the meantime, many empirical biologists naively assumed that adaptations can evolve at any level of the biological hierarchy without requiring special conditions. When the need for higher-level selection was recognized, it was often assumed that higher-level selection easily trumps lower-level selection. This position became known as “naïve group selectionism” and it was a genuine problem recognized by George C. Williams in the 1950’s. Williams was reacting to biologists such as W.C. Allee at the University of Chicago and he started writing his classic book, Adaptation and Natural Selection (Princeton UP, 1966), years before Wynne-Edwards’s book was published.

V.C. Wynne-Edwards

Wynne-Edwards proposed that animals evolve to regulate their population size to avoid overexploiting their resources. He was aware that his hypothesis required group selection—in this sense he was not naïve—but he assumed without justification that between-group selection easily trumps within-group selection. He was not a theoretical biologist and cited Sewall Wright for support, even though Wright had written little on the subject (not to be confused with Wright’s shifting balance theory, which addresses a different set of issues). Wynne-Edwards then provided an encyclopedia of examples across the animal kingdom that he interpreted in support of his hypothesis.

The reaction to Wynne-Edwards’s book in 1962, along with the publication of Adaptation and Natural Selection in 1966, caused multilevel selection theory to occupy center stage in evolutionary biology for the first time. Note that this is about two decades after the so-called Modern Synthesis. New mathematical and computer simulation models were constructed, which suggested that the conditions for between-group selection prevailing against within-group selection were quite restrictive. It was easy to poke holes in Wynne-Edwards’s empirical examples. And other ways to explain the evolution of apparently altruistic behaviors “without invoking group selection” became available, as we will shortly discuss.

Here is how I entered the field. I was offended by the idea that altruism can’t evolve when I encountered it at the University of Rochester as an undergraduate student. One of Wynne-Edwards’s putative examples of group selection was vertical migration in zooplankton. A common pattern is for adults to migrate during the day, while the young remain near the surface. This is probably a response to size-dependent predation, but Wynne-Edwards interpreted it as a form of mass parental care—adults migrating so they don’t compete with their offspring for food. I was studying vertical migration in zooplankton and thought that Wynne-Edwards’s hypothesis could not be dismissed out of hand. Far from homogenizing zooplankton populations, ocean currents and wave action concentrate them into patches. By vertically migrating, adults were very likely to horizontally separate themselves from their offspring. I included this speculation in my undergraduate thesis, but went on to other topics in graduate school.

In my final year of graduate school, a new article on vertical migration prompted me to dust off my old idea. By then I had picked up some theoretical skills and the model of within- and between-group selection that I developed went far beyond vertical migration in zooplankton. My main innovation was to define a group as the set of individuals influenced by the expression of a given trait. In fact, this is how groups are usually defined in natural language. When we say “my family,” “my class,” “my church,” “my bowling club,” we are defining groups in terms of relevant activities. In any model of social evolution, it is necessary to determine the fitness of individuals, which requires defining the set of neighbors influencing a given individual’s fitness. Thus, my concept of trait-groups made explicit something that is implicitly assumed in all models of social evolution.

My model showed that while between-group selection did not invariably trump within-group selection, neither could it be categorically dismissed. The relative importance of within- and between-group selection needed to be determined on a case-by-case basis. I knew immediately that this was—or should be—a game-changer for the group selection controversy. I was so fired up that I wrote to the great E.O. Wilson requesting a meeting, in hope that he would sponsor an article for the Proceedings of the National Academy of Science (PNAS). On my way to Harvard to meet with Wilson, I made a side trip to Stony Brook to meet with George C. Williams. My first words after striding into his office were “I’m going to convince you about group selection.” His first response was to offer me a post-doc on the spot! I had other plans, but we became good friends despite our very different positions on group selection. This is how science should be.

Ed Wilson did sponsor my article for PNAS, which was the beginning of my relationship with him. My next problem was what to do for my PhD. My previous research topic no longer held any interest for me! My thesis advisor was a free-spirited aquatic ecologist named Don Hall. He reasoned that if my article on group selection was good enough for Ed Wilson and PNAS, it was good enough to be a Ph.D. thesis. I therefore probably have the shortest Ph.D. thesis in the history of evolutionary biology (11 pages), although I did write several other publications on other topics as a graduate student that were not included in my thesis.


We gather that Wynne-Edwards was something of an isolated, even tragic, figure. Did you know him personally? If so, what was he like? What do you think his place in the history of science will be in the long run.

David Sloan Wilson

I never met him but we corresponded on numerous occasions. He was a vigorous man and I don’t think that he was crushed by the controversy. I regard him as less pivotal than the simplified history makes him out to be. He did not play any role in the construction and testing of theoretical models, which was the main action. His second book in 1986 [7] did not reflect much understanding of everything that had happened to revive multilevel selection theory, which we will shortly discuss.

If we turn our attention away from the person and focus on his hypothesis, there is now solid evidence that some consumer species do regulate their population size to avoid overexploiting their resources. This is not invariably the case, but it is sometimes the case. The best examples are parasites and diseases that reduce their virulence to avoid killing their hosts, which were showcased—and correctly interpreted—by none other than George C. Williams when he started to pioneer the subject of “Darwinian medicine.” Thus, the categorical rejection of Wynne-Edwards’s hypothesis does not reflect well upon his critics.

