Breeding-Policy Sep12
Breeding-Policy Sep12
Guidelines
For
Healthy
And
Responsible
Breeding
Breeding
Policy For The
British Shorthair
Sep-12
Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Summary
Acknowledgments
1
These can be found at Appendix 4 Contributing Breeders
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Contents
Summary ...................................................................................... 2
Acknowledgments .................................................................. 2
Contents ....................................................................................... 3
1. Introduction ..................................................................... 6
4. Type ......................................................................... 21
4.6. Coat.......................................................................... 22
5. Health ...................................................................... 25
7.3. Tortie and White and Van Pattern Tortie and White. 59
8. Smoke ............................................................................ 61
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
1. Introduction
It is very easy to breed a cat, in fact if you share your life with a female cat and
do nothing to prevent it you will soon have a litter of kittens and a very good
mother taking care of everything. However, if one makes a decision to become a
breeder of pedigree cats all sorts of responsibilities accompany that decision.
GCCF
BAC Breeders
Registration Breeding
SOP
Policy Policy
Breeding decisions become easier if breeders develop and manage a breeding
program within which they consider the key points of cat breeding, which are:
2
Governing Council of the Cat Fancy
3
SOP
4
BAC
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
In order to achieve the desirable goals, whilst managing the risks the breeders
system should give consideration to:
1.2. Exclusions
This document is not intended to guide breeders as to the day to day welfare and
care of the breeding animal.
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2.1. Origin
The Romans introduced large numbers of cats to the United Kingdom as working
cats to help reduce the rodent population. These cats interbred with the native
wild cats of Great Britain to produce the native domestic shorthaired cat. The
accepted date of origin of the British Shorthair as we know it today is 1870. 5
Despite being popular at the Crystal Palace cat show of 1871, by the end of World
War II along with many other British cat breeds, the British Shorthair numbers
were devastated. A number of careful breeders worked to recreate and restore
numbers using a combination of British Shorthairs, Persians, Russian Blues,
Burmese and other pedigree and non pedigree shorthair varieties.
Whilst the diversity helped with the health of the breed, breeders found that the
‘foreign type’ was persistent in the background and breeders found that they had
to keep falling back on the Persian occasionally to keep type. Always, the
objective was to reach a point where the British would breed true without any
need for outcrosses. (The longhair gene remains recessive amongst some British
Shorthairs today) 6
The British cat is compact, well balanced and powerful, showing good depth of
body, a full broad chest, short strong legs, rounded paws, tail thick at base with
rounded tip. The head is round with a good width between small ears, round
cheeks, firm chin, large round and well opened eyes and a broad short nose. The
coat is short and dense. A muscular cat with an alert appearance and in perfect
physical condition
The British Shorthair is one of the largest breeds of cat. It is chunky and
substantial; the male is much larger than the female. The face is round with full
cheeks and the nose is short and broad. The chin is deep and strong. The ears
are small and rounded and set so as to blend with the round contour of the head.
The eyes are large and round. The head is set on a short thick neck. The body is
cobby with a short level back. The chest is deep and the shoulders are strong.
The legs are short and strong with round paws. The tail is thick and of medium
length.
5
Robinson
6
Genetic tests for the LH gene are now available)
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Within these descriptions we can read that this is a cat whose beauty is all about
balance, size and roundness, balanced by strength and physical health.
The British Shorthair is a hugely popular show cat and with the wide range of
colours and patterns available it is one of the biggest breed sections at GCCF cat
shows.
2.4. Registration
The British Shorthair is the most popular pedigree cat, with more kittens
registered with the GCCF each year than any other breed.
Registered breed numbers have risen by 48% over the past 10 years to almost
6,800 in 2008.7
British Shorthairs are an easygoing breed of cat. They have a stable character
and take well to being kept as indoor-only cats, making them ideal for apartment
7
Latest GCCF data published
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
living. They are not terribly demanding of attention, though they will let their
owner know if they feel like playing. They are not hyperactive preferring to sit
close to their owners rather than on them.
They like attention and enjoy being petted. They are not a very vocal breed but
will meow to communicate with their owners. British Shorthairs like to follow
people from room to room, as they often wish to be with their owner and see
what is going on. Some do not mind being cuddled, but most prefer to keep four
paws on the ground and be patted rather than picked up. 8
2.6. Longevity
British Shorthairs are very healthy and long lived. You can expect a British to live
from 14 to 20 years.
2.7. Recognition
The British Shorthair is recognized by all the exhibiting bodies across the
continents.
8
Extract from Wikipedia
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
As we can see from the background information the British Shorthair is a popular
and robust cat, deserving dedicated and ethical breeders with the time to
establish good breeding systems and practices. And whilst breeding cats is ‘just a
hobby’ it is not possible to breed responsibly without a basic understanding of
genetics.
We will cover some aspects of the genetics of our breed, but much more
information can be found in works such as Robinson’s Genetics for Cat Breeders
and Veterinarians and of course the Internet is a valuable source of materials for
those seeking further information.
For some experienced breeders much of this information is already well known
and understood, but as a BAC we have responsibility to help the novice breeder
establish a basic understanding.
The phenotype is basically, that which you can see, phenotypes result from an
individual’s genes as well as the impact of environmental factors and the possible
effect that the latter may have upon the former. The genotype is the genetic
make-up of a cell, or an individual.
That fabulous rusty coloured (phenotype) cat that you may admire as chocolate,
may well in fact be a very happy black (genotype) cat enjoying the sun
(environmental)
Unlike many breeds where origins can be traced back to individual named cats,
the origins of the British lie very firmly with Felis Silvestris or Felis Lybica, a
mackerel tabby ancestor, with just a bit of help along the way.
By the beginning of the 20th century standards had been written for many of the
colours and type was generally as seen today, round compact cats with large
round heads and eyes, coats described as short and deep, fine texture with dense
undercoat.
(By 2009, the BAC determined that type had been successfully fixed and outcrosses no
longer permitted.)
In common with the Asian breed the BSH has one of the largest numbers of gene
variations of any breed of pedigree cat recognised by GCCF.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Whilst we understand much about how these genes work, there are so many that
even the scientists struggle to understand and complex polygenes 9 make the
effects of major genes even harder to get to grips with. Sometimes we have to
accept that the evidence for how these genes and polygenes work together lies in
the ability to breed with consistent results.
It will help if we explain that there are genes for colour and genes for pattern.
The genetic penny drops for many novice breeders when they are able to
separate the genes that govern colour from those that govern pattern
3.1.1.Coat Colours
Essentially there are four basic colours for coat or pigment in the cat, these are
Black, Chocolate, Cinnamon and Red.
And just when it seems simple, we have to consider the effect of the dilute gene
on the basic gene colours. And it always helps breeders to understand the rules of
colour dominance.
Figure 1 shows the dominance hierarchy and the effect of the dilute gene.
Dominant Dilute
Black
Blue
Dominant over Chocolate and
Cinnamon Dominant over Lilac and Fawn
Chocolate Lilac
Dominant over Cinnamon Dominant over Fawn
Cinnamon Fawn
Figure 1
9
Genes with small but cumulative effects on the expression of a characteristic
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Red is the sex-linked orange gene, ‘O’ and ‘O’ determines if there will be any red
or cream (dilute) hair. ‘O’ completely replaces the black pigment gene.
3.1.2.Coat Patterns
It seems as we have more coat colours than we have because of the genes for
‘pattern’ which modify the colours. The main genes for pattern in the British are:
To help us get to grips with the genes for colour and pattern we need to know
some basic terms.
These terms will crop up throughout the document and so are well worth
understanding. 11
10
This list is repeated in the Glossary
11
A list of the key genes at work to produce the many patterns of British Shorthair can
be found at appendix 1.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Any ‘best practice’ breeding system must include the planning of the actual
breeding program, this is where a breeder decides his or her objectives, seeking
to maintain good breed type, and make improvements. The breeder will make a
decision to mate a certain queen to a given stud, often with great hopes for the
outcome.
As a part of this policy document we will ask our breeders to consider health,
selection of parents and inbreeding.
Colourpointed 438 72 15
Tortoiseshell 16 405 68
White 23 4 1
13
Assumptions: The average litter size is 4; the average queen will have 3 litters in 2
years. There is on average one stud cat for every 5 queens.
15
There is a table of British Registrations available on the BAC website
16
The calculated fathers of these tortie kittens have been shared between self and
red/cream.
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3.3. Inbreeding
Well it certainly is, the media are full of tales of ‘mutant cats and dogs’ and if one
‘Google’s’ ‘Inbreeding cats and dogs, the first four resources offered all refer to
‘problems’ However, as well as being ‘emotional’ it is also an extremely complex
topic. Therefore, in this section we will seek to provide basic information for
novice breeders and share some best practice advice from more experienced
breeders.
Further reading and information can be found both in Robinsons Genetics For Cat
Breeders and Veterinarians and also, the GCCF breeding policy 17 provides a
section on inbreeding.
In line with our ethos of keeping this simple, it is virtually impossible for breeders
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of the British Shorthair to avoid inbreeding. Data provided by Robinson tells us
that if we wished to prepare a 20 generation pedigree for two cats with no
duplicated ancestors we would need in excess of two million cats. To get this into
perspective we only have to see that the above table is suggesting that we have
somewhere in the region of 1000 queens and 200 studs currently at work in the
UK. We must also accept that of the possible 200 stud cats available, many of
these will be at closed stud, limited stud, or simply the wrong colour.
17
Available free of charge on the GCCF website.
18
Robinson’s Genetics for Cat Breeders & Veterinarians.
19
Sourced from RGCB&V
20
To a geneticist it is all inbreeding.
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Cousin
So, we are saying that inbreeding is unavoidable, and to a certain extent we must
accept that despite the ‘bad press’ inbreeding is, if correctly managed, important.
After all without it we would not have the cat that we all know and love today.
The important point is ‘correct management’ and if we can ascertain some form
of best practice we can hope to influence the decisions of both experienced and
reputable breeders, who will hopefully mentor novice breeders and thus help to
keep a healthy balance of inbreeding within the breed as a whole.
When we mate two cats together their offspring will each be given one gene from
each parent, when these genes are different, the offspring will be said to be
‘heterozygous’ for the inherited characteristics.
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These genes will combine to determine (to a greater or lesser extent) the
following characteristics:
Health
Reproductive
Ability
Therefore, it would seem to make sense that breeders will select superb
specimens of their breed to mate together and thus produce even more wonderful
cats. However, to produce cats which closely meet the desired standard, breeders
often mate together animals which are related and which share desirable physical
characteristics. Some breeders create a line which will ‘Breed True’ and breeders
can predict not just how the offspring will look, but that the offspring will, largely
be uniform within a litter.
All of this sounds fine so far until we remember that it is not only the ‘phenotype’
of the cat that is ‘breeding true’, behind the scenes the ‘genotype’ may also
becoming uniform.
Once again, if the breeder is responsibly managing all four aspects of the above
inherited characteristics, and can be reasonably certain that all the kittens from
these ‘inbred’ matings will be sound, healthy, long-lived and able to produce,
without difficulty, sound and healthy kittens. Then all will be well.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Where it starts to go potentially wrong is when any one or any group of breeders
start to breed for certain characteristics such as ‘coat’, – breeding a daughter
back to her father in the hope that:
Is fine unless both cats also carry the gene for disease, then sadly the
outcome for the offspring could be very bleak. 21
It is not just the risk of genetic disease that comes with inbreeding. The more
that we use inbreeding to ‘fix’ desirable traits the more likely it is that kittens will
also inherit the same set of genes for the immune system from both parents, and
be born with potentially compromised immune systems. Persistent inbreeding will
build up over successive generations as the offspring become genetically more
uniform, an increase in sickness may result.
21
Please note that this diagram is illustrative. The genes for PKD and HCM are both
dominant mutations. Therefore, cats only need to carry one copy in order to develop
disease. However, it is thought that kittens born with two copies of the HCM mutation are
more likely to develop serious disease and PKD is almost certainly a deadly double
domiant.
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The following ‘traffic light’ might be useful for breeders able to calculate the
coefficient. 22
% Coefficient
• ACCEPTABLE
• Over 8 Generations (where only 5 generations 1-17%
1-25%
• BE CAREFUL
26-39% • Over 8 Generations (where only 5 generations 17-27%)
• STOP
40-50% • Over 8 generations (Where only 5 generations 28-33)
Figure 4
The truth is that there are no hard and fast rules that can help us determine a
‘safe level’ of inbreeding. What we can do, both as individuals and as a group is
to ‘listen’ to our cats. There is no perfect coefficient ‘number’ that can declare one
mating safe and another risky – one sure fire way of knowing that we are going
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too far is too look out for signs of Inbreeding Depression, some of which are :
Of course, any of the above signs may occur randomly, or indeed may be
traceable to a single animal, which could then be removed from a breeding
program. True inbreeding depression, describes an ongoing state of affairs that
will continue unless responsible breeders step in and make a genetic correction.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
There is a risk that all of this sounds very depressing, but in truth many
experienced breeders naturally manage their breeding programs holistically,
taking all the characteristics into consideration. These, our best breeders, will not
hesitate to remove an otherwise superb physical specimen from their breeding
programs if a genetic defect is suspected or found.
The good news is that the BSH does not show signs of dangerous
inbreeding at a breed level.
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4. Type
4.1. Overall
The Breed Standard looks for, a round face with full cheeks and good breadth of
skull with round underlying bone structure. The head should be set on a short
thick neck. A good way to describe a ‘good head’ is a broad apple shape with
chubby cheeks.
Importantly, the head should always be in balance with the body, an oversized
head is just as undesirable as a head that is too narrow.
The eyes are large, round and should be an important facial feature. Top breeders
aim for a sweetness of expression as well as eye colour.
