breeding policy british shorthair · 2018. 7. 1. · draft british shorthair breeding policy –...

Jun-18

British Shorthair Breed Advisory Committee

Breeding

Policy For The British Shorthair

Guidelines

For

Healthy

And

Responsible

Breeding

Draft British Shorthair Breeding Policy – Genetics Committee – after BAC Seminar

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Summary

When an individual or group makes a decision to ‘breed’ an animal for any

purpose that decision carries a degree of responsibility. This British Shorthair ‘breed specific’ policy seeks to safeguard the genetic health, distinctive type and integrity of our wonderful breed, by providing

our breeders with sufficient information to inform their breeding programs or systems, such that they understand those responsibilities relative to

the genetics of the breed and seek to protect and improve all aspects of the most popular pedigree cat in the UK today.

This policy is intended to support and accompany the British Shorthair Registration Policy.

Acknowledgments

• The GCCF Breeding Policy

• Steve Crow and the Asian Breeding Policy

• Dr Diane D. Addie and her Invaluable Website.

• Contributing British Shorthair Breeders1

• Robinson’s Genetics for Cat Breeders and Veterinarians

• Photographers: Alan Robinson and Robert Fox

1 These can be found at Appendix 4 Contributing Breeders


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Contents Summary ...................................................................................... 2

Acknowledgments .................................................................. 2

Contents ....................................................................................... 3

1. Introduction ..................................................................... 6

1.1. Purpose of this Document........................................... 7

1.2. Exclusions .................................................................. 7

2. Background to the British Shorthair Cat ..................... 8

2.1. Origin .......................................................................... 8

2.2. Breed Description ....................................................... 8

2.3. The Show Cat ............................................................. 9

2.4. Registration ................................................................ 9

2.5. Characteristics and Temperament .............................. 9

2.6. Longevity .................................................................. 10

2.7. Recognition .............................................................. 10

3. Breeding System Information ..................................... 11

3.1. Genetic Make Up ...................................................... 11

3.2. Size and variety of gene pool .................................... 14

3.3. Inbreeding ................................................................ 15

3.3.1. Best Practice Advice and Recommendations ........ 20

4. Type ......................................................................... 21

4.1. Overall ...................................................................... 21

4.2. The Head .................................................................. 21

4.3. Nose ......................................................................... 21

4.4. Chin .......................................................................... 22

4.5. Ears .......................................................................... 22

4.6. Coat .......................................................................... 22

4.7. Eyes ......................................................................... 23

4.8. Body ......................................................................... 23


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4.9. Legs & Paws ............................................................ 23

4.10. Tail ........................................................................... 24

4.11. Size and Weight ....................................................... 24

5. Health ...................................................................... 25

5.1. Genetic Defects ........................................................ 25

5.1.1. PKD ...................................................................... 25

5.1.2. HCM ..................................................................... 25

5.2. Blood Groups and Neonatal Isoerythrolysis .............. 26

5.2.1. Recommendations ................................................ 28

5.3. Brachycephaly .......................................................... 29

5.4. Weight ...................................................................... 29

5.5. Infectious Disease .................................................... 30

6. The Self Coloured British Shorthair ...................... 31

6.1. The White Self .......................................................... 33

6.2. Black and Blue Self .................................................. 37

6.3. Chocolate or Lilac Self .............................................. 40

6.4. Cinnamon or Fawn Self ............................................ 42

6.5. Red or Cream Self .................................................... 45

6.6. Tortoiseshell (all colours) ............................................ 49

7. Non-Self/Non Tabby ..................................................... 52

7.1. Bi-colour ........................................................................ 52

7.2. Van Pattern (Preliminary) ............................................ 56

7.3. Tortie and White and Van Pattern Tortie and White . 59

8. Smoke ............................................................................ 61

9. Tabbies and Spotties ................................................... 64

9.1. Ticked Tabby ................................................................. 67

9.1.1. Ticked Tabby (All colours) Non Silver (Preliminary)67

9.1.2. Silver Ticked Tabby (Preliminary) .......................... 71

9.2. Mackerel and Classic Tabbies (non Silver) ............... 72


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9.2.1. Mackerel Tabby .................................................... 73

9.2.2. Classic Tabby. ...................................................... 75

9.2.3. Spotted Tabby (non-silver) .................................... 78

9.2.4. Silver Tabby, (All colours) ..................................... 79

9.2.5. British Silver Spotted Shorthair. ............................. 81

10. Tipped – Silver Series ............................................... 83

10.1. Black Tipped ............................................................. 83

10.2. Golden Tipped including all Non-silver Series Tipped.87

11. Colourpointed ........................................................... 92

11.1. Colourpointed, Non-silver, without white spotting ..... 92

11.2. Colourpointed -Silver Series ................................... 101

11.3. Colourpointed & White (Any colour) Preliminary ..... 104

11.4. Silver Colourpointed & White (Any colour) Preliminary106

12. British Shorthair Variant .......................................... 107

13. Eye Colour .............................................................. 109

14. Record Keeping ...................................................... 112

15. Recommendations .................................................. 114

Appendix 1 References .......................................................... 115

Appendix 2 Glossary .............................................................. 116

Appendix 3 Key Genes ........................................................... 117

Appendix 4 Contributing Breeders ....................................... 117


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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.

Fortunately in the UK we have a well respected registration body, the GCCF2

which has an established framework that determines the Registration Policy for

each breed of cat and an official Standard of Points3. To accompany these, the

British Shorthair Group Committee (Breed Advisory Committee) 4 has compiled

this ‘breed specific’ Breeding Policy, the aims of which are to provide the

dedicated and ethical breeder with advice and guidance such that the cats that

they breed, whether for companion, show or breed purposes, are healthy, and

conform to the Breed Standard.

Breeding decisions become easier if breeders develop and manage a breeding

program within which they consider the key points of cat breeding, which are:

• Maintenance and perpetuation of the Breed

• Improvements in quality of the phenotype (appearance) as

measured against the Breed Standard

There is a trade off in any successful breeding program, in order to increase

homogeneity (similarity) we must reduce random variation. Therefore, breeders

must use a robust process of selective breeding.

2 Governing Council of the Cat Fancy 3 SOP 4 BAC

GCCF

BAC

Registration

Policy

Breeding

Policy

Breeders

SOP


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In doing so we run the risk that:

• Undesirable, and even harmful, genetic anomalies will begin to

express themselves

• extremes of physical type may begin to appear

• Inbreeding becomes attractive

In order to achieve the desirable goals, whilst managing the risks the breeders

system should give consideration to:

• The maintenance of a robust, healthy and viable breeding

population of cats.

• Properly documented monitoring of the outcomes of any

mating that produce anomalies.

• The wide range of genetic cat tests that are available.

1.1. Purpose of this Document

The purpose of this document is to provide breeders with some guidance as to

‘best practice’ in breeding the British Shorthair Cat and to ensure that the Breed

Standard, as set out within the Standard of Points, is at the forefront of any

breeding program, together with due regard for the health and integrity of the

breed.

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. Background to the British Shorthair Cat

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

2.2. Breed Description

The breed standard states that;

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

Extract from the British Shorthair Cat Club;

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.

2.3. The Show Cat

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

2.5. Characteristics and Temperament

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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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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3. Breeding System Information

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.

One of the very first terms to get to grips

with is - Phenotype and genotype

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)

3.1. Genetic Make Up

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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Whilst we understand much about how these genes work, there are so many that

even the scientists struggle to understand and complex polygenes9 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.

Red is confusing because it isn’t really a colour in its

own right, it is actually a modifier of the other three

colours, but because the result is a distinct colour then it is

easier if we consider it as a colour

White is not a colour, it is a ‘masking’ gene that masks anything.

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.

Figure 1

9 Genes with small but cumulative effects on the expression of a characteristic

Dominant

Black

Dominant over Chocolate and

Cinnamon

Chocolate

Dominant over Cinnamon

Cinnamon

Dilute

Blue

Dominant over Lilac and Fawn

Lilac

Dominant over Fawn

Fawn


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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.

Because Red is replacing Black, Red is also a dominant colour. It is easiest if we

think of Red and Black as being of equal dominance.

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:

• The Tabby genes

• The Siamese gene

• Silver undercoat genes

• Bi-colour and white spotting genes

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

Table 1- Basic Terms in Genetics

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.

Genes Are 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

Chromosomes Genes are located on the chromosomes. All cats have 38

chromosomes in each body cell, 19 from each parent.

Chromosomes appear in pairs. With the exception of the sex

chromosomes each pair is the same size and shape.

Locus Is the exact position on the chromosome where the gene is

located

Allele Refers to the different forms of a gene

Homozygous Refers to having the same 2 alleles at a given locus on a

chromosome

Heterozygous Refers to having two different alleles at a given locus on a

chromosome

Dominant

Gene

Is one which is capable of expressing its trait even when carried

by only one member of a chromosome pair.

Recessive

Gene

can only express itself when both members of the chromosome

pair contain the gene10

Wildtype The phenotype of the most typical or normal form.


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3.2. Size and variety of gene pool

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.

In common with other pedigree breeds, we do not have an unlimited choice of

parent cats. Figure 2 shows an estimate13 of the number of British Shorthair

queens and studs contributing to the total number of kittens registered in 200915

Breed Kittens

Registered in

2009

Estimated

Queens

Estimated

Stud Cats

Self excluding

Red and Cream

2248 375 81

Silver Tabby Inc

Smoke

762 127 25

Colourpointed 438 72 15

Tortoiseshell16 405 68

Bi-Colour & Tortie

and White

322 54 11

Non Silver Tabby &

Red & Cream Self

302 50 17

Tipped (Silver & non

Silver)

204 34 7

Cinnamon & Fawn 29 5 1

White 23 4 1

Other 682 114 23

TOTALS 5,414 903 181

Figure 2 – Estimated Breeding Population

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

Robinson’s Genetics’ section on inbreeding opens with

“Inbreeding is an emotional topic for some people”

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 policy17 provides a

section on inbreeding.

