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Susan Tran
Professor Haas
Writing 39C 5pm
24 April 2016
Dog Domestication: On the Origin of Dogs
To begin with the study of the domestication of dogs, we must first acknowledge Charles
Darwin for his work in evolutionary theory. Without Darwin, studies of animals may have never
been as increasingly advance as it is today. Darwin innovated evolution leading us to be able to
hypothesize that dogs are evolving species from wolves. 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). To this day, scientists don't have the
exact answers to how and when dogs became domesticated. There are many hypothesis of the
origin of the domestication of dogs and in this review I will be going over a few different
hypotheses. I will begin the review with Franois Vallas "Evidence for domestication of the dog
12,000 years ago in the Natufian of Israel." I will follow it with Wayne Roberts genetics
discovery of geographic origins of dogs. Then I will continue with Yan Lis hypothesis of
enhanced excitatory synaptic plasticity to start off and end with Belyaevs behavioral studies on
foxes.
Franois Valla, under the department of zoology 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, giving Francois a solid evidence that
dogs were domesticated about 12,000 years ago. In order to prove his argument, Francois looked

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at the puppys remaining jaw and compared

its jaw to other wolves and dogs in
Nutafian, Israeli. In his investigation,
Francois found that the puppy was either a
dog or a wolf, not a jackal. While
comparing the jaws, Franois 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. Tooth
Figure 1A human skeleton with a puppy.

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, Francois found that recent wolves in Israeli also had overlap
making it a less effective way of deciphering wolves form dogs. Which lead Francois to
conclude that wolves, by the early Natufian, been subject to some degree of selective breeding
by man (610). Francois research helped him prove his hypothesis of the domestication of dogs
12,000 years ago but other researchers have found evidence elsewhere.
Robert Wayne is an Evolutionary Biology professor at UCLA and Bridgett vonHoldt is
an assistant professor of Ecology and Evolutionary Biology at Princeton University. In their
article, Evolutionary Genomics of Dog Domestication, Wayne and vonHoldt explain the
origins of domestication of dogs through the understanding of the genetic changes during

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domestication. Wayne and vonHoldt argue that there are two primary modes of evolution within
dogs; first being mutations within dogs that cross breed and second selective breeding. Wayne
suggests 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). Wayne
also mentions 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 Francois, Wayne has a more diverse geographic research, Wayne researched a
wider range of dogs and looks more closely at DNA. Wayne found that the MtDNA sequence
data suggest that East Asia was a center for dog domestication (9). Wayne concludes that
admixture mapping can be a method to solving the origin of dog 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). The 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 willingness to be
tamed. 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

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memory through influencing short- and/or long-term potentiation (3119). Yan Li suggests 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 leads
to Yan Lis argument, strengthen in learning ability should help the skill of reading human
communicative behaviors (31120). Yan li concludes with the hypothesis that changes in
synaptic plasticity will help behavioral shirt and that prolonged human interaction can help the
human-canine relationship.
Dmitry K Belyaev was a Russian geneticist in the 1950s, famously known for his studies
on the domestication of silver foxes, closely related to dogs. 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. Such foxes are quite tame, not as a result of
training or taming, but due to prolonged selection for a tame genotype (Belyaev 303). 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.

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In conclusion, the question of when, where, and how dogs were domesticated dont have
a definite answer. Francois, hypothesized that dogs were domesticated about twelve thousand
years ago in Natufian, Israeli. While Wayne hypothesized that East Asia was the center for dog
domestication. Both authors had their own research that led them to their conclusion but it
doesnt come to the same conclusion, which means that anything is possible for dog
domestication. As for Yan Li and his team, they hypothesized that the domestication of dogs was
able to happen because of wolves willing to be tamed. 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 starts with the characteristic of being tamed. Belyaev dives deeper through
his research on foxes. Both Yan and Belyaev argues that the willingness to be tamed helps
advance the domestication of dogs. There is still a large gap for the origin of the domesticated
dog but with advancing technology and the love for dogs, the definite answer can be found.

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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.