"Would I lay down my life to save my brother? No, but I would to save two brothers or eight cousins." — JBS Haldane


One of the main reasons why group selection fell from favor in the ‘60s was the development of alternative, individual-selection-based theories to explain altruism: especially W.D. Hamilton’s notion of “kin selection” [8] and Robert Trivers’s concept of “reciprocal altruism” (based on game theory).[9] George C. Williams famously argued that, given the existence of these ideas, it is more parsimonious to reject group selection.

As we understand it, your position is that kin selection and game-theoretic reciprocity are forms of group selection, so Williams was wrong. We gather that this claim remains highly controversial.

Could you briefly (if such a thing is possible!) give us your reasons for believing this? Where, exactly, did Williams go wrong?

David Sloan Wilson

George got some things very right and other things very wrong. He was right to forcefully assert the logic of multilevel selection, which is: Group-level adaptations require a process of group-level selection and should never be invoked otherwise. He was wrong to conclude on the basis of empirical evidence that between-group selection is invariably weak, compared to within-group selection. Once we abandon this sweeping empirical claim, we arrive at the reasonable conclusion that the balance between levels of selection needs to be determined on a case-by-case basis.

I know from my conversations with George that he wrote Adaptation and Natural Selection to educate a broad biological audience in basic concepts from population genetics. One of these is the concept of average effects, which is essentially the “gene’s eye view” of evolution further popularized by Richard Dawkins. Consider two alleles, A and a, at a single genetic locus. They exist in three genotypic combinations, AA, Aa, and aa. They also exist in different combinations with genes at other loci and in different social groupings in a multi-group population. It is possible to average the fitness of the two alleles across all of these contexts to achieve a bottom-line estimate of which gene has the highest fitness and therefore increases in frequency in the total population. Williams called this an “accounting method” and it is indeed one of the most useful tools in the population genetics toolbox.

A problem arises, however, when average effects are taken to be an argument against group selection. When altruism evolves in a group selection model, the gene for altruism is more fit than the gene for selfishness, all things considered. It achieved its advantage on the strength of fitness differences between groups and despite opposing fitness differences within groups. The fact that it has the highest average effect, and therefore increases in frequency in the total population, is not an argument against group selection! Yet that is exactly how “the gene’s eye view” of evolution became interpreted. This confusion began with Williams and was amplified by Dawkins.


As we understand it, you believe that one of the main reasons the idea of group selection has been so maligned over the years has to do with something called the “averaging fallacy.” Could you explain in simple terms what this means?

David Sloan Wilson

The concept of average effects that I just described is an example of the averaging fallacy. By itself it is a useful accounting method, but it becomes a fallacy when used as an argument against group selection. N-person game theory provides another example. Individuals employing different social strategies are assumed to interact in groups of size N. Cooperators are less fit than defectors within any given group, but groups with more cooperators are more fit than groups with fewer cooperators. The logic of multilevel selection is there for anyone to see, but it is obscured when average payoffs are calculated for each strategy, similar to the average effects of genes. Once again, there is nothing wrong with this procedure; it only becomes a fallacy when taken as an argument against between-group selection.

IllustrationIn this fashion, every theory of social behavior that was developed as an alternative to group selection, including kin selection theory (=inclusive fitness theory), evolutionary game theory (including reciprocal altruism), and selfish gene theory, turns out to include the logic of multilevel selection theory within its own structure. They all assume that social interactions take place within groups that are small compared to the total population. The behaviors labeled “cooperative” or “altruistic” are selectively disadvantageous within these groups and evolve only by virtue of fitness differences between groups. My concept of trait-groups helped to make this clear, along with a statistical method for partitioning selection into within- and between-group selection that was developed by George Price and convinced W.D. Hamilton that kin selection is a type of group selection. Hamilton announced his “conversion” in an article written in 1975, the same year that my PNAS article was published. Oren Harman’s The Price of Altruism provides a book-length account.

So, all major theories of social evolution include the basic logic of multilevel selection and their differences are primarily a matter of perspective. This has become known as “Equivalence” and it has become accepted by most authors of peer-reviewed articles. In 1975, it was almost mandatory to say that one’s ideas did not invoke group selection. Today, it is almost mandatory to say something like this:[10]

In earlier debates, biologists tended to regard kin and multilevel selection as rival empirical hypotheses, but many contemporary biologists regard them as ultimately equivalent, on the grounds that gene frequency change can be correctly computed using either approach. Although dissenters from this Equivalence claim can be found, the majority of social evolutionists appear to endorse it.

That’s quite a change! It demonstrates that the group selection controversy is essentially over. In my latest book, Does Altruism Exist? (Yale UP, 2015), I say that I offer a “post-resolution” account.

Stephen Jay Gould


Stephen Jay Gould famously believed that the Modern Synthesis had undergone a “hardening” after World War II, which led to a decrease in healthy diversity of viewpoints within evolutionary biology and a growing intolerance for heterodox opinions such as yours.[11]

Thankfully, that appears to be changing now. As the late theoretical biologist Robert Rosen put it not long before his death, there is

… a real sea change in science today; a general increase in conceptual temperature which is liquefying outmoded doctrines which have hovered around absolute zero for the past half-century or more.[12]

Would you agree with Gould that the “hardening of the Modern Synthesis” was real? If so, what role do you believe it played in the reception of Wynne-Edwards’s and your own work?

Do you agree with Rosen and us that the situation has changed drastically over the past few years, for the better?