In line with our ethos of keeping this simple, it is virtually impossible for breeders

of the British Shorthair to avoid inbreeding. Data provided by Robinson18 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.

Within these constraints, breeders dedicated to producing healthy kittens will

always have to consider to what extent their chosen cats are related.

The following table provides an explanation of some of the terms used in

breeding.

Term Simple Explanation % Inbreeding coefficient

calculated19

Inbreeding This is the mating of closely related cats. Such as Father to Daughter, Mother to Son and Brother to Sister

25%

Line-

breeding

This term is not used by geneticists20, and is described as

a ‘less intense’ form of inbreeding.

But it is still breeding within a family line and may include Half Brother to sister, Aunt to Nephew,

Grandparent to Grandchild.

Single First Cousins

Single First Cousin to Second Cousin and Single Half First

12.5%

6.25%

3.13%

1.56%

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

Single Second Cousin

Some breeders would consider that mating cats that are related by at least one common ancestor within the first

three generations of their parents as line-breeding.

Outcrossing This is when the two parents are unrelated.

Realistically, in pedigree animals, this condition almost

impossible to meet. Especially within a ‘closed’ gene pool such as the British.

Breeders, often refer to an outcross when common

ancestor does not occur behind either queen or stud within a four /five generation pedigree. (i.e. when any

inter-relatedness cannot easily be seen)

Breed True Two parents with a desired phenotype produce

offspring of that same phenotype exclusively

Grading Up A rapid improvement of the quality of breeding

stock by mating inferior with superior strain.

Back Cross The mating a son or daughter to one of the

parents. Either to continue a ‘Grading Up’ process

or to capture a recessive gene

Inbreeding Depression

The loss of Hybrid Vigour.

Figure 3

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.

In order to understand why breeders would wish to ‘inbreed’ we must go back to

our basic understanding of inheritance, the gene is the basic physical and

functional unit of heredity. Genes, which are made up of DNA, act as instructions

to make molecules called proteins.

Every cat has two copies of each gene, one inherited from each parent. Alleles

are forms of the same gene with small differences in their sequence of DNA

bases. These small differences contribute to each cat’s unique physical features.

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:

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.

This is caused by a switch in the genetic makeup from a heterozygous state, to

one which is ‘homozygous’

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.

Health

Temperament

Reproductive Ability

Coat Colour & Type


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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.

The GCCF recommend that breeders calculate the inbreeding coefficient – If we

look at Figure 3, we will see that we can add a ‘percentage’ value to the offspring

of any mating we do. The calculation works on a ‘path’

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.

Short Crisp Coat

Short Crisp Coat

Short Crisp Coat

PKD/HCM PKD/HCM PKD/HCM


19

The following ‘traffic light’ might be useful for breeders able to calculate the

coefficient.22

% Coefficient

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

too far is too look out for signs of Inbreeding Depression, some of which are23:

• Males showing signs of reduced ‘sex drive’ or fertility

• Females failing to come into heat or failing to conceive

• Falls in average litter size

• Low birth weight

• An increase in stillbirths

• High kitten mortality

• An increase in abnormal kittens

• Kittens failing to thrive or grow into healthy good sized adults

• Signs of poor immune systems leading to an increase in infectious

diseases.

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.

22 Taken from GCCF Breeding Policy 23 Taken from various sources including Robinson

1-25%

• ACCEPTABLE• Over 8 Generations (where only 5 generations 1-17%

26-39%

• BE CAREFUL• Over 8 Generations (where only 5 generations 17-27%)

40-50%

• STOP• Over 8 generations (Where only 5 generations 28-33)


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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 following advice might help inform a

‘good breeding program’

3.3.1.Best Practice Advice and Recommendations

• Breeders should avoid inbreeding as much as possible.

• Stud cat owners should ask for copies of the pedigree of any queen

wishing to be mated.

• Novice breeders should always take advice from a mentor.

• Breeders should only use inbreeding as a part of their breeding system

when they are certain that they have the ability and experience to

recognize early signs of inbreeding depression. (and the courage to make

any necessary corrections)

• Kittens from closely related parents should not be sold to novice breeders.

• Remember breeders computer programs can only feedback on data input

– if only 4-5 generations are available for a breeding coefficient calculation

– the program cannot calculate the inbreeding of any missing generations.

• Ensure that you understand the pedigree of any cat you might wish to buy

for breeding or use at stud, avoid kittens with pedigrees where any one

ancestor is over represented.

• Avoid over use of ‘fashionable’ stud cats

• Breeders may work in groups and agree to avoid any inbreeding within say

three or four generations of a pedigree.

• Importing new bloodlines will refresh the gene pool. However, check the

pedigree to ensure that the overseas breeder is not breeding from recently

purchased UK stock.

• Breeders to accept collective responsibility for the breed health and

integrity.

The good news is that the BSH does not show signs of dangerous inbreeding at a breed level.


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4. Type

This section is intended to add additional information to inform breeders as to the

phenotype24 we should all be aiming for.

4.1. Overall

The British Shorthair cat is compact, well balanced

and powerful, showing good depth of body, a full

broad chest, with generous, strong shoulders standing

four square on short thick legs, with strong hips. He

has rounded paws, with a tail thick at the base with a

rounded tip. A muscular cat with an alert appearance

and in perfect physical condition.

We are looking for a cat that has an overall body

shape that is a series of curved or rounded shapes

from his head and paws to the tip of his tail.

4.2. The Head

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

The Breed Standard calls for a nose that is short,

broad and straight. In profile, a rounded forehead

should lead to a short straight nose with a nose

break that is neither too pronounced nor too

shallow.

Almost all experienced breeders of the British

Shorthair will know that ‘short, broad and straight’

seems so simple and yet the reality is that getting

the British Shorthair nose and forehead right is

frustratingly difficult.

24 The ‘visible’ cat


22

Keeping the nose short, with a balanced break is just one of the juggling acts

most breeders have to perform.

Irrespective of fashion the extreme Brachycephalic look is not a feature of

the British Shorthair25.

4.4. Chin

The Breed Standard is very clear, a strong, firm and deep

chin is essential.

Any deviation from this to be considered a fault.

The bite MUST be level, the tip of the chin to line up with

the tip of the nose in the same vertical plane.

Breeders seeking to emphasize the roundness to the head

seek to create a slightly rounded chin and powerful jaw.

4.5. Ears

Breed Standard asks for, small, rounded at the

tips. Set far apart, fitting into (without distorting)

the rounded contour of the head.

External ear to be well covered with fur, internal

furnishings not to be excessive.

Getting those ears right presents a real challenge

to our breeders, why oh why do ears want to be

large and sitting high on the head?

Experienced breeders remove the ear tufts for

showing to accentuate the roundness and

neatness.

4.6. Coat

The SOP26 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


23

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

Breed Standard says,

large, round and well-

opened.

Set wide apart with no

tendency to Oriental

shape. No squint.

For some breeders it is all in the eyes – not just colour - but shape and set.

4.8. Body

The SOP says - Cobby type with

short level back.

Low on legs with broad deep chest.

Equally massive across the shoulders

and the rump.

Medium to large, but not rangy.

We are looking for cat with hips and

shoulders of the same width.

A well balanced cat will also have a firm body, without sagging. Breeders are

looking for good bone and a sound muscular body.

4.9. Legs & Paws

The Standard asks for; Short strong legs.

Paws round and firm. Toes carried close, five on each

forefoot (including dew-claw) and four on each back

foot.

That gentle roundness carries right through even to

the feet.


24

4.10. Tail

According to the SOP the tail should be thick and of medium length, thicker at the

base with rounded tip.

4.11. Size and Weight

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.


25

5. Health

5.1. Genetic Defects

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

Polycystic Kidney Disease – PKD, a gene mutation which causes enlarged

kidneys composed of dilated cystic channels, resulting in early kidney failure and

death. This mutation has been found in the British Shorthair. Fortunately simple,

effective DNA tests are available. These can be undertaken by your veterinary

practice or simple swab tests can be carried out at various laboratories27

5.1.2.HCM

Hypertrophic cardiomyopathy (HCM). Sadly, according to the Feline Advisory

Bureau a familial incidence of HCM has been reported in the British Shorthair

breed. Whilst this may be due to an inherited gene defect, as yet the defective

gene has not been identified in this breed. (See screening scheme info on

www.fabcats.org/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

http://www.fabcats.org/hcm/index.html


26

5.2. Blood Groups and Neonatal Isoerythrolysis

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.

It is estimated that 92 % of the UK domestic shorthair feline population are blood

group A. However, only 47% of the British Shorthair population are group A, the

remainder being blood group B or the very rare AB. In recent years blood group

‘mismatch’ has been identified as being responsible for some cases of fading

kitten syndrome.28

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

type in their own homes using non-invasive inexpensive swab tests.29

The following detail is extracted from data provided by UC Davis.

The A and B blood groups are genetically determined.

UC Davis has proposed the following ‘naming’ system:

Nomenclature Blood

Group

Genetic Make Up Note

A A A/A, A/a or A/b Dominant to B

a AB Probably less than 1% of

all cats.

b B b/b

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.


