Captive breeding

Captive breeding is the process of

maintaining plants or animals in controlled
environments, such as wildlife reserves,
zoos, botanic gardens, and other
conservation facilities. It is sometimes
employed to help species that are being
threatened by human activities such as
habitat loss, fragmentation, over hunting
or fishing, pollution, predation, disease,
and parasitism.[1] In some cases a captive
breeding program can save a species from
extinction, but for success, breeders must
consider many factors—including genetic,
ecological, behavioral, and ethical issues.
Most successful attempts involve the
cooperation and coordination of many
institutions.

History
USFWS staff with two red wolf pups bred in captivity

The Arabian Oryx is one of the first animals

reintroduced via a captive breeding program.

Captive breeding techniques began with

the first human domestication of animals
such as goats, and plants like wheat, at
least 10,000 years ago.[2] These practices
were then expanded with the rise of the
first zoos, which started as royal
menageries in Egypt[3] and its popularity,
which led to the increase in zoos
worldwide. The first actual captive
breeding programs were only started in the
1960s. These programs, such as the
Arabian Oryx breeding program from The
Phoenix Zoo in 1962, were aimed at the
reintroduction of these species into the
wild.[4] These programs expanded under
The Endangered Species Act of 1973 of
the Nixon Administration, which focused
on protecting endangered species and
their habitats to preserve biodiversity.[5]
Since then, research and conservation
centers have been housed in zoos, such as
the Institute for Conservation Research at
the San Diego Zoo founded in 1975 and
expanded in 2009,[6] which have
contributed to the successful conservation
efforts of species such as the Hawaiian
Crow.[7]

Coordination
The breeding of species of conservation
concern is coordinated by cooperative
breeding programs containing
international studbooks and coordinators,
who evaluate the roles of individual
animals and institutions from a global or
regional perspective. These studbooks
contain information on birth date, gender,
location, and lineage (if known), which
helps determine survival and reproduction
rates, number of founders of the
population, and inbreeding coefficients.[8]
A species coordinator reviews the
information in studbooks and determines
a breeding strategy that would produce
most advantageous offspring.

If two compatible animals are found at

different zoos, the animals may be
transported for mating, but this is
stressful, which could in turn make mating
less likely. However, this is still a popular
breeding method among European
zoological organizations.[9] Artificial
fertilization (by shipping semen) is another
option, but male animals can experience
stress during semen collection, and the
same goes for females during the artificial
insemination procedure. Furthermore, this
approach yields lower-quality semen,
because shipping requires extending the
life of the sperm for the transit time.

There are regional programmes for the

conservation of endangered species:

Americas: Species Survival Plan SSP

(Association of Zoos and Aquariums
AZA, Canadian Association of Zoos and
Aquariums CAZA)
Europe: European Endangered Species
Programme EEP (European Association
of Zoos and Aquaria EAZA)
Australasia: Australasian Species
Management Program ASMP (Zoo and
Aquarium Association ZAA)
Africa: African Preservation Program
APP (African Association of Zoological
Gardens and Aquaria PAAZAB)
Japan: Conservation activities of
Japanese Association of Zoos and
Aquariums JAZA
South Asia: Conservation activities of
South Asian Zoo Association for
Regional Cooperation SAZARC
 South East Asia: Conservation activities
of South East Asian Zoos Association
SEAZA

Challenges
Genetics

The objective of many captive populations

is to hold similar levels of genetic diversity
to what is found in wild populations. As
captive populations are usually small and
maintained in artificial environments,
genetics factors such as adaptation,
inbreeding and loss of diversity can be a
major concern.
Domestication (Adaptation)

Adaptive differences between plant and

animal populations arise due to variations
in environmental pressures. In the case of
captive breeding prior to reintroduction
into the wild, it's possible for species to
evolve to adapt to the captive environment,
rather than their natural environment.[11]
Reintroducing a plant or animal to an
environment dissimilar to the one they
were originally from can cause fixation of
traits that may not be suited for that
environment leaving the individual
disadvantaged. Selection intensity, initial
genetic diversity, and effective population
size can impact how much the species
adapts to its captive environment.[10]
Modeling works indicate that the duration
of the programs (i.e., time from the
foundation of the captive population to the
last release event) is an important
determinant of reintroduction success.
Success is maximized for intermediate
project duration allowing the release of a
sufficient number of individuals, while
minimizing the number of generations
undergoing relaxed selection in
captivity.[11] Can be minimized by reducing
the number of generations in captivity,
minimizing selection for captive
adaptations by creating environment
similar to natural environment and
maximizing the number of immigrants
from wild populations.[12]