David Sloan Wilson

I think that we need to take a nuanced view of so-called “hardening.” Consider the sensory organs of any species. They have evolved to be highly selective at perceiving and processing information from the environment. In the case of our species, we can see only a narrow segment of the light spectrum and hear only a narrow segment of the sound spectrum, and we can’t perceive electrical currents or magnetic forces at all. It is necessary to be selective, because attending to everything would lead to paralysis. The same is true for scientific ideas, which means that there is a positive side to “hardening.” Putting some things in the foreground and others in the background can be very useful. That said, it is also necessary to periodically revisit and reconfigure such “hardenings” and there is a tendency for them to become frozen into rigid dogmas. Major “hardenings” in the history of evolutionary thought include the distinction between “Darwinian” and “Larmarkian” inheritance, the ideas associated with the Modern Synthesis in the 1940s, and the consensus against group selection in the 1960s. In each case, I agree with Gould and Rosen that a major loosening was in order and that decades shouldn’t be required. Really, the group selection controversy should have been settled by the 1980s and the fact that it took longer does not reflect well upon the scientific process as actually practiced.

David Sloan Wilson is third from the left

The term “Extended Evolutionary Synthesis (EES)” describes current efforts to loosen past hardenings.[13] I am an enthusiastic proponent of the EES and the This View of Life website is devoting a series of articles to it, beginning with this interview with Kevin Laland, who is heading a major grant from the John Templeton Foundation to fund research on the EES.


As everybody knows, Richard Dawkins popularized Hamilton’s idea that individual genes are the main, if not exclusive, units of selection in his book, The Selfish Gene (Oxford UP, 1976). Since you obviously believe Dawkins is mistaken on this point, how do you account for the fantastic popularity of his books and of the idea of the “selfish gene” (genic selection), in particular?

David Sloan Wilson

First, I would like to praise Richard Dawkins in many respects. He’s a gifted writer and thinker who has turned legions of people on to evolutionary theory. He’s right about lots of things, even if he’s wrong about group selection. One way to rephrase your question is to ask if the “everything is caused by our selfish genes” element of Dawkins’s thought is responsible for some of his popularity. The answer is almost certainly yes—in part because of its shock value (a smart rhetorical ploy) and in part because it resonates with a broader individualistic worldview that we will be discussing later.

While on the subject of Dawkins, I am much more critical of his stance on religion, which has seriously tarnished his reputation.


The units/levels of selection debate is a highly convoluted and controversial affair.[14] In some ways, it seems a lot closer to philosophy than to ordinary empirical science.

If your hierarchical theory is correct, then selection takes place at multiple levels. But how can we determine which units at which levels are the actual basis for selection in a given case? Is there really any real empirical substance to the units of selection debate?

David Sloan Wilson

It is sad that this question has to be asked. When we operate within the framework of MLS theory, measuring fitness differences within and between groups is ordinary empirical science. The work of Omar Eldakar (my former graduate student) on sexual conflict in water striders provides an example.[15]

Males differ greatly in their aggressiveness toward females. Omar created pools containing six males and six females, with the proportion of aggressive males varying from 100% to 0% and various mixes in between. Within every pool containing both types, the aggressive males copulated with the females more than the docile males, which was perfectly easy to quantify. However, females laid more eggs in pools with docile males than pools with aggressive males because they weren’t being harassed. This was also easy to quantify.

In a second experiment, Omar allowed free movement between the pools. Females that entered a pool with aggressive males left as soon as they could. Aggressive males were free to leave also, but the result of everyone moving was a considerable degree of clustering of females around docile males. Thus, free movement creates the variation among groups necessary for group selection to act. This was also easy to quantify.

To summarize: Selfishness beats altruism within groups (favoring aggressive males); altruistic groups beat selfish groups (favoring docile males); and in this case the balance between levels of selection maintains both types of male in the population. All of this can be quantified and many other examples can be cited.

What gives the levels of selection debate its “convoluted and controversial” aspect, as you put it, is the confusion between theories that invoke different causal processes, such that one can be right and the other wrong, and theories that describe the same causal processes from different perspectives and deserve to coexist to the extent that they provide useful insights. The concept of Equivalence does a good job of resolving this confusion. Once we become clear on Equivalence, we can do normal science within each framework and become adept at translating among frameworks.


Thank you. That was extremely helpful.

Some have claimed that where one stands on the group selection controversy is not unrelated to where one stands politically—to put it crudely, that “neo-liberals” love old-fashioned individualistic natural selection, while communitarians and socialists prefer group selection. Do you think there is anything to this?

David Sloan Wilson

Yes, indeed. Just like other people, scientists have worldviews that influence what they find plausible and would like to be true. That’s why it is important for a scientific community to include a diversity of worldviews. As long as personal biases lead to testable hypotheses, they become grist for the scientific mill.

Charles Darwin

Cultural biases are more difficult to correct when they are shared by most members of a scientific community. With the benefit of hindsight, we can see the imprint that Victorian culture had on Darwin and most of his peers on topics such as the mental inferiority of women and the superiority of European culture. No one questioned these assumptions in their everyday lives, so they had no reason to question them as scientists. I think that the swing toward individualism in evolutionary theory during the second half of the twentieth century needs to be seen against the background of a more general cultural swing that included a position known as methodological individualism in the social sciences, and of course neoclassical economic theory. When UK Prime Minister Margaret Thatcher said “There is no such thing as society, only individuals and families,” she was reflecting this trend. I hope that historians and sociologists of science pay close attention to this period.


Let’s shift gears a bit now and talk about how evolutionary theory impacts (or ought to impact) human self-understanding.

