PROS
AND CONS OF INBREEDING
SOK
2003-2, Copyright 1996, 2001, 2003 Sarah Hartwell
Inbreeding
is the mating together of closely related cats, for example mother/son, father/daughter,
sibling/sibling matings and half-sibling/half-sibling. It is the pairing of
animals which are more closely related than the average population. For breeders,
it is a useful way of fixing traits in a breed - the pedigrees of some
exhibition cats show that many of their forebears are closely related. For
example, the name of Fan Tee Cee (shown in the 1960s and 1970s) appeared in more
and more Siamese pedigrees, sometimes several times in a single pedigree, as
breeders were anxious to make their lines more typey. Superb specimens are
always much sought after for stud services or offspring (unless they have
already been neutered; cloning may solve that problem in the future) having won
the approval of show judges.
To
produce cats
which closely meet the breed standard, breeders commonly mate
together animals which are related and which share desirable characteristics.
Over time, sometimes only one or two generations, those characteristics will
become homozygous (genetically uniform) and all offspring of the inbred animal
will inherit the genes for those characteristics (breed true). Breeders can
predict how the offspring will look. "Line-breeding" is not a term
used by geneticists, but comes from livestock husbandry. It indicates milder
forms of inbreeding. Line-breeding is still a form of inbreeding i.e. breeding
within a family line and includes cousin/cousin, aunt/nephew, niece/uncle and
grandparent/grandchild. The difference between line-breeding and inbreeding may
be defined differently for different species of animals and even for different
breeds within the same species. It is complicated by the fact that a cat's
half-brother might also be her father!
However,
inbreeding holds potential problems.
The limited gene-pool caused by continued
inbreeding means that deleterious genes become widespread and the breed loses
vigour. Laboratory animal suppliers depend on this to create uniform strains of
animal which are immuno-depressed or breed true for a particular disorder e.g.
epilepsy. Such animals are so inbred as to be genetically identical (clones!), a
situation normally only seen in identical twins. Similarly, a controlled amount
of inbreeding can be used to fix desirable traits in farm livestock e.g. milk
yield, lean/fat ratios, rate of growth etc. In human terms, inbreeding is
considered incest; cats do not have incest taboos.
Outcrossing
is when the two parents are totally unrelated. In pedigree animals, this often
means where a common ancestor does not occur behind either parent within a four
or five generation pedigree. In animals with a small foundation gene pool, this
condition is difficult to meet.
First
I will define some terms used by animal breeders. In general I've avoided
specialist terms, but you will meet these terms outside of this article.
"Homozygous"
means having inherited the same "gene" for a particular trait from
both parent e.g. for fur length. Barring random mutation, 100% of the offspring
of a homozygous individual will inherit that gene. Inbreeding increases
homozygosity by "fixing" a particular trait. Purebred animals display
a high degree of homozygosity compared to mixed breeds and random-bred animals.
The idea of purebred animals is that they should "breed true". When
one purebred is mated with another of the same breed, the offspring will have
uniform characteristics and will resemble the parents.
"Heterozygous"
means having inherited a different gene for a particular trait from each parent.
For example one gene of long fur (recessive) and one gene for short fur
(dominant). 50% of a heterozygous individual's offspring will inherit one form
and 50% will inherit the other. Carefully controlled "out-crossing"
increases heterozygosity for selected traits by introducing new genes into the
hybrid offspring.
"Heterosis"
is the scientific term for hybrid vigour. It is possible that there are
"bad" genes which produce less vigourous individuals when in the
homozygous state because good genes have been bred out along with the
undesirable characteristics; theoretically the bad genes could be bred out, but
in practice this doesn't seem to happen. The other theory is simply that you
simply need to have a mixture of two different genes to get the desired effect
as they somehow complement each other; highly inbred animals lack this diversity
and have poorer immune systems.
"Sex-linked"
refers to a trait which is passed on, or determined by, a particular gender. In
Abyssinian cats there are several versions of the red colour. One is sex-linked
i.e. a male cat only needs one copy of the gene, but a female needs two copies
of the gene to produce the red colour.