4.3. Nose
24
The ‘visible’ cat
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Keeping the nose short, with a balanced break is just one of the juggling acts
most breeders have to perform.
4.4. Chin
The bite MUST be level, the tip of the chin to line up with
the tip of the nose in the same vertical plane.
4.5. Ears
4.6. Coat
The SOP 26 calls for a coat that is short, dense and crisp. A soft and / or overlong
and fluffy coat is incorrect.
The above statement makes it all sound so simple, yet the British Shorthair coat
is unique, no other cat breed has this wonderful coat. Read through Judge’s
reports and find words such as cracking, crisp, scrunchy, dense and firm. Judges
25
Shortened muzzle and face
26
Standard of Points
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
also refer to coats as being, too soft, too long, lying flat and lacking in density. So
how do we know when we have that coat right? Judges will press the coat feeling
for a firmness and density of undercoat, when held between two fingers the coat
should not stand higher than the fingers by very much. If the coat is stroked
backwards towards the head, a ‘good’ coat will remain standing erect.
4.7. Eyes
For some breeders it is all in the eyes – not just colour - but shape and set.
4.8. Body
A well balanced cat will also have a firm body, without sagging. Breeders are
looking for good bone and a sound muscular body.
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4.10. Tail
According to the SOP the tail should be thick and of medium length, thicker at the
base with rounded tip.
The male British is noticeably larger, broader and rounder than the female, with
weights ranging between 6-7 kilos. But the female is also a big cat with weight,
when in condition, around 5-6 kilo’s.
The British is a slow growing cat and should be given time to mature. The adult
will not reach full maturity until three to four years of age.
The British Shorthair may be a big cat but it should not be a fat cat.
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5. Health
Just as genes determine all the good characteristics they also contribute to many
‘bad’ characteristics. We only have to look over our shoulder at some of the
unfortunate attributes that have slipped into some of the pedigree dog breeds.
But closer to home, the Munchkin Cat has been produced as a result of deliberate
breeding of a genetic mutation. Genes also contribute to disease and as breeders
seek to fix type they may inadvertently fix disease.
Fortunately, there are tests available for some devastating genetic diseases and
breeders can make informed choices. Reputable breeders should be aware of the
genes that can adversely affect the British. Where testing is possible, these
should be performed routinely and affected animals removed from breeding
programs.
Section 6 of the GCCF Breeding Policy gives details of the well known genetic
anomalies; we will just mention two of these:
5.1.1.PKD
5.1.2.HCM
The GCCF section on genetic anomalies states that the majority of HCM is
genetic. Unfortunately, at the moment testing in the British is limited to scanning
with ultrasound, which can be unpleasant for the animal and expensive. Amongst
British Breeders in Europe, annual HCM tests have become routine, but it has not
yet become custom and practice for British Breeders here in the UK. Our current
recommendation is that any breeder with reason to suspect an inherited HCM
condition in any of their cats, should seek a scan and if the animal is affected
remove him or her from the breeding population. Any progeny should also be
scanned and monitored.
27
A list of laboratories can be found on the BAC website – some of these offer
discounts to breeders as members of the breed clubs
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Blood Group Mismatch is an issue for the British section. As a BAC we have
decided to include a section on Neonatal Isoerythrolysis as it is the kittens of our
breed that are most likely to be affected by blood group mismatch.
There are experienced breeders in the UK today who have bred cats for many
years who do not know the blood group of their cats, who add new stock to their
lines and yet have never lost a kitten due to blood group mismatch. There are
others who have sad stories of entire litters have lost to a fading kitten syndrome
that has been diagnosed as Neonatal Isoerythrolysis.
Fortunately today it is possible for breeders to test their breeding cats for blood
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type in their own homes using non-invasive inexpensive swab tests.
A genetic mutation associated with the B blood group in most cats has been
identified and a DNA test has been developed. Animals can be tested at an early
age from a buccal swab.
The DNA blood group test identifies cats that have the B serotype (two copies of
b allele) and also that are B carriers (one copy of the b allele, A or AB serotypes).
Because the DNA test cannot distinguish between a Type A versus a Type "AB"
cat, the non-b allele is reported as "N". The cat could have a Type A or Type AB
serotype.
28
Data courtesy of Dr Diane Addie and FabCats
29
DNA Laboratories can be found online via search engines. Many cat clubs operate a
discount scheme with accredited laboratories.
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Test
Blood group status
Result
Once a breeder knows the blood type of all their breeding cats they need to
understand that Neonatal Isoerythrolysis represents the most serious problem to
any A offspring of a B mother. This risk increases with every mating that may
result in A group kittens. The effect of the antibodies is accumulative.
“Queens with blood group A have less anti-type B, antibody than blood group B
queens have anti-group A, antibody, so the type B kittens of this mating might
survive. However, if the mating is repeated, the queen will build up anti-B
antibody and eventually one fourth of her kittens could die of Neonatal
Isoerythrolysis.”
Figure 5 shows how to calculate any risk (note we have ignored AB as it is so rare
and is covered by Dr Diane Addie in full)
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A, B
Ab Stud B
, or B Ab or Ab
B or A
Stud Queen Stud
Queen
Quee
It is the antibodies from B group dams that represent the greatest risk to A kittens
Figure 5
Figure 5 shows that a B group stud can be mated to any queen and an A queen
can be mated to any stud. Breeders need to manage the matings and or kittening
of their B group queens. Records of any fading kittens in Ab to Ab matings should
be kept.
5.2.1. Recommendations
30
These are available to Veterinary practices from the Oxford Cat Clinic.
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5.3. Brachycephaly
5.4. Weight
The British can be a lazy cat, and may be prone to weight gain. Overweight stud
cats can find the mating act difficult. Overweight queens may have difficulty in
getting pregnant and giving birth.
31
Wikipedia
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Table 2
Viral Bacterial Parasites
Feline Calicivirus (FCV) Feline Chlamydia Fleas
Feline Panleukopenia Campylobacter Ear Mites
(Distemper)
Feline Leukemia Virus Bordetella Bronchiseptica Hookworms,
(FeLV) Ringworms and
Tapeworms
Feline Rhinotracheitis Toxoplasma Coccidia
(Feline Herpesvirus)
Feline Immunodeficiency Streptococcus Tri Trichomonas
Virus
Corona (FCoV) and Staphylococcus Giardia
Feline Infectious
Peritonitis
Ringworm
Key: Diseases in Green have vaccines available. Blue, have treatments available.
Red, these diseases have no vaccines or treatments in common use.
Our best recommendation is that all our breeders find a good Vet, one that is
prepared to be a partner on our journey to produce happy healthy British
Shorthairs. Breeding is teamwork and the support and care of a good Vet is
invaluable.
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In order to understand the genes for coat pattern at play in the British self we
must first understand the Agouti (A) gene - the natural “wild” gene that is the
basis of the tabby cat. The base agouti pattern is bands of black on a yellow or
orange background. This dominant, wild-type (A) causes the agouti shift
phenomenon which causes the hairs to be black pigmented at the tips and orange
pigmented at the roots this is overlaid with one of the tabby patterns.
There are three tabby patterning genes, which give rise to the patterns that we
know: Mackerel (Mc) with the basic stripes, Classic (mc) a mutation of (Mc)
which blotches the pattern, Ticked (T), a form that removes most of the stripe
pattern, Spotted (Sp) a gene that breaks the pattern into spots.
Working with the tabby patterning genes we have the Wide-banding (Wb) gene.
This little understood, and much argued over, gene has the effect of pushing the
darker, pattern colour in the cat up away from the hair base towards the tip,
turning the normal tabby patterns into a Shaded or Tipped cat.
Now that we understand the basis for the ‘original’ pattern of the BSH ancestor,
32
we must now accept that genes can mutate overtime.
When the dominant Agouti gene (A) mutated into the weaker Non-Agouti gene
(a)33 what actually happened was that the gene became faulty and no longer
works properly, so that the little genetic trick known as the ‘agouti shift’ 34 simply
doesn’t happen, or at least not properly in (aa) cats.
In the absence of the ‘agouti shift’ the basic tabby cat turns into a self black
because the whole hair shaft is allowed to fill with eumelanin 35 meaning that the
whole animal appears black, although often in certain light the underlying tabby
pattern may still just be discernible, especially in kittens. Other genes work to
change this black pigment to other colours.
Something important for breeders of the self coloured British to bear in mind is
that whilst not all cats are Agouti, they are all Tabbies, the loss of the Dominant A
gene does not affect the base tabby pattern. Thus those underlying patterns we
see in kittens can be stripes, spots or indeed ticked.
Another point to note at this early stage is that the (A) to (a) mutation has little
effect on the (O) allele 36 This explains why it is almost always possible to see the
tabby markings on red and cream non agouti cats.
Our beautiful British Self coats are created in the embryo. Very early on, all the
particles responsible for the colour of the coat flow into the hair shafts. This is a
very complex process, not just technically, but also impacted by genes and
polygenes that are not yet fully understood. What seems to be important to the
‘self’ is the speed of this process, the faster the colour pigments pour into the
shaft the more even the hair colour becomes.
32
Mutations are changes in the DNA sequence of a cell's genome
33
Also known as the Hypermelanistic gene
34
Far too complicated for this section
35
Black pigment
36
The gene responsible for Red and Cream
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
In the British Self we are looking for a coat with hairs that do not fade out as they
get closer to the skin. This can be helped by selective breeding of individuals that
have the most solid coats. What we are probably playing with are those
polygenes that control the depositing of the melanin in the hair shaft and the
length of time that the process takes.
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Introduction
It is estimated that the White Self represents less than half a percent of total
kitten registrations.
It will help at the outset if we understand that White is not a colour in the sense
that Black is a colour – we need to think of a white cat as one that has not got a
colour and thus appears white.
Keeping this very simple, what happens is, that as the embryo forms, the W gene
interrupts some of the normal cellular activity, and those cells, for pigment or
colour, that multiply and spread over the embryo, effectively get stopped in their
tracks, so the skin of the newborn kitten has no pigment and the kitten is white.
A completely white cat could carry the genes for red/black, agouti/non-agouti,
dilute/non-dilute, etc, but despite any hidden genetic codes for coat colour and
pattern the cat may have, the cat will always be visually white.
Pure white cats often have a small patch of colour on the head at birth. This spot
is evidence that the process of distributing cells for melanin actually started, but
was then halted by the W gene. This is why this spot may reveal what colour the
kitten carries. These patches usually disappear as the cat matures.
37
Epistatic
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Eyes
As the white coat may be masking many coat colours or patterns including the
40
Himalayan gene the source of the eye colour in the white cat may not always
be known. A blue eye colour, for example, may be as a result of have de-
pigmentation of the iris of one or both eyes, or may be as a result of the
Himalayan gene.
Due to the lack of pigmentation in the skin, white cats are more sensitive to
sunlight and are more likely to affected by sunburn and develop skin cancer.
Deafness
It would be irresponsible not to mention the deafness that is associated with the
gene for White (W).
38
See White Spotting Gene and Section on Bi-Colour British
40
See Appendix 1 and Section on Colourpointed.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
There is much data available giving statistics, the following data paints a picture.
During our research for this section we came across several papers each arguing
the case both for and against the existence of a gene for deafness in white cats.
In the absence of a definitive position we must settle for the generally accepted
position which is;
• There may be a gene for deafness which may be found one day.
• The deafness may be caused by the W gene halting the normal migration
of pigment cells that causes both the eye to be blue and the degeneration
of the canal of Corti, (the part of the cochlear duct that leads to deafness)
• Studies have shown that it is the same stem cells that cause the lack of
pigment in the blue eye and the absence of a cell layer in the inner ear
that is found in deaf cats.
• White cats irrespective of eye colour may be deaf, but it is more common
in blue-eyed white cats than in yellow-eyed.
• The deafness is caused by the W, gene working with polygenes.
• Odd-eyed cats are often deaf on the ear which is situated on the same
side as the blue eye.
What all this tells us is, that we breeders need to be very careful when dealing
with the W gene, and studies have proved that there is a considerable fall in the
incidence of deafness in pure bred cats, so it is clearly possible to breed a healthy
hearing White cat.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
41
One breeder interviewed suggested that using cats with perfect V shaped blazes
seemed to produce a higher number of kittens with single eye colour.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Introduction
Black aa B- D- oY/oo ww
Blue aa B- dd oY/oo ww
B- = BB or Bb or Bbl
To arrive at the coat that defines the superb cat that is the British Black we must
go back to the Non-agouti (a) gene the recessive gene mutation that turns the
basic tabby cat into a self black. The pigment granules in each hair of the cat's
coat contain Eumelanin 43 which is black. Black melanin or eumelanin granules are
thought to be oval in shape and absorb almost all light.
The above genetic explanation of colour does not explain the superb texture of
the British Black and Blue coat. The plush, dense coat sets the standard for all the
other British Breeds’.
42
Data provided by the GCCF using 2009 registration data
43
Eumelanin is, in the absence of colour modifier genes, black
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The blue colour of a British Shorthair can range from a gentle pale grey to a dark
charcoal. The colour should be solid and even throughout the body. Any ghost
tabby rings on the tail are forgiven only in immature cats.
Eyes
Breeders seek to achieve dramatic effects with stunning eye colour of copper,
orange or deep gold. Eyes should be large and round, giving rise to a sweet, open
expression.
Nose leather follows coat colour. Black for the Black British and Blue for the Blue.
The sight of the British Black at the top of his or her game on the show bench
always causes a stir and high awards are usually met with great approval, the
black needs to be jet black, with no paling to the roots, the coat must be dense
and plush. Eye colour needs to be as good as it can be. This fabulous coat will
show any dust so good grooming is vital. Kittens may show some rustiness, but
this will not be expected in the adult cat.
There should be no tabby markings in either cat on the adult show bench.