Genetic diversity

One consequence of small captive

population size is the increased impact of
genetic drift, where genes have the
potential to fix or disappear completely by
chance, thereby reducing genetic diversity.
Other factors that can impact genetic
diversity in a captive population are
bottlenecks and initial population size.
Bottlenecks, such as rapid decline in the
population or a small initial population
impacts genetic diversity. Loss can be
minimized by establishing a population
with a large enough number of founders to
genetically represent the wild population,
maximize population size, maximize ratio
of effective population size to actual
population size, and minimize the number
of generations in captivity.[11]

Inbreeding

Inbreeding is when organisms mate with

closely related individuals, lowering
heterozygosity in a population. Although
inbreeding can be relatively common,
when it results in a reduction in fitness it is
known as inbreeding depression. The
detrimental effects of inbreeding
depression are especially prevalent in
smaller populations and can therefore be
extensive in captive populations.[13] To
make these populations the most viable, it
is important to monitor and reduce the
effects of deleterious allele expression
caused by inbreeding depression and to
restore genetic diversity.[13] Comparing
inbred populations against non-inbred or
less-inbred populations can help
determine the extent of detrimental
effects if any are present.[14] Closely
monitoring the possibility of inbreeding
within the captive bred population is also
key to the success of reintroduction into
the species' native habitat.

The Speke's Gazelle was the focus of a captive

breeding program centered on determining the effect
of selection on reducing genetic load.

Outbreeding

Outbreeding is when organisms mate with

unrelated individuals, increasing
heterozygosity in a population. Although
new diversity is often beneficial, if there a
large genetic differences between the two
individuals it can result in outbreeding
depression. This is a reduction in fitness,
similar to that of inbreeding depression,
but arises from a number of different
mechanisms, including taxonomic issues,
chromosomal differences, sexual
incompatibility, or adaptive differences
between the individuals.[15] A common
cause is chromosomal ploidy differences
and hybridization between individuals
leading to sterility. The best example is in
the orangutan, which, prior to taxonomic
revisions in the 1980s would be commonly
mated in captive populations producing
hybrid orangutans with lower fitness. If
chromosomal ploidy is ignored during
reintroduction, restoration efforts would
fail due to sterile hybrids in the wild. If
there are large genetic differences
between individuals originally from distant
populations, those individuals should only
be bred in circumstances where no other
mates exist.

Behavior Changes

Captive breeding can contribute to

changes in behavior in animals that have
been reintroduced to the wild. Released
animals are commonly less capable of
hunting or foraging for food, which leads
to starvation, possibly because the young
animals spent the critical learning period
in captivity. Released animals often
display more risk-taking behavior and fail
to avoid predators.[16] Golden lion tamarin
mothers often die in the wild before having
offspring because they cannot climb and
forage. This leads to continuing population
declines despite reintroduction as the
species are unable to produce viable
offspring. Training can improve anti-
predator skills, but its effectiveness
varies.[17][18]

Salmon bred in captivity have shown

similar declines in caution and are killed by
predators when young. However, salmon
that were reared in an enriched
environment with natural prey showed less
risk-taking behaviors and were more likely
to survive.[19]

A study on mice has found that after

captive breeding had been in place for
multiple generations and these mice were
"released" to breed with wild mice, that the
captive-born mice bred amongst
themselves instead of with the wild mice.
This suggests that captive breeding may
affect mating preferences, and has
implications for the success of a
reintroduction program.[20]
Chatham Island Black Robin on Rangatira Island, New
Zealand.

Human mediated recovery of species can

unintentionally promote maladaptive
behaviors in wild populations. In 1980 the
number of wild Chatham Island Black
Robins was reduced to a single mating
pair. Intense management of populations
helped the population recover and by 1998
there were 200 individuals. During
recovery scientists observed "rim laying"
an egg laying habit where individuals laid
eggs on the rim of the nest instead of the
center. Rim laid eggs never hatched. To
combat this land managers pushed the
egg to the center of the nest, which greatly
increased reproduction. However, by
allowing this maladaptive trait to persist,
over half the population were now rim
layers. Genetic studies found that this was
an autosomal dominant mendelian trait
that was selected for due to human
intervention[21]

Successes
A cheetah at the De Wildt Cheetah and Wildlife Centre.