The landmark work in this area, of course, was E.O. Wilson’s Sociobiology (Harvard UP/Belknap Press, 1975). In that book, if memory serves, E.O. Wilson explicitly embraced Hamilton’s kin selection and abjured Wynne-Edwards’s group selection. And yet very recently, he has signed on as co-author with you and others in a series of controversial papers intended to make group theory intellectually respectable once again. Has he ever told you what made him change his mind?

David Sloan Wilson

Actually, I have a different impression. Ed was as supportive of group selection as anyone could be when he wrote Sociobiology. Of course, he also lauded Hamilton’s theory at a time when its relationship with group selection wasn’t even clear to Hamilton. Ed’s current critique of kin selection theory with Martin Nowak is complicated and I have faulted them, along with Richard Dawkins, for failing to absorb the message of Equivalence. However, when it comes to human self-understanding, we definitely must go beyond kin selection theory.

Human evolution is all about a shift in the balance between levels of selection. In most primate societies, members of the same group cooperate to a degree, but are also each other’s main rivals. To the best of our current knowledge, our ancestors managed to suppress the potential for disruptive self-serving competition within groups, so that between-group selection became the dominant evolutionary force. Almost everything that is distinctively human, including our ability to cooperate with unrelated individuals, our capacity for symbolic thought, and our ability to transmit large amounts of learned information across generations, are forms of physical and mental teamwork that required between-group selection to evolve. Books that develop this thesis include Maynard Smith and Szathmary’s The Major Transitions in Evolution, Deacon’s The Symbolic Species, Boehm’s Moral Origins, E.O. Wilson’s The Social Conquest of Earth, and Turchin’s Ultrasociety.[16]


With respect to the evolution of human social practices and institutions, your work has ranged over many topics. But let us take as an example religion, which was the focus of your book Darwin’s Cathedral (University of Chicago Press, 2002).

How, exactly, does a “Darwinian” explanation of religion work? How does one go about shedding light on a human social phenomenon like religion by invoking evolutionary theory, in general, and group selection, in particular?

David Sloan Wilson

Religions puzzle the secular imagination because their beliefs depart so flagrantly from factual reality and result in practices that seem so wasteful. It’s easy to understand why people make blankets, but why do they burn them in sacrifice to gods for whom there is no verifiable evidence? This question has two potential answers.

First, religious beliefs and practices might be just as irrational and wasteful as they seem and persist as byproducts of psychological and social processes that are useful in non-religious contexts.

Second, despite appearances, religious beliefs and practices might have their own logic and utility after all.

Émile Durkheim

Émile Durkheim was an early proponent of the latter view. He famously defined religion as:[17]

A unified system of beliefs and practices relative to sacred things … which unite into one single moral community called a Church, all those who adhere to them.

Durkheim also stressed the importance of symbolic thought in the organization of human societies:[18]

In all its aspects and at every moment of history, social life is only possible thanks to a vast symbolism.

Nevertheless, over a century of scholarship in the humanities and social sciences has not led to a consensus on the “secular utility” of religion, as Durkheim put it. The tradition of functionalism that he initiated peaked in the mid-twentieth century and is currently disparaged in many quarters. When I was writing Darwin’s Cathedral at the turn of the twenty-first century, the most authoritative theory of religion was a byproduct theory inspired by economics, which held that gods are imaginary beings that people invent to bargain with for goods that can’t be had, such as rain during a drought or everlasting life.

Evolutionary biologists are accustomed to studying whether a given trait qualifies as an adaptation vs. a byproduct, the unit of selection, and so on—all questions that can be asked for cultural evolution in addition to genetic evolution. When this theoretical toolkit started to be applied to religion, it established a consensus that did not previously exist: Appearances notwithstanding, most enduring religions have an impressive degree of secular utility at the level of the religious community, much as Durkheim posited. Religions are also replete with byproducts, just as biological adaptations are, but the view of religion writ large as a byproduct has been authoritatively rejected. This is an excellent demonstration of the “added value” that evolutionary theory brings to the human social sciences.


We would like to return now to this question of the “softening” of the Modern Synthesis that we alluded to earlier.

It seems to many observers today that evolutionary biology is in a state of flux—even, perhaps, of long-overdue reassessment. You yourself participated in a widely advertised conference that billed itself as presenting an “extended synthesis.”[19] Your interlocutor in the upcoming TBS Focused Civil Dialogue on evolution, Denis Noble, is involved with a group that calls itself the “Third Way of Evolution.”[20]

For the rest of this interview, we would like to begin the process of exploring what these notions really amount to—a process which will be continued in the subsequent statements to be submitted by you and Noble.

First, we would like to ask you a fundamental philosophical question—one which most practicing scientists shy away from as obscure and remote from their day-to-day concerns—but one which we believe is not only absolutely critical to our self-understanding as animals and as rational beings, but also of potential practical importance for science itself. We are thinking of the “problem of teleology.”

It has been argued by Mary Jane West-Eberhard [21] and others that biological matter is inherently adaptive and that every instance of phenotypic change (e.g., due to genetic mutation) must be understood against the backdrop of active and adaptive response and compensation to the change on the part of the organism, at least if a new viable form is to result. It this is true, then while variation may be “random” at the genetic level, it is non-random and adaptive at the phenotypic level. This means that the neo-Darwinian theory of natural selection does not dispense with teleology, but rather silently presupposes it—it is the inherent adaptive capacity of the organism that is doing all the explanatory work, not the random-variation-and-selective-retention schema.

What do you say to this? Do you think there is such a thing as internal or immanent teleology (purpose, goal-directedness), conceived of as an objective feature of “the living state of matter”?

