Muir’s experiments reveal a tremendous naiveté in the idea that creating a good society is merely a matter of selecting the “best” individuals. A good society requires members working together to create what cannot be produced alone, or at least to refrain from exploiting each other. Human societies approach this ideal to varying degrees, but there is always an element of unfairness that results in some profiting at the expense of others. If these individuals are allowed to breed, and if their profiteering ways are heritable, then selecting the “best” individuals will cause a cooperative society to collapse. It’s a good thing that the early eugenicists did not have their way!
Muir’s experiments also challenge what it means for a trait to be regarded as an individual trait. If by “individual trait” we mean a trait that can be measured in an individual, then egg productivity in hens qualifies. You just count the number of eggs that emerge from the hind end of a hen. If by “individual trait” we mean the process that resulted in the trait, then egg productivity in hens does not qualify. Instead, it is a social trait that depends not only on the properties of the individual hen but also on the properties of the hen’s social environment.
Ever since Muir’s experiments were published, I have been using them to illustrate the concept of multilevel selection and as a parable for thinking about human social evolution. However, their implications for animal breeding practices are important in their own right. Very few domestic animals are housed as individuals. This means that selection for bullying traits might be taking place even when it isn’t intended, resulting in decreased productivity from the human perspective and increased suffering from the animal perspective. Parenthetically, plant breeders face a similar problem. A corn plant that produces big ears by suppressing the productivity of its neighbors is little different than a hen that produces many eggs by bullying her cage mates.
My interview with Bill is centered on animal husbandry practices, but we also touch upon the parallels for human genetic and cultural evolution.
DSW: Welcome, Bill, to This View of Life.
WMM: Thank you for this opportunity to comment on those experiments. I have been working on the concept of individual vs. group productivity since I was a graduate student and found parallels in insects, plants, and many domesticated animal species. However, the idea of using those concepts on humans never crossed my mind, as with Darwin, I consider the concept of eugenicists against our moral and ethical principles.
DSW: I don’t propose that we breed people like farm animals in any sense! Let’s begin with that great experiment of yours. In my introduction, I observed how surprising it is against the background of the early proponents of human eugenics. How surprising is it against the background of Animal Science and what gave you the idea to do it?
WMM: It was not very surprising to me at all. In fact, I expected the results. Earlier experiments with Tribolium castanium (flour beetle) during my PhD thesis at Purdue University showed that selecting for beetles that could survive on minimal amounts of food in a group environment did so at the expense of others. The selected population evolved to grow faster so they could pupate first and eat the slower growing pupae, i.e. they became cannibalistic and the winning strategy was to grow the fastest with the slower fueling the growth of the survivors.
After graduation and teaching for a few years at the University of Kentucky, my first job as an animal breeder was back at Purdue University as a poultry geneticist. The curious result with beetles motivated the experiments with poultry. The only result that was really surprising to me was the amazing rate of response of the group selected birds, which showed a realized response greater than the selection differential, i.e. the realized heritability was greater than 1 for the first few generations. All of these results were beyond “classical theory” taught at the time, but was found to be due to associative effects, i.e. the trait “group productivity” capitalized on social effects, of which there are many. Cumulatively those social effects were much more important than that of the individual. I have since demonstrated the same finding when selecting for growth rate in quail and swine.
DSW: Tell me more about your educational background, which appears to have been unorthodox for an animal breeder. You anticipated your results, but were they more surprising to your colleagues with a more standard animal breeding training? Were standard animal breeding practices unknowingly selecting for antisocial traits in chickens and other farm animals?