Silver Tipping
One of the topics of discussion during review of this breeding policy and the
seminar held in April 2011 was the Silver Tipping seen on the coats of some
British Blues.
Despite some detailed research we have as yet been unable to determine the
genetic nature of the ‘tipping’ seen on some blue coats or to ascertain as to when
the British Blue coat first started to show this tipping.
If we go back to our introduction we can read that after both wars many other
breeds of cats known to have this Silver Tipping were used to re-establish the
British.
There are some renowned Blue breeders who are actively breeding to keep their
lines free of this tipping, whereas others feel that it is a not unattractive price to
pay for an otherwise good for type cat with a pale blue coat.
39
Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Introduction
The chocolate British Shorthair is a huge favourite and very desirable companion
animal. The chocolate colour mutation was first seen in the Siamese breeds,
where it was thought to be ‘just a poor or strange’ Seal 44 Records for the
Chocolate Siamese appear in the 1880’s and they were first registered by the
GCCF in 1950.
It was not until the 1980’s that breeders started to bring the colourpointed
pattern into the British Shorthair and with it the Chocolate and Lilac colours.
The Chocolate Self did not gain championship status at GCCF shows until 2009.
The chocolate colour is a true mutation of the gene for Black and can be found by
gene tests. In chocolate cats, the round or oval black pigment granules
(eumelanin) which absorb almost all of the light, are a bit flattened. This means
more light can be reflected and the coat looks paler. This gene for Chocolate is
recessive.
44
Seal is the colour description for the Black Colourpointed
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The breed standard for Chocolate calls for any shade of rich chocolate in a sound
coat with chocolate or pink nose leather and paw pads.
The Lilac cat should be a Frosty grey with a distinctive pinkish tone, giving an
overall lilac appearance.
The Chocolate and Lilac series are always popular on the show-bench. Judges are
looking for chocolate coats that are the mid-warm tone of a chocolate Labrador,
which are also sound to the roots, something that breeders find hard to achieve.
• Experienced breeders of the chocolate colour find that using black self cats
within their breeding programs helps to produce chocolate or lilac coats
with more desirable tones to the colour.
• Novice breeders should appreciate that this colour is relatively new to the
breed and still needs work to get sound coats.
• Rufous genes appear to be reducing barring and making colours warmer in
other breeds. Selective breeding should help to see whether this can also
be true in chocolates and lilacs.
• Some breeders advise breeding chocolate cats not carrying the
colourpointed gene
• Some trial matings are showing that chocolate coats in cats carrying
cinnamon are lighter in shade.
• Experienced breeders are experimenting with the effect of using cats
known to have an underlying non agouti ticked base pattern to see if there
is an improvement in the quality of the coats.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Cinnamon is the third true cat colour. Just as the Chocolate ‘b’ gene is flatter and
longer than the oval shaped Black ‘B’ the Cinnamon (b¹) granules are even longer
and thinner than those in the Chocolate, causing the hairs to be a lighter and
warmer brown colour.
The dilute gene causes the Cinnamon colour to fade to Fawn. Fawn is similar in
tone to lilac but is paler, perhaps more beige in colour.
And whilst the above table shows that the Cinnamon gene for colour looks an
awful lot like the black gene for colour on paper, the truth is that little b¹ gene
has travelled a long way into the British and has potentially bought some
interesting companions.
In order to explain, we need to go back into the history, firstly of the Cinnamon
colour, and secondly how it has arrived onto the British show bench today.
Essentially the Cinnamon colour was first identified in the Sorrel Abyssinian. This
cat had been labeled as ‘red’ until the late 1960’s when tests proved that the
genotype of the Red Abyssinian was in fact that of a Cinnamon Tabby. The genes
responsible for the Abyssinian coat pattern are dominant to most other genes,
hiding recessives such as non-agouti, other colours and other tabby patterns.
It was a passion for the richness of the Cinnamon colour that tempted a group of
breeders to seek to introduce the colour into the British early in the 1990’s. Much
discussion over ‘where to go’ to outcross ensued, with the Abyssinian and the
Oriental being the main contenders. Eventually it was decided to go out to the
46
Oriental .
45
Interesting reading includes - The Abyssinian Cat, by Vicky Hakin Smith
46
Malsome British Shorthairs have some background information.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The main reason for choosing the Oriental over the Abyssinian was the fact that
the Abyssinian ‘genotype’ included the dominant ticked agouti gene and the
Oriental Cinnamon Self is a non agouti cat.
47
The Oriental section had originally out-crossed to the Abyssinian but selective
breeding of the recessive ‘a’ gene produced a cat with all the magnificent colour
48
of the Abyssinian in a non ticked form.
Those of you still paying attention will note that, although the agouti gene may
have bowed out leaving his recessive cousin in charge – the underlying non-
agouti tabby pattern passed through the subsequent generations may well be
‘ticked’ 49
47
More information can be found via the ‘The Oriental Cinnamon and Fawn Breeders Group’
48
Just out of interest see both cats at shows – note the effect the ticked agouti gene has on the
colour.
49
The effect the underlying (non-agouti) pattern may have on the British Coat is a ‘hot topic’ in the
British section.
50
Photo of Mewzishun Symphonia taken by Alan Richardson in 1996. This cat shows the Oriental
Cinnamon type as used by the British/Cinnamon project.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The breed standard for Cinnamon calls for a warm cinnamon brown and for Fawn
a warm rosy mushroom.
This wonderful,
short legged,
cobby girl with
neat ears shows
just how much
has been
achieved by
some of our
ground breaking
breeders.
• Novice breeders should appreciate that this colour is relatively new to the
breed and should seek to understand fully the pedigree of any breeding
animal.
• Getting ‘type’ right is imperative. This cat’s ancestor has, large tall ears
and a long nose – selection will be key as this breed progresses.
• Experienced breeders are removing the colourpointed gene from the
phenotype of some lines – to ‘fix’ the Cinnamon and Fawn Self.
• As this breed moves into Championship Status, it is likely to be adopted by
a greater number of breeders. Remember, to protect the colour, that
warm burnt sienna, cinnamon colour, was after all what the last 20 years
was all about.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Those budding geneticists amongst us, will have remembered that the (A) to (a)
51
(agouti to non-agouti) mutation has little effect on the (O) allele This explains
why it is almost always possible to see the tabby markings on red and cream non
agouti cats. So it is not surprising that breeders are still working hard to get the
52
Red Self through the Preliminary Status stage. Whereas the dilute form, the
British Cream is a very popular cat both on and off the show-bench.
Because the genetics of the red and cream series start to get a bit more
complicated it is not surprising that when they popped up in Tortoiseshell litters in
the 1800’s breeders were often uncertain how to actually breed the cream self.
It was not until the 1920’s that breeding programs became established.
53
Having explained eumelanin we now need to get to grips with the only other
feline pigment – phaeomelanin the pigment for red. Essentially what the ‘O’ gene
54
does is get rid of any eumelanistic pigment from the hair shaft.
We must also remember that in the dominance stakes black and red are equal.
And whilst we practice pronouncing ‘phaeomelanin’ we must also take a step back
to our biology lessons and recall the rules that determine whether we are male or
female or more importantly whether our cats are male or female.
Females, have a pair of chromosomes that appear identical and are X shaped.
Therefore, we refer to females as:
XX
Males also have a pair of chromosomes, but one of the males chromosomes is
smaller and looks like a Y or an incomplete X with the right lower ‘leg’ missing.
XY
55
The reason this is important information is because the genes for colour are
carried on the X chromosome and once the penny drops that girls have two XX
places to carry or inherit colour/pigment and boys only have one to carry or X
inherit colour/pigment we can begin to understand why we can have tortie’ girls
but not boys. The ‘O’ gene sits on the X chromosome and we call it a ‘sex
linked’ gene
51
The gene responsible for Red and Cream
52
Preliminary status breeds compete in Assessment classes
53
The pigment for black
54
Black, Chocolate, Cinnamon
55
Important to note this refers to the genes for colour – NOT the genes for pattern.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
XX Black on both X
Chromosomes
Black Female
XX Red on both X
chromosomes
Red Female
Figure 6
• If any cat carries two copies of the dilute gene then cats will be blue,
cream or dilute tortie.
• We can substitute chocolate or cinnamon for black in Figure 6and the rules
still apply.
For those of you still with us, we will just do a quick reminder of how gender is
inherited and how that affects this ‘O’ gene.
In order to be a boy the male cat has to have one copy of the Y chromosome,
which he can only get from his father. This is balanced by an X from his mother.
Therefore, an orange/red male always gets his gene for colour from his mother.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The good news is that Breeders are making progress and this colour group -
together with the non-silver series of tabby – is one of the few British Shorthair
varieties that had an increase in registrations in 2009.
Leather should be brick red for the red and pink for the cream.
56
Appendix 3
57
Ticked Tabby
Ticked Tabby (All colours) Non Silver (Preliminary
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
A beautiful
youngster
showing an
underlying non
agouti spotty
pattern
A lovely
rich red girl
– so rarely
seen
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Just when things were starting to get simple, we learn that the tortoiseshell
pattern is not genetically a pattern – it is actually something that happens when a
cat is born with two colours showing.
If neither parent has the gene for ‘red’ then the colour of the female kitten will be
58
determined according to the rules for colour inheritance
59
If the father is red or cream, then his daughter can only receive that colour
from him and so she will have one X chromosome with the red colour carried. If
60
What happens next is complicated, but important to all tortie breeders.
Shortly after the mating, the original cells start dividing like mad. Quite quickly, the process
determines that there is altogether too much ‘stuff’ coming from both those XX’s – and so the
process shuts down or inactivates one of the X chromosomes in each cell.
This shutdown is random and happens early in the embryo. But meanwhile some
of the cells for colour from each of the two XX’s have ‘made it through’ we can
61
see this visibly in skin cells as they produce either red or black pigment and as
the skin cells multiply they form patches of varying sizes and shapes. It is
thought that the red and black cells multiply equally and actually compete with
each other. It is the resulting ‘mosaic’ of colour that determines the distribution of
red and black.
What this means is, that to all intents and purposes, the arrival in a litter of a
beautifully marked tortie is largely due to good fortune.
58
Robinson’s genetics have a table showing the likelihood of colour
inheritance (thankfully)
59
Note: These rules also apply to bi-colours, colourpointed or tabbies with
red.
60
And not completely understood as yet
61
Or one of the black/blue mutations.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
A Black/Red
(dominant
tortie) kitten,
showing
beautifully
mingled colour
distribution.
Despite the
‘naughty tortie’
reputation, this
delightful cat
has the 4th
highest
registration
numbers.
The tortoiseshell is a mixture of the base colour with red or cream as appropriate,
evenly intermingled, with both colours clearly defined over the whole animal but
without any obvious patches of any colour. A small blaze on the face is
permissible.
Judges will fault, uneven balance of colour, unbroken legs and paws, tabby
markings and brindling.
The tortie is shown in the following combinations: Black, Chocolate and Cinnamon
with Red and Blue, Lilac and Fawn with Cream.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Just for the record we will settle the argument as to whether this section is really ‘one for
the girls’ or not. According to Robinson’s Genetics one in every 3,000 male cats born is a
‘tortie’. Those of you with enough time to invest in understanding the reasons for something
so rare will find the answers in Robinson’s under the section ORANGE.
62
These have not yet been identified, but breeders are enjoying selecting for them nevertheless
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
7. Non-Self/Non Tabby
7.1. Bi-colour
The Bi-Colour represents the 5th most popular of British Shorthairs registered,
making up almost 6% of the total registered.
Those of you who think that the sentence above suggests that all we have do is
mate the self white with any of the self colours and hey presto! we will get Bi-
colours just haven’t been paying attention. The gene for White is W, which always
delivers a completely white cat.
The gene responsible for our beautiful Bi-Colour is known as the ‘S’ gene.
The S gene, known as the Spotting gene or the Piebald gene is a Semi-dominant
63
Gene and produces variable results. A cat may have almost no visible white
spots or may be completely white, with almost any variation in between.
In the British it is the S gene that is needed to produce both the Bi-Colour and
. 65
Van Pattern 64 The S/s gene is extremely variable, the following diagram may
help.
Figure 7
63
Producing an intermediate phenotype in the heterozygous condition
64
52
Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The British Bi-colour cat falls very neatly into the mid-point and although we
cannot yet test for this gene we can be fairly certain that our Bi-colours may fall
into either genetic ‘camp’
This
wonderful
picture just
may help
with all the
genetic
information
in this
section.
Unlike the tortie, the British breeder is able to influence the Bi-colour through
selective breeding. This is because by breeding only the cats with a lot of white
on them you can increase your chances of getting cats with more white. This is
possible because every gene that we know about has all those little polygenes
hanging around, it is these polygenes that help to influence the amount of white
inherited and it is widely accepted that selective breeding can make good use of
polygenes.
After mating, and the egg is fertilized, cells change 66 and multiply, and begin to
form what will be the ‘finished’ kitten. During this embryonic stage some cells
called melanoblasts start to move, or migrate from the top (neural crest) of what
will be the kitten, down towards the bottom of the embryo.
66
Differentiation
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
These cells, become the cells that produce melanic pigment (colour).
If these cells do not contain either the white gene (W) 67 or the spotting gene (S)
they will cover the whole embryo and the result will be a fully coloured kitten.
If the cells contain one or two copies of the spotting gene, (SS or Ss) a defect
occurs and the migration of cells fails to complete and the parts of the kitten
furthest from the neural crest does not get a fair share of ‘colour cells’ and the
result is white patches – this is why so many breeders refer to the white pattern
as ‘falling to the bottom’ when in fact it is actually the colour that fails to fall!!!
Eyes
Colour in line with the breed standard for the Self British. Incorrect rim colour or
flecks will be faulted.