27

Results reported as:

Test

Result Blood group status

N/N Cat is Type A or Type "AB"

N/b Cat is a carrier of B factor; serotype could be Type A or Type

"AB"

b/b Cat is Type B

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.

On her website Dr Diane Addie makes the following statement regarding Ab

Queens mated to Ab studs and thus potentially carrying B group kittens -

“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.”

Despite much research amongst UK BSH breeders we have no evidence that

kittens from Ab to Ab matings are fading from NI. However, record keeping is

vital and if breeders experience any fading kitten in Ab to Ab matings blood tests

may help to inform future matings.

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)


28

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

1. Breeders should use available technology to inform their breeding

programs. (ie, Blood Group Tests)

2. Inexperienced Breeders with B group queens should seek to mate those

queens to B group studs.

3. Kits to test blood group using placental blood are available.30

30 These are available to Veterinary practices from the Oxford Cat Clinic.

Ab, or B Stud

Ab Queen

Offspring will be either blood group A, Ab or B Risk Increases

For B group kittens

A, or Ab Stud B

Quee

Offspring will be either blood group Ab or B

Any non B kittens may be at risk

B Stud B

or A Queen

Offspring will be either blood group B or Ab Not at Risk

It is the antibodies from B group dams that represent the greatest risk to A kittens

Figure 5


29

5.3. Brachycephaly

Brachycephalic is a word that is hard to say, meaning short headed or broad

headed, wherein the length of the cranium is shorter than the width. This gives

the top and sides of the cranium a round shape, often referred to as 'apple-

head'.31 Whilst we are very happy to have that lovely apple-head description, if

one were to Google ‘Brachycephalic’ all sorts of distressing articles and images

appear, especially in relation to poor breeding practice in the dog world.

As the British BAC, our remit is

limited to the welfare of the British

Shorthair, and whilst we are

attracted to a short, broad nose we

are also very proud of the

reputation that the British

Shorthair has for health and for

being as natural as possible for a

pedigree cat.

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.

Keeping your breeding ‘team’ at

the right weight takes patience

and skill, but the health benefits

to your cat are worthwhile

31 Wikipedia


30

5.5. Infectious Disease

There is much information available for breeders regarding infectious diseases

both viral and bacterial. The feline, unlike the canine, is not a pack animal, and

there is no doubt that the feline physiology cannot cope with the stress that can

accompany some cattery environments.

We recommend that breeders acquaint themselves with the following diseases,

such that they understand symptoms, treatments, impact and remedy.

Table 2

Viral Bacterial Parasites

Feline Calicivirus (FCV) Feline Chlamydia Fleas

Feline Panleukopenia (Distemper)

Campylobacter Ear Mites

Feline Leukemia Virus (FeLV)

Bordetella Bronchiseptica Hookworms,

Ringworms and Tapeworms

Feline Rhinotracheitis (Feline Herpesvirus)

Toxoplasma Coccidia

Feline Immunodeficiency Virus

Streptococcus Tri Trichomonas

Corona (FCoV) and Feline Infectious

Peritonitis

Staphylococcus Giardia

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.


31

6. The Self Coloured British Shorthair

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,

we must now accept that genes can mutate32 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 eumelanin35 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) allele36 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


32

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.


33

6.1. The White Self

Introduction

A beautiful cat with a

sparkling non-silver white

coat and dramatic eye

colour, the white self is not

often seen on the show

bench today, although

when one does appear it

generates a lot of interest.

In the early days of the

British Shorthair they were

very popular working cats

often found in Flour Mills!

It is estimated that the White Self represents less than half a percent of total

kitten registrations.

With a legacy of hearing problems and a need to understand the dominance of

the genes for white coats the white self cat is a breed for the experienced and

ethical breeder.

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.

Genes for Coat

The gene (W) that causes a cat to have a white coat is a dominant masking

gene37. A white cat can be a homozygous white (Ww. 2 white genes) or

heterozygous (Ww. one white gene/one nonwhite gene).

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


34

It is, in rare cases possible to have a cat with

a naturally white coat without this gene, as an

extreme form of white spotting38, some small

non-white patch usually remains, if only during

kitten-hood.

This sweet baby aged 11 days carries the

mark she was born with. The pigment mark

shows how the pigment started to ‘migrate’

before the W gene caused the process to stop.

Eyes

Breeders are seeking to achieve the following eye colours:

• Deep Sapphire Blue

• Orange, preferably copper, deep orange or deep gold

• Odd eyed, one each of the above two striking colours

As the white coat may be masking many coat colours or patterns including the

Himalayan gene40 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.

Nose and Paw Leather

Bright Clean Pink for the British White

The Show Cat

Breeders are looking for a white ‘self’ which should have a dense pure white coat

un-tinged with yellow. Kittens may have slight blue/black markings on their

heads which should disappear when they reach maturity.

Specific Health – Skin Care

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.


35

There is much data available giving statistics, the following data paints a picture.

• Deafness in the general cat population is extremely rare

• 5% of the general cat population is White

• Estimates are that about 50% of these cats are deaf in one or both ears

and when one or two eyes are blue the chances of deafness rises to nearer

80%

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.

Best Practice and Responsible Breeding

• GCCF registration of white male stud cats requires a BAER test.

• The registration policy for the White self restricts full registration to

matings between White and Self/tortie.

• Responsible breeders, of white cats will breed only from cats that have

undergone the BAER test which tests for bilateral hearing.

• Caring and ethical breeders can promote awareness of the test on their

websites to raise the profile of the test in cats and hopefully steer people

away from breeders that may not have the interests of the unborn kitten

at heart.

• Breeders should offer copies of their BAER test certificates to people

buying their kittens.


36

Recommendations White Self

• All White British Shorthairs on the active register irrespective of gender

should be BAER tested.

• White x White breeding should be discouraged (research suggests that

cats that carry two copies of the W gene (WW = homozygous) are more

likely to produce deaf offspring.

• As any cat can be covered by the white gene, Breeders should mate cats

as close to the breed standard as possible to produce whites that are as

good as they can be.

• Breeders looking to achieve a higher incidence of odd eyed whites in their

breeding programs might like to consider experimenting with Bi-colours

with UNEAVEN facial markings41

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.


37

6.2. Black and Blue Self

Introduction

The British Blue almost needs no introduction

being the most loved of all pedigree cats in the

UK today.

Together with the Chocolate and Lilac series, the

British Black and Blue account for almost 42% of

all BSH registrations42

Genes for Coat

Colour Agouti Black Dilute Orange White

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 Eumelanin43 which is black. Black melanin or eumelanin granules are

thought to be oval in shape and absorb almost all light.

The B (black/brown) gene has a

lightening effect on eumelanin. The D

(dense/dilute) gene affects the density

of pigment in the hair and affects the

eumelanin, the effect is a washed out

version of the original colour and the

Jet Black of the British Black becomes

the light to medium blue that is so

desirable.

Black is a dominant gene for

colour

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


38

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 Show Cat

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.

The British Blue carries the

banner for the breed and

competition is very tough.

Breeders work very hard

to achieve the balance of

coat; texture and colour,

eyes; shape and colour,

ears; size and set; body;

size and weight and

because this is such a

popular colour, judges all

know just how good this

cat can be. The SOP calls

for a light to medium blue

coat in the show cat and

heavy silver tipping is a

fault.

There should be no tabby markings in either cat on the adult show bench.

Best Practice, Responsible Breeding and Recommendations

• Breeders seeking to breed from black or cats with other genetically

dominant traits should ensure that they understand how the genes for

colour and dilution are inherited. Breeding from a black cat that does not

carry the genes for dilute or chocolate/cinnamon will only produce black

cats.

• Just because the British Blue enjoys huge popularity both on and off the

show bench – breeders should not compromise on type or quality.


39

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 is much debate amongst established breeders as to whether what we see

today is a relatively new phenomenon or something that has ‘always’ been there.

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.

As genetic information becomes more available we will, hopefully, learn more

about the genetic make-up of our most popular colour of British. In the

meanwhile breeders are free to choose the polygenic phenotype of their breeding

programs, whilst our Judges have to make their own interpretation of the

definition ‘heavy silver tipping’


40

6.3. Chocolate or Lilac Self

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’ Seal44 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.

Genes for Coat


Chocolate aa bb or bbl D- oY/oo ww Lilac aa bb or bbl dd oY/oo ww

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.

When we add two copies of the dilute gene to

the Chocolate cat the result is a delicious lilac

shade.


Nose leather and paw pads should be a Pinkish lilac.

44 Seal is the colour description for the Black Colourpointed


41

The Show Cat

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.


42

6.4. Cinnamon or Fawn Self

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.

Genes for Coat


Cinnamon aa b¹ b¹ D- oY/oo ww Fawn aa b¹ b¹ dd oY/oo ww

Although we are attempting to keep the genetic explanations as simple as we

can, the truth is feline genetics are complex, and the genes that we do

understand and can test for work in complex ways with genes that we cannot test

for and can only really ‘prove’ by experiment and testing our theories.

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.

History of the Cinnamon colour45

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.

Cinnamon arrives in the British Section

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

Oriental46.

45 Interesting reading includes - The Abyssinian Cat, by Vicky Hakin Smith 46 Malsome British Shorthairs have some background information.


43

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.

The Oriental section had originally out-crossed to the Abyssinian47 but selective

breeding of the recessive ‘a’ gene produced a cat with all the magnificent colour

of the Abyssinian in a non ticked form.48

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

The first outcrosses started, and what a task these

brave breeders set themselves – the Oriental50 is a

beautiful cat, but the phenotype is almost as far from

the British as it is possible to get.