NATURAL
OCCURRANCE OF INBREEDING
This
is not to say that inbreeding does not occur naturally. A feral colony which is
isolated from other cats, by geographical or other factors, can become very
inbred especially if a dominant male mates with his sisters, then with his
daughters and grand-daughters. When he is deposed it will most likely be by his
own son or grandson which therefore continues the inbreeding. The effect of any
deleterious genes becomes noticeable in later generations as the majority of the
offspring inherit these genes. Scientists have discovered that cheetahs, even if
living in different areas, are genetically very similar. Possibly disease or
disaster drastically reduced cheetah numbers in the past creating a genetic
bottleneck. All modern day cheetahs may be descended from a single surviving
family unit hence their genetic uniformity.
In
the Cheetah,
the lack of genetic diversity makes them susceptible to disease
since they lack the ability to resist certain viruses. Extreme inbreeding
affects their reproductive success with small litter sizes and high mortality
rates. Some scientists hope that the appearance of the "King Cheetah",
characterised by its blotched markings, means that the Cheetah can develop a
healthier gene-pool through mutation (provided man doesn't wipe them out in the
meantime). Mankind has contributed to the loss of diversity by sport-hunting
cheetahs and reducing the number of available mates. Hunters preferred to shoot
unusual specimens (i.e. genetically diverse ones) for the trophy room and this
previously included long-haired "woolly cheetahs" and grey/blue
cheetahs indicating much greater genetic diversity in the past.
Despite
the hopes of scientists, some cheetah populations are showing further signs of
inbreeding. Confined to ever-smaller areas such as wildlife reserves, the
populations have become genetically isolated from each other. In one population
there is an increasing frequency of misaligned jaws and kinked tails. Continued
inbreeding will reinforce, or fix, these traits. Ultimately, they could reduce
the cheetah's hunting effectiveness - the kinked tail will reduce its agility
and cornering ability and the misaligned jaws may not be able to hold onto prey.
The
wolf
was once widespread throughout North
American, but many of the remaining
packs are isolated and have become inbred. The isolation/inbreeding problem has
become so acute that conservationists have taken wolves from one area and
introduced them into a another area to revitalise the gene pool. In some areas,
the choice of mates is so reduced that wolves have resorted to mating with
domestic dogs - an extreme form of outcrossing called hybridization. A similar
situation in Scotland where the Scottish Wildcat mates with domestic cats is
threatening to wipe out the wildcat as domestic cat genes become more widespread.
These are two cases where outcrossing (following, and followed by, inbreeding)
may lead to extinction of a species (analogous to loss of type in domestic
breeds).
Another
animal
suffering from the effects of inbreeding is the Giant Panda. As with the
Cheetah, this has led to poor fertility among Pandas and high infant mortality
rates. As Panda populations become more isolated from one another (due to humans
blocking the routes which Pandas once used to move from one area to another),
Pandas have greater difficulty in finding a mate with a different mix of genes
and breed less successfully.
It
is almost inevitable that the Giant Panda will become extinct even if cloning
techniques become available since the gene pool is now probably too impoverished
for the species' long term viability. It may, therefore, be considered that all
purebred animals will ultimately become unviable through inbreeding and that
breeders must work carefully to maintain type while slowing down the detrimental
effects of selective breeding.
Natural
isolation
and inbreeding have given rise to domestic cat breeds such as the Manx
which developed on an island so that the gene for taillessness became widespread
despite the problems associated with it. Apart from the odd cat jumping ship on
the Isle of Man, there was little outcrossing and the effect of inbreeding is
reflected in smaller-than-average litter sizes (geneticists believe that more
Manx kittens than previously thought are reabsorbed due to genetic abnormality),
stillbirths and spinal abnormalities which diligent breeders have worked so hard
to eliminate.