67
It seems that the W gene also interferes with melanic migration – almost as soon as it starts
around the neural crest – it is thought that this melanic ‘defect’ is related to the deafness that is
associated with the W gene.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
• The Bi-Colour gene does not have anything to do with the White Masking
gene (W)
• Mating your Bi-Colour to a white cat will not add any more white to your
Bi-colour program 68
• Using Bi-Coloured cats in a breeding program does not cause lockets
• It is clear from the high expectations from the Breed Standard that the Bi-
Colour is not a cat for the novice breeder. Getting the pattern and self
colour right for the show bench calls for hard work and dedication. As we
have read, the show quality coat can be selectively bred, and using a dam
or sire with poor markings can set back a breeding program.
• Working with Bi-Colours teaches breeders a huge amount about just how
diverse cat genes can be – those at the top of their game, must have
patience in abundance and a sense of humour.
68
Geneticists are confident that the W gene sits at a separate locus from the S gene. Experienced
Breeders (anecdotally) report using White (W-) cats with Bi colour (S-) does result in high grade Bi-
colour – see section on Van Pattern.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The Van Patterned British Shorthair is a rarely seen attractive cat, that can be
bred in all the British self colours with white. The phenotype is a white cat with
colour confined to the head and tail, although up to three small patches of colour
on the body are acceptable.
It is generally accepted that selective breeding can exert influence over the tricky
69
Spotting gene and his polygenic mates , which is why Breeders of the Van
Pattern British have sought to get this (probably) dominant variety recognised in
its own right.
Because both the British White (W) is an established breed, as is the Bi-Colour
(SS or Ss) it seems sensible to expand upon the genetics of the Van at this point
as (at least, in phenotype) the Van seems to fall part way between the two
genotypes.
Some breeders of both White and Bi-Colour have long assumed that the Spotting
gene is just a different form 70 of the White gene. This is understandable because
in practice the two genes work in such a similar way.
Much work has been done to prove that there are in fact two separate genes in
two separate places 71 The following figure seeks demonstrate evidence for there
being two separate genes in two places.
69
Birman Breeders have done wonders with Gloving!
70
An allele
71
Loci
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
1 2 3
Self White Van-Patterned Self White
Sire Dam Male Kitten
aa Ww ss B- aa ww SS B- aa Ww Ss B-
D- dd Dd
Figure 8
3 4 5
Self White Blue Self Blue Self
Male Kitten Female female Kitten
aa Ww Ss B- aa ww ss B- aa ww ss B-
Dd dd dd
Figure 9
In Figure 9 we mate a non agouti, heterozygous Self White male (1) masking
black to a non-agouti, homozygous blue Van Patterned female (2) and we get a
Self White Kitten (3)
If we look at kitten (3) we can see that he has inherited his W gene from his dad
and his w gene from his mother. Because it sits at different locus from the W
gene he gets an S gene from his mother and the s from his father. He gets the B,
black gene from both his parents so he is genetically black. He gets the dilute
gene from his mother. When he grows up and we mate him to cat (4) they have
a blue self kitten, who gets her father’s w gene paired with her mother’s w gene,
she gets an s, a B and a d from each of them and so we get a non-agouti, non
white, non spotted, dilute black cat, in phenotype and genotype a Blue Self.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Eyes
The breed standard is looking for a cat with one self colour with white. He patches
should be clear and well defined with no tabby markings. A predominantly white
cat with small patches of colour on the head, but not below the level of the eye or
beyond the base of the rear of the ears. The colour on the head should preferably
be separated by a vertical white blaze. The tail should be fully coloured. For
perfection, no colour on the body or legs but up to three small thumb prints of
colour on the body would not fault an otherwise superb show cat.
• Until the Van Pattern is established and can breed true, and we can be
certain that that high white gene for Van (probably homozygous for
spotting SS) does not start to bring a hearing problem into the Bi-Colour,
it is recommended that only experienced breeders, with experience of both
White Self and Bi-Colour breeding, undertake a Van breeding program.
• If novice breeders wish to join a breeding program they should be
mentored.
• Breeders should keep detailed records of all matings and outcomes, both
positive and negative. Keeping good records of ALL kittens born to a
breeding program are valuable aid in informing decisions.
• Breeders finding evidence of best practice that would suggest a review of
the registration policy should gather evidence and approach their breed
club. 72
72
Anecdotal evidence suggests that mating a self white to tri-colour produces ‘good Van’s.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
7.3. Tortie and White and Van Pattern Tortie and White
The Tortie and White is another very popular cat on the show bench causing
heads to turn. The tortie and white can be bred in any of the usual tortie colours.
Called the Tri-colour, by breeders, genetically, the tortie and white is ‘just what it
says on the box’ but our old friend the white spotting gene SS or Ss has an
interesting effect on the tortie pattern.
Remember the ‘migration’ that the cells for pigment make 73, with the red and
black competing for space – resulting in a mingled cat? And then add the ‘lazy’
factor that the white spotting gene SS or Ss brings 74, causing the migrating cells
for colour to stop, before they reach the bottom of the cat?
It seems that the spotting gene causes fewer pigment producing cells. Therefore,
less competition between them as they migrate into position. Cells reach an area
and then multiply where they are to create a patch of colour.
73
See Tortie Section
74
See Bi-Colour Section
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Symmetry of design is desirable, with colour on the top of the head, the ears,
cheeks, back, tail and flanks.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
8. Smoke
Rarely seen on the British Show-bench, the Smoke is a ‘self’ non-agouti (aa) cat
with the addition of the Dominant Melanin Inhibitor (I) gene and whilst we may
not often see a fine example of the British Smoke, the impact of the Inhibitor
gene is all around us and it would be wise if we get to grips with it sooner rather
than later.
Some Facts
• The I gene is called the Inhibitor gene because it ‘inhibits’ the feeding of
melanin or colour into the growing hair 75.
• It seems to have more effect on the part of the hair shaft with the least
pigment, which is the part of the hair nearest the body.
• The gene seems to have a stronger effect on the production of the
pigment phaeomelanin 76 more than the pigment eumelanin 77.
• It has a ‘variable’ effect in aa cats (that do not have the agouti gene)
What this results in is a self coloured cat with a silver undercoat and breeders are
looking for a cat with as much contrast as possible between a good strong self
colour and a shimmering clear silver/white undercoat.
78
The problem is that the Silver (I) gene is extremely unpredictable in the
absence of the agouti (A) gene and breeders rarely get what they would like.
The problems are many and varied, the inhibitor gene may actually not inhibit
very much at all and then the little bit it does inhibit might not be a glorious silver
white colour and instead may be a murky dark grey colour that blends into the
‘self’ coloured hair shaft. To add insult to injury, the (I) gene may also cause the
‘self’ colour to be washed out or rusty. These Cats with little contrast between
under coat and top coat colours may look self coloured. And if that isn’t enough,
the smoke cat would love to be a tabby and ghost markings abound. At this point
we can blame all the various, nameless, polygenes yet to be identified, but the
fact remains, breeding a good smoke is tough going.
This is possibly a sensible place to ‘discuss’ the registration policy regarding cats
with ‘Silver’ in the pedigree. To experienced breeders of cats incorporating the
Silver gene, it can often be irritating to receive GCCF paperwork over-stamped
with ‘Silver in Pedigree’. And yes, as a BAC we understand that the difference
between a good Brown Spotty and a Black Silver Spotty is very evident even to
novices. But it quite simply isn’t that simple. In the wrong hands, a poor smoke
may well be bred as a self. The over stamping of Silver is intended to be
informative. To those who know what they are doing, it should be of no
consequence whether the beautiful British Blue they are breeding from has an
ancestor who was a Silver, nor should it matter if the registration paperwork
makes reference to that ancestry. Colourpointed Breeders are grateful for this
75
Not like the S gene that affects melanin getting to skin cells
76
Orange
77
Black
78
Inhibitor
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Dilute B aa B-/bb-/bb¹ C- dd I
oY/oo
Dominant O aa C- D- I OY/OO
Dilute O aa C- dd I OY/OO
Dom/Tortie aa B-/bb-/bb¹ C- D- I Oo
Dilute/Tortie aa B-/bb-/bb¹ C- dd I Oo
Eyes
The top coat should be one of the recognised self or tortie colours, over a silver
undercoat. When the cat is still it will look solid-colored, but when the coat is
parted the silver should flicker through creating an ephemeral effect.
• The British Smoke can be a lovely sight on the show bench. Often
produced as a ‘by-product’ of a silver program, this cat can be overlooked.
Any show quality variety needs work.
• There are some superb examples of the Smoke cat to be found on the
Persian show benches, with breeders devoting entire breeding programs to
this variety. Never forget, we all have much to learn from experienced cat
breeders, irrespective of breed.
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In the section on The British Self, we said that in order to understand the genes
79
for the ‘Self’, we must first understand the Agouti (A) gene. We then went on
to deliver a couple of paragraphs about ‘agouti’ and ‘tabby’, throwing in the odd
‘Mc’s’ for good measure. In doing so we made it all sound oh! so simple. For
those of you wanting an easy life turn back now; because, despite our best
efforts, it’s going to get a teensy bit complicated now.
Part of the problem is that we live in very exciting times for the discovery of
genetics and our understanding is being constantly challenged and as breeders
we are necessarily in the business of experimentation and boundary pushing. Far
better then, that we are as informed as much as possible about the ‘genotype’
that makes up our beautiful cats.
Hopefully, by now, we know that a self cat is non agouti (aa) but rather
confusingly a self cat is still a ‘tabby’, this is because all cats are tabbies. And this
is one of those ‘light bulb’ moments for most cat breeders, when the
understanding dawns, that there are TWO genes that make a cat a TRUE TABBY
as we know it.
So all cats have the ‘T’ gene for pattern, but only tabbies have the A gene for
Agouti. It is almost as if the A gene turns on a magic switch that lets us see the
pattern. Before we go any further, it would be sensible to agree on the symbols
we shall use for this section.
The Agouti gene (A) is nice and easy, but the (T) (tabby) gene is not quite so
simple and often causes confusion. This is because of the relationship between
the Ticked, Classic/Mackerel and Spotted Tabby genes. Recent studies 80 have
suggested that there are in fact three separate genes at different loci 81
Therefore, we feel it will be easier if we discard the use of T except for the Ticked
Tabby and use Mc and Sp 82 instead 83.
79
The natural “wild” gene that is the basis of the tabby cat.
80
Genetics Society of America, 2009.
81
A locus is the exact position on the chromosome where the gene is located
82
See Robinsons Genetics section on Tabby and Non-Agouti.
83
This convention has also been adopted by the GCCF and the Asian BAC.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
19 Pairs of Chromsomes
Locus for Locus for Locus for
Locus for Locus for Locus for
Spotted Inhibitor Masking
Agouti Genes Tabby Genes Modifier Genes Colour Genes
Genes Genes
2 alleles 2 alleles 2 alleles 2 alleles
2 alleles 2 alleles
Figure 11 84
If we look at Figure 11, we should remember that all the genes along the
chromosome start life at the moment of conception.
This will really help us understand that the genes for Tabby Pattern and the
Genes for Agouti colouring are working at the same time and affecting each
other.
85
As the hair grows in the hair follicle eumelanin
is produced by cells and dropped into the hair
shaft. When the Agouti gene is present it causes a
protein to join the process and as this protein
builds up in the cell it causes the production of
eumelanin to falter and suddenly the cell switches
86
to produce phaeomelanin , this process is
known as the Agouti shift.
84
There are many other Genes of Course
85
Black Pigment
86
Red pigment
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Whilst the Agouti gene is busy creating these banded hairs, if the cat has also
inherited the tabby genes Mc or mc, these genes set up in competition with the
Agouti gene, stealing patches of skin and follicles and reducing the amount of
Agouti protein produced in these patches. The shift from black to red on the hair
shaft does not happen. Instead these patches of hair show as groups or patterns
of black hair beautifully offset by the Agouti coloured, ‘ground colour’ hairs. These
patterns are of course the well known Tabby patterns.
According to Robinson the Agouti shift, switches pigment between eumelanin and
phaeomelanin. However, there is an argument that the pigmented hair shaft does
not shift from eumelanin to phaeomelanin, but instead the Agouti protein cause
the hair granules to thin and shred, it is this weakening in the granules that
causes the banded hair colour to fade to the yellow/orange colour alternately.
The UCLA Davis website does refer to the Agouti shift but does not confirm the
mechanism for the colour change.
Finally on this point, current opinion suggests that the colour of the Agouti band
can be influenced by ‘Rufus’ type polygenes These Rufus factors have not been
identified but breeders have been able to select for them to reproduce warmer,
richer background colors in the Tabby. Selective breeding in the Brown Tabby is
producing a rich redder background colour.
87
We have used many sources and resources for our research; wherever and whenever opinion
differs we have fallen back on Robinson’s Genetics for Cat Breeders and Veterinarians as our base
reference. However, it must be noted that the latest edition was produced in 1999. Where there has
been compelling evidence that evidence has changed we have sought to validate opinion and have
found the UCLA Davis website invaluable.
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If you decided to play a game of ‘scissors, paper, stone’, against a Brown Ticked
Tabby, you would lose. This beautiful cat is, genetically, at the top of the
domesticated feline tree.
Whilst it is generally accepted that Felis Silvestris or Felis Lybica, both striped
cats are the original UK wildcat species, it is less well known that the first
domesticated or companion cats were probably ticked tabbies, imported as pets.
These early imports were crossed with the British Shorthair to create the cat
known today as the Abyssinian.
The ticked coat can be seen in the Somali and Singapura breeds as well as the
Abyssinian. At first glance the entire cat will appear to be one colour until one
gets closer and notices that in fact each individual hair has alternating bands of
colour giving rise to a flecked appearance.
