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Susan Tran
Professor Haas
Writing 39C 5pm
08 May 2016
Dog Domestication: On the Origin of Dogs Literature Review
Charles Darwin is famously known for his theory of evolution and creating the term
natural selection. Natural selection is process of survival of the fittest, the species that can
adapt well to new environments will survive and successfully reproduce. Another term that
Darwin coined to oppose natural selection is artificial selection, also known as selective
breeding. Artificial selection is the breeding of animals based on desired traits. Leslie Irvine,
professor of sociology at the University of Colorado, in her book If You Tame Me, states that
humans were able to breed dogs however they wanted to in a short amount of time (12). For the
past century, humans have been breeding the domesticated dogs based on artificial selection and
for the past decades there have been evidence that these artificial breeding are taking a toll on
dogs. So when did the domestication of dog occur in order for humans to begin artificial
breeding in dogs? To this day, scientists don't have the exact answers to how or when dogs were
domesticated. This review will cover the hypotheses of several scientific researchers concerning
the origin of the domestication of dogs. I will begin the review with Franois Vallas "Evidence
for domestication of the dog 12,000 years ago in the Natufian of Israel," followed by Wayne
Robert and Bridgett vonHoldts genetics discovery of geographic origins of dogs. Last, I will
cover Yan Lis hypothesis of enhanced excitatory synaptic plasticity to start off and end with
Belyaevs behavioral studies on foxes.

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One of the earlier research on canine

domestication was by Franois Valla in
1978. Franois Valla, a zoologist at the
Hebrew University states, A domestic
animal is one whose breeding is largely
controlled by man. Evolution of a
domesticated species, therefore, results
mainly from artificial selection (608). In
northern Israeli, Mallaha, researchers found
a puppy buried completely with a human
(See Figure 1), giving Valla a solid evidence
that dogs were domesticated about 12,000
Figure 1 Human skeleton with a puppy remains.

years ago. In order to prove his argument,

Valla looked at the puppys remaining jaw and compared its jaw to other wolves and dogs in
Natufian, Israeli. In his investigation, Valla found that the puppy was either a dog or a wolf, not
a jackal. While comparing the jaws, Valla comes to the conclusion that three size classes of the
wolf-dog line exist in Israel Upper Pleistocene wolves, recent wolves and domestic dogs. He
found that Upper Pleistocene wolves are larger in size compared to the domestic dogs as shown
in. Tooth crowding and smaller size have been used as criteria for the identification of dogs.
Dogs jaws are slightly shorter than their wolf progenitors but through more detailed research,
Valla found that recent wolves in Israeli also had overlap making it a less effective way of
deciphering wolves form dogs. Which lead Valla to conclude that wolves, by the early Natufian,
been subject to some degree of selective breeding by man (610). Valla research helped him

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prove his hypothesis of the domestication of dogs 12,000 years ago but other researchers have
found evidence elsewhere. Vallas study between a dogs jaw and a wolfs jaw indicates that dogs
were already evolving to develop canine human relationship.
Although Valla concluded that dogs were domesticated 12,000 years ago, his research
may be flawed because of technological advancement that has happened within the last two
decades. In 2012, Robert Wayne, an evolutionary biologist at UCLA, and Bridgett vonHoldt,
ecologists and evolutionary biologist at Princeton, recorded their study of canine domestication
in Evolutionary Genomics of Dog Domestication, explaining the origins of domestication of
dogs through the understanding of the genetic changes during domestication, arguing that there
are two primary modes of evolution within dogs: first, due to crossbreeding and second selective
breeding. They suggest that small dogs were domesticated from Middle East because small dogs
have similar IGF1, a gene that determines body size sequence to wolves from the Middle East
(12). They also mention that IGF1 is a major contributor of the size diversity in dogs,
accounting for about fifty percent of the genetic variation in size (6). The differences in
domestic dogs are abundance and most likely reflect the unique history of selection in dogs
which involved the appearance of discrete mutations followed by fixation through intense
inbreeding. Although it may appear to be an abundance of diversity of domestic dogs but in
reality, has limited genetic basis (12). Unlike Valla, Wayne and vonHoldt have a more diverse
geographic research, they researched a wider range of dogs and looks more closely at DNA.
They found that the MtDNA sequence data suggest that East Asia was a center for dog
domestication (9). Wayne and vonHoldt concludes that admixture mapping can be a method to
solving the origin of dog domestication. Their conclusion demonstrates that there has been