It takes passion and a great deal of patience and

many many years to convince all those polygenes to

make the shift from the Oriental to the British and we

should be proud of those dedicated breeders who

break the first ground


Nose leather and paw pads should be a pink or

cinnamon brown.

A lovely example of the Fawn Coat.

Now, does that coat look as if it has an

underlying non-agouti ticked base?

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.


44

The Show Cat (Preliminary Status)

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.


45

6.5. Red or Cream Self

Those budding geneticists amongst us, will have remembered that the (A) to (a)

(agouti to non-agouti) mutation has little effect on the (O) allele51 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

Red Self through the Preliminary Status52 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.

Having explained eumelanin53 we now need to get to grips with the only other

feline pigment – phaeomelanin the pigment for red. Essentially what the ‘O’ gene

does is get rid of any eumelanistic 54pigment 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.

Each cat has two chromosomes for ‘SEX’ or gender –

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

The reason this is important information is because the genes for colour55 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 X to carry or

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.


46

The following table may help:

Example XY Genotype Phenotype

XX Black on both X

Chromosomes

Black Female

XX One black & one Red X Chromosome

Dominant Tortie Female

XY Black on X

Chromosome

Black Male

XY Red X Chromosome Red Male

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.

A female cat gets an X from each of her parents.

Genes for Coat

Colour Agouti Black Full

Coat

Dilute Orange White

Red aa C- D- OY/OO ww Cream aa C- dd OY/OO ww


47

This picture shows a delicious cream female

at the top of her game.

The delicate cream

tones that make this girl blend into her blankets

are exactly what breeders are trying to

achieve. However, it is

not easy to produce this pale, balanced,

colouration.

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.

The Show Cat

The cream colour should be an even-toned cream, with as few markings as

possible and no white hairs. Paler shades are preferred. Kittens are forgiven any

ghost tabby markings.


• A popular mating to produce good examples of the cream is blue-cream

females mated to blue or cream males.

• It is thought that producing cream kittens from a red or dominant tortie

parent carrying dilute may contribute to a ‘hot’ cream. However, some

breeders have found the reverse to be true!

• Because the Red-Self is still at the preliminary stages, breeders are still

working ‘against’ the genes that make the agouti markings appear on non-

agouti cats. Some of the breeding ‘trials’ underway include:

• Working with ‘Rufousing polygenes’ 56which seem to blur the contrast

between the pattern and the ground colour.

• Breeding self cats with an underlying base pattern that is ticked tabby

rather than classic, spotted or mackerel57

56 Appendix 3 57

Ticked Tabby (footnote continued)


48

A beautiful youngster

showing an underlying non agouti spotty

pattern

A lovely rich red girl

– so rarely seen

Ticked Tabby (All colours) Non Silver (Preliminary


49

6.6. Tortoiseshell (all colours)

This is a very popular variety of the British Shorthair, especially amongst

breeders who find the multi-colour useful in breeding programs and who also

appreciate just how difficult it is to produce a ‘correctly’ patterned ‘tortie’ for the

show-bench

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.

A female cat inherits an X chromosome from each of her parents.

If neither parent has the gene for ‘red’ then the colour of the female kitten will be

determined according to the rules for colour inheritance58

If the father is red or cream,59 then his daughter can only receive that colour

from him and so she will have one X chromosome with the red colour carried. If

her mother is also red or cream she will have two X X and she will be a red or

cream female. If the dam is Black, Blue, Chocolate, Lilac, Cinnamon or Fawn,

then we will have a tortie female.

What happens next is complicated,60 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

see this visibly in skin cells as they produce either red or black61 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.

Genes for Coat

Tortie Colour

Agouti Black Full Coat

Dilute Orange White

Dominant aa B-/bb-/bb¹ C- D- Oo ww

Dilute aa B-/bb-/bb¹ C- dd Oo ww

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.


50

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.


Pink and/or the colour corresponding with the coat colour.

The Show Cat

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.


• The tortie is also what is called an obligate heterozygote. Therefore, she

cannot breed true. And as we have learned that the distribution of colour

is largely out of the breeders hands, we should seek to breed our torties’

using the best for health, temperament and type parents that we possibly

can. In doing so even if we fail to achieve that oh so desirable mingled

coat we will at least produce kittens that are otherwise correct for eye

colour, coat etc, that will be happy, healthy kittens that are a credit to our

breeding.

• Breeding from an otherwise ‘poor’ tortie with good mingling will probably

just end in disappointment.


51

• Experienced Breeders might like to experiment with the Rufous

polygenes,62 to see if they can see an improvement in the richness of the

tortie reds.

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


52

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.

A well bred Bi-Colour is an extraordinarily pretty cat causing much admiration.

They can be shown in any of the recognized ‘self’ colours with the addition of

white.

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

Gene63 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

Van Pattern64. The S/s gene is extremely variable, the following diagram65 may

help.

63 Producing an intermediate phenotype in the heterozygous condition 64

Van Pattern (Preliminary) 65 Thanks to Wikipedia

Figure 7


53

In Figure 7 we could guess that:

• Cats with high to medium grade spotting (numbered 7-9) would be

homozygous for the S gene, (SS)

• Cats with low to medium grade spotting (numbered 2 – 4) would be

heterozygous for the S gene (Ss)

• Cats with medium grade spotting (numbered 5-6) could be either SS or

Ss.

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.

At this stage it is worth a shortened and simplified explanation of the geneticist’s

best guess as to how the white pattern is ‘formed’

After mating, and the egg is fertilized, cells change66 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


54

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!!!

Genes for Coat

Bi- Colour Agouti Black Full Coat

Dilute Orange

Spotting

Dominant B aa

B-/bb-/bb¹ C- D- Oo Ss

Dilute B aa B-/bb-/bb¹ C- dd Oo Ss Dominant O

aa

C- D- OY/OO Ss

Dilute O aa

C- dd OY/OO Ss

Eyes

Colour in line with the breed standard for the Self British. Incorrect rim colour or

flecks will be faulted.


Pink and/or the colour corresponding with the patches.

The Show Cat

Judges are looking for patches of one self-colour

and white. The patches must be clear and well

defined, without tabby markings. At least one

third and not more than half the coat to be

white. No scattered white hairs. The coloured

patches must be sound and any blue must not

have silver tipping.

Symmetry of design is desirable, with colour on

the top of the head, the ears, cheeks, back, tail

and flanks. Tail to be fully coloured.

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.


55

Exploding Some Popular Bi-Colour Myths (Just for Fun)

• 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 program68 • 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.


56

7.2. Van Pattern (Preliminary)

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.

Essentially the Van

Pattern British is an

high grade Bi-Colour,

and if we go back to Figure 7 we can see

that the Van Pattern

is a grade 8 or 9

suggesting that the

genotype for the Van

pattern is usually the

dominant piebald or

spotting gene SS.

It is generally accepted that selective breeding can exert influence over the tricky

Spotting gene and his polygenic mates69, 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 form70 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 places71 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


57

Time for a little lesson in the inheritance of genes!

Figure 8

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.

1

Self White Sire

aa Ww ss B-D-

2

Van-Patterned Dam

aa ww SS B-dd

3

Self White Male Kitten

aa Ww Ss B-Dd

3

Self White Male Kitten

aa Ww Ss B-Dd

4

Blue Self Female

aa ww ss B-dd

5

Blue Self female Kitten

aa ww ss B-dd


58

Genes for Coat

Colour Agouti Black Full Coat

Dilute Orange Spotting

Dominant aa B-/bb-/bb¹ C- D- oY /oo Ss or SS Dilute aa B-/bb-/bb¹ C- dd oY/oo Ss or SS Dominant O

aa

C- D- OY/OO Ss or SS

Dilute O aa

C- dd OY/OO Ss or SS

Eyes

The breed standard calls for eye colour to be

deep gold or copper or orange. (there is

anecdotal evidence of deafness in blue eyed

van patterned cats, this breed standard is a

wise example of selective breeding)


Pink or corresponding body colour

The Show Cat

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.


59

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 make73, 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 brings74, 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.

Genes for Coat

Tortie Colour

Agouti Black Full Coat

Dilute Orange Spotting

Dominant aa B-/bb-

/bb¹ C- D- Oo SS or Ss

Dilute aa B-/bb-

/bb¹ C- dd Oo SS or Ss

73 See Tortie Section 74 See Bi-Colour Section


60

The Show Cat

A good show girl will have patches of

base self-colour, with either red or

cream and white. The patches must be

clear, well defined and balanced without

mingling or tabby markings. For the

non-Van patterned at least one third,

and not more than half the coat to be

white, with patches of colour on top of

the head, ears, cheeks, back, tail, legs

and flanks. Tail to be fully coloured.

(Van patterns to have markings as per Van pattern.

Symmetry of design is desirable, with colour on the top of the head, the ears,

cheeks, back, tail and flanks.


Pink or corresponding body colour

Best Practice and Recommendations.

• Although we have seen in the ‘Tortie’ section that Breeders cannot

influence that tortie mingling, it seems that the ‘S’ gene can be ‘biddable’

so have some fun and just see if the bigger the white areas, the bigger

and better defined the separate patches of blue and cream become.

Oh! – And don’t forget that type matters!!


61

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 hair75. • 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 phaeomelanin76 more than the pigment eumelanin77. • 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.

The problem is that the Silver78 (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

75 Not like the S gene that affects melanin getting to skin cells

76 Orange 77 Black 78 Inhibitor


62

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

information, as it can be extremely difficult for even the most experienced

breeder to discern the Silver, especially in the Tabby colourpointed series.

Genes for Coat

The Smoke may be bred in any of the self or tortie colours.