If so, do you think the neo-Darwinian framework completely accounts for it (perhaps in terms of Ernst Mayr’s notion of “teleonomy,”[22] or in some other way)?

If so, what do you say to West-Eberhard and similar critics?

David Sloan Wilson

There is a lot of ground to cover here. First, an essential book on this topic is Jablonka and Lamb’s Evolution in Four Dimensions.[23] Eva Jablonka is listed as a member on the Third Way of Evolution website. I use it as a first text in most of my courses and agree with just about everything in it. In general, I endorse the need to study evolutionary theory and complex systems theory in conjunction with each other. After all, organisms are complex systems living in environments that are also complex systems.

Stuart A. Kauffman

However, anyone who invokes complexity must provide a way to navigate through it. Merely citing dozens of factors that interact with each other is not helpful and leads to what can be called “combinatorial paralysis.” In addition, there is an ironic tendency of some thinkers to treat evolutionary theory and complex systems theory in an either/or fashion, as if to say “Darwin is dead! Long live Complexity!” Stuart Kauffman’s Origins of Order suffered from this problem.[24] The challenge is to study the two bodies of theory in conjunction with each other—evolution operating on complex systems.

A recent essay of mine titled “Two Meanings of Complex Adaptive Systems”[25] addresses some of the confusion associated complex systems thinking. This key term lumps two very different meanings: 1) A complex system that is adaptive as a system; and 2) A complex system composed of agents that follow adaptive strategies. A social insect colony is an example of CAS1 and an ecosystem is an example of CAS2. Many treatments of Complex Adaptive Systems fail to distinguish between these two meanings or to specify the conditions required for a system to qualify as CAS1. This amounts to a form of naïve group selectionism in authors who are otherwise very sophisticated. A complex system by itself can produce a lot of pattern, but this pattern is no more likely to be functionally adapted to a given environment than a point mutation. System-level selection is required to produce a complex system that is adaptive as a system (CAS1)!

An exception is when a Complex Adaptive System is designed by a past evolutionary process to be anticipatory or itself an evolutionary process—what William H. Calvin [26] and Henry Plotkin [27] call “Darwin machines.” Epigenetic inheritance systems, forms of social learning found in many species, and forms of symbolic thought that are distinctively human (the 2nd, 3rd, and 4th dimensions of evolution discussed by Jablonka and Lamb) are all Darwin machines that evolved by genetic evolution (the 1st dimension). In addition, genetic inheritance mechanisms are a product of genetic evolution—they didn’t begin as sophisticated as they are now! Thus, Wagner and Altenberg’s concept of “the evolution of evolvability” has merit.[28]

All of these call for a “loosening” that is represented by the term “Extended Evolutionary Synthesis” and is being competently developed by people such as Jablonka, Lamb, and Kevin Laland.[29] But notice that the term “Extended Evolutionary Synthesis” is carefully chosen to ring in the new without announcing a break in the past. Jablonka and Lamb make the same point when they say that if Lamarckian mechanisms of inheritance do exist, the basic concept of adaptation and natural selection will remain much the same (e.g., giraffes will still evolve long necks to browse tall trees).

In another recent essay of mine titled “Intentional Cultural Change,”[30] I make the point that even if we stick to the traditional view of genetic evolution as a purposeless process (“blind variation and selective retention”), it clearly results in organisms that behave in a purposeful fashion—intelligently looking for food and mates, avoiding predators, and so on. Ernst Mayr thought it was important to call this “teleonomy” rather than “teleology,” but I have always thought that the similarities are more interesting to focus upon than the differences. We know since James Mark Baldwin [31] that purposeful animal behaviors can double back to influence the evolutionary process in a kind of indirect Lamarckism. Some of the recent discoveries about anticipatory mutations, epigenetic systems, and developmental systems duplicate what we have long known about learning.

Human cultural evolution indubitably has an intentional component, but it also has a very substantial blind component. Moreover, intentional planning has a way of converting to blind variation when intentions collide and result in unforeseen consequences. Cultural evolution needs to become more intentional and needs to focus directly on the solution to global problems. The intentional pursuit of lower-level interests, such as individual wealth or national self-interest, will usually be disruptive at higher scales. That’s the basic message of multilevel selection theory, which is profoundly antithetical to the concept of the invisible hand in economics.[32]


West-Eberhard is only one of many investigators who are now working under the general banner of “evolutionary developmental systems” theory (or “evo-devo”). Another similar recent development is the burgeoning field of “epigenetics”—the study of modifiable and heritable non-DNA genetic markers.

In brief, proponents of evo-devo and epigenetics are renewing a former emphasis on the whole organism in evolutionary studies, while de-emphasizing to some extent the traditional role of the neo-Darwinian model of random-variation-and-selective-retention.

What is your attitude towards these recent developments?

David Sloan Wilson

A classic article by Niko Tinbergen titled “The Methods and Aims of Ethology” is worth describing in this regard.[33] Tinbergen pioneered the study of animal behavior (ethology) and shared the Nobel Prize in Medicine with Konrad Lorenz and Karl von Frisch in 1973. Before the field of ethology was established, it wasn’t obvious that a behavior such as aggression could evolve in the same way as a physical trait such as a deer’s antlers. In the process of describing ethology as a branch of biology, Tinbergen wrote that four questions need to be addressed for all products of evolution:

  1. What is their functional basis (if any)?
  2. What is their physical mechanistic basis?
  3. How do they develop during the lifetime of the organism?
  4. What is their historical evolution (phylogeny)?

