Artificial selection & domestication

 People decide who breeds and who doesn´t  try to influence specific “traits” that they
want to change in future generations

Why separate “artificial” from “natural” selection?

 Like sexual selection, artificial selection is just a special case of natural selection 
expect here it´s people making decisions as to why breeds and who doesn’t
 Why should it matter?  it doesn´t really matter if you think about natural selection
from its 3 main principles
o Variations
o Heritability
o Reproduction
 But if you think about animals being “well adapted” or being the “fittest” then it makes a
difference because people can make some very weird decisions as to who breeds and who
doesn´t  and this has consequences
 Would this cow be successful in the wild if it
produces 10x more milk than its offspring need?
 Artifical selections is a useful concept to consider
what we see in our domestic aniamls  they are a
compromise between their evolutionary history
driven by natural selection, and recent chaanges
brought about by our selective breeding

Today´s dogs are descended from wolves. Now it´s your turn to start with wolves and breed
them until you have chihuahuas. How would you do this? What traits will you select for
breeding? What changes would you expect?
 Less fearful
 Less aggressive
 Smaller
 Changes in coat colour
 Changes in body shape
 Respond to commands
 bark

Pleiotropy
 Occurs when one gene influence two or more seemingly
unrelated phenotypic traits. Therefore, mutation in a pleiotropic
gene may have an effect on several traits simultaneously due to
the gene coding for a product used by many different cells
 Why would these wildly different traits be linked?  the clue
is that they aren´t really radical changes in the phenotype of the
 dog, but rather most these represent the “puppy” stage of development
 So by selecting for “tameness” a few genes that regulate development from being a
playful puppy to a wary adult can affect a whole bunch of things that also change as the
animal matures from youth to adulthood

 Artificial selection often focuses on extreme traits  for production or just for show
 Artificial selection can focus on simply finding differences and magnifying them
 But how do you magnify these effect from one generation to the next?

How to exaggerate traits

 You are an animal breeder who want to increase milk yield in a cow, skin folds in a dog
or weird feathers around the head of a pigeon. But how to do this?
 First you need to mate the “mutant” to another dog and
produce offspring  then what?
 To quickly create a new breed or exaggerate a trait you
need to inbreed
What is inbreeding?
 Inbreeding is the breeding of individuals that are closely related genetically
 Inbreeding will remove genetic variation between “alleles” for the trait you are interested
in because the small number of animals you breed from will not have much genetic
variation at that allele
 This will result in the trait you are interested in becoming a permanent and unvarying part
of the breed or population
 How inbreeding creates genetic
uniformity
 Consequences of inbreeding:
o Benefits  rapid creation
of a variety or breed, with
the defining trait of that
breed fixed into the
population
o Problems  genetic
fixation of defects 
expression of deleterious
recessive effects 
inbreeding depression 
smaller gene pool available
for future selection

Problems with inbreeding: Genetic fixation of defects

 Here we considered inbreeding as a useful way to make sure
all animals in the breed carried the gene for the trait we
wanted (green hair)
 But if the animal we breed from also has a genetic defect in
another gene, by inbreeding we might (by accident) create a
breed where all the animals also carry that genetic defect

Problems with inbreeding: expression of deleterious

recessive effects
 Genes come in pair and are expressed in different
ways. Often one form of the gene (allele) is
expressed in preference to another (dominant
allele > recessive allele)
 So, disease can be hidden in recessive alleles
unless you get 2 of them: then you are in trouble
 Inbreeding makes this more likely
Problems with inbreeding: inbreeding depression
 Inbreeding depression is the reduced survival and reproduction in a individual or
population as a result of inbreeding
 It´s cumulative effect of potentially many genetic problems created from problem genetic
fixation of defects & expression of deleterious recessive effects

Problems with inbreeding: smaller gene pool available for future selection
 Many alleles provide larger phenotypic
variation to select amongst for natural
selection or artificial selection
 Thus, species can adapt quicker to
environmental change, or breeders can create
new breeds or exaggerate traits more easily
 Fewer alleles provide little phenotypic variation to select amongst for natural selection or
artificial selection
 Thus, species will adapt slower to environmental change, or breeders can´t improve or
change traits in their breeds

But isn´t inbreeding “wrong”: should we really be doing it?