WMM: In many ways life is a random walk, so too was my education. During the great “experimentation” days of quantitative genetics in the 1960-70’s, everyone in academia was testing quantitative genetics theory, which means there was great doubt that the infinitesimal model was correct because of the many assumptions. The infinitesimal model assumes an infinite number of genes each with small effects such that the entire process can be modeled by statistics with means and variances, rather than genes with Mendelian segregation ratios. We knew the model was wrong in detail, but was it right in practicality. Of interest was long term selection and how many generations we could select animals before the beast ran out of genetic variation. The only way to answer that in real time was to use a model organism with a short generation interval. One such model organism was Tribolium castanium because it had a generation interval of a month and was easy to rear. My major professor, Dr. Earl Bell, was a pioneer in use of the beetle to study quantitative traits, such as pupae weight, egg number, and development rate. As a graduate student I was interested in how competition influenced the outcome of long term selection and set my experiments up to examine the results, i.e. when selecting for survival on limited amount of feed, did it matter how many individuals were in the group given that the total amount of food was adjusted to give a constant amount per individual. The results showed that classical theory based on non-interacting individuals was wrong. The number of individuals in the group did matter, profoundly.
However, it was hard to convince animal breeders that these “bugs” had anything to do with animal breeding. In fact the state of the art for animal breeding was called BLUP, or Best Linear Unbiased Prediction. Statistical theory showed that this method would give the optimal response to selection, above all others, because it combined information from all sources and weighted them optimally. However, in order to use the theory phenotypes on individuals were needed. So pig breeders could measure individual growth rate and make optimal rate of progress, or so it seemed. In contrast chicken breeders in the layer industry (eggs) could not measure performance of individuals in group cages because one could not tell from which bird the egg came from. To address this problem, they redesigned the testing facility to put birds in individual cages. Now they had individual performance data, but unfortunately they did not make much progress .
This was about the time I started my research at Purdue. I saw that they set themselves up for what is called a genotype x environment interaction, i.e. the environment of selection (single bird cages) was different from the environment of production (colony cages). I reasoned that competition was the culprit; however, few believed me, and because birds could be beak trimmed to remove their “weapons” the problem could be largely addressed by management. However, I did not accept the management solution because it was costly and compromised the wellbeing of the bird; i.e. it hurt and continued to hurt. I suggested that group selection would solve the problem. Few believed me and several laughed openly because everyone knew that “group selection” has been debunked, i.e. early myth-busters claimed it could not happen, too many problems. Few bothered to examine what the problem with group selection theory was, only that the “experts” said it would not work. (Group selection is now referred to by the more general term “multi-level selection” of which group selection is a special case.)
However, I saw that all the theoretical issues with group selection could be overcome in a breeding situation, i.e. we could control the group size and composition (relatives, and no migrants) as well as who mated to whom. When I started the experiment, my major professor said he thought it was a major risk, “I was gambling my tenure on the experiment”, he was not sure it would work convincingly enough to publish and gain the recognition needed for tenure.
When I first presented the work at an international poultry wellbeing conference, the poultry company whose birds I used to demonstrate the results were outraged because I did the experiment was conducted in full light and did not beak trim the birds. I told them that this was the future, change was needed, and the theory they were using was wrong as it did not include indirect genetic effects. But I was a young pup and they did not believe me, nor did the company change, so they did not flourish. To me this is another example of group selection theory operating at the breeder level.
Another company who was doing sire family (half-sibs) selection in group housing did flourish and is now the largest poultry egg producer in the US, this is largely due to improvements in livability due to group (family) selection. The company was trying to mimic maize (corn) breeding that uses a combination of inbreeding (to create family lines) and ear to row planting whereby the row was the unit of selection, not the individual. Plant breeders were by coincidence using group selection, not because they were interested in reducing competition, but because there is too much variability among individual plants. They found that the average yield of a row was much more stable and assumed this result was because of the law of averages, which is true, but was also due to co-selecting groups that interacted well within families. Thus when the chicken breeders mimicked the maize breeding scenario, they also unknowingly adapted a selection method that guaranted competitive behavior will be reduced and animal welfare improved.
Ironically the company was on the cusp of changing their breeding program to individual housing because they wanted to use BLUP. I warned them not to do that and explained that what they were doing was group selection on a grand scale and this was the reason for their success. I think they heeded my warning as they flourished and now advertise that they are using and have been using group selection to produce their birds.