Ever since, “Tinbergen’s four questions” have been cited as a compact description of a fully-rounded evolutionary approach.

Against this background, evo-devo and epigenetics provide exciting new answers to Tinbergen’s “mechanism” and “development” questions. Evo-devo is especially important because development is arguably the most neglected of Tinbergen’s four questions, having been largely pushed into the background by the Modern Synthesis. However, more attention to the mechanism and development questions does not require less attention to the function question! I therefore disagree with the suggestion that good old-fashioned “natural selection thinking” needs to be de-emphasized. Reasoning on the assumption that heritable phenotypic variation exists, without needing to know about the physical or developmental basis of the variation, will always be one of the most powerful tools in the evolutionary toolkit.

Incidentally, I recently conducted an interview with Richard Lenski that provides an overview of his research on long-term evolution in E. coli from the perspective of Tinbergen’s four questions. I titled the interview “Evolutionary Biology’s Master Craftsman” to emphasize the idea that an evolutionist is like a carpenter or a plumber, who shows up on a worksite (a particular topic of interest), sizes up the job, and pulls out the appropriate tools to get the job done. The most important tools in the evolutionary toolkit are conceptual tools, not physical tools. Tinbergen’s four questions did an excellent job of describing Lenski’s fully-rounded evolutionary approach. Whatever we mean by the “Extended Evolutionary Synthesis” or “Third Way of Evolution,” it does not require a radical departure from Tinbergen’s four questions.


Some who would agree with the recent emphasis on the whole organism at the expense of neo-Darwinism’s relentless preoccupation with genes and replication would say that it still does not go far enough.

After all, it seems clear that in some sense “metabolism” (shorthand for the ensemble of functions which sustain any living being in existence) is conceptually prior to replication. There are many ways to show this, but take for instance this simple thought experiment.

Somehow or other, the first living cell came into existence. Call this Process G (for “genesis”). Now, imagine a population of cells which spontaneously undergo Process G, die off, and are replaced with another population of cells brought into existence by means of Process G. Such cells would be utterly incapable of replication, hence a fortiori incapable of evolution via natural selection. But they would clearly be alive. So, evolution is not necessary for life.

Also, consider the fact that replication is a highly regulated functional system. It is empirically well-established that replication in living systems could not exist without metabolism, whereas we just showed that metabolism could perfectly well exist without replication (and indeed must have done so at the origin of life).

All of this, if correct, would mean that we ought first to endeavor to understand life itself in a far more fundamental way than we presently do, before we can hope to achieve a deep understanding of the evolutionary process. Call this idea the “New Organicism.”

Would you care to comment on the New Organicism?

David Sloan Wilson

It’s true that there must be something that qualifies as an organism with a metabolism before evolution has something to act upon, but evolution enters the picture so early that I don’t see much point in separating them. Moreover, replication is not necessarily a big deal. It can involve the proto-organism fragmenting into parts, for example. An important point is that there can be evolution without replicators. Whole systems can replicate (the concept of a hypercycle, as I understand it) without gene-like entities within the system. Most of what you describe under the title of the “New Organicism” strikes me as similar to what I already associate with the literature on the origin of life. I’m not an expert in this area, however.


Some authors take the New Organicism still farther than West-Eberhard and the other adepts of evo-devo and epigenetics do.

James A. Shapiro

For example, James A. Shapiro [34] has recently claimed that the genome is most likely a “read-write” mechanism, and not a “read-only” mechanism as usually supposed, thus upending Francis Crick’s “central dogma of biology” (the idea that the causal arrow goes uniquely from nucleic acids to proteins, and never the reverse). The proposal, as we understand it, is that bacteria (at least) are capable of modifying their own genomes in an adaptive manner, in response to their changing physiological requirements.

If well confirmed, this phenomenon (which Shapiro also refers to as “natural genetic engineering”) would not only give further support to objective teleology and the New Organicism, it would basically vindicate Lamarck over Darwin.

Are you familiar with Shapiro’s work? What do you say in response to his claims?

David Sloan Wilson

I am not intimately familiar with Shapiro’s work but what he describes sounds plausible to me. This would be an example of a relatively blind process of evolution resulting in a system that possesses its own capacity for adaptive change. We already have a good intuition about this at the behavioral level with myriad forms of phenotypic plasticity. In theory, there is no reason why comparable mechanisms can’t exist for developmental programs, replication machinery, anticipatory mutations, and so on.

On Lamarck vs. Darwin, what most people think they know is based on the same kind of simplified and patriotic history that I described for the group selection controversy. The real history is admirably summarized by Jablonka and Lamb in Evolution in Four Dimensions.[23] To begin, Darwin was Lamarckian! More important, if anything requires loosening, it is the so-called central dogma of biology and the prohibition against thinking about Lamarckian forms of inheritance.


In closing, we would like you to tell us—in bulleted list format, if you like—what you consider to be the five strongest arguments in support of the consensus view within evolutionary biology that “neo-Darwinian is enough,” as well as the five weakest arguments that critics of neo-Darwinism commonly advance.

David Sloan Wilson

In a less oppositional spirit, I will list 10 points that stake out the productive middle ground.

  1. We should never forget the power of evolutionary theory as originally formulated by Darwin and Wallace. Organisms vary. Their differences often make a difference in terms of survival and reproduction. Offspring tend to resemble their parents. Given these three conditions, the properties of organisms change over time. They become well adapted to their environments. This will always be the centerpiece of evolutionary theory, no matter how many syntheses come and go.