As
mentioned, some feral colonies become highly inbred due to being isolated from
other cats (e.g. on a remote farm) or because other potential mates in the area
have been neutered, removing them from the gene pool. Most cat workers dealing
with ferals have encountered some of the effects of inbreeding. Within such
colonies there may be a higher than average occurrence of certain traits. Some
are not serious e.g. a predominance of calico pattern cats. Other inherited
traits which can be found in greater than average numbers in inbred colonies
include polydactyly (the most extreme case reported so far being an American cat
with 9 toes on each foot), dwarfism (although dwarf female cats can have
problems when try to deliver kittens due to the kittens' head size), other
structural deformities or a predisposition to certain inheritable conditions.
The
ultimate result of continued inbreeding is terminal lack of vigour and probable
extinction as the gene pool contracts, fertility decreases, abnormalities
increase and mortality rates rise. On the other hand, too much outcrossing will
cause loss of type and therefore the loss of a distinct breed.
SELECTIVE BREEDING
Artificial
isolation (selective breeding) produces a similar effect. When creating a new
breed from an attractive mutation, the gene pool is initially necessarily small
with frequent matings between related cats. Some breeds which resulted from
spontaneous mutation have been fraught with problems such as spasticity (cerebellar
hypoplasia) in Devon Rexes, skeletal problems in Scottish Folds and the effects
of a semi-lethal gene (aka deferred lethal gene) in Manxes and the lethal gene
in Ojos Azules. Problems such as hip dysplasia and patella luxation are more
common in certain breeds and breeding lines than in others, suggesting that past
inbreeding has distributed the faulty genes. Selecting suitable outcrosses can
reintroduce healthy genes, which might otherwise be lost, without adversely
affecting type.
An
example of a lethal gene which determines a breed trait is the blue-eyed Ojos
Azules. The gene is lethal in the homozygous form causing stillbirth, cranial
deformities, white fur and a small curled tail. In the heterozygous form, the
Ojos Azules are blue eyed non-white cats. Breeders must therefore breed
blue-eyed cats (heterozygous) to non-blue-eyed cats (lacking the gene for the
eye colour, but having the conformation) in order to get a roughly 50/50 split
of blue-eyed and non-blue-eyed kittens while avoiding deformed dead kittens.
The
more that inbreeding is used to get rid of undesirable traits or to fix a
desirable trait, the more likely it is that individuals will also inherit the
same set of genes for the immune system from both parents, and be born with less
vigourous immune systems. The immune system problem is compounded over
successive generations as the animals become genetically more uniform (like the
cheetah). According to one theory, immunodeficiency may be caused by a simple
lack of heterozygosity in the genes that control the immune system. This is why
random-bred cats are generally so robust.
Zoos
engaged in captive breeding programs
are aware of this need to outcross their
own stock to animals from other collections. Captive populations are at risk
from inbreeding since relatively few mates are available to the animals, hence
zoos must borrow animals from each other in order to maintain the genetic
diversity of offspring. In sheep, centuries of selective breeding to improve the
quality of wool has caused an important trait to be lost. Ancestral sheep could
breed more than once per year. Modern sheep breed once per year. Only recently
has the importance of the lost gene been realised (i.e. to increase meat yield),
but to reintroduce it from primitive sheep would reduce other qualities
selectively bred for over centuries.
Most
laboratory mice
are becoming so highly inbred that they would probably not
survive outside of a sterile laboratory due to poor immune systems (they are
generally killed before this becomes a problem in the laboratory situation) and
some strains become extinct due to reproductive failure. Many are selectively
bred to exhibit defects which will kill them.
Inbreeding
holds problems for anyone involved in animal husbandry - from canary fanciers to
farmers. Early Turkish Vans were reported to be temperamental, a problem
apparently rectified by the importation of new stock. Attempts to change the
appearance of Burmese cats in America to produce a cat with a rounder head
resulted in cats with congenital problems. Siamese cats have become
progressively finer-boned as breeders strive to emphasise the foreign look,
resulting in frailer cats in some breeding lines.
In
the dog world,
a number of breeds now exhibit hereditary faults due to the
over-use of a particularly "typey" stud which was later found to carry
a gene detrimental to health. By the time the problems came to light they had
already become widespread as the stud had been extensively used to "improve"
the breed. In the past some breeds were crossed with dogs from different breeds
in order to improve type, but nowadays the emphasis is on preserving breed
purity and avoiding mongrels.