Eyes

Large and round, should be copper, orange or deep gold in colour.

Nose Leather and Paw Pads

Corresponding with coat colour and mingled with pink for torties.

The Show Cat

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.

Recommendations and Best Practice

Smoke Agouti Black

Full Coat Dilute Inhibitor Orange

Dominant B

aa B-/bb-/bb¹ C- D- I

oY/oo

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


63

• 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.


64

9. Tabbies and Spotties

In the section on The British Self, we said that in order to understand the genes

for the ‘Self’, we must first understand the Agouti (A)79 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 studies80 have

suggested that there are in fact three separate genes at different loci81

Therefore, we feel it will be easier if we discard the use of T except for the Ticked

Tabby and use Mc and Sp82 instead83.

The following table may be useful for future reference.

Tabb

y

Pattern

Former

Gene

code

Updated

Gene code

Notes

Ticke

d

Ta

Ta If this gene is present the cat will

show as a ticked tabby regardless of any

other tabby genes they may have.

Mack

erel

T Mc Can only be seen when Ta is not

present. Dominant over mc

Class

ic

tb mc Recessive to Mc

Spott

ed

Ts

Sp

This is a modifier gene, effective only

when Ta is absent. Sp breaks up the

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.


65

patterns of Mc and mc.

Figure 10

It is difficult when producing an introduction to the Agouti cat such as this, to

know just how much detail we need to go into, too much would take up far too

much space and time. Whilst, too little may mean that we do not provide enough

of a basis to allow a proper understanding.

Figure 1184

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.

As the hair grows in the hair follicle eumelanin85 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

to produce phaeomelanin86 , this process is

known as the Agouti shift.

The result is hair shafts with alternating bands of

84 There are many other Genes of Course 85 Black Pigment 86 Red pigment

Genome or Blueprint for the Finished Cat

19 Pairs of Chromsomes

Locus for Agouti Genes

2 alleles

Locus for Tabby Genes

2 alleles

Locus for Spotted

Modifier Genes

2 alleles

Locus for Colour Genes

2 alleles

Locus for Inhibitor Genes

2 alleles

Locus for Masking Genes

2 alleles


66

yellow and black colour, ending with black tips.

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.

At this point it seems prudent to mention something discovered during our

research regarding the Agouti shift87.

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.


67

9.1. Ticked Tabby

9.1.1.Ticked Tabby (All colours) Non Silver (Preliminary)

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.

Genes for Coat

Ticked Agouti Colour Coat Dilute Tabby Orange

Dominant B A- B-/bb-/bb¹

C- D- Ta- oY/oo

Dilute B A- B-/bb-

/bb¹ C- dd Ta- oY/oo

Dom/Tortie A- B-/bb-/bb¹

C- D- Ta- Oo

Dilute/Tortie A- B-/bb-/bb¹

C- dd Ta- Oo

A- can be: AA or Aa D- can be: DD or Dd

C- can be: CC or Cs Ta- can be: TaTa or Tata

Any Tabby Pattern may also be carried see Figure 10

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.


68

This cat shows a ticked pattern showing the stripes

and markings on the head, legs and tail.

This cat is likely to be Heterozygous for the Ticked

Gene Tata

This cat has no tabby markings on either his body or

tail – he is likely to be Homozygous for the Ticking Gene

TaTa.

Eyes

Copper, orange or deep gold

The Show Cat

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.

Necklaces may be broken or unbroken but a lack of these is not a fault.

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.

Markings –the ticking should be of the appropriate colour affording a good

contrast with the base colour. The Base Colour should be even in all areas other

than the lips and chin, which may pale significantly, but should not be white and

should not extend down the chest. Tarnishing or discolouring of the silver base

colour is undesirable.

Figure 14

Figure 12

Figure 13


69

This variety of British Shorthair arrived unexpectedly in 2007, in a litter born to

an experienced breeder88 of Tabby British Shorthair.

The following section is a little lesson in genetics to help explain how this sweet

baby arrived:

A lilac British Shorthair Stud cat imported from Sweden with a genotype of:

Colour Agouti Tabby Black Full Coat Dilute Orange White

Lilac

Sire aa Ta89 bb C- dd oY ww

Was mated to

Colour Agouti Spotty Black Full Coat Dilute Orange White

Choc’-

Spotty

Dam

A-

Sp

bb

C-

Dd

oo

ww

The resulting kitten Inherited:

Genes From Dad From Mom Note

Agouti aa A Mum’s dominant agouti gene prevailed

Tabby Ta - Although non-agouti, the sire’s ticked tabby gene is

dominant for underlying base

pattern.

Spotted - Sp The mother’s spotted gene

will override any other Mc or mc genes.

Black bb bb Chocolate

Full Coat C C Full Coat

Dilute d D 1 copy of dilute so kitten is Chocolate

Orange o o No red gene to have

White w w No white gene to have

88 Mrs Celia Leighton (Porteous) 89 Ticked tabby


70

The only really surprising thing about this entirely natural production of a

delightful chocolate ticked tabby is that it hasn’t happened more often.

Best Practice and Recommendations

• Whilst breeds are in the preliminary stages it is best leaving the

development to experienced breeders and those breeders working in

breeding groups.

• When working with the gene for ticking, try to understand the pedigree

of the cat, remember that any cat that is Homozygous for ticking can

only throw Ticked kittens. (When the cat is Tata, heterozygous, the cat

is most likely to have bars and markings on the legs, tail, face, etc)

• You may be able to tell what colour a tabby is by looking at the color of

its stripes and its tail tip.

• The agouti shift, describes a process where the switch between dark

and light pigment happens suddenly. But there are also variations in

just how often the ‘switch’ turns on and off, Ticked tabby breeders are

looking for as many bands of colour on each hair shaft as possible, this

can vary between 4 and 6, but up to 8 have been reported90.

• The Wide Banding91 gene can play havoc with the ticked bands,

causing the dark hair to the tips – avoid mating shaded or tipped cats

in a Ticked breeding program.

• Breeders should experiment for the Rufusing gene, it is evident from

the great variety of colours in the Agouti ground colour that polygenes

are able to produce coats that range from rich creamy oranges through

to pale washed out greys. Mating cats with good ground colour to cats

with good ground colour is advisable.

• Remember that although this variety is new to the British, other

breeders have extensive experience of working with the genes for

Ticking, personal knowledge transfer is invaluable.

• Breeders may find that despite their best efforts, the ticked gene will

cause the agouti colour on the muzzle and throat to pale almost to

white.

90 Chaos and Confusion genes may be at work see Tipped section. 91 Theoretical at present


71

9.1.2. Silver Ticked Tabby (Preliminary)

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.

Despite the British input to other breeds,

the Silver Ticked is not yet a common

sight in the British Section.

However, they are being bred and indeed

shown – A rather lovely tortie-silver has

appeared at several GCCF shows.

92

Eye Colour

Green or Hazel.

Genes for Coat

As the non-silver Ticked plus the dominant (I). The gene

for Silver (I) changes the rich Agouti ground colour to a

cooler sparkling silver.

The Show Cat

In silvers the colour of the ticking may lack warmth and

this should not be penalised.


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

results93

92 Photo by Milt Webb 93 Of course our gene pool is tiny as yet

Figure 15 An American

Silver Ticked Tabby


72

9.2. Mackerel and Classic Tabbies (non Silver)

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

stripes flashed throughout the ticked coat94. 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 shift95. 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).

The face of the British Tabby can

be beautiful, and the markings

should be clearly defined and

dense. 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.

Muzzle and Throat

It is worth mentioning the colour of the chin and lips of the tabby. Sometimes

there can be a reduction in the pigment in this area, such that the ground colour

can appear almost white. Whilst we can expect that the ground colour will pale

from the chin to the throat, breeders should try not to use cats that show ‘white’

in this area in their breeding programs.

94 Probably for camouflage 95 If indeed it is phaeomelanin!

Brown Classic tabby at 6 weeks


73

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).

The stripes are always vertical, and gently

curving, on the body, ringing the tail, and

lining the legs. The Mackerel pattern can be

found in any of the self colours, but the

pattern of the stripes will always remain the

same.

Genes for Coat

Mackerel Tabby

Agouti Colour Pattern Dilute Tabby Pattern

Orange

Dominant B A- B-/bb-

/bb¹

C- D- Mc- spsp

tata

oY/oo

Dilute B A- B-/bb-

/bb¹

C- dd Mc- spsp

tata

oY/oo

Dominant O A- C- D- Mc- spsp

tata

OY/OO

Dilute O A- C- dd Mc- spsp

tata

OY/OO

Dom/Tortie A- B-/bb-

/bb¹

C- D- Mc- spsp

tata

Oo

Dilute/Tortie A- B-/bb-/bb¹

C- dd Mc- spsp

tata

Oo

A- = AA or Aa Mc- = McMc or Mcmc C- = CC or Ccs

Eyes

In copper, orange or deep gold, with no trace of green

The Show Cat

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,


74

may be complete or broken with the tip of the tail being the same colour as the

markings.

Markings of the appropriate colour to be clearly defined and dense in colour,

without brindling, affording a good contrast with the ground colour. The colour

should reach as near the roots as possible.

Ground Colour, should be even, in all areas other than the lips and chin, which

may pale significantly but should not be white.

Best Practice and Responsible Breeding and Recommendations

• 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!


75

9.2.2.Classic Tabby.

Also known as the blotched tabby, this tabby pattern

is recessive to the Mackerel pattern. The ticked hairs

alternate with the solid hairs making the pattern that

is so much loved by breeders. The recessive, mutant,

mcmc causes the dark pigmented hairs to form

thicker stripes, which travel horizontally across the

cat's body, often swirling and creating spirals.