The British Ticked Tabby can be bred in all self colours including tortie.
Ticking results when the agouti gene (A) causes the individual hairs to have
bands of light and dark pigmentation. As the hair starts to grow, the agouti gene
allows full pigmentation into the hair shaft, it then causes a sudden stop and the
hair shaft goes lighter, the process then turns back to full pigment for a bit
longer. As the hair reaches its full length this process known as the Agouti shift
stops. The result is a hair shaft that has dense colour at the tip, then a band of
yellow to orange, then a band of dense pigment, fading to yellow or orange at the
base.
What is special about the Ticked feline is that they carry a dominant gene for
Ticking (Ta) and when Ta is present it prevents other feline tabby patterns from
developing. However, like all tabbies, it may have tabby markings on the face and
agouti hairs on the body.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Figure 12
Figure 13
Eyes
Figure 14
The coat should be evenly ticked with two or three bands of colour extending well
down the hair shaft ending with the base colour at the roots. This ticking may be
heavier along the spine line and extend down the length of tail. The face, legs
and abdomen clearly show the base colour and on the head the ticking may be
more dense, or even solid in kittens, which may clear to an ‘M’ on the forehead.
Other facial markings are as per standard British Classic Tabby standard.
Apart from the underside of the abdomen, which should be spotted, the body
should be free of spots stripes or blotches.
The tail may be ringed with complete or broken rings or have a continuation of
the darker colour of the spine line, the tip of the tail being the same colour as the
markings. In adults the legs may or may not be barred.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
A lilac British Shorthair Stud cat imported from Sweden with a genotype of:
Was mated to
Choc’-
Spotty A- Sp bb C- Dd oo ww
Dam
88
Mrs Celia Leighton (Porteous)
89
Ticked tabby
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The only really surprising thing about this entirely natural production of a
delightful chocolate ticked tabby is that it hasn’t happened more often.
90
Chaos and Confusion genes may be at work see Tipped section.
91
Theoretical at present
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The Silver Ticked tabby cat is essentially the agouti cat the Ticked tabby with the
addition of the Inhibitor gene for Silver (I)
This colour can be seen in other breeds such as the Abyssinian and the Somali. In
fact all the silver Abyssinians bred in Britain today are descended from an
outcross to a British Silver Spotty in the 1960’s.
92
Eye Colour
Green or Hazel.
Remember that it is possible to breed silvers with every colour of ticking, and
some of the paler varieties in other breeds can be very pretty.
If we peep over our shoulders into the Asian Breeding Policy, it suggests that
mating non silver to non silver and silver to silver ticked produces the best
results 93
92
Photo by Milt Webb
93
Of course our gene pool is tiny as yet
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The basic ticked coat is quite common and can be seen in mice and rabbits. The
feline has developed a secondary system of pigmentation which creates dark
94
stripes flashed throughout the ticked coat .
These stripes or patches are caused by genes that ‘fight’ with the Agouti gene
(A), and effectively ‘stop’ the agouti protein from performing the agouti shift, in
some patches of the skin, thus preventing the eumelanin to phaeomelanin
95
shift . Hair shafts in these patches only have the darker colour. This leads to an
impression of one dark-coloured pattern superimposed upon another pattern.
The genes responsible for these dark patterns are either Mackerel (MC) which is
dominant over (mc).
On the neck and upper chest there should be unbroken necklaces, the more the
better. The edges of the ears to be the same colour as the markings with a
central patch of ground colour resembling a thumb-print.
94
Probably for camouflage
95
If indeed it is phaeomelanin!
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
9.2.1.Mackerel Tabby
The non-silver series Mackerel Tabby is not often seen on the British Show bench.
However, the Mackerel pattern is probably the ancestral striping pattern, which is
seen in the African wild cat (Felis libyca) and the European wild cat (Felis
silvestris).
Eyes
The SOP calls for a narrow unbroken central spine line, on either side of which is
a broken spine line from which the narrow lines which form the Mackerel Pattern
run vertically down the body; these lines should be as narrow and numerous as
possible. The tail rings, which should be as narrow and numerous as possible,
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
may be complete or broken with the tip of the tail being the same colour as the
markings.
Ground Colour, should be even, in all areas other than the lips and chin, which
may pale significantly but should not be white.
• Some Breeders believe that the Mackerel Tabby makes the best ‘base cat’
for Spotted Breeding Programs – believing that Spots are more numerous
when they break the Mackerel Pattern rather than the Classic Pattern
• Conversely – some breeders believe that the Mackerel pattern in the
British doesn’t actually exist – being instead poorly marked Spotties!
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
9.2.2.Classic Tabby.
The Classic tabby pattern without silver, is not seen very often on the show bench
which is a great shame, because, the combination of pattern and agouti colour
presents a super playground for breeders as we can see in other non- British
96
breeds on the show-bench
As we have learned throughout the sections, genes are never as simple as they
might seem, they really do work with groups of mates, either as yet unproven
genes or polygenes. Experienced breeders see something in a cat that they really
like and selectively ‘breed’ for that trait, in doing so they not only select the
‘obvious’ genes for Classic Tabby, they are also selecting 97 some ‘best friend
genes’ that enhance or subtly alter the phenotype.
This is probably the right time to present some of these hypothetical modifier
genes.
96
Have a wander around the show-benches and have a guess at the genes underneath
the coats.
97
Often without really knowing it
98
See section on Silver Tabbies.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The reason the Classic tabby appears so infrequently probably lies in it’s recessive
qualities, in the scissors, paper, stone game – the classic tabby loses out to the
hugely popular Silver series, the not so sought after Mackerel pattern, the
99
Spotty and of course the Ticked tabby.
The Classic, comes in all the colours of the self, Black, but just to confuse we call
it the Brown tabby, Blue, Chocolate, Lilac, Cinnamon, Fawn and of course Red
and Cream.
The Brown tabby in the wild has a sort of murky appearance, the agouti banding
is evident, but the pale bands are cool and greyish.
Generations of
breeding the
pedigree tabby has
produced a cat with
rich dark browns
for the eumelanin
pigment and warm
orangey tones for
the phaeomelanin.
1. Pattern – On the forehead there should be a letter 'M' giving the impression of
a frown. There should be an unbroken stripe running back from the outer
corner of the eye and narrow lines on the cheeks. On the neck and upper
chest there should be unbroken necklaces, the more the better. The edges of
the ears to be the same colour as the markings with a central patch of ground
colour resembling a thumb-print.
A series of lines runs from above the 'M' marking, over the top of the head
and extends to the shoulder markings. The shoulder markings form the
outline of a butterfly, when viewed from above. Both upper and lower 'wings'
should be clearly defined with the central areas broken by small areas of
ground colour. On the back there should be an unbroken line running down
the spine from the butterfly to the tail, and there should be a stripe on either
side of this, running parallel to it. These stripes should be separated from
each other by stripes of ground colour. On each flank there should be an
'oyster-shaped' patch which should be surrounded by one or more unbroken
rings.
The tail should have complete rings, as numerous as possible, with the tip of
the tail being the same colour as the markings. The legs should be barred
evenly with bracelets from the body markings to the toes which are spotted.
Markings on the hind legs extend (when adult) from the soles of the feet, up
99
See section on Spotted Tabby.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
the back of the leg, to the hock. The abdominal region should also be spotted.
Ground colour and markings should be evenly balanced and both sides of the
cat should have identical markings.
2. Markings – These are the dark pigmented areas, creating the ‘pattern’ these
markings must be solid in colour and sound to the roots they may be black or
various shades of brown. The depth of pigment should be the same from the
facial drawings through to the tip of the tail. And if that isn’t enough, breeders
must not allow any stray agouti hairs to drift into the markings, so no ticked
or banded hairs.
3. Ground Colour – This is the agouti ticked ground colour against which the
pattern is presented. The warmer and richer breeders can get this ground
colour the better.
4. Type – Of Course!
• There can be no doubt that breeding classic tabbies is a job for those with
patience, but as we can learn the benefits of exploiting all these polygenes
can be just so rewarding.
• As we learn more about cat genetics we can begin to see that sometimes
we cause a conflict for ourselves. Silver tabby breeders have over the
years worked really hard on getting that wonderful classic tabby pattern to
stand out clearly against a crisp silver undercoat. In doing so they have
bred out polygenes for Rufousing, because these polygenes probably
cause the silver to tarnish, in doing so any non-silver Mackerel or Classic
tabbies in the litter will be short of those lovely colour enhancing
Rufousing chaps and the results may be poorly coloured well patterned
kittens.
• Breeding programs for show quality non silver tabbies and silver tabbies
100
may benefit from some separation .
100
Experienced breeders will already have separate breeding programs.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The Spotted tabby is exactly what it sounds like and comes in the same colours
as the Classic tabby. This popular variety of British Shorthair offers breeders the
same challenges as the Classic tabby, with spots on!
Note: The genes for coat are as for Mackerel tabby, but spsp becomes Sp-
101
The pattern is often less obvious in the dilute cats
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The Silver Tabby and Spotted British Shorthair is the second most popular
variety, with more than 14% of registrations being this wonderful cat.
Dominant B A- B-/bb-
/bb¹
C- I- D- Mc- spsp oY/oo
tata
Dilute B A- B-/bb-
/bb¹
C- I- dd Mc- spsp oY/oo
tata
Dominant O A- C- I- D- Mc- spsp OY/OO
tata
Dilute O A- C- I- dd Mc- spsp OY/OO
tata
Dom/Tortie A- B-/bb-
/bb¹
C- I- D- Mc- spsp Oo
tata
Dilute/Tortie A- B-/bb-
/bb¹
C- I- dd Mc- spsp Oo
tata
A- = AA or Aa Mc- = McMc or Mcmc C- = CC or Ccs Sp- = spsp I=II or Ii
Eyes
We have all heard about the influence of polygenes on coat colour and pattern.
But polygenes also come together to help us with eye colour. The Black Silver
Tabby should have green or hazel eyes, whilst all other colour of silver tabby
should have copper orange or deep gold.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
The Silver series British Shorthair benefits from a breeder group that has
developed an extremely high standard. But popularity within a breed can
sometimes lead to inexperienced breeders producing kittens that ‘let the side
down’. In safe hands the Inhibitor gene can sparkle, but it also has a tendency to
tarnish, and there are three schools of thought as to why.
1. The Inhibitor gene does not work alone and has a bunch of polygene
friends, sometimes some of these friends do not pull their weight and the
result is a poor silver colour.
2. Breeding cats can be Homozygous for Silver (II) or Heterozygous (Ii) and
effectively carrying the genes for Brown. Some breeders believe that it is
the genes for non-silver that ‘show through’ and spoil the show.
3. When a non-silver parent is used in a silver breeding program, if that
non-silver parent carries a large dose of those lovely (if you want them)
Rufousing genes, then those Rufousing genes can cause the silver to
tarnish.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
This stunning cat is hugely popular. Genetically, breeders take the Silver Tabby,
either Mackerel or Classic, add the Spotty gene and the result is quite simply
stunning.
See the table in the Silver tabby section and substitute sp for Sp.
Because the Spotty gene Sp- sits at a separate location from Mc- or mcmc, unless
your breeding cats are Homozygous for Spots, litters can be full of both tabby
and self kittens.
Whilst we cannot be
certain if the breeder was
hoping for more spotted
kittens, the Queen will
love them all equally.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
In the 1970’s a Chinchilla (longhair) was mated to a British Silver Tabby 102. A
breeding program ensued, which included the British Blue and other cats from the
silver series, culminating in the development of the Black Tipped as we know it
today. This was recognised as a distinct British type in 1978.
The ‘Tippy’ can be bred in any of the British colours. However, it is the gorgeous
Black Tipped that is seen out most often.
At this point it
would be lovely if
we could just say
that genetically
the Black Tipped
is just a slightly
complicated
version of the
Silver Tabby,
throw in a pretty
picture and move
right on.
We can see that this looks an awful lot like the Silver Tabby series and yet the
Tippy looks so different. This ‘difference’ that breeds true has led breeders and
102
The pioneer of this program was Norman Winder of the Peerless prefix
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
It seems that this hypothetical Wideband gene can only work with agouti cats as
it is the agouti shift that causes the colour change on the hair shaft – that the
Wideband gene then forces into a wider band.
It would also appear that when teamed up with either the Inhibitor gene I-
(Silver) or the Rf- genes for rufousing the Wb- gene is Incompletely Dominant.
Firstly, the cat must be a hetrozygote (two different copies) for the allele in
question – and is a condition that expresses itself by showing a phenotype
(appearance) that lies somewhere in between the phenotype of the two parental
phenotypes.
So, for example, a Black Tipped cat with a good expression of the Wideband gene
(probably Homozygous for Wb) is mated to is mated to a Brown Tabby –
The offspring will be heterozygous for Wideband, Rufousing and Inhibitor. The
resulting litter may well include Silver Shaded cats with pretty orange noses!!!
With the following genotype A- B- Tata or tata Mc- Sp- or spsp Wbwb Ii Rfrf
Sometimes we can distinguish those cats that are homozygous from those that
are heterozygous by studying these little phenotypical clues.
We know that there are many genes that we cannot yet test for, but that we can
‘see’ such as the Inhibitor gene (I) – and it is widely accepted that these genes
are present and one day we will have tests for them.
However, The more ‘theoretical’ genes e.g. Rufousing require a greater act of
faith, and it is important that we as breeders understand the counter argument.
The argument for the presence of genes such as Wideband and Rufousing is the
exploitation of polygenic traits by careful selection.
The first breeders of the Black Tipped bred cats which had significantly heavier
tipping than we see today. By selecting parent cats with less tipping and clear
undercoats, subsequent generations of Black Tipped cats have evolved with wider
intermediary (yellow) agouti bands.