Those
involved with minority breeds (rare breeds) of livestock face a dilemma as they
try to balance purity against the risk of genetic conformity. Enthusiasts
preserve minority breeds because their genes may prove useful to farmers in the
future, but at the same time the low numbers of the breed involved means that it
runs the risk of becoming unhealthily inbred. When trying to bring a breed back
from the point of extinction, the introduction of "new blood" through
crossing with an unrelated breed is usually a last resort because it can change
the very character of the breed being preserved (as noted by cat fanciers when
Russian Blues were crossed to Blue Point Siamese after World War II). In
livestock, successive generations of progeny must be bred back to a purebred
ancestor for 6 - 8 generations before the offspring can be considered purebred
themselves.
In
the cat fancy,
breed purity is equally
desirable, but can be taken to ridiculous lengths. Some fancies will not recognise "hybrid" breeds such as the
Tonkinese because it produces variants (yet Manxes are recognised and also
produce variants). Breeds which cannot produce some degree of variability among
their offspring risk finding themselves in the same predicament as Cheetahs and
Giant Pandas. Such fancies have lost sight of the fact that they are registering
"pedigree" cats, not "pure-bred" cats, especially since they
may recognise breeds which require occasional outcrossing to maintain type!
The
breed purity debate
goes along these lines: should a breed be based on genotype
(what genes it inherited) or phenotype (appearance, despite an out-cross four
generations ago) A Tabby-point Siamese is phenotypically Siamese, but because
the tabby patter was introduced from non-Siamese cats, genotype-followers
consider it "not Siamese" and are worried it will pollute their
purebred breeding lines. In some registries, Exotic Longhairs are identical to
Persians, but may not be bred with Persians. Likewise, some registries do not
allow chocolate or lilac Persians to be classed as Persians because the colours
were introduced from Siamese cats (via the Himalayan breed) umpteen generations
ago and are therefore "tainted". Those "tainted" genes may
be bundled with whole lot of healthy genes; by not outcrossing "purebreds"
to "tainted" cats, the opportunity to increase the heterozygosity of
the immune system is lost.
One
formula to reduce inbreeding
and slow down the loss of vigour is to line-breed
for 2-3 generation and then out-cross to an unrelated line (or occasionally
another breed) to get back hybrid vigour and genetic diversity. However with the
emphasis on breeding for type and competitiveness on the showbench (and when
making a sale), the typey studs get used more and more often and there is less
and less chance of finding a truly unrelated line.
See
The
Pros and Cons of Cloning
for further discussion on inbreeding hazards should cloning of typey animals
become permissible.
IMPLICATIONS
OF INBREEDING FOR THE CAT BREEDER
Most
cat breeders are well aware of potential pitfalls associated with inbreeding
although it is tempting for a novice to continue to use one or two closely
related lines in order to preserve or improve type. Breeding to an unrelated
line of the same breed (where possible) or outcrossing to another breed (where
permissible) can ensure vigour. Despite the risk of importing a few undesirable
traits which may take a while to breed out, outcrossing can prevent a breed from
stagnating by introducing fresh genes into the gene pool. It is important to
outcross to a variety of different cats, considered to be genetically
"sound" (do any of their previous offspring exhibit undesirable traits?)
and preferably not closely related to each other. Outcrossing is made difficult
by the amount of inbreeding in previous generations - it becomes hard to find
cats which are not related, sometimes several times over.
How
can you tell if a breed or line is becoming too closely inbred? One sign is that
of reduced fertility in either males or females. Male Cheetahs are known to have
a low fertility rate. Failure to conceive, small litter sizes and high kitten
mortality on a regular basis indicates that the cats may be becoming too closely
related. The loss of a large proportion of cats to one disease (e.g. enteritis)
indicates that the cats are losing/have lost immune system diversity. If 50% of
individuals in a breeding program die of a simple infection, there is cause for
concern.
