Domestication Genes
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National Geographic has an article "Taming the Wild" by Evan Ratliff on the now famous under-the-radar experiment by a renegade Soviet geneticist, who had been previously sent to the Gulag by the Lysenkoists, to breed a domesticated silver fox that would be a amiable as a dog

In fact, says Anna Kukekova, a Cornell researcher who studies the foxes, "they remind me a lot of golden retrievers, who are basically not aware that there are good people, bad people, people that they have met before, and those they haven't." These foxes treat any human as a potential companion, a behavior that is the product of arguably the most extraordinary breeding experiment ever conducted. ...

One number I've never seen in accounts of this experiment is what percentage of these domesticated silver foxes breed true. Do 99% of new kits grow up to act like Labradors or do a sizable percentage have to be shipped off to a fur farm?

Miraculously, Belyaev had compressed thousands of years of domestication into a few years. But he wasn't just looking to prove he could create friendly foxes. He had a hunch that he could use them to unlock domestication's molecular mysteries. Domesticated animals are known to share a common set of characteristics, a fact documented by Darwin in The Variation of Animals and Plants Under Domestication. They tend to be smaller, with floppier ears and curlier tails than their untamed progenitors. Such traits tend to make animals appear appealingly juvenile to humans. Their coats are sometimes spotted-piebald, in scientific terminology-while their wild ancestors' coats are solid. These and other traits, sometimes referred to as the domestication phenotype, exist in varying degrees across a remarkably wide range of species, from dogs, pigs, and cows to some nonmammalians like chickens, and even a few fish.
Belyaev suspected that as the foxes became domesticated, they too might begin to show aspects of a domestication phenotype. He was right again: Selecting which foxes to breed based solely on how well they got along with humans seemed to alter their physical appearance along with their dispositions. After only nine generations, the researchers recorded fox kits born with floppier ears. Piebald patterns appeared on their coats. By this time the foxes were already whining and wagging their tails in response to a human presence, behaviors never seen in wild foxes. ... The Soviet biology establishment of the mid-20th century, led under Joseph Stalin by the infamous agronomist Trofim Lysenko, outlawed research into Mendelian genetics. But Dmitry Belyaev and his older brother Nikolay, both biologists, were intrigued by the possibilities of the science. "It was his brother's influence that caused him to have this special interest in genetics," Trut says of her mentor. "But these were the times when genetics was considered fake science." When the brothers flouted the prohibition and continued to conduct Mendelian-based studies, Belyaev lost his job as director of the Department of Fur Breeding. Nikolay's fate was more tragic: He was exiled to a labor camp, where he eventually died. ...
Not all domestication researchers believe that Belyaev's silver foxes will unlock the secrets of domestication. Uppsala University's Leif Andersson, who studies the genetics of farm animals-and who lauds Belyaev and his fellow researchers' contribution to the field-believes that the relationship between tameness and the domestication phenotype may prove to be less direct than the fox study implies. "You select on one trait and you see changes in other traits," Andersson says, but "there has never been proven a causal relationship."

To understand how Andersson's view differs from that of the researchers in Novosibirsk, it's helpful to try and imagine how the two theories might have played out historically. Both would agree that the animals most likely to be domesticated were those predisposed to human contact. Some mutation, or collection of mutations, in their DNA caused them to be less afraid of humans, and thus willing to live closer to them. Perhaps they fed off human refuse or benefited from inadvertent shelter from predators. At some point humans saw some benefit in return from these animal neighbors and began helping that process along, actively selecting for the most amenable ones and breeding them. "At the beginning of the domestication process, only natural selection was at work," as Trut puts it. "Down the road, this natural selection was replaced with artificial selection."

Where Andersson differs is in what happened next. If Belyaev and Trut are correct, the self-selection and then human selection of less fearful animals carried with it other components of the domestication phenotype, such as curly tails and smaller bodies. In Andersson's view, that theory understates the role humans played in selecting those other traits. Sure, curiosity and lack of fear may have started the process, but once animals were under human control, they were also protected from wild predators. Random mutations for physical traits that might quickly have been weeded out in the wild, like white spots on a dark coat, were allowed to persist. Then they flourished, in part because, well, people liked them. "It wasn't that the animals behaved differently," as Andersson says, "it's just that they were cute." ...

These perspectives might also apply to the evolution of phenotypical racial differences. Some differences in looks might have just been unselected for side effects of traits that were selected for by the environment. Or, as Darwin suggested, sexual selection or, among children, what Judith Rich Harris calls selection for cuteness might have played major roles.

But delving into the DNA of our closest companions can deliver some tantalizing insights. In 2009 UCLA biologist Robert Wayne led a study comparing the wolf and dog genomes. The finding that made headlines was that dogs originated from gray wolves not in East Asia, as other researchers had argued, but in the Middle East. Less noticed by the press was a brief aside in which Wayne and his colleagues identified a particular short DNA sequence, located near a gene called WBSCR17, that was very different in the two species. That region of the genome, they suggested, could be a potential target for "genes that are important in the early domestication of dogs." In humans, the researchers went on to note, WBSCR17 is at least partly responsible for a rare genetic disorder called Williams-Beuren syndrome. Williams-Beuren is characterized by elfin features, a shortened nose bridge, and "exceptional gregariousness"-its sufferers are often overly friendly and trusting of strangers. ....

"They didn't select for a smarter fox but for a nice fox," says Hare. "But they ended up getting a smart fox." This research also has implications for the origins of human social behavior. "Are we domesticated in the sense of dogs? No. But I am comfortable saying that the first thing that has to happen to get a human from an apelike ancestor is a substantial increase in tolerance toward one another. There had to be a change in our social system."

I'm not sure that the friendliest dogs are the smartest dogs. If Golden Retrievers don't distinguish between humans in terms of their intentions, which keeps them from biting your kid's friends but also makes them lousy guard dogs, that doesn't seem too smart.

There is also much else in the article, such as on nature-nurture adoption experiments with silver foxes. The keepers have also been breeding an Evil Twin breed of extremely nasty foxes. What happens when Nasty Fox is raised by a Nice Fox and vice-versa?

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