Therefore, we should keep our minds open to the question – how are breeders
achieving the wide range of tipped bands that we see?
• Are we dealing with complex polygenes that are happy to let us clear
undercoats and minimize the expression of black pigment to produce our
Tippies
• Is there an Incompletely Dominant Wideband gene?
• Or has the agouti gene mutated to allow better and better suppression of
pigment production?
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Eyes
Black Tipped - Green. In all other colours Copper, Orange or Deep Gold. The
skin outlining the eyes to be appropriate to the colour of the tipping.
This silver cat, most often in the form of the Black Tipped, can be shown in any of
the standard British colours including tortie.
The legs should show tipping which will decrease towards the paws, but which
should be as even as possible. The chin, stomach, chest and undertail should be
as pale as possible.
Nose Leather - Brick red preferred, outlined in colour appropriate to the colour
of the coat tipping.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Under the GCCF we breeders in the UK do not have to worry about the distinction
between Silver Shaded or Silver Shell (more lightly Tipped) instead Tippy
breeders can happily breed for personal preference, with the emphasis on
evenness of colour.
To what extent Tippy breeders can influence their preferences by using genes for
pattern, is still the subject of much debate - Let us remind ourselves what the
agouti shift does to affected hairs – remember that the hairs that make up the
darker ‘pattern’ remain a solid colour – but in Agouti cats there are areas of coat
wherein the hairs have bands of colour or tipping – and the Wideband gene can
only work on hair shafts that have these bands of colour –
If we look back at the table for genes for coat on page 81 we can see that the
Black Tipped cat like all other agouti (and non agouti) cats has a base pattern, of
either Ticked, Mackerel/Classic tabby or Spotted. It is important that we picture
this – because it is this base pattern that is affected by the Wideband gene.
Some breeders believe that the Tippy sparkle that can be observed in some lines
is due to as yet unidentified ‘sparkle’ genes such as the Glitter in the Bengal. The
Tippy sparkle can best be seen in October when the new undercoat comes
through!
• Breeding show quality Silver Series Tipped – is quite simply not easy. In
the British Section there are a few dedicated breeders producing kittens of
excellent type – but the gene pool is very small and it is difficult for
breeders to make improvements without making a sacrifice of some sort.
• As litters are born breeders should look to see if they can detect the base
pattern of their kittens and keep records as to which base pattern results
in the best Tippy pattern.
• The Ticked gene as a base pattern is causing some excitement amongst
breeders and there is a sense that the ticked base pattern may offer the
best opportunity for evenness of ticking. (Breeders working with the ticked
gene should note that when the cat has two copies of the ticked gene it
will override all other agouti patterns)
• Black Tipped Breeders may wish to consider keeping a Black only line -
undoubtedly – cats that are Homozygous for silver (II) produce the best
Black Tipped coats – free of tarnishing.
• Breeding Tipped to other colours of BSH may help type – but first
generation will not make show cats – they will probably be too dark. But
hopefully second generation will pull back.
• Just for fun Tippy breeders could experiment with the other Tipped
colours!
• Some Tippy breeders would welcome more definitions of ‘Tipped’ such as
Shaded, Cameo, Shell etc.
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
As each BAC prepares a breeding policy for the breeds within their remit, it
provides a fresh opportunity to study the registration policy and the standard of
points in conjunction with the breeding policy.
For us in the British BAC it is the Non-silver series of Tipped British that is
presenting us with some of the most interesting genetic puzzles – but first we will
explain the genetic make-up of the Golden Tipped as seen on our show benches.
To that end we will start with the show cat!
This is the non-silver version of the British Tipped in which the hair is a rich
golden apricot colour, sound to the roots with black/dark brown tipping. The coat
on the back, flanks, head, ears and tail should be tipped with colour. The tipping
should be evenly distributed and it is the even distribution rather than the degree
of tipping which is of paramount importance. The legs should show tipping which
will decrease towards the paws, but which should be as even as possible. The
chin, stomach, chest and undertail are a pale apricot colour.
Tipped
A- B- C- D- ii Ticked oY/oo
Ta-
Mackerel
Mc- tata
Classic
mcmc
tata
Spotted
Sp- tata
A- = AA or Aa B- = BB or Bb or Bbl C- = CC or Ccs
D- = DD or Dd I – II or Ii Wb- can be: WbWb or
Wbwb
If we look carefully at the above table we can see that the only difference
between the Golden Tipped and the Silver Tipped is the fact that the Dominant
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Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar
Inhibitor gene (I-) is in its ‘switched off’ state. This absence of a gene in the
dominant state is sometimes referred to as being ‘recessive’.
What this tells us is that we have a cat that has a genotype that is
indistinguishable from the Brown Tabby and yet looks nothing like his genetic
twin.
If we just confirm that which we know about this beautiful cat – he is: 104
But how we get from the Brown Tabby to the Golden is a subject of much conjecture.
Five Theory’s
104
This is also true for the Black Tipped – but add the Inhibitor Gene
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Many breeders believe that it is in fact the Inhibitor Gene (ii) that forces the tabby
pattern to the tips of the coat – but that in its ‘recessive’ state it does not restrict the
melanin or colour from the hair shaft – a theory that suggests the Silver gene is in a
sort of sleeping rather than switched off state – (There are anecdotal accounts of
Golden cats producing Silver offspring)
The coats of Golden cats look very similar to the Silver series Tipped, with dark
tipping or shading. However, the ground colour or undercoat is a warm cream which
combined with black tipping produces a wonderful 'golden' appearance. This warm
creamy-red undercoat proves that the Silver gene is not present (and we know that it
cannot be carried)
The Golden Tipped, seems to strengthen the argument in favour of a separate gene or
set of polygenes, for ‘widebanding’ independent of the Silver Inhibitor Gene.
Although, it must be pointed out that the Silver (I-) gene does seem to have the
effect of widening the wideband to create the ‘Shell’ effect of the Chinchilla, whereas
the Golden’s ‘tips’ bleed more deeply into the hair shaft to produce an effect that is
more of a black smudge than a tip.
But Golden breeders must always remain grateful to those past Chinchilla breeders
and the newer BSH Black Tipped breeder’s, because it is the selective breeding over
many many generations for clear, wide Silver bands that has set the stage for the
appearance of the Golden trump card, Rufousing!
Rufous genes are the second theoretical genes that are thought to be a critical part of
what makes the Golden so special. Rufism is a term describing the genetic or
polygenetic factors that breeders have been able to select for that allows them to
produce ever warmer ground colours in non-silver agouti cats.
Essentially, the amount of rufousing determines whether a brown tabby cat will have
a warm brown pigment or a colder grey colour.
The impact that these Rufous genes can have is very important when breeding for
colour. And it certainly seems as if the effect is accumulative the more one breeds for
Rufism the warmer the tones on Brown and Red cats.
So, we have either a Black Tipped with the Silver gene switched off and a good dose
of Rufousing or we have a Brown Tabby, with a big dose of widebanding that allows
us to exploit that Rufousing!
All this now sounds quite simple! – but actually breeding the Golden is more
complicated than it might first seem – which is why we do not see many examples on
the Show bench and there are two main reasons for this:
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‘golden’ litter only to find that they are looking at a litter of very sweet, but
poorly marked Brown Classic tabbies or Spotties.
• Golden breeding is not for the short term breeder. Persian breeders have
taken many years to achieve the beautiful Golden Persians we see today.
• Homozygous seems to be the answer, Homozygous for Agouti, Wideband and
Rufous – will all help to breed true!
• It seems that the greater the Rufousing dose, the less contrast we get
between ground and marking colour.
• Understand your base pattern - On neonate tipped kittens we can usually see
signs of tabby/mackerel/spotted/ticked patterns.
• Keep meticulous records – these kittens have a habit of changing – many is
the ‘Brown Spotty’ kitten born to Golden parents – that slowly turn into lovely
Golden adults.
• Try to run some kittens on to understand your lines.
• Keep Golden series separate from Brown or Silver unless you need to ‘go out’
for genetic diversity – in which case be prepared to spend time ‘undoing’ any
effects of Incomplete dominance!
• Remember that Golden kittens can be born to non Homozygous for Silver
Black Tipped parents –
• Avoid the red- gene – novice breeders already confuse well rufoused Spotties
from Tipped to Spotty matings with Tortie/Tabbies.
Earlier in this section we alluded to puzzles, conjecture and theoretical genes, none of
which are for the faint hearted –
Unlike the long term Persian breeding programs that have produced the Silver and
Golden longhairs – the Tipped British is still in its infancy. Whereas the Chinchilla
Persian is very likely to be Homozygous for Agouti, Black, Silver and Wideband –
generations of ethereal cats breeding true, we cannot be certain without testing what
the British Golden gene pool looks like – and of course whilst we can test for the
Agouti gene we cannot yet test for the Tabby base pattern.
There is a great deal of argument about the relationship between Wideband and Silver
– The ‘Golden’ suggests that the two are separate. There is also an argument that
says we cannot have Wideband without the Agouti shift, because there is no ‘band’ to
widen - but the Smoke is a non-agouti cat, with the Silver gene and there is a ‘band’
before the colour kicks in – not very wide – but it is there all the same.
Golden Tipped breeders are intrigued by the dilute Golden, the really rather beautiful
Blue Golden The creation of the gentle shades of warm, but pale cream tipped with
the softest tones of pale blue combine to produce one of the most gorgeous cats
imaginable. And this colour-way works much better within the Golden program than
the Black tipped.
What all this adds up to – is that we have de-stabilized the genetics that the Persian
and Abyssinian breeders have spent many generations to ‘fix’ – and we now have to
put our time in to get these beautiful Tipped cats to a point where we can predict
what we will get in our litters.
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More recently breeders are adding the colourpointed gene into the Golden Mix with
some intriguing results. Kittens with very definite, golden ears, noses and testicles!!!
– can this all be down to the agouti shift?
Another issue for breeders of the non-silver Tipped series, is Type and Coat. This
combination of genes just loves to have either long fluffy coats or very short almost
oriental coats – and getting round faces, eyes and cheeks is hard won!
The Golden tipped and the newer, as yet, unrecognized Blue Golden represents a
great deal of genetic interest to the dedicated breeder. They are a lovely sight on the
show bench and breeders producing good examples deserve a hearty congratulation.
Now, anybody still hungry for genetic challenges can do some research on the late
colour change phenomenon!!! 106
The following extract gives Robinson’s opinion on the Wide Band debate
The presumed effect of the gene is to widen the agouti band on the hairs. In addition, the
gene is said to make the tabby pattern less distinct or blurred. The overall effect is a tabby
of a rich golden hue. Examination of hairs from golden tabbies reveals that the hair shafts
are nearly completely yellow with a black tip, with a suggestion of pale blue at the base.
This is just as would be expected for a wide band phenotype. The breeding data to
substantiate the existence of the wide band gene are slim; but the gene has been
theorized as a dominant (either complete or incomplete) and has been provisionally
symbolized by Wb However, the apparent difference between silver tabbies and chinchilla
silvers, and thus brown tabbies and goldens, could be easily explained by polygenetic
effects on the quantitative expression of the agouti and inhibitory proteins. As the
level of these inhibitory proteins increases pigment production is reduced. A chinchilla
golden is simply brown tabby with such high amounts of agouti protein production that
the agouti-shift occurs very early during hair growth. This inhibitory effect is so strong
that it causes the shade of yellow pigment seen in the agouti band to change to a lighter
colour characteristic of golden cats.
105
Was she a Ticked based Colourpointed?
106
More incomplete dominance perhaps!
107
Where there is a genetic ‘argument’ we the BAC will ‘present’ Robinson’s view
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11. Colourpointed
The Colourpointed British Shorthair is a huge success story. During the 1980’s a
group of breeders decided to
cross the Colourpointed
Persian with the British Blue
and the result today is a
beautiful cat that has the
third highest 108 number of
registrations, with only the
self and the Silver Tabby
series ranking higher.
The colourpointed in the UK is registered in all the colours of the British Shorthair
except the Tipped Series. 109
The genetic codes behind this beautiful cat are the same as those in all the other
sections except for a variation of the gene on the C 110 Locus – so we will
concentrate on this gene.
However, before we start to examine gene codes we need to sort out some
misunderstandings regarding the common names for this gene.
The colourpointed Persian carries what we commonly call the Siamese gene. The
longhaired colourpointed cats were originally bred using Siamese breed stock.
What this means is that the genes that we use in the British Shorthair are the
same as those in BOTH the Persian and the Siamese.
Also known as the Himalayan gene we shall refer to the gene in our beautiful cat
as the Siamese gene.
The ‘C’ gene governs the expression of ‘full coat’ and is a dominant gene. The
Siamese pattern is believed to be a natural random temperature sensitive
mutation that occurred hundreds of years ago in Siam. 111 It is thought that all
colourpointed cats have descended from this original mutation.
108
8% of British Shorthair Kitten Registrations 2009
109
April 2011
110
Also called the Albino Locus – we shall call it the Full Coat Locus
111
Now Thailand
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There are five alleles (forms) of this gene: Full Colour C, Burmese cb, Siamese cs,
blue-eyed albino ca and albino c. 112
The mutations of C are all a form of Albinism and pointed cats are what we call
partial albinos.
The Pointed mutations are temperature sensitive mutations, which mean that the
C gene, is only active at low temperatures.
The other genetic mutations of the C gene as shown in the following table.