There is now another major layer breeding company trying to introduce multi-level selection into their breeding strategy, but by using multilevel BLUP, which I expanded to include indirect genetic effects. Unfortunately for multilevel BLUP to work, the genetic parameters must be measured very accurately. I showed using another model organism (Japanese quail) that family selection produced better responses than multilevel BLUP selection for those reasons.
DSW: Fascinating! Your passing reference to “group selection theory operating at the breeder level” hints that there is a parallel story to be told about human cultural evolution, but we’ll save that for the end of this interview. Returning to animal breeding practices, when viewed from the perspective of Multilevel Selection (MLS) Theory, the need for between-group selection and to avoid the disruptive effects of within-group selection should apply to any domestic animal that is raised in groups. Is this what is observed?
WMM: This depends on the relationship among individuals within the group. Obviously if all individuals are clones, which can be produced by inbreeding, then individual selection is the same as group selection. Which is to say that in family groups, the individual carries many of the same genes that produce competitiveness among others. So its own performance automatically includes its own indirect genetic effects. If the group is composed of unrelated individuals, then individual selection only increases its direct effects. Group selection based on random formation of groups does not work very well, but at least things will not get worse. Individual selection in random groups almost always makes for increased competition..
DSW: How widely have these results become known and have they resulted in changes in animal breeding practices?
As discussed above, they have become widely known among poultry breeders and most are using some form of multi-level selection. The results were explored by some swine breeders, and used very successful by Newsham Hybrids. They found the same results I did, i.e. very rapid improvements in wellbeing. But unfortunately the company was sold and the new owners got rid of the group selected lines because they called them “a bunch of pacifists who got the crap beat out of them when mixed with non-group selected pigs”, i.e. they could not stand up to others. Since mixing breeds is common they decided it was better to have aggressive pigs. This thinking is now starting to change, however they still have not adopted multi-level selection.
The reason for not adopting was due to a series of papers examining competitive effects in swine that were misinterpreted. They estimated the indirect genetic effects and found those effects to be a small part of the total genetic variance and concluded they could be ignored. The fallacy in this thinking is that each pig affects all other pigs in the group and as the group size increases, so does the impact of indirect genetic effects. In groups of 16, an indirect genetic effect is 15 times more important than the direct effect. In essence, more gains can be made by reducing indirect genetic effects (competition) than by increasing direct performance, as I showed with poultry. In the next several years it is my goal to correct this misinformation and demonstrate the power of multi-level selection. The timing is about right as animal welfare concerns are becoming critical in Europe and the USA.
DSW: Were your results more surprising for your colleagues with a more standard training in animal breeding?
WMM: Yes, absolutely, most could not believe the rapid responses observed in the first few generations and concluded there must be a major gene. They could not conceive that the infinitesimal model could produce the same thing. Even recently I presented the work at a prestigious European university, and when I remarked the heritability of the trait was greater than 1 in the initial generations, the immediate response was that I clearly did not know what I was talking about because, by definition, the heritability of a trait cannot exceed 1. Again, they were caught in old school thinking. With MLS in the first 2 generations of selection there was more response to selection than the SD. This is exactly what is meant by the whole is greater than the sum of the parts. In totality (as a group), we got out more than we put in (as individuals). This is the magic of multi-level selection, changing the unit of selection gives more than it takes. So reducing indirect genetic effects (competition) can be much more impactful than increasing direct (performance of the individual). Old school thinking is based on performance of the individual (direct effects).
DSW: I’m told that raising hens in cages is outlawed in Europe and that there is also a market niche in the US for raising hens under more “humane” conditions, which usually means larger enclosures. However, larger enclosures do not prevent aggressive hens from dominating other hens, denying them access to food, and so on. Thus, the selective environment under so-called humane housing conditions can result in some of the same pathologies as for cage rearing. Is my reasoning correct, and if so, what can be done to provide humane selective environments?