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

breeds on the show-bench96

Genes for Coat

As for Mackerel tabby, just substitute Mc- for mcmc.

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 selecting97 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.

‘Best Friend Genes’ (Hypothetical)

1. Rufous genes – Everybody wants to be in with this gang (except perhaps

silver breeders)98 – this lovely group are thought to make the agouti pale

colours warmer and richer and dark colours darker – what more could any

tabby breeder want from her mates? It seems that Rufous modifiers can

get more intensive over successive generations. Fabulous news for

breeders’ intent on selection over many generations.

2. The Tabby Pattern Size modifier - Breeders may recognize this group of

‘friends’ who seem to alter the Classic Tabby Pattern by creating pale

areas within the blotched markings.

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.


76

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

Spotty99 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.

See how the

colours on these

sweet litter

brothers differ from

each other.

The Show Cat

Breeders are after four things:

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.


77

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

may benefit from some separation100.

100 Experienced breeders will already have separate breeding programs.


78

9.2.3.Spotted Tabby (non-silver)

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!

Genes for Coat.

Spotted Tabby

Agouti Colour Pattern Dilute Tabby Pattern

Orange

Dominant

B

A- B-/bb-

/bb¹ C- D- Mc-

mcmc Sp- tata

oY/oo

Dilute B A- B-/bb-

/bb¹ C- dd Mc-

mcmc

Sp-tata

oY/oo

A- = AA or Aa Mc- = McMc or Mcmc C- = CC or Ccs Sp- = SpSp or Spsp

Note: The genes for coat are as for Mackerel tabby, but spsp becomes Sp-

Naturally occurring Spsp is the spotting or modifier gene. Sp is dominant over Mc

– and mcmc and causes the basic

tabby patterns to break into spotted

patterns. Mackerel tabby patterns

will have more, smaller spots which

run in vertical lines. The Classic

tabby will have larger spots that are

scattered more randomly.

A beautifully marked brown spotty

baby

(Now do we think these spots are Mackerel or Classic based?)

The Show Cat101

The pattern on the body and legs

should consist of numerous, well

101 The pattern is often less obvious in the dilute cats


79

defined, oval, round or rosette-shaped spots which follow the tabby pattern in

distribution.

9.2.4.Silver Tabby, (All colours)

The Silver Tabby and Spotted British Shorthair is the second

most popular variety, with more than 14% of registrations

being this wonderful cat.

Silver Tabby Breeders aim to produce the Classic and

Spotted Patterns in all the usual Self British Colours.

The Silver tabby is essentially the same cat as the Classic

tabby with the addition of the dominant Inhibitor gene, (I)

introduced in the Smoke section. But this time the Inhibitor

gene gets to come out to play with the Agouti gene (A) and

with quite spectacular results.

In the Silver Tabby, agouti hairs have colour or pigment at

the tip, but the remaining hair shaft is stripped of agouti

colour by the I gene, leaving a sparkling silver grey. The

solid hairs have normal colouration. Breeders are seeking

the greatest contrast between the now almost-white agouti

hairs and the markings made by the fully pigmented patches of hair.

Genes for Coat

Silver

Tabby

Agouti Colour Pattern Silver Dilute Tabby

Pattern

Orange

Dominant B A- B-/bb-

/bb¹ C- I- D- Mc- spsp

tata oY/oo

Dilute B A- B-/bb-

/bb¹ C- I- dd Mc- spsp

tata oY/oo

Dominant O A- C- I- D- Mc- spsp tata

OY/OO

Dilute O A- C- I- dd Mc- spsp tata

OY/OO

Dom/Tortie A- B-/bb-

/bb¹ C- I- D- Mc- spsp

tata Oo

Dilute/Tortie A- B-/bb-

/bb¹ C- I- dd Mc- spsp

tata Oo

A- = AA or Aa Mc- = McMc or Mcmc C- = CC or Ccs Sp- = spsp I=II or Ii


80

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.

The Show Cat

Just to keep us on our toes the British Breed Standard for the Classic Tabby

pattern is more exacting than that in the American Shorthair. The pattern

requirements for the Silver Tabby are as for the non-silver tabby. The key to the

show cat is clear, sharp markings, against an untarnished silver backdrop.


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.

The good news is that all of the above can be dealt with by careful breeding

selection. But, as with all breeding programs, it is not a quick fix, breeding a high

quality line takes years of patience.


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9.2.5.British Silver Spotted Shorthair.

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.

Genes for Coat

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.

The Show Cat


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Judges are looking for SPOTS, lots of them, and please make them as dramatic as

possible.

3 A beautiful Queen snoozing on show afternoon.


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10. Tipped – Silver Series

10.1. Black Tipped

In the 1970’s a Chinchilla (longhair) was mated to a British Silver Tabby102. 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.

But, that would

be taking the

easy way out.

If we start with the Simple Genes for Coat:

Black

Tipped

Agouti Black Full

Coat

Dilute Silver Tabby

Pattern

Orange

A- B- C- D- I- Ticked Ta- Mackerel Mc- tata Classic mcmc tata Spotted Sp- tata

oY/oo

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

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


84

geneticists to refer to a hypothetical gene commonly referred to as the Wideband

gene (Wb). In essence it seems that the Wideband gene works with the agouti

gene and causes a hiccough in the agouti shift, such that the on /off switch turns

the eumelanin switch on, then ‘off’ suddenly but leaves the phaeomelanin switch

‘on’ for longer, causing a wider than normal agouti band. The Silver Inhibitor

gene then joins the party causing the hair shaft to lose the colour, thus leaving

an almost totally silver cat with black (or other colour) tipped hair shafts.

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.

At this point it may help if we understand Incomplete Dominance –

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?


85

Because it makes discussion easier – we will embrace the theoretical Wideband

gene throughout this breeding policy! But we will also accept that even the

Wideband genes will have help from those polygenes!

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.

The Show Cat

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 colour is restricted to

the ends of the hairs in

the form of tipping and

the undercoat is so pale

as to appear white. 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.

Heavily tipped cats must

not be penalised as long

as the tipping is even and

free from tabby

markings: in fact, they are preferable to cats with so little tipping that they are

almost white.

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 and Paw Pads

Nose Leather - Brick red preferred, outlined in colour appropriate to the colour

of the coat tipping.

Paw Pads - Appropriate to the colour of the coat tipping.


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Some Extra Genetic Information

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 83 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.

We know that the Chinchilla is predominantly based on a Classic pattern

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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10.2. Golden Tipped including all Non-silver Series Tipped.

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.

Overtime, as new generations of kittens are born, breeders gain an increasing

awareness of the genetics at play in their cats and new genetic information

becomes available or is updated.

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!

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.

Eyes - Green. The skin outlining the eyes to be black/dark brown.

Nose Leather - Brick Red, may be outlined in black/dark brown.

Background and Genes

Genes for coat

Golden

Tipped

Agouti Black Full Coat Dilute Silver Tabby

Pattern

Orange

A- B- C- D- ii Ticked

Ta-

Mackerel

Mc- tata

Classic

mcmc

tata

Spotted

Sp- tata

oY/oo

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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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

• Genetically Black, (with

Chocolate and Cinnamon

Possible)

• He could carry dilute, so blue,

lilac and fawn offspring

possible.

• He is Agouti for sure, but he

may be Homozygous AA or

Aa.

• Somewhere he will have a

tabby base pattern (we know

this because all cats do) – and

that will probably have been

quite obvious to his breeder

when he was a kitten – but

without carrying out some

test-matings we just don’t

know what he carries

But how we get from the Brown Tabby to the Golden is a subject of much conjecture.

Five Theory’s

• There is a gene for Golden

• The Inhibitor gene forces a tabby pattern to the coat tips, but when the silver

gene is not passed on to offspring, the result is a ‘Golden’ tipped instead of

‘Silver’ tipped cat.

• A Wide-band gene, exploited by Chinchilla breeders, causes a wider agouti

band that allows us to see more of the ‘yellow’ pigment.

• Years and Years of breeders working on producing the silver and golden

Persians have perfected a polygenic mix of both agouti and inhibitor genes.

• Rufousing Polygenes working in conjunction with any of the above!

A Gene for Golden?

Some breeders working with the Golden gene do

believe that there is a separate gene for the Golden

colour -

Can this Golden gene be separated from the Agouti

gene? –

Perhaps this is something that breeders could work

on?

104 This is also true for the Black Tipped – but add the Inhibitor Gene


89

Golden and Inhibitor Gene

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)

Wide-banding, Rufousing and Polygenes

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:

1. Many Golden Tipped breeders breed Golden’s as a ‘by-product’ of a Black

Tipped program – and whilst the exploitation of Wideband is very good for

both silver and non-silver series, the Golden badly needs that dose of

Rufousing, which is really counter-productive to Silver breeding.

2. The Wideband and Rufousing genes are incompletely dominant and as such

just don’t breed true in the early generations of a breeding program. There

are many breeders who have been present at the birth of a much anticipated


90

‘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.

The Genetic Puzzle

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.

Tipped breeders, along with Colourpointed breeders are increasingly becoming

interested in the properties of the Ticked tabby or the Ticked base coats.

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.


91

.

This little Blue Ticked Tabby appeared in a litter.

The Golden Tipped sire’s genes were AA, Tata B

Dd ii WbWb – so homozygous, for Agouti, and

Wideband, Black, carrying dilute. Non Silver and

hetrozygous for Ticked.

The Dam was a simple lilac colourpointed105.