Table 4
C – Full Coat
Genotype Description Phenotype
C/C Homozygous - Solid Coloured Solid colour
C/- Heterozygous - Solid Coloured Solid colour
carrying a single copy of a
recessive.
c/c Albino with pink eyes White with pink eyes
ca/ca Albino with blue eyes White with pale blue eyes
cs/cs Siamese Colourpointed Colourpointed, Siamese pattern
cb/cb Burmese/Sepia Burmese pattern
cb/cs Tonkinese – Co-dominant Tonkinese pattern
Himalayan/Burmese
So in summary:
• The full colour form (allele) C, is Wildtype, dominant, and produces a full
expression of the coat colour
• The Siamese form cs, is recessive to C, co-dominant with the Burmese form,
cb and dominant to the blue-eyed albino ca and albino c. The albinism
produced by this form is more temperature sensitive than cb, causing the
classic Siamese pattern with blue eyes.
• The Burmese form cb, is recessive to C, co-dominant with the Siamese form
Cs
, and dominant to the blue-eyed albino ca and albino c. This form of the
gene produces a slight albinism. Black becomes dark brown. It is best known
in the Burmese breed, which has green or green/gold eyes.
• The blue-eyed albino form ca, is recessive to C, cb and cs but dominant
over the albino allele. Cats will show an almost complete albinism, with
washed out blue eyes and pigment free, translucent white coat.
• The albino form c, is recessive to all the other forms producing a pink eyed
complete albinism.
Whilst complete albinism is very rare in the domestic cat, partial albinism is
common. These partial albino patterns are temperature sensitive mutations.
112
Some scientists dispute that both c and ca exist – and that there is only one
recessive for pure albino.
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The C allele or form of this gene is a gene with a job to do, and is usually kept
busy producing an enzyme called Tyrosinase, and all the other forms of C have
become lazy and have become ‘loss of function’ alleles. When a gene form loses
function an enzyme or other protein may no longer be produced, or may be
produced in a different form or at lower levels or indeed may become
nonfunctional.
What has happened with our Cs gene is that Tyrosinase is still produced, but the
enzyme they produce does not work at core body temperature. Instead pigment
is only produced at below-average body temperatures.
Over the next few weeks and months, pigment continues to form on the coolest
parts of the body. Eventually colour creeps, from the feet up through the legs,
and the mask bleeds out from a smudgy nose to form that famous colourpointed
mask.
This is a complex topic, not yet fully understood, but we will do our best.
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For those people who share their lives with colourpointed cats, they can see them
playing and hunting in the same way as other cats. For Breeders and exhibitors
the spectre of Nystagmus or a squint is never far away.
Many breeders understand that these eye problems are linked to the
colourpointed gene, which is correct; it seems that the mutated Tyrosinase found
in temperature-sensitive albino cats does not act to pigment the eyes. This is why
all cats with true colourpointed genes will have blue eyes.
The absence of pigment or melanin means that the eye does not develop properly
in the womb. As the foetus develops, some parts of the eye, including the crucial
optic nerves become disorganized and the result leaves the cat with problems
with focusing and depth perception.
We humans, with our eyes facing forwards have what is known as binocular
vision – our eyes are wired to our brains via our optic nerves – and this set up
allows us to integrate the things seen by each eye into a single image.
Most birds and reptiles have their eyes on either side of their heads and have
panoramic vision – the effect of this is that, each side of the brain only ''knows''
about the visual input from one eye,
Most cats have their eyes wired to their brains in the same way that we do but
this is not true of the colourpointed cats – their optic nerve fibres do not cross
properly in the brain 113 and many end up connected to the wrong side of the
brain, in a way that is much more similar to the birds and reptiles.
What we cannot know is the exact extent to how the visual fields of our cats are
affected, and how they compensate for the mix-up in their wiring – but we can be
certain that they cannot have full binocular vision and this may impact on their
perception of distance.
In addition to this criss-cross wiring problem, partial albinism can also cause
other vision problems including: Congenital Strabismus and Nystagmus.
Congenital Strabismus
In simple terms this means a squint, and we will deal only with the squint as it
occurs in the colourpointed cat.
When a cat with a Siamese gene has an eye (or two) that points down and over
towards the nose this is called a convergent squint. When the eye or eyes point
outward, away from the nose it is called a divergent squint.
(At one time these squints were commonly seen in the Siamese breed.)
This is a permanent condition, which cannot be cured and usually appears in the
kitten at about 6-8 weeks of age.
We know that the primary cause, is an excessively high proportion of optic nerve
fibres which cross over at the optic chiasm. This causes a reduced medial visual
field. It is thought that when the kitten develops a congenital 114 permanent
squint, it is a muscle imbalance caused by the kitten’s eyes attempting to correct
the distortion of the visual field.
113
Dr. R. W. Guillery, Neurophysiologist - University of Chicago
114
Condition present at birth or soon after.
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Nystagmus
It is probably true to say that virtually everyone with albinism has Nystagmus, so
it is small wonder that it crops up in our colourpointed cats with their major genes
for partial albinism.
It is probably safe to say that whatever the ‘form’ of Nystagmus we see in our
colourpointed cats it is usually due to the cross wiring of the optical nerves 115
There is much that we do not know about Nystagmus in cats 116 but it is likely to
be at least in part Polygenic, and kittens that show sign of a severe and or
permanent congenital Nystagmus should not be used in Breeding programs.
Most kittens and cats with severe or permanent early onset Nystagmus, will not
have eye, brain or other health problems and are unlikely to be affected by
constantly moving images, as the young brain can adapt to the condition.
Additionally neither the condition nor the animal’s vision is likely to worsen as the
animal grows up. The condition is not painful and kittens will benefit from the
normal play stimulation that most kittens enjoy.
However, they are quite likely to experience some of the following symptoms.
Breeders letting affected animals go to companion homes may wish to apprise
new families of the condition.
115
If an otherwise normal eyed cat, colourpointed or not develops a sudden late onset
Nystagmus Veterinary opinion must be sought.
116
The GCCF genetics committee are working with the BAC’s to launch an investigation
into Nystagmus with the University of Bristol, Langford.
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Whilst it is important for breeders to understand the major genes that they are
working with, such that they can ensure that they always act in the best interest
of their breed, there are no contra indications to responsible breeding of the
partial albino cat. Scientists, who have studied these visual issues, have been
surprised that despite the very unusual wiring of those optical nerves, the
average colourpointed cat appears to be able to operate pretty much as cats with
normal eye/brain wiring.
Points Colour - Mask, ears, legs and tail should be a clearly defined colour,
matching on all points.
We can see from the earlier sections that no matter what colour British Shorthair
one chooses to breed there are always pitfalls and the Holy Grail of colourpointed
breeding is the clear coat.
Although the standard of points says that an otherwise good exhibit should not be
penalised for shading, it also refers to ‘light shading’
The problem for colourpointed breeders is that the genes that cause pigment to
form in areas of the body that are cooler due to reduced circulation keep working
throughout the lifetime of the cat, and so as the cat ages and the circulatory
system slows down as it does with all mammals the colourpointed cat begins to
shade beyond the ‘points’ – at first it will be the shoulders and flanks and then
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gradually the whole upper body will eventually shade. Because this shading is
related to pigment, the Seal and Blue series will be most affected, followed by
chocolate.
Neutering can add to the natural slowing of the body metabolism and exacerbate
the shading on the body.
The result is that when we do see older colourpointed cats on the show bench
they will most likely be lilac!
It is frustrating for breeders who will try to campaign their young cats early
knowing that they will inevitably shade and will almost certainly be faulted for
something that is largely not polygenic.
Of course there are some lines that have managed to breed clearer coats, this
may be due to a higher metabolic rate in some lines of cats – but we only have to
look at the ratio of colourpointed kittens shown in the various colours and
compare that to the number of adult cats shown, especially in the darker colours
to know that this is an issue shared by most colourpointed breeders.
The other major headache for the British Colourpointed breeder is eye envy.
Compared to the Siamese benches with their rich array of shades of blue,
deepening to almost navy, the British colourpointed eye is often a disappointing
wish washy blue.
This is a polygenic factor and almost entirely due to the success many British
breeders have had with the orange eye.
The deeper colours of the orange eye are produced by polygenes for an increased
volume of pigment producing cells (melanocytes) and a deeper pigment.
Whereas the green eye has less pigment producing cells but also produces a lot of
pigment.
Now, this is all quite complicated but the colourpointed eye is set up in the same
way as the orange and green eye and so also has melanocytes. But the melanin
production is impeded by the fact that the eye is always warm and the
temperature never gets cold enough to produce an orange or green eye.
So what we see in the colourpointed eye is actually blue light that gets refracted
from the lens of the eye. And the less clutter in the eye, ie melanocytes, the more
blue light gets refracted and hey presto we get a deeper blue eye.
The Mask
Colourpointed breeders are seeking to produce a cat that is fully masked and
where the margins between the mask and body colour remain distinct.
Tabby colourpointed although not fully masked should still have an obvious
demarcation between the face and rest of the body.
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If we go back to the table on page 89 Table 4 we can see that the alleles for the pink
eyed albino c and the blue eyed albino ca are recessive to the genes for the
Siamese pattern cs
There is a hypothesis that suggests that if we were to have a cat that was
s a
genetically c /c we would have blue eyed colourpointed cats with a higher
contrast of colour between coat and points with a wishy washy blue eye. If any of
our lines in the UK carried this combination of genes we might see clearer coats,
but not necessarily good eye colour, breeders may occasionally see a blue eyed
albino appear in a litter if they cross back into their own lines. Although the c and
ca genes cannot be tested for and indeed some doubt whether they both exist,
any breeder suspecting that their lines may hold such a combination may wish to
keep detailed records and share these with the Colourpointed Club.
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In previous sections we have covered the genetics behind the genes for Smoke
and Silver and also for Colourpointed. Here we are looking at cats bred with an
amalgamation of these two genes.
It is worth remembering that the Silver Inhibitor gene (I) is dominant whilst the
Colourpointed (Cs) gene is recessive.
In the UK the Smoke or Silver Tabby Pointed British Shorthair is rarely registered
with the GCCF.
However, in Europe breeders are working increasingly both with Silver and Wide-
banding in their Colourpointed Breeding programs.
Essentially, this Series reflects the simultaneous presence of both the Inhibitor
gene and the Siamese Partial Albino gene, either with or without The Agouti gene.
The Smoke or Silver Tabby Pointed Cat can come in the full range of solid and
tortie point colours.
History
It seems that the first recorded Silver pointed cats in the UK were bred in
Scotland in the 1940’s. These cats, known as Silverpoint Siamese were
introduced to the cat fancy in the 1960s. It is not clear whether some of these
cats were in fact Silver-pointed, but they were tabby pointed and in 1966, the
Tabby Point Siamese was granted recognition by the Governing Council of the Cat
Fancy (GCCF).
Smoke Pointed
The undercoat of the pigmented point colour of the Smoke Colourpointed should
pale to silver at the roots. As with the British Smoke there is a great deal of
variation in the effect of the Inhibitor gene in these cats, and breeders should
select for those with the greatest degree of contrast.
With the dominant colours, Seal and Chocolate the points may appear darker
than the non-silver series and the dilute blue and lilac colours may appear colder
with a steel effect. And other pointed breeds with the Inhibitor gene, report that
body shading is often more intense than in non- smoke individuals.
These cats display the main Tabby patterns on their ‘Points’ The stripes in the
dominant colours of seal and chocolate, may appear darker than the non-silvered
tabby point. Between the tabby markings the coat will appear to be silvery white.
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Again, body shading may be darker than the non-silvered cat . The dilute colours;
blue, lilac, cream and fawn may appear to have very pale points.
Silver eye ‘goggles’ may show around the eyes of the smoke and silver tabby cats
and full pigmentation may not develop until the cat reaches her second birthday.
The Problem
At the time of writing there is no test for the Inhibitor gene. If we remind
ourselves of the fact that when the Inhibitor gene (I) is present the full
development of pigmentation in the hair does not occur, and the non-agouti or
'Self' Cat turns into a Smoke, and an agouti cat becomes either a Silver Tabby or
Silver Tipped. We know that the (I) gene has a hugely variable impact, some cats
show a strong sparkling white base, whilst others show hardly any contrast and
the breeder may not see the expression of silver at all.
It is this problem with the Silver gene that has caused the GCCF to over-stamp
litter registrations with ‘Silver in Pedigree’
In the Red-series Silver ground colour becomes Ivory and this is compounded by
the effects of the Siamese (Cs) gene
During the research for this section, we interviewed several well known European
breeders – during these interviews one breeder commented that she felt that as
soon as a test for the (I) gene is found we will have to re-register at least 25% of
all colour pointed British as Silver series.
117
EMS Breed Numbering System
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There are a great number of colourpointed breeders today both here and in
Europe breeding from cats directly descended from Beeblebrox Plum Crazy 118 –
and if his Silver gene slipped through the net – how many others?
There are a number of colourpointed lines with incomplete masks around the
muzzles, with dark shaded coats and brindling of the points – and right now the
only way to determine whether your colourpointed cat is secretly Silver is to test
mate out to a self cat and evaluate the results -
118
Many Pedigrees still show this cat as red-pointed today.
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In previous sections we have covered the major genes for the Bi-Colour (White
Spotting (S) and the Colourpointed (Cs) British.
Whilst new to the British Section in the UK, the Colourpointed and White genes
are not strangers to each other and can be seen in both the Ragdoll and Snow
Shoe breeds. The Birman (a pointed cat) also shows an expression of White
Spotting (S) in the form of mittens on all four feet.
The British Colourpointed & White may be bred and shown in a mitted variety, Bi
Colour and high white/Van pattern and although all patterns are acceptable, the
pattern preferred by breeders is the Bi Colour. 119
In all three patterns, breeders are looking for the White Spotting (S) gene to fall
and mask the points colour on the feet and legs, leaving four white feet. The tail
and mask will show pigment as the Colourpointed gene allows pigment to seep
into the colder points and breeders hope that that lovely white Bi-Colour blaze
will make it’s V pattern neatly in the centre of the face.
The tonal shading that often appears in the coat of a Colourpointed, takes the
form of Bi Colour markings, this area is shaded only and not solid colour.