 It can definitely create problems, and in nature close relatives avoid mating with each
other, but it does occur in the wild when population are forced through “bottlenecks”
 A genetic “bottleneck” is when a population is squeezed so that very few individuals
breed for a time, then the population expands again

New varieties, breeds or species

 Natural selection:
o Selects for the best adapted individuals based on the current environment
o There is no goal, there is nothing that evolution is aiming for
o It´s a mistake to think evolution has been trying to make people
o A bacteria is just as “evolved” as we are. It´s simply adapted to a different set of
conditions
 Artificial selection:
o Exaggerate a current trait (e.g., milk production) or selectively breed for a
mutation (e.g., fancy pigeon breeds), based on what people want
o There is a goal, there is something we are aiming for
o We want a blue rose, or a fluffy chicken or a high producing milk cow

Domestication and domesticated species

 Domestication = is sustained multi-generational relationship in which one group of
organisms assumes a significant degree of influence over the reproduction and care of
another group to secure a more predictable supply of resources from that second group
 Domestication should not be confused with taming. Taming is the conditioned behavioral
modification of an animal to accept the presence of humans
 Domestication is the permanent genetic modification of a bred lineage that leads to an
inherited predisposition towards humans
 In other words, for “taming” you need to train each individual by teaching them to not be
afraid, but through domestication you “tame” a population by changing their genes to suit
your needs

Which process is important during domestication?

 Natural selection:
o The best suited phenotypes to the local environment are “selected”, in that those
animals with the right traits are more likely to survive and reproduce. These traits
are passed onto their offspring
 Sexual selection
o Occurs when females choose males based on some measure of quality. Results in
males having ornaments or weapons to demonstrate their quality to females
 Artificial selection
o Humans choose who breeds and who doesn´t. May involve separating different
breeding lines, or inbreeding to produce animals with the traits we want

How to domesticate a species

 When the first animals were beginning their domestication, what were people doing to
make this happen? To what extent was it natural selection, sexual selection or artificial
selection? And how did this change over time as animals become more integrated into
society?
 Stage 1: humans create a new ecological niche

 Stage 1: natural selection

o Humans create new environmental conditions that other species need to adapt to
o Thus, those individuals that can use these new resources will benefit
o Natural selection operates around the human settlement and selects those animals
that aren´t afraid of people and can live in this zone
 o Animals that are afraid will move out into the general population  creates some
genetic segregation
o Thus, natural selection initially operates on the fearfulness or animals in this area
o Over generations, the animals near human settlement will have a genetic
predisposition to be less fearful of humans

Parapatric speciation
Poor movement of individuals within a population created poor gene flow, leading to
divergence across the population range and eventually different species

Sympatric speciation
Is the evolution of a new species from a surviving ancestral species while both continue to
inhabit the same geographic region

So, we can think about the domestication of animals as a form of sympatric “speciation”
There is a focal point in the population where selection will be pushed in a different direction

Stage 2: humans start selective culling

 At some point people realize that females are useful for producing offspring, and their
hunting change from hunting everything, to selectively hunting young males for food
 This is both within the settlement area (i.e., selectively keeping females and culling young
males) and within the wild population (i.e., hunting males)
 Natural selection will keep operating as people and animals (especially the females
around their settlements) become more closely associated. Animals will be selected for
domestication traits (e.g., lack of fear of humans)
 But the culling of surplus males from the area means that there will be less males, less
male-male competition, and less sexual selection. Over generations the difference
between males and females will get less and less. Males get smaller, horns get smaller (or
disappear)

Stage 3: Humans start managing their animals and isolating them from
wild populations
 The next stage is that people start managing their animals, by controlling predation of
their livestock, by continuing selective culling of young males and increasingly using
females as the basis of their herds, and by restricting their chances of breeding with their
“wild” cousins, and providing supplementary food
 Gene flow between captive and wild
animals is reduced, accelerating their
adaptation to local “captive” conditions by
natural selection
 Thus, animals become much easier to
handle than their wild siblings
 Sexual selection is further reduced as
females have little choice in sexual partners. Thus, difference between males and females
is reduces
 By controlling predation and providing food the stabilizing effect of natural selection on
extreme forms is partly relaxed
 Thus, we start to see new color forms (e.g., white or spotted)

Genetic isolation of early domesticates and wild ancestors

 Gene flow from wild animals needs to be reduced or stopped for changes resulting from
domestication selection to become fixed in the captive population
 Historically this has occurred in 3 ways:
1. The captive animals were fenced and access to breeding with wild animals was
limited
2. Wild populations were hunted to extinction
3. Captive animals were exported to areas outside of their former range, meaning there
were no wild animals in the surrounding environment to breed with

What about the predators?

 For cats and dogs these stages were not characterized in the same way  Cats and dogs
were not “herded” and hunted like prey spices were
 Instead, domestication was primarily natural selection for tameness, as their wild
ancestors exploited the opportunities provided by the human settlements
 Dogs were genetically isolated from wolves at an earlier stage than cats were from their
wildcat ancestors  thus, dogs have changed more relative to their ancestors than cats
have
Stage 4: humans start selectively breeding animals for certain traits
 Horses for speed, strength, companionship and etc.
 Dogs for hunting, herding, livestock guardian, companionship and etc.
Animals undergoing domestication will evolve to local conditions:
landraces (lantraser)
 This will occur not only during the initial domestication process, but also as these animals
are dispersed and traded over wider area
 These sub-populations will often be genetically isolated from each other and so will
diverge through natural selection (adaptation to local conditions) and artificial selections
(as local people want different things from their animals  e.g., milk vs. meat, or hunting
dog vs. livestock protection)