WMM: Yes that is correct, Europe is ahead of the USA in mandating animal welfare laws, the USA is close behind, California has banned battery cages too. Unfortunately, changing housing with the intent of improving welfare of the bird has proven disastrous. The one commodity that all will fight for in addition to food is territory. In battery cages all share the same territory, but in floor pens, with 10 or 100 times more space per bird, there is a blood bath, mortality increases dramatically. What was missing was places for birds to get away from others. Increasing the space only allowed the birds to truly express their competitiveness. Again, there is a management and genetic solution, but this time the management solution is in the interest of animal wellbeing, but at great cost in terms of housing. The genetic solution is still group productivity, it addresses the core issue of animals getting along with each other in confined spaces, and sharing that space. This is the next step in domestication of animals reared for food.
DSW: This has been a fascinating look at the animal breeding industry! To close, I’d like to return to the human case. Many years ago, after I gave a lecture that featured your chicken experiments, a woman rushed up to me and said, “That first experiment describes my department! I have names for those three hens!” What she meant was that her department had adopted a policy that rewarded members for their individual accomplishments and not for their collaborative efforts. Genetic evolution had not taken place, of course, but something else had that led to the same outcome. That “something” was the selection and cultural transmission of practices in behaviorally flexible individuals. You speculated along the same lines when you described companies that went out of business because they didn’t adopt the right poultry breeding practices. Nobody suggests that we breed people like farm animals any more, but what if the same ideas apply (roughly) to human cultural evolution? Have you thought along those lines?
WMM: Yes, indeed, in humans, because of our ability to treat diseases, we have transformed Darwinian evolution, or survival of the fittest genetically, to social evolution, which is survival of the fittest group within a society. Nations evolve as a group, those nations that cooperate internally are among the greatest competitors internationally, such as the USA. Ironically those countries that were based on the grand ideas of community sharing, i.e. communism, ultimately failed or are failing, because of cheating, i.e. within group social selection, which is the bane of group selection. In that system, some individuals cheat, end up preying on that society, and become very rich at the expense of the commoner. In such a society the idea of shared community good soon breaks down. Similarly companies with the greatest internal dynamics, associated with cooperation and collaboration, are the survivors. Survival of the strongest individuals in a society (company or nation) leads to predatory behavior, loss of a community goal, and ultimately failure.
DSW: Thanks so much for this TVOL interview. It’s a great addition to our series of articles titled “Truth and Reconciliation for Social Darwinism”.
WMM: I am very happy to have this opportunity to share what I have learned about breeding in a social setting and those linkages to the human situation. We can learn much from animals. While we no longer evolve genetically, we have the intellect to evolve socially and should take the lessons learned from genetic evolution and apply them to society for social evolution to advance. Multi-level selection is powerful, but requires safeguards to prevent cheating and breakdowns of that path. Ironically, between group (company) selection based on capitalism is the only way to keep the system honest. In a capitalistic society, those groups (companies) where cooperation fails will soon be out of business. This process is social selection at the group (company) level. The trust me attitude of an autocratic socialistic society cannot provide adequate safeguards to allow social evolution to keep evolving. As with genetic selection, there can be no evolution without failure, i.e. survival of the fittest groups means that some groups must fail, and be allowed to fail, such that new groups can be formed to continue the process.
Articles in this series:
Truth and Reconciliation for Social Darwinism by David Sloan Wilson and Eric Michael Johnson
The Case for Rescuing Tainted Words by David Sloan Wilson
Social Darwinism: Myth and Reality by Paul Crook
Social Darwinism: A Case of Designed Ventriloquism by Adriana Novoa
When the Strong Outbreed the Weak: An Interview with William Muir by David Sloan Wilson
Was Hitler a Darwinian? No! No! No! by Robert J. Richards and David Sloan Wilson
Was Dewey a Darwinian? Yes! Yes! Yes! An interview with Trevor Pearce by David Sloan Wilson
Toward a New Social Darwinism by David Sloan Wilson and Eric Michael Johnson