Highly
inbred cats
also display abnormalities on a regular basis as "bad"
genes become more widespread. These abnormalities can be simple undesirable
characteristics such as misaligned jaws (poor bite) or more serious deformities.
Sometimes a fault can be traced to a single stud or queen which should be
removed from the breeding program even if it does exhibit exceptional type. If
its previous progeny are already breeding it's tempting to think "Pandora's
Box is already open and the damage done so I'll turn a blind eye". Ignoring
the fault and continuing to breed from the cat will cause the faulty genes to
become even more widespread in the breed, causing problems later on if its
descendants are bred together.
One
breed which was almost lost
because of inbreeding is the American Bobtail.
Inexperienced breeders tried to produce a colourpoint bobtailed cat with white
boots and white blaze and which bred true for type and colour, but only
succeeded in producing unhealthy inbred cats with poor temperaments. A later
breeder had to outcross the small fine-boned cats she took on, at the same time
abandoning the rules governing colour and pattern, in order to reproduce the
large, robust cats required by the standard and get the breed on a sound genetic
footing.
|
|
PROS
|
CONS
|
|
INBREEDING
(Mating
of closely related individuals)
|
Produces
uniform or predictable offspring.
Hidden (recessive) genes show up and can be eliminated.
Individuals will "breed true" and are "pure."
Doubles up good genes.
Eliminates unwanted traits.
|
Doubles
up on faults and weaknesses.
Progressive loss of vigor and immune response.
Increased reproductive failures, fewer offspring.
Emphasis on appearance means accidental loss of "good" genes
for other attributes.
Genetically impoverished individuals.
|
|
LINE-BREEDING
(Mating
of less closely related individuals)
|
Avoid
inbreeding of very closely-related cats, but cats are still
"pure".
Produces uniform or predictable offspring.
Slows genetic impoverishment.
|
Require
excellent individuals.
Does not halt genetic impoverishment, only slows it down.
|
|
OUTCROSSING
(Mating
of unrelated individuals within the same breed)
|
Brings
in new qualities or reintroduces lost qualities.
Increases vigor.
Cats are still "pure".
|
Less
consistency and predictability of offspring.
May have to breed out unwanted genes accidentally introduced at same
time.
May be hard to find individuals which are true outcrosses.
|
|
HYBRIDIZATION
(Mating
of unrelated individuals of different breeds)
|
Brings
in new qualities or reintroduces lost qualities.
Increases vigor, may improve immune system and reproductive capacity.
Introduces totally new traits e.g. color. Fur type.
May result in new breeds.
The offspring are considered "impure" for many generations.
|
Unpredictable
- new traits may not all be desirable.
Must choose outcross breed whose qualities complement or match own
breed.
May take years to eliminate unwanted traits/loss of type.
May take years to get consistent offspring.
Produces many variants not suitable for use in breeding program.
|
CONCLUSION
Inbreeding
is a two-edged sword. On the one hand a certain amount of inbreeding can fix and
improve type to produce excellent quality animals. On the other hand, excessive
inbreeding can limit the gene pool so that the breed loses vigour. Breeds in the
early stages of development are most vulnerable as numbers are small and the
cats may be closely related to one another. It is up to the responsible breeder
to balance inbreeding against crossings with unrelated cats in order to maintain
the overall health of the line or breed concerned.
FOOTNOTE:
THE CHEETAH
In
January 2003, India announced plans to clone cheetahs to help restore the Indian
sub-continent's now extinct cheetah population. India plan to clone cheetahs
from Iran where about 50 Asiatic Cheetahs remain.
Cloning
creates genetically identical individuals. However, cheetahs are already so
highly inbred that individuals are already almost genetically identical so the
impact of clones on the population will be to increase numbers rather than
further decease genetic variation. If it uses the leopard's own egg cells it
would introduce the leopard's mitochondrial DNA (the DNA found in an egg cell)
into the cheetah population.
Home
Siamese Oriental
Shorthair Kittens
Kittens 2 Planed
Nesten Offspring New
Owners News
Studs Studhouse
Our Aim Neuters
Healthy Breeding Who
we are Otazu Updates
Links