The De Wildt Cheetah and Wildlife Centre,

established in South Africa in 1971, has a
cheetah captive breeding program.
Between 1975 and 2005, 242 litters were
born with a total of 785 cubs. The survival
rate of cubs was 71.3% for the first twelve
months and 66.2% for older cubs,
validating the fact that cheetahs can be
bred successfully (and their endangerment
decreased). It also indicated that failure in
other breeding habitats may be due to
"poor" sperm morphology.[22]

Wild Tasmanian devils have declined by

90% due to a transmissible cancer called
Devil Facial Tumor Disease.[23] A captive
insurance population program has started,
but the captive breeding rates at the
moment are lower than they need to be.
Keeley, Fanson, Masters, and McGreevy
(2012) sought to "increase our
understanding of the estrous cycle of the
devil and elucidate potential causes of
failed male-female pairings" by examining
temporal patterns of fecal progestogen
and corticosterone metabolite
concentrations. They found that the
majority of unsuccessful females were
captive-born, suggesting that if the
species' survival depended solely on
captive breeding, the population would
probably disappear.[24]

In 2010, the Oregon Zoo found that

Columbia Basin pygmy rabbit pairings
based on familiarity and preferences
resulted in a significant increase in
breeding success.[25]

Methods Used in Captive

Breeding
Every known individual of the California condor
population has been captured and then bred using
research from microsatellite regions in their genome.

To found a captive breeding population

with adequate genetic diversity, breeders
usually select individuals from different
source populations—ideally, at least 20-30
individuals. Founding populations for
captive breeding programs have often had
fewer individuals than ideal because of
their threatened state, leaving them more
susceptible to challenges such as
inbreeding depression.[26]
To overcome challenges of captive
breeding such as adaptive differences,
loss of genetic diversity, inbreeding
depression, and outbreeding depression
and get desired results, captive breeding
programs use many monitoring methods.
Artificial insemination is used to produce
the desired offspring from individuals who
don't mate naturally to reduce effects of
mating closely related individuals such as
inbreeding.[26] Methods as seen in panda
pornography allow programs to mate
chosen individuals by encouraging mating
behavior.[27] As a concern in captive
breeding is to minimize the effects of
breeding closely related individuals,
microsatellite regions from an organisms
genome can be used to determine
amounts of relationship among founders
to minimize relatedness and pick the most
distant individuals to breed.[26] This
method has successfully been used in the
captive breeding of the California condor
and the Guam rail. The maximum
avoidance of inbreeding (MAI) scheme
allows control at a group level rather than
an individual level by rotating individuals
between groups to avoid inbreeding.[26]

New technologies
Assisted reproduction
technology (ART): Artificial
insemination

Getting captive wild animals to breed

naturally can be a difficult task. Giant
pandas for example lose interest in mating
once they are captured, and female giant
pandas only experience estrus once a year,
which only lasts for 48 to 72 hours.[28]
Many researchers have turned to artificial
insemination in an attempt to increase the
populations of endangered animals. It may
be used for many reasons, including to
overcome physical breeding difficulties, to
allow a male to inseminate a much larger
number of females, to control the paternity
of offspring, and to avoid injury incurred
during natural mating.[29] It also creates
more genetically diverse captive
populations, enabling captive facilities to
easily share genetic material with each
other without the need to move animals.
Scientist of the Justus-Liebig-University of
Giessen, Germany, from the working group
of Michael Lierz, developed a novel
technique for semen collection and
artificial insemination in parrots producing
the world's first macaw by assisted
reproduction[30]

Cryopreservation
Animal species can be preserved in gene
banks, which consist of a cryogenic
facilities used to store live sperm, eggs, or
embryos in ultracold conditions. The
Zoological Society of San Diego has
established a "frozen zoo" to store frozen
tissue from the world's rarest and most
endangered species samples using
cryopreservation techniques. At present,
there has been more than 355 species,
including mammals, reptiles, and
birds.Cryopreservation can be performed
as oocyte cryopreservation before
fertilization, or as embryo cryopreservation
after fertilization. Cryogenically preserved
specimens can potentially be used to
revive breeds that are endangered or
rextinct, for breed improvement,
crossbreeding, research and development.
This method can be used for virtually
indefinite storage of material without
deterioration over a much greater time-
period relative to all other methods of ex
situ conservation. However, cryo-
conservation can be an expensive strategy
and requires long term hygienic and
economic commitment for germplasms to
remain viable. Cryo-conservation can also
face unique challenges based on the
species, as some species have a reduced
survival rate of frozen germplasm,[31] but
cryobiology is a field of active research
and many studies concerning plants are
underway.