We could have expected this kitten to show some

widebanding and certainly much more tabby

markings – but apart from her facial markings

and one necklace – this little girl had no tabby

markings at all –

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

Précis from Robinsons Genetics on Wide Band107

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


92

11. Colourpointed

11.1. Colourpointed, Non-silver, without white spotting

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 highest108 number of

registrations, with only the

self and the Silver Tabby

series ranking higher.

So successful is this breed

that it boasts its own club,

which holds a championship

show annually.

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 C110 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.

Basically the C gene is required for the production of any pigment.

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


93

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 gene code for our colourpointed cats is cs (Siamese)

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

carrying a single copy of a

recessive.

Solid colour

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

Himalayan/Burmese

Tonkinese pattern

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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Temperature Sensitive Mutations

In order to understand the mutation we need to go back to the Wildtype form of

the gene. The C gene is the Wildtype, the term Wildtype is sometimes used to

refer to the most common phenotype (or genotype) found in the natural

population.

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.

Because of the uniform warmth in the

mothers womb, Colourpointed kittens

are born with almost pure white coats

on a pink skin.

Where blood circulation is at its most

efficient keeping the kitten warm the

coat will remain white. Parts of the

body with lesser blood circulation will

begin to activate pigment.

By the time the kitten is 3-5 days of

age, colour will begin to form on the

ears, feet and tail. The darker the

base colour the more quickly the

colour will form. With the red/cream

series being the last to show their true

colours.

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.

Specific Health - Eyes

Colourpointed cats are undoubtedly popular. And on the whole these

temperature-sensitive partial albino genes are thought to be a beautiful and

largely benign mutation. However we would be derelict in our duty if we did not

touch on some of the issues behind those lovely blue eyes.

This is a complex topic, not yet fully understood, but we will do our best.


95

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 brain113 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 congenital114 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.


96

Affected kittens can be expected to have a near normal quality of life.

We also know that this permanent condition is relatively uncommon. We know

that the Siamese breeders have to all intents and purposes bred it out. This

means that it is likely to be a polygenic trait and affected kittens should not be

used within a breeding program.

Occasionally, it is possible to notice the colourpointed eye drifting in either a

convergent or divergent squint, which will correct as the cat refocuses – this can

happen on the show bench. This is not to be confused with the permanent

congenital squint

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.

Nystagmus is an uncontrolled movement of the eyes, usually from side to side,

but sometimes the eyes swing up and down or even in a circular movement.

There are two types of Nystagmus recognised:

1. Congenital Nystagmus, that first appears in the early months of life.

2. Acquired Nystagmus, when the condition develops later in life.

However, colourpointed breeders report a sort of mid-way partial Nystagmus that

comes and goes, usually when the cat is in a stressful state. 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 nerves115

There is much that we do not know about Nystagmus in cats116 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. • Nystagmus may be made worse by emotional and physical factors, such as

tiredness, unfamiliar surroundings, nervousness and stress.

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.


97

• Depth perception may be reduced. Affected kittens may seem a little

clumsier than normal and may need watching if climbing higher objects.

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.

Some Extra Information on the Colourpointed Cat

• The Siamese gene is recessive – both parents must either be

colourpointed or carry the colourpointed gene in order to get

colourpointed kittens.

• The Blue eye is more sensitive to bright sunlight.

• The gene that gives rise to the Siamese pattern is the same as pointed

rabbits and goats found in the Himalayas.

The Show Cat

Points Colour - Mask, ears, legs and tail should be a clearly defined colour,

matching on all points.

Body Colour - There should be a good

contrast between points and body

colour. Light shading, if present,

should tone with the points. Shading

in a mature cat should not penalise an

otherwise good exhibit.

Eye Colour - A definite clear blue;

deeper shades preferred.

Nose Leather and Paw Pads -

Appropriate to coat colour.

Showing Colourpointed Cats - Shading and Eye Colour.

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


98

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.

Fortunately, all this seems to be polygenic so plenty of work to do on the

colourpointed 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.


99

The ideal mask extends from

above the eyes down through

the chin and stretches beyond

the eyes evenly on both sides.

Sometimes we see a cat that

is hooded, where the mask

extends over the head and

sometimes, often on

chocolate points we see

incomplete masks, often

around the mouth and

whisker pads.


• Colourpointed breeders should try as much as possible to understand the

major Siamese genes as discussed in this policy. • The colourpointed British has inherited a potential for long soft coats from

the Persian ancestors and a desire to have large ears and a long nose from

the Siamese ancestory – balancing

these factors to maintain good British

type is a challenge! • Breeders may wish to work with the

Colourpointed cat club to see if the

standard of points that covers all

colours of the Colourpointed is

weighted against the darker colours

on the show bench. • Experienced Breeders of the

colourpointed and tabby colourpointed

British might benefit from talking to

some of the Siamese Breeders about

their experiences of the dilute modifier

genes – too complicated to go into

here but very interesting, and

deserving of further research. • The colourpointed Queen may require some careful management, like her

Siamese cousins, she can be a precocious caller, often calling from early

January through to December, and coming back on call within weeks of

having her kittens. She may also have very large litters. Breeders will

need to manage these situations carefully. • Also in common with the Siamese the colourpointed cat is deeply

affectionate, which some people who like the British tempererment may

find needy. This affection which may translate into a deep pair bond with

her human owner, over whom she may feel possesive, may mean that

some cats may not wish to share their lives with other cats. This trait may

run in some lines.


100

One last thing for the Experienced Colourpointed Breeder

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

genetically cs/ca 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.


101

11.2. Colourpointed -Silver Series

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.

Silver Tabby (Lynx) Pointed

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.


102

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’

If we accept that Silver can be tricky to detect in the

self cat, it doesn’t take a quantum leap to understand

that with the colourpointed it can get really difficult

and with the Red series colourpointed it can become

nigh-on impossible.

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.

Recommendations and Best Practice

Those colourpointed breeders interested in further research behind their

pedigrees might like to make a visit to Pawpeds – and search for a cat born in

1982 named Beeblebrox Plum Crazy – this cat was originally registered in the UK

as a Red Pointed and as such fathered a good number of offspring – he was later

re-registered as a Red Silver Smoke and Pawpeds have him as BRI ds 33.117

117 EMS Breed Numbering System

16– A lovely Chocolate Smoke Pointed Female


103

There are a great number of colourpointed breeders today both here and in

Europe breeding from cats directly descended from Beeblebrox Plum Crazy118 –

and if his Silver gene slipped through the net – how many others?

We have spoken with a number of UK breeders who have experimented with

Silver in their colourpointed lines and who have decided that in the absence of a

test for the Inhibitor (I) there are no benefits to be had.

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.


104

11.3. Colourpointed & White (Any colour) Preliminary

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.

The Show Cat

Points Colour - The face

should have an inverted V of

white, starting on the

forehead, but not exceeding

the inner aperture of the eye,

and extending down covering

the nose and whisker pads.

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.

Eye Colour - A definite clear blue: deeper shades preferred.

Nose leather/paw pads-appropriate to coat colour or pink.

119 The SOP does not specify the amount of white on the body.


105

Some Early Findings From Breeders

The following are the observations from breeders of Colourpointed and White.

• Breeders working on this pattern are finding that after several

generations they can

regularly produce

kittens with that

perfect face blaze.

• The eyes are always

blue as in the

traditional

Colourpointed. But

some breeders are

reporting a deeper

shade of blue

amongst some of

those individuals

with White.

• When breeding for

pattern the base

colour of the

Colourpoints does

not affect the outcome.

• It doesn't matter whether you mate two Colourpointed and White cats

or a Colourpointed to a C/P and White – it seems that the litter will

usually get a mixture of both patterns120

• Two C/P and Whites together are more likely to throw a Van Patterned

C/P and White,

The Polygenic Properties of White Spotting (S)

Within the section on the Bi-Colour we touch upon the Polygenic121 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


106

11.4. Silver Colourpointed & White (Any colour) Preliminary

Please refer to the earlier Colourpointed sections.


107

12. British Shorthair Variant

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 Phenotype122 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.

One genetic legacy is the gene for long hair.

The length of fur in the feline is governed by the Long hair gene which is known

as L. In the Wild Form123 the dominant form of L codes for short hair, whilst the

recessive l codes for long hair.

To understand how hair length in the cat is determined we need to understand

the cycle of hair growth.

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.

In simple terms the

Catagen phase is a short

stage that effectively stops

the hair growing.

The final stage is the

Telogen phase, during

which the hair or fur is in its

resting state, neither

growing or shedding

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.

122 Appearance 123 Typical Phenotype

Catagen phase

Telogen phase

Anagen phase

http://en.wikipedia.org/wiki/Domestic_longhaired_cat


108

The British Shorthair will

be either Homozygous

for Short Hair and have

two copies of L so LL or

will be Heterozygous so

Ll – any breeder mating

two Heterozygous cats

together stands a good

chance of those

recessive l’s meeting and

the result will be the

really rather beautiful

Long haired British

Variant

The British Long Hair

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.

Long Hair Mutations (extract from UC Davis)

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 1 (M1): present in Ragdolls

Mutation 2 (M2): present in Norwegian Forest Cats

Mutation 3 (M3): present in Maine Coons and Ragdolls

Mutation 4 (M4): present in all breeds of long hair cats, including Ragdolls, Maine

Coons, and Norwegian Forest Cats.

17 - Exquisite British Longhair from Europe


109

13. Eye Colour

The most common colour for wild, non colourpointed, cats is hazel somewhere on

the scale between orange and green.