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copious amount of selective breeding that has made it difficult to find the exact origin of canine
domestication.
After a few generations, the less fearful wolves would be on the path to becoming
domestic dogs, and the humans would probably have been on their way to becoming a species
that likes to live with dogs (Grandin). Yan, Li, et al, authors of Domestication of the Dog from
the Wolf Was Promoted by Enhanced Excitatory Synaptic Plasticity: A Hypothesis, researchers
at Yunnan Laboratory of Molecular Biology of Domestic animals and Kunming Institute of
Zoology, hypothesized that dogs evolved to be domesticated because of their unique social
cognitive capabilities, which we hypothesized to be a by-product of selection for tameness
(3115). The researchers tested dogs and wolves for stress response within their genes, seeing
changes in the synaptic plasticity. The synaptic plasticity is associated with learning and memory
abilities. Glutamate is the major excitatory neurotransmitter in the brain that regulates many
kinds of behaviors and emotions and plays a key role in cognitive ability, including learning and
memory through influencing short- and/or long-term potentiation. They suggest that glutamate
may have helped the domestication of wolves because changes in synaptic plasticity are thought
to be associated with changes in learning and memory abilities (3119). Which lead to their
argument, strengthen in learning ability should help the skill of reading human communicative
behaviors (31120). They conclude with the hypothesis that changes in synaptic plasticity will
help behavioral shift and that prolonged human interaction can help the human-canine
relationship. With prolonged human-canine relationship, dogs can naturally evolve and shift their
behavior to further deepen the relationship, similar to the relationship we have with dogs today.
Parallel to Lis hypotheses, Dmitry K Belyaev, a Russian geneticist in the 1950s,
famously known for his studies on the domestication of silver foxes, believed that foxes were

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tamed due to prolonged selection for a tame genotype (Belyaev 303). Belyaev studies
special bred foxes on farms and among the foxes, thirty percent show aggressive behavior
towards man, twenty percent showed fear, forty percent were aggressively fearful, and ten
percent were curious. During his study, Belyaev found that the type of behavior is formed during
their first two months, defensive behavior is hereditary, and there is phenotypic and genotypic
correlation to females time of onset of their reproductive activity. Overtime, because of selective
breeding for tameness, foxes naturally inherit the taming genotype but with the changes in
behavior it also was accompanied by changes in reproductive function. Thirty to forty percent of
females dont reproduce or eat their offspring. Concluding that the change in reproductive
system is associated with domestication. The new reproductive system has made changes to the
new generation of tamed foxes, such as molting, position of tail, brown spots, and drooping of
ears characteristics of dogs. Belyaev concludes that The process of domestication in all animals
species seems to have resulted in the same kinds of homologous variations as a result of selection
for the single important characteristic of tame behavior.
In conclusion, the origin of canine domestication cannot be traced back to the exact
moment because of selective breeding. Valla concluded that dogs were domesticated about
twelve thousand years ago in Natufian, Israeli based on jaw comparison between wolves and
dogs. Wayne and vonHoldt concluded that selective breeding has led to large numbers of genetic
differences in dogs, making it difficult to tack the origin of canine domestication. As for Yan Li
and his team, they hypothesized that the domestication of dogs was able to happen because of the
selection for tameness. Yan Li did research on the synaptic plasticity in wolves and dogs,
showing that the more they strengthen their learning and memory, they can learn that humans are
most of the time harmless. Similarly, Belyaev argues that the domestication of dogs started with

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the genotype selection for being tamed. Belyaev dives deeper through his research on foxes.
Both Yan and Belyaev argues that the selection for tameness helps advance the domestication of
dogs. In the past century, selective dog breeding has become a common occurrence within our
society. Selective dog breeding has been for appearance or purebreds because of the rise of dog
shows. Although it is a great success for humans to be able to artificially breed dogs, it has taken
a great toll on dogs because it can lead to genetic disorders and health issues.

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Work Cited
Belyaev, Dmitri K. "Destabilizing selection as a factor in domestication." Journal of Heredity
70.5 (1979): 301-308.
Davis, Simon JM, and Franois R. Valla. "Evidence for domestication of the dog 12,000 years
ago in the Natufian of Israel." (1978): 608-610.
Grandin, Temple, and Catherine Johnson. "Why Cats and Dogs Think Differently About You."
Animals Make Us Human: Creating the Best Life for Animals. Boston: Houghton Mifflin
Harcourt, 2009. N. pag. Print.
Irvine, Leslie. If You Tame Me: Understanding Our Connection
with Animals. Philadelphia: Temple UP, 2004. 12-32.
Wayne, Robert, and Bridgett vonHoldt. "Evolutionary Genomics of Dog Domestication."
Mammalian Genome 23.1/2 (2012): 3-18. Academic Search Complete. Web. 19 Apr.
2016.
Yan, Li, et al. "Domestication of the Dog from the Wolf Was Promoted by Enhanced Excitatory
Synaptic Plasticity: A Hypothesis. Genome Biology & Evolution 6.11 (2014): 31153121. Academic Search Complete. Web. 20 Apr. 2016.