An example of the use of cryoconservation

to prevent the extinction of a livestock
breed is the case of the Hungarian Grey
cattle, or Magya Szurke. Hungarian Grey
cattle were once a dominant breed in
southeastern Europe with a population of
4.9 million head in 1884. They were mainly
used for draft power and meat. However,
the population had decreased to 280,000
head by the end of World War II and
eventually reached the low population of
187 females and 6 males from 1965 to
1970.[32] The breed's decreased use was
due primarily to the mechanization of
agriculture and the adoption of major
breeds, which yield higher milk
production.[33] The Hungarian government
launched a project to preserve the breed,
as it possesses valuable traits, such as
stamina, calving ease, disease resistance,
and easy adaptation to a variety of
climates. The government program
included various conservation strategies,
including the cryopreservation of semen
and embryos.[32] The Hungarian
government's conservation effort brought
the population up to 10,310 in 2012, which
shows significant improvement using
cryoconservation.[34]
Cloning

The best current cloning techniques have

an average success rate of 9.4 percent,[35]
when working with familiar species such
as mice, while cloning wild animals is
usually less than 1 percent successful.[36]
In 2001, a cow named Bessie gave birth to
a cloned Asian gaur, an endangered
species, but the calf died after two days. In
2003, a banteng was successfully cloned,
followed by three African wildcats from a
thawed frozen embryo. These successes
provided hope that similar techniques
(using surrogate mothers of another
species) might be used to clone extinct
species. Anticipating this possibility, tissue
samples from the last bucardo (Pyrenean
ibex) were frozen in liquid nitrogen
immediately after it died in 2000.
Researchers are also considering cloning
endangered species such as the giant
panda and cheetah. However, cloning of
animals is opposed by animal-groups due
to the number of cloned animals that
suffer from malformations before they
die.[37]

Interspecific pregnancy

A potential technique for aiding in

reproduction of endangered species is
interspecific pregnancy, implanting
embryos of an endangered species into
the womb of a female of a related species,
carrying it to term.[38] It has been used for
the Spanish Ibex[39] and Houbara
bustard.[40]

Ethical Considerations
With successes such as in 1986 when a
population only of 18 black ferrets left in
the world was brought back up to 500 in
the wild, and when the Arabian oryx was
brought back from extinction in 1972 to a
population of 1,000 in the deserts of the
Middle East, captive breeding programs
have proven successful throughout
history.[41] While captive breeding sounds
like an ideal solution for preventing
endangered animals from facing serious
threats of extinction there are still reasons
to believe that these programs can
occasionally do more harm than good.
Some detrimental effects include delays in
understanding optimal conditions required
for reproduction, failure to reach self-
sustaining levels or provide sufficient
stock for release, loss of genetic diversity
due to inbreeding, and poor success in
reintroductions despite available captive-
bred young.[42] Although it has been
proven that captive breeding programs
have yielded negative genetic effects in
decreasing the fitness of captive-bred
organisms, there is no direct evidence to
show that this negative effect also
decreases the overall fitness of their wild-
born descendants.[43]

There is reason to demand for the release

of animals from captivity programs for
four main reasons: a lack of sufficient
space due to overly successful breeding
programs, closure of facilities due to
financial reasons, pressure from animal
rights advocacy groups, and to aid the
conservation of endangered species.[44]
Additionally, there are many ethical
complications to reintroducing animals
born in captivity back into the wild. For
example, when scientists were
reintroducing a rare species of toad back
into the Mallorcan wild in 1993, a
potentially deadly fungus that could kill
frogs and toads was unintentionally
introduced.[45] It is also important to
maintain the organism’s original habitat, or
replicate that specific habitat for species
survival.

See also
Breeding in the wild
 European Endangered Species
Programme (EEP)
Ex-situ conservation
Panda pornography
Species Survival Plan or SSP
World Conference on Breeding
Endangered Species in Captivity as an
Aid to their Survival or WCBESCAS
Zooborns

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doi:10.1111/1365-2664.12449 .
43. Araki, H; Cooper, B; Blouin, M. S
(2009). "Carry-over effect of captive
breeding reduces reproductive fitness
of wild-born descendants in the wild" .
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doi:10.1098/rsbl.2009.0315 .
PMC 2781957 . PMID 19515651 .
44. Waples KA, Stagoll CS (1997). "Ethical
Issues in the Release of Animals from
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45. "Captive Breeding Introduced
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External links
Fraser, Dylan J (2008). "How well can
captive breeding programs conserve
biodiversity? A review of salmonids" .
Evolutionary Applications. 1 (4): 535–86.
doi:10.1111/j.1752-4571.2008.00036.x .
PMC 3352391 . PMID 25567798 .
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