The British Shorthair comes with four distinct eye colours:

They can be sorted into two categories:

• The Polygenic eye colours:

• The Major Gene eye colours:

The Polygenic eye colours:

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.

The Major Gene eye colours:

When eye colour is linked to a major gene it means that breeders have to work

with that colour.124

• 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.

124 For example it is not possible to breed a colourpointed cat with an Orange eye.


110

HOW EYES GET THEIR COLOUR

There are two key factors that influence eye colour in the cat:

• Pigmentation

• Blue Light Refraction

The iris has two layers of cells or melanocytes,

which produce pigment.

1. The Stroma - an outer layer made up of a

bunch of loosely constructed cells.

2. The Epithelium – situated under the Stroma

consisting of tightly packed cells.

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

When there is an absence of melanocytes, as in the un-pigmented white cat the

eye will appear blue125 - all that blue light reflecting and refracting just bounces

back -

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!

125 Or in rare cases pink

Epithelium

Stroma


111

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 -


112

14. Record Keeping

Keeping good records is one of the cornerstone principles of a sound breeding

management system.

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.

As a minimum the following records should be kept:

Purchase Cat Records Contact

Records

Breeding Records

Record details

of all cats

purchased

Pedigree Name Name of Sire

Pedigree

records

Colour Address/Phone

number/email

address

Name of Dam

Breeder

records

Gender Relationship Inbreeding co-

efficient

Cost to

purchase

including travel

Date of

Birth/Purchase/Sale

Fellow Breeder Cost

Vaccination

records

Breeder Stud Cat

owner

Dates matings

observed

Health Care/Veterinary Used your

stud

Describe the

mating – how was

the stud/how was

the dam.

Temperament Cat Club Calculate pinking

up date

Performance as stud Judge Calculate due date

Performance as Dam Show

Management

Show Records Bought kittens

from You

Registration Number Sold kittens to

you

Genotype

DNA Tests

Nutrition


113

Birthing Records

Kitten Records

Litter Records

Book Keeping

Records

Date and time

labour starts

Date and time of birth Success of

litter against

predetermined

criteria (did

you get what

you hoped

for!!)

Money In – by

type

Describe queens

behaviour/any

discharge

Birth weight Overall health

and vigour

Money Out – by

type

Time of

contractions

Any birth notes, ie

Breech etc

Number of live

kittens born

Time of first and

subsequent

kittens

Identification details Number of

kittens lost in

nesting

Time of last

kitten

Daily weight records

Make a note of

all the placentas

Nutrition

Birth weights Health records

Assistance

required by

dam/medications

Worming

Fleas

Vaccination

Show potential

Sale records/contracts

Fortunately there are a number of packaged software programs to help with

much of this information and simple spreadsheets can be used to keep and

calculate much of the required information.


114

15. Recommendations

To be successful a breeding program must be undertaken in a planned,

considered and structured way, within a system or framework of accepted best

practice. However, a breeding program or system can only be as good as the

breeder who operates the system.

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.

© British Shorthair BAC - 2010


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Appendix 1 References

We have referred to a greater or lesser extent to the following publications. We

have also made much use of other widely available materials and are, as ever,

grateful to those who have gone before us.

GCCF Breeding Policy

Asian Breeding Policy

The Official Standard of Points (GCCF)

British Registration Policy (GCCF)

Robinsons Genetics for Cat Breeders and Veterinarians

Judges Reports (GCCF Website)

Wikipedia

Feline Advisory Bureau

DEFRA Cat Welfare Policy

PAWPEDS (www.pawpeds.com)


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Appendix 2 Glossary

Allelle refers to the different forms of a gene

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.

Chromosomes - Genes are located on the chromosomes. All cats have 38

chromosomes in each body cell, 19 from each parent. Chromosomes appear in

pairs. With the exception of the sex chromosomes each pair is the same size and

shape.

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

Heterozygous refers to having two different alleles at a given locus on a

chromosome

Homozygous refers to having the same 2 allelles at a given locus on a

chromosome

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

SOP – The Standard of Points is established by Breeders and Breeder Groups,

who present them via their Breed Clubs to the Breed Advisory Committee. The

SOP is agreed by the GCCF Executive & finally by the full GCCF Council made up

of representatives from affiliated clubs. Any changes to a published SOP follows

the same procedure. The SOP is used by judges as the base criteria against which

to award placings and awards. It is the SOP that ultimately protects the Breed

integrity.


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Appendix 3 Key Genes

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.

Non- agouti or “hypermelanistic” (a) - a recessive gene mutation that

turns the original “wild” tabby cat into a self black by overlaying the agouti base

colour with melanic pigment, making the whole animal appear black, although

often in certain light the underlying tabby pattern may still just be discernible.

Other genes work to change this black pigment to other colours.

Inhibitor (I) – a dominant gene that suppresses the development of pigment

in the hair of the coat, typically producing hairs that are fully coloured only at the

tip and have a silvery white base. It has greater effect on the lighter pigment in

an agouti cat, removing the yellow colour and turning the base colour white or

“silver”. In the case of a non-agouti cat the inhibitor removes colour from the

base of the hair-shaft to produce a silvery white hair with a coloured tip, i.e a

Smoke. This allele appears to interact with other genes to produce various

degrees of tipping, ranging from deeply tipped silver tabby to lightly tipped silver

shaded tabby.

Tabby patterning genes –

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.

Wide-banding (Wb) – this has been hypothesized either as a gene

(Robinson) or more probably a group of genes (Joan Wasselhuber, who coined

the term “wide-banding genes”): increasing evidence for their existence has led

to wide acceptance. Undercoat width genes determine the width of the undercoat

whether or not the cat has a silver inhibitor gene. The term “undercoat” used

here refers to part of the hair shaft closest to the body, and includes both guard

hairs and the shorter hairs often referred to as “undercoat” hairs. The variability

seen in the undercoat widths in cats points to the polygenetic nature of wide-

banding genes. If a single gene it is likely an incompletely dominant gene

mutation, the effect serving to push 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. Precisely how the agouti, inhibitor and wide-banding genes

interact on a molecular level is not clear - one possibility is that the wide-banding

genes influence the agouti protein production to remain high so that eumelanin


118

pigment remains inhibited or down-regulated; another possibility is that the wide-

banding gene encodes for a second inhibitory protein that also down-regulates

eumelanin.

Long-hair (l) – a recessive gene mutation which produces a semi-long haired

cat. (still found today in the British owing to breeding legacy)

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)

White spotting or piebald spotting gene, S/s, - has variable expression,

so that an SS cat often has more extensive white patching than an Ss cat. It is

this gene that creates the familiar white blaze across the face, a white bib, tuxedo

pattern, or dappled paws. A hypothetical Sb allele ("gloving gene") may cause the

mittens in Birman and Snowshoe breeds. Some researchers believe that there are

separate white spotting genes for distinct forms of white pattern, such as the

white locket that some cats have on their neck or bellies.

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.

Orange (O) – this is a mutation on the X chromosome and is thus sex-linked.

The gene eliminates all melanin pigment (black and brown) from the hair fibres,

replacing it with phaeomelanin, a lighter compound appearing yellow or orange

depending on the density of pigment granules. The O allele is also epistatic over

the non agouti genotype; that is, the agouti to non-agouti mutation does not

have a discernible effect on red or cream coloured cats, resulting in these self-

coloured cats displaying tabby striping independent of their genotype at this

locus. This explains why you can usually see some tabby pattern on red, cream

and apricot coloured non-agouti cats, even if only on the head/face. Rufus

polygenes, as yet unidentified, affect the richness of the orange gene’s

expression.

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


119

“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.

Rufus – (polygenes) as yet unidentified, can affect the richness of the

orange gene's expression.

Genes not at ‘work’ in the British Section but Useful to Know!

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.

Jun-18

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.

Adrian Miller

Teknocat

Andrea Zerrilli

Willowood

Mr and Mrs A Brown

Kolinga

Ann Stubbs

Positively

Ann Greatorex

Mewzishun

Brenda Hollandt

Pridycats

Bron Keeling

Gooderick

Caroline Prout

Fergan

Corinne Ash

Jococa

Dee Wiley

Purrissimo

Denise Conway

Pampurred

Doreen Goadby

Dorothy Parry

Pinemartin

Brenda Wolstenholme

Cassandra Moss

Regents

Celia Leighton

Porteous

Christine Clayton

Malmo

Elena Nikitenko

Bastet-a-tet

Jane Best

Leonitara

Jean Townsend

Skellum

Jean Wolfe

Bumpkin

Jenny Hemming

Hinderslyne

Joanne Hewitson

Countrystyle

Julia Oughtibridge

Alejandro

June Payne

Poshpets

Hayley Wild

Apatchicats

Helen Maclay

Kruzinkal

Jackie Steele

Steeleaway

Judith MacArthur

Old Possum’s

Julie Caffel

Bluerain

Karen Holloway

Windymeadow

Lesley Tricker

Lizz Benson

Limatz

Ludmila Pankova

Photographer

Marion Wade

Rodwaye

Marilyn Storey

Skidoosh

Michelle Simpson

Althalus

Monica Acton

Idlebeck

Patrick Booth

Kolinga Cougar

Sandie Holloway

Rayanshel


118

Sandra Schürmans

Aus-der-Mondberg

Sue Lorton Hobbs

Loraston

Ruta Towse

Miletree

Shirley Bullock

Suzanne Dalton

Donyosabuk

Suzanne Griss

Grisselian

Terri Callahan

Calla Cats

Tamila Aspen

British Empire Cats

Tammy Kingwell

Purrade

Veldes Carnell

Misskin

Viv Clerkin

Silvercloud

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