  2. Tinbergen’s four questions concerning function, mechanism, development, and phylogeny are an admirable description of a fully-rounded evolutionary perspective. The most productive research programs will address all four questions in conjunction with each other.
  3. Darwin based his theory on the concept of heredity and knew nothing about genes. Yet, evolutionary science became highly gene-centric during the twentieth century, as if the only way for offspring to resemble their parents is by sharing genes. Evolutionary science needs to return to its roots by including all mechanisms of inheritance.

  4. Illustration
    George C. Williams
    The study of evolution in relation to human affairs lags behind the study of evolution in the biological sciences by almost a century. Basic literacy in evolutionary theory needs to be taught in the human-related disciplines, in addition to developments that are new for biologists. Also, biologists must become literate about the human-related disciplines to begin studying human cultural evolution.
  5. George C. Williams [35] was right to insist that adaptation at any level of a multi-tier hierarchy requires a process of selection at that level and tends to be undermined by selection at lower levels. Williams and others were wrong to claim that higher-level selection is invariably weaker than lower-level selection. Instead, the balance between levels of selection must be evaluated on a case-by-case basis. This should be part of basic training in evolutionary science.
  6. Multilevel selection is especially important for the study of evolution in relation to human affairs because we are such a group-selected species. Moreover, the solution to nearly every important problem in modern life requires expanding the scale of functional organization by managing the process of cultural multilevel selection.

  7. Behaviors that qualify as intentional can evolve by a process of blind variation and selective retention that does not qualify as intentional. The intentional behaviors can then double back to influence evolutionary processes. This basic insight has many applications, including human cultural evolution and phenotypic plasticity in all organisms. Developmental programs, mechanisms of genetic replication, and mechanisms of genetic change can be “directed” in the same way as more familiar forms of phenotypic plasticity.
  8. Organisms are complex systems inhabiting environments that are also complex systems. Hence, evolutionary theory needs to be studied in conjunction with complex systems theory. This is challenging, however, because the study of complexity is inherently complex. Merely invoking complexity is not helpful because it leads to combinatorial paralysis. Complex systems by themselves do not result in functional design. A process of selection is required, no less than for simple systems.

  9. Scientific inquiry is itself a highly managed process of cultural evolution that works well under some circumstances and breaks down under other circumstances. The more we explicitly regard science as a cultural adaptation, the better we can make it work.
  10. The concept of Equivalence, which emerged largely from the group selection controversy, is relevant to other controversies, as well. Sometimes theories are different because they invoke different causal processes, such that some can be right and others wrong. Alternatively, theories can be different by viewing the same causal processes from different perspectives, deserving to coexist to the extent that they provide useful insights. Arguing equivalent theories against each other is a waste of time. Instead, it is necessary to become literate about each theory and adept at translating from one to the other. Decades of futile debate can be saved by becoming mindful about Equivalence.


Finally, could you tell us about your plans for the immediate future? Do you have a new book in the works? Any new projects to advance the cause of evolution?

David Sloan Wilson

Very few people realize how much evolutionary science is needed to solve the problems of modern life. That is my current passion, which I pursue largely through the auspices of the Evolution Institute. It is also the subject of my next book, titled The New Social Darwinism: Completing the Darwinian Revolution and Improving the Quality of Life. It will be completed in a couple of months and published by Vintage/Anchor books in 2017.


Thank you very much for taking the time to participate in this Interview, and in the upcoming Focused Civil Dialogue with Professor Noble. We appreciate it very much indeed, and we are sure that our readers will profit greatly from your insights.

David Sloan Wilson

Thank you! Your questions have been better-informed than almost any other interview I have taken part in. Congratulations for your own literacy and for increasing the literacy of others.

Join the Dialogue!

Share your thoughts in the comments section below …while the comments are still open!


1.  T. Dobzhansky, “Nothing in Biology Makes Sense Except in the Light of Evolution,” American Biology Teacher, 1973, 35(3): 125–129.

2.  E.O. Wilson, Sociobiology: The New Synthesis. Cambridge, MA: Harvard University Press/Belknap Press, 1975.

3.  E.g., D.S. Wilson, “Evolution for Everyone: How to increase acceptance of, interest in, and knowledge about evolution,” Public Library of Science (PLoS) Biology, 2005, 3: 1001–-1008; and D.S. Wilson, G. Geher, J. Waldo, and R. Chang, “The EvoS Consortium: Catalyzing Evolutionary Training in Higher Education (Introductory article to a special issue devoted to EvoS,” Evolution: Education and Outreach, 2011, 4(1): 8–10.

4.  S. Wright, “Genic and Organismic Evolution,” Evolution, 1980, 34: 825–843; p. 841.

5.  D.S. Wilson and E.O. Wilson, “Rethinking the theoretical foundation of sociobiology,” Quarterly Review of Biology, 2007, 82: 327–348.

6.  E. Sober and D.S. Wilson, Unto Others: The Evolution and Psychology of Unselfish Behavior. Cambridge, MA: Harvard University Press, 1998; M. Borrello, Evolutionary Restraints: The Contentious History of Group Selection. Chicago: University of Chicago Press, 2010; and O.S. Harmon, The Price of Altruism. New York: Norton, 2010.

7.  V.C. Wynne-Edwards, Evolution through Group Selection. Oxford: Blackwell Scientific, 1986.

8.  W.D. Hamilton, Narrow Roads of Gene Land, Vol. 1: Evolution of Social Behaviour. Oxford: Oxford University Press, 1996. [Contains the seminal papers on kin-selection theory from the early 1960s—Eds.]