Landraces
 A landrace is a domesticated, locally adapted, traditional
variety of animal that has developed over time, through
adaptation to its natural and cultural environment of
agriculture and pastoralism, and due to isolation from
other population of the species
 Its morphologically distinctive and identifiable (i.e., has
particular and recognizable characteristics or properties)
 Its genetically adapted to, and has a reputation for being
able to withstand, the conditions of the local
environment, including climate, disease, pests, and
cultural practices
 It’s not the product of formal breeding programs, and may lack systematic selection,
development, and improvement by breeders
 Its maintained and fostered less deliberately than a standardized breed, with its genetic
isolation principally a matter of geography acting upon whatever animals that happened
to be brought by humans to a given area
 It has a historical origin in a specific geographic area, will usually have its own local
name(s), and will often be classified according to intended purpose
 At the level of genetic testing, its heredity will show a degree of integrity, but still some
genetic diversity

Stage 5: humans intensify selective breeding, often for novel traits

 Here artificial selection completely takes over, as people intensify food production in
farm species, speed, or strength in horses, ands size, shape, and color in companion
species
 Breeding lines are established and kept separate, creating “purebreds” using inbreeding
and other techniques
 Phenotypic changes accelerate, with most exaggeration of traits occurring during this time
The history of domestication depends on the species and the goal: but has
similar selection processes
 Prey species being bred for meat will have a different domestication history to prey
species used for transport, and they will have a different history to predator species being
bred for companionship or work
 But with all domesticated mammals we need to interact with and handle them  thus,
there are many similarities that domestication will “breed” into them
 This often comes unintentionally, with natural selection acting in the new environment to
favor those individuals with low fear/stress levels in the presence of humans

Do these principles apply to non-mammal species?

 For example, insects, that we might want to domesticate as food livestock?
 We don’t need to cuddle them, but they will still benefit by channeling their resources
into growth and reproduction, rather than stress hormones and “fight or flight” responses
when people handle them
 Also, they will be kept in intensive rearing systems  here they will need to be very
social
 Thus, the same factors will apply as with our domestic livestock:
1. Natural selection for domestication traits
2. Reduction in sexual selection (if reproduction is controlled by humans)
3. Artificial selection for specific production behavioral traits

The domesticated phenotype

 Regardless of the species domesticated, the following changes in appearance are almost
universal
o Less frightened of people
o More social
o Smaller (changes in body shape)
o Shortened faces
o More vocal
o Juvenile behaviors (begging, licking)
o Reduced differences between males and females
o New fur colors and patterns

Paedomorphosis: retention of juvenile traits

 The big question is why do we get such pleiotropic effects
when we select for tameness in animals?
 One idea is that the gene(s) selected for when we select for
tameness are those that affect hormonal development as
animals mature from juveniles to adults
 Thus, selection acts to delay or stop this maturity  this has
many effects on the body
 So, by selecting mammals for tameness, we are selecting for delayed maturity  This
brings with it all of these physical and psychological juvenile characters into the adult
animal
 Just by changing one or a few genes for specific hormonal pathways

Reindeer: in an early stage of domestication

 Reindeer are semi-domesticated, what changes have occurred between the wild and
managed reindeer, or are they the same? What evidence is there that reindeer are in the
early stages of domestication?
 What should we expect if we compare a “wild” reindeer to a reindeer from a managed
herd?
 shorter flight distance (less fearful of humans)
 Changes in head or body shape/size
 more vocal
 higher diversity of fur colour
 reduced differences between males and females
o female reindeer also have antlers (unlike other deer), so there are probably natural
selection reasons for this. Also, reindeer herders do not control mating to any
large extent, meaning that sexual selection can still operate

Dogs, cats & horses: in a late stage of domestication

 Until very recently, dogs, cats and horses were bred for function (and sometimes just for
their form)
 In the past 50-100 years this has changed with breed
standards becoming the most important aspect of
defining the breed. Thus ‘kennel clubs’, ‘cat fanciers’
and horse racing promotes the exaggeration of certain
traits that are not always in the best interest of the
animals.
 Breed standards or the ‘bloodlines’ of pedigree animals
have resulted in some very poor outcomes for animal
welfare…
 In dogs we have created all sorts of medical issues with
our selection for certain traits

So which dog at the animal shelter is the healthiest?

 German shepherd = hip dysplasia, elbow dysplasia, lupus
(immune system) pancreatic insufficiency spondylosis
deformans, lifespan ca 7-10 years
 Cavalier king Charles spaniel = heart valve failure,
syringomelia (paralysis), hip dysplasia, entropion (eye
problem), patella luxation (lameness), lifespan ca 7-10 years