Symmetry of design is
desirable.
Remainder of mask, ears and tail should be clearly defined and matching in
colour.
Body Colour - There should be a good contrast between coloured Points and body
colour. Shading if present should tone with the coloured Points. Heavy shading in
a mature cat should not be penalised in an otherwise good exhibit. Bib, chest and
under body to be white. Legs should be white or have tonal shading, small
patches of colour on the legs are permissible. All four feet must be white.
119
The SOP does not specify the amount of white on the body.
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The following are the observations from breeders of Colourpointed and White.
Within the section on the Bi-Colour we touch upon the Polygenic 121 properties of
the White Spotting (S) gene – Although it is often said that the Bi-Colour pattern
is completely random – we can see that the classic Bi-Colour pattern can be
worked at over generations. These Polygenic traits of (S) are proving interesting
to the Colourpointed and White breeders. The following paragraph is an extract
from the records of one such breeder.
“Certain matings of mine breed true every time, perfect markings, others are
more random. However, after previously breeding Bi-Colours, and having worked
on the Colourpointed and White, I believe that you will produce very similar
markings in the kittens as you have in the parents with White Spotting.
To within a certain percentage, say the same markings to within 10 %, meaning
more or less white either way and the markings in the same place as the parents
give or take 10%.either way - Sometimes mitted kittens are produced,
but the colour always stops where it should on the ankle, I have not known of any
C/P and White kitten developing even one pigmented paw!”
120
One experienced breeder reports when any mating contains a C/P and White she
will always have a kitten with White in the litter
121
Genes working alongside major genes that can be exploited in breeding programs
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The British Shorthair we know today has a long and interesting history, we have
learned how after the major world wars breeders worked hard to restore the
breed by going out to the Persian breeds, we have read that the British Tipped
and the Colourpointed breeds have been derived from the Persian and Chinchilla
breeds.
And whilst breeders have worked for generations to get the Phenotype 122 as close
to the breed standard as possible the hidden Genotype or unseen genetic makeup
of our beautiful breed has been carried from generation to generation.
The length of fur in the feline is governed by the Long hair gene which is known
as L. In the Wild Form 123 the dominant form of L codes for short hair, whilst the
recessive l codes for long hair.
Anagen is the active growth phase of hair follicles. During this phase, the cells in
the root of the hair are dividing rapidly, adding to the hair shaft. Hair stays in this
active phase of growth for a genetically predetermined period of time. At the end
of the Anagen phase a
signal causes the follicle to
go into the Catagen phase.
The L gene works as the switch between the Anagen (growth) phase and the
Catagen (stop) phase - In the shorthaired cat, L steps in and creates the shorter
coat we know in the British, in the mutated recessive l form the switch from
Anagen to Catagen is delayed, the hair keeps growing and the longer haired cat
results.
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Appearance
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Typical Phenotype
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Although currently registered with the GCCF as a Variant of the British Shorthair
and as such cannot be shown, there is a good deal of interest in this very pretty
cat.
Variously known throughout the world as The British Long Hair, The British Semi-
Long Hair, The Britanica, The Lowlander and a bit confusingly the Highlander this
cat is described by most registries to have the same standard of points as the
British Shorthair, but with a Semi-Long coat standing away from the body.
One gene – fibroblast growth factor5 (FGF5) – determines hair length. Short hair
is a dominant trait determined by the wild-type form of FGF5. Long hair is a
recessive trait. Four mutations in FGF5 have been identified that are associated
with long hair in cats. Long-haired cats can carry two copies of the same
mutation (homozygote recessive) or have two different mutations, one on each
chromosome (compound heterozygote). Three of the mutations are fairly breed
specific, while the fourth is present in all long-haired cat breeds and crossbreds,
as follows:
Mutation 4 (M4): present in all breeds of long hair cats, including Ragdolls, Maine
Coons, and Norwegian Forest Cats.
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The most common colour for wild, non colourpointed, cats is hazel somewhere on
the scale between orange and green.
These eye colours are not linked to major genes and breeders are able to
manipulate polygenes to produce the desired colour and intensity.
• Orange/Gold/Copper
• Green
Breeders breeding for eye colour will know that within each of those colours there
is great deal of variation in the range of colours, both in depth of colour and hue.
And the shades of orange can merge into green and vice-versa leaving cats with
the hazel eyes of their wild counterparts.
The good news is that selective breeding can influence these polygenes so, within
reason, eye colour can be stabilized. So when we see outstanding eye colour time
and time again in a line – we know that this is thanks to hard work and planning.
When eye colour is linked to a major gene it means that breeders have to work
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• Colourpointed Blue
• Blue
Kittens are always born with blue eyes. The final eye colour begins to develop
between 6 and 7 weeks of age. It will be at least 3 or four months before
breeders will be able to see the much longed for colour.
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For example it is not possible to breed a colourpointed cat with an Orange eye.
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There are two key factors that influence eye colour in the cat:
• Pigmentation
• Blue Light Refraction
Stroma
Just like the coat colour, the pigmentation of the iris is caused by melanin and the
intensity of the colour depends on the numbers of melanocytes in the eye.
When melanocytes levels are low the eye colour will be in the green spectrum, as
the number of melanocytes increase, possibly due to selective breeding, the eye
colour will intensify to orange
Melanocytes don’t all produce pigment at the same rate – sometimes they can be
quite lazy – so when a cat has fewer, but lazy melanocytes the eye will be a weak
green. A black tipped, with stunning black-rimmed deep-green eyes will not have
many melanocytes but those she has will be working hard at producing pigment.
Using this logic we can understand that a cat with an iris packed full of
melanocytes all working as hard as possible will result in a cat with fabulous
copper eye colour.
The blue of the colourpointed cat is different again – the colourpointed eye could
have eyes packed full of melanocytes or not – they could be the lazy sort or not –
it won’t matter – because those melanocytes simply won’t work in the iris of a cat
that cannot produce pigment unless core body temperature falls below normal -
Additionally the light that is taken in through the cats eyes is refracted and
reflected and it is the number of fibres milling about in the Stroma level that
determines the levels of blue light that interacts with the pigmented cells – this
combination of blue coloured light playing over the pigmented cells determines
the eventual eye colour of the cat.
Fewer fibres in the colourpointed eye will result in a deeper clearer blue – the
breeder’s best chance of producing good blue eye colour is to consistently mate
cats with good blue eyes to cats with good blue eyes!
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Or in rare cases pink
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Finally, behind the retina we have the Tapetum Lucidum. This is reflective layer of
tissue that sends light back through the eye. and helps cats see in low light - it
also causes eye shine in flash photos. This layer is missing from the eyes of most
blue eyed white cats.
ODD EYES
These occur when the cat has different genes affecting each eye – if pigment cells
do not reach the eyes because they are halted by the White gene (W) the cat will
have blue eyes. However, if pigment cells reach one eye the cat will have one
orange eye and one blue eye -
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This section will cover the importance of keeping accurate breeding and cattery
records.
We will use the term Cattery, to help remind us that even if our cats are our
much loved pets that live in our homes, as soon as we implement a breeding
program of selected animals, a well managed system of housing and husbandry
must be planned.
Running a cattery requires a full time commitment to health and welfare. It will
require time, space, patience and financial support and critically it will require the
breeder to keep accurate and long term records.
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15. Recommendations
Good breeders will set themselves goals and objectives. They will seek to
understand the needs of the feline and ensure that they do not overcrowd or fail
to meet the needs of the animals in their care.
Breeders will need patience, creating a line needs time, and an ability to see
one’s offspring objectively. It takes courage to neuter animals with undesirable
characteristics.
Breeders can often recognize cats bred by other breeders, they can recognize a
‘line’ by its look, this not happenstance, it is the product, often of many years, of
careful planning, selection, heartache and luck.
The breeder that we all admire, will be selecting kittens that they like the look of,
that have good health and temperament. They will seek to mate their queens to
excellent stud cats in the hope that those selected queens will produce easily,
strong sound kittens that have no genetic abnormalities but do have the kind of
beauty that the breeder can be proud of. It is this balancing of genes for good
and bad that is the true test for any breeder. If we get it right, we should be able
to predict the outcomes, we should be able to produce kitten after kitten that
have such distinctive attractive qualities that unmistakably stamp them as ‘ours’
Using this breeding policy in conjunction with the standard of points and the
registration policy, breeders should be able to work with their clubs and each
other to protect and improve this wonderful breed that is so very much loved.
The GCCF draft business plan for 2011-12 includes a scoping exercise for a
“GCCF Breeder Scheme” – which will be promoting responsible breeding.
We recommend that our British Breeders support this initiative - In the final
analysis it is we, the breeder that is the custodian of the British Shorthair Cat.
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Appendix 1 References
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Appendix 2 Glossary
Baer Test - The BAER test is based on the electrical response of the brain to
auditory stimuli. When a sound enters the ear, tiny electrical impulses are
generated by components of the auditory pathway. The signals can be picked up
by recording electrodes positioned on the head and are, in turn, passed into a
computer. The test offers quick, non invasive and accurate assessment of an
individual’s hearing status.
Blood Groups - Blood consists of red blood cells, white blood cells and plasma.
The red cells carry oxygen and their well-being is essential for life. Red blood
cells, like most other components of the body, are made of protein, and proteins
can be recognised and destroyed by the body’s immune system. The cat has
essentially two blood groups: A and B (although there is a third, very rare, group,
AB, which is a combination of the two). The red blood cell proteins are A in one
group, B in the other (see figure 1). In fact, the proteins on the surface of type A
and B red blood cells are only slightly different, but of course the immune system
can detect the difference.
Dominant Gene is one which is capable of expressing its trait even when
carried by only one member of a chromosome pair.
Genes - are the individual units of inheritance. Each gene (or combination of
genes) determines a characteristic that govern the eventual shape, size, sex,
colour, pattern and hair length of the individual animal. Genes work in pairs – but
the pairs do not have to be identical
Locus - is the exact position on the chromosome where the gene is located
Recessive Gene can only express itself when both members of the chromosome
pair contain the gene
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Agouti (A) – the natural “wild” gene that is the basis of the tabby cat. The
base agouti pattern is bands of black on a yellow background; in the cat this is
overlaid with one of the tabby patterns.
Mackerel (Mc) – the basic striped tabby pattern that overlays the agouti base
(ie “wild” form)
Classic (mc) – a mutation of the mackerel allele recessive to all other tabby
patterns which gives a blotched pattern with the characteristic “butterfly” motif
across the shoulders and “oysters” on flanks.
Ticked (T) – an incompletely dominant gene which removes most of the stripe
pattern leaving the ticked agouti base pattern on the body with minimal
overlaying stripes on legs, chest (necklace) and face.
Spotted (Sp) – current thinking is that it is likely that a specific single gene
causes the spotted tabby pattern, breaking up the mackerel or classic pattern into
elongated or rounder spots respectively.
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Chocolate (b) and Cinnamon (b1) – two mutations of the basic black non-
agouti gene which modifies black into dark brown or medium brown respectively
The white masking gene, (W). The "W" gene prevents the normal
replication and migration of pigment producing cells during embryologic
development. As a result, WW and Ww cats have a greatly reduced number of
melanocytes and appear white, no matter what other colour genes it may carry.
Only a cat that is homozygous recessive (ww) will express normal pigmentation.
Also called Dominant white (more properly "epistatic white" since it occurs on a
different gene to the black-based or red-based colours)
Siamese (Himalayan) gene (cs) - The Siamese colored kittens are born
completely white. The color of the mask doesn't appear until after a week. This is
due to that the cells of Siamese colored cat cannot develop any pigment if the
temperature is too high. When the kittens are in the womb of their mother's they
are warm and comfortable, and no pigment at all is developed. When they are
born it becomes cool enough for the pigment to develop on the cooler
extremities, like for instance ears, legs and tail. This temperature dependant
pigment production is called acromelanism. Acromelanism is present in other
animals, for instance in rabbits.
Dilute (d) – a recessive gene which reduces and spreads out the pigment
granules along the hair-shaft and turns a black to blue, chocolate to lilac,
cinnamon to fawn and red to cream.
Dilute modifier (Dm) – a dominant gene which serves to modify the action of
the dilute gene (it has no effect on undiluted colours), it lightens and
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“caramelises” the colour turning blue into brownish-grey, lilac and fawn into pale
taupe (in all three cases known as Caramel) and cream into a warmer pinkish-
cream tone (Apricot)
Polygenes – these are collections of genes which modify the effect of the main
dominant and recessive genes above. A build up of polygenes creates a bigger
effect, for example a collection of certain polygenes increases the length and
density of the long-hair gene to create the Persian, and a build-up of polygenes
serves to enhance the effect of the main colour genes, turning the effect of the
orange gene from the sandy colour of the ginger domestic tom to the rich vibrant
red of the Red Persian, British or Asian Self. It is likely that a group of polygenes
is the reason for variation in the degree of tipping in the Shaded Tabby/Burmilla,
the polygenes working to create the band-width in interaction with the inhibitor
gene (when present) resulting in the range of pattern from tipped to heavily
shaded.
Burmese Colour Restriction (cb) – a mutation on the albino allele one step
up from the Siamese (Himalayan) gene. This reduces the amount of pigment
produced in the coat but, because it is thermo-sensitive, the pigment is darker at
the points and (slightly) lighter on the body; the action of the Burmese gene
causing a genetically black cat to turn sepia brown, a red cat to turn pale
tangerine.
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Appendix 4 Contributing Breeders
Just as the British Shorthair would not be the cat it is today without the love, tireless
energy and sheer hard work that is invested by the breeders, a Policy such as this
could not be produced without help.
Our heartfelt thanks to the following contributors and all of our breeders who are
doing such a magnificent job on what is the best cat in the world.
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