9.  R.L. Trivers, Social Evolution. Menlo Park, CA: Benjamin/Cummings, 1985.

10. J. Birch and S. Okasha, “Kin Selection and Its Critics,” BioScience, 2014, 65(1): 22–32; p. 28.

11. S.J. Gould, “The Hardening of the Modern Synthesis,” in M. Grene, ed., Dimensions of Darwinism: Themes and Counterthemes in Twentieth-Century Evolutionary Theory. Cambridge: Cambridge University Press, 1983; pp. 71–93.

12. Personal letter from Robert Rosen to James Barham, 1996.

13. [This term was popularized by a meeting of 16 of the world’s top theoretical biologists at the Konrad Lorenz Institute in the Vienna suburb of Altenberg in July of 2008; many of the papers presented at this conference were later published as M. Pigliucci and G.B. Müller, eds., Evolution: The Extended Synthesis. Cambridge, MA: MIT Press, 2010; see also the popular account of the meeting in S. Mazur, The Altenberg 16: An Exposé of the Evolution Industry. Berkeley, CA: North Atlantic Books/Wellington, New Zealand: Scoop Media, 2010.—Eds.]

14. See, for example, E.A. Lloyd, “Units and Levels of Selection,” in D.L. Hull and M. Ruse, eds., The Cambridge Companion to the Philosophy of Biology. Cambridge: Cambridge University Press, 2007; pp. 44–65.

15. O.T. Eldakar, M.J. Dlugos, G.P. Holt, D.S. Wilson, and J.W. Pepper, “Population structure influences sexual conflict in wild populations of water striders,” Science, 2009, 326(null): 816; and O.T. Eldakar, D.S. Wilson, M.J. Dlugos, and J.W. Pepper, “The role of multilevel selection in the evolution of sexual conflict in the water strider aquarius remigis,” Evolution, 2010, 64(11): 3183–-9.

16. J. Maynard Smith and E. Szathmary, The Major Transitions in Evolution. New York: W.H. Freeman, 1995; T.W. Deacon, The Symbolic Species. New York: Norton, 1998; C. Boehm, Moral Origins: The Evolution of Virtue, Altruism, and Shame. New York: Basic Books, 2011; E.O. Wilson, The Social Conquest of Earth. New York: Norton, 2012; and P. Turchin, Ultrasociety: How 10,000 years of war made humans the greatest cooperators on Earth. Storrs, CT: Baresta Books, 2015.

17. É. Durkheim, The Elementary Forms of Religious Life. New York: Free Press, 1995; p. 44. [First published in French in 1912—Eds.]

18. Ibid; p. 229.

19. D.S. Wilson, “Multilevel Selection and Major Transitions,” in M. Pigliucci and G.B. Müller, eds., Evolution: The Extended Synthesis. Cambridge, MA: MIT Press, 2010; pp. 81–93. [See, also, Note 13, above—Eds.]

20. Third Way of Evolution website.

21. M.J. West-Eberhard, Developmental Plasticity and Evolution. Oxford: Oxford University Press, 2003.

22. E. Mayr, “The Multiple Meanings of Teleological,” in idem, Toward a New Philosophy of Biology. Cambridge, MA: Harvard University Press/Belknap Press, 1988, pp. 38–66.

23. E. Jablonka and M. Lamb, Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life. Cambridge, MA: MIT Press, 2006.

24. S.A. Kauffman, The Origins of Order: Self-Organization and Selection in Evolution. New York: Oxford University Press, 1993.

25. D.S. Wilson, “Two meanings of complex adaptive systems,” in D.S. Wilson and A. Kirman, eds., Complexity and Evolution: Toward a New Synthesis for Economics. Cambridge, MA: MIT Press, 2016 (in press).

26. W.H. Calvin, “The brain as a Darwin machine,” Nature, 1987, 330: 33–34.

27. H. Plotkin, Darwin Machines and the Nature of Knowledge. Cambridge, MA: Harvard University Press, 1994.

28. G.P. Wagner and L. Altenberg, “Perspective: Complex Adaptations and the Evolution of Evolvability,” Evolution, 1996, 50: 967–976.

29. K.N. Laland, T. Uller, M.W. Feldman, K. Sterelny, G.B. Müller, A. Moczek, E. Jablonka, and J. Odling-Smee, “The extended evolutionary synthesis: its structure, assumptions, and predictions,” Proceedings of the Royal Society of London, B, 2015, 282: 20151019.

30. D.S. Wilson, “Intentional cultural change,” Current Opinion in Psychology, 2016, 8: 190–193.

31. J.M. Baldwin, “Development and Evolution,” Philosophical Review, 1903, 12(4): 442–451.

32. D.S. Wilson and J.M. Gowdy, “Human ultrasociality and the invisible hand: foundational developments in evolutionary science alter a foundational concept in economics,” Journal of Bioeconomics, 2015, 17: 37–52.

33. N. Tinbergen, “On aims and methods of ethology,” Zeitschrift für Tierpsychologie, 1963, 20: 410–433.

34. See, e.g., J.A. Shapiro, “The Basic Concept of the Read–Write Genome: Mini-Review on Cell-Mediated DNA Modification,” BioSystems, 2016, 140: 35–37; idem, “Physiology of the Read–Write Genome,” Journal of Physiology, 2014, 592: 2319–2341; and idem, “How Life Changes Itself: The Read–Write (RW) Genome,” Physics of Life Reviews, 2013, 10: 287–323.

35. G.C. Williams, Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Princeton, NJ: Princeton University Press, 1966.

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