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Genetics

Farm Fox Theory Bites the Dust

A renowned experiment regarding dog domestication takes a major blow.

In my last posting, I noted that many students of the dog seemed to be edging away from the standard model of dog domestication, also known as the dumpster diver hypothesis, or theory. Little did I know that the entire edifice was about to collapse of its own deceptions. Based on the long-running Siberian Farm Fox Experiment that was started by Dmitry Belyaev and has continued long after his death, that model claims to show definitively that by breeding for tameness alone, a group of wild wolves can within, say, 10 generations, produce a population of dogs possessed of the behavioral and phenotypic characteristics that separate them from their wild cousins.

They include, among other traits, an addition to tameness, sometimes described as a reduced fear response to humans, and hyper-sociability; piebald coats, curly tails, and shorter, broader muzzles with crowded teeth. They are said to be juvenilized versions of their ancestor species, eager to play and grovel for attention all day—thus this model is also sometimes referred to as domestication by neoteny.

Juvenile-appearing traits were first noted by Charles Darwin as being common markers of domestication but not named until botanists grouped them together as a syndrome and applied it to plants early in the 20th-century. The term and concept were applied to animals in the last quarter of the 20th-century, when the Farm Fox Experiment captured the imaginations of, and became the “truth” about, dog evolution—to be challenged at your peril.

There were people who sought to uphold scientific standards. At no small risk to their reputation, they brought problems with the dumpster-diver hypothesis to light. Marc Bekoff’s Psychology Today blog, Animal Emotions, on this subject provides a rich web of links. From my first encounter with the dumpster-diver hypothesis, I questioned the shifting terms for the trait being bred for, from tame to calm to bold to solicitous of attention to a reduced flight response. I also remain unconvinced that dogs are juvenilized wolves.

Just as troublesome was the insistence on leaving humans out of the domestication process on the grounds that they could not possibly have foreseen where engaging with wolves would lead. Finally, I said that argument by analogy was the weakest sort. My arguments and those of other critics fell on deaf ears. I discussed the situation in my last posting, giving credit where it is due.

What might prove the fatal blow against the standard model, and perhaps Domestication Syndrome itself, when applied to animals, came December 3, 2019, in an online article, “The History of Farm Foxes Undermines the Animal Domestication Syndrome,” by Kathryn A. Lord, Greger Larson, Raymond P. Coppinger [posthumously], and Elinor K. Karlsson, writing in the journal, Trends in Ecology and Evolution. They report several serious revisions to the canned history of the experiment and then use those to re-examine the farm fox experiment’s claims. With an abundance of caution, they write: “Our results suggest that both the conclusions of the Farm-Fox Experiment and the ubiquity of domestication syndrome have been overstated.” Later in the paper, they are more emphatic, “The Farm-Fox Experiment, however, does not validate the Domestication Syndrome.”

What made the Siberian Fox Experiment fall so low so fast? In brief, in 1959, Soviet Geneticist Dmitry Belyaev began an experiment for the Institute of Cytology and Genetics at the Russian Academy of Sciences, Novosibirsk, Siberia, designed to prove his theory that breeding for tameness alone would soon produce a population of tame foxes manifesting the traits Darwin had associated with domestication. After 10 generations of intensive breeding, Belyaev announced results, documented in photographs and film.

Journalists and scientists around the world took note, especially, it seems, from English speaking countries, and began making the pilgrimage to Siberia. Some scientists were convinced they would find the gene or genes responsible for domestication. Belief in direct correspondence between a single gene and specific behavior was common at the time—and still is in some quarters.

Mark Derr
Navajo dogs vie for a piece of cantaloupe.
Source: Mark Derr

Raymond Coppinger, an evolutionary biologist at Hampshire College with an interest in dogs, made Belyaev’s work the cornerstone of his theory of canine evolution. Essentially that holds that dogs self-domesticated at the garbage dumps of Mesolithic villages among people making the transition from hunting to agriculture and animal husbandry.

The problem is that dogs appeared when people were still hunters and gatherers—thousands of years before the advent of agriculture. Coppinger also believed that natural selection at dumpsites favored animals who would neither run from humans nor stand to fight them. They licked people and they groveled for attention. In short, they were neotenic versions of their wild progenitors.

Coppinger died before this paper was completed. Although recognizing him is considerate, there surely should be an editor’s note explaining why he would be a party to an article calling for reconsideration of a theory that he had promoted for nearly 40 years. Lord, the first author of the article, has said in an interview that Coppinger had begun working on material from the International Fox Museum and Hall of Fame on Prince Edward Island, Canada, showing doglike foxes being bred and raised from the late 19th-Century.

I was more surprised to see Greger Larson listed as a co-author. Co-leader of an international effort to solve the mystery of dog evolution, he has for years argued that self-domestication and neoteny provided the answer. He has routinely cited Belyaev’s experiment in glowing terms. (For example, listen to our 2013 interview by Ira Flatow on Science Friday.)

Now in calling for a review of the underpinnings of the Farm Fox Experiment and Domestication Syndrome he has challenged his own orthodoxy. But he continues to insist the foxes are of great worth to science for studying the genetics of behavior. Belyaev did on at least one occasion indicate that his foxes had been bred in captivity for a number of generations, but he neglected to say they were from Canadian stock that had been bred since the late 19th-Century for appearance and behavior by humans. They looked and acted like his tame foxes, who in fact were descended from them. From that unraveling come others showing that traits associated with domestication are neither uniform nor universal; rather, they arise at different times in the domesticated population, if they appear at all.

Elinor Karlsson, the senior author of this article and the leading geneticist tracing Belyaev’s foxes back to Canada—from what one sees in the few follow-up stories that have appeared in the popular press—seems a scientist willing to follow the data wherever it leads. I wonder how many journalists and scientists who have built careers on discredited claims for the Siberian foxes will take up the challenge to reconsider what they believe to be true to define common terms and concepts.

I wonder as well whether anyone will take on the task of writing the history of the Siberian Foxes. These discussions are for another time and place. There is a more immediate need to ask and whether Belyaev and his followers deliberately misled the international community, and why the scientists and science journalists so fervently embraced his work and explanations. I would hope that in working its way out of this mess, the community would become more open in its use of sources from other fields. That would involve judging each source on its merit, and in some cases giving an explanation for its use or exclusion.

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