As advances in DNA research are made, new information and coding with regard to color genetics will change.   While the basic principle remains the same the newly identified genes will have to be incorporated into a new article on Brittany Color Inheritance.   For Example "B" used for Brown, and "K" used for dominant BlacK.    At this point in time research has found that all Tricolor dogs are of the make up recessive Black (k/k) and if they are the makeup of K/k or K/K the dominant expresser...then Tricolor is not allowed.  The mutation for Brown has been identified in three different forms identified as "b" for brown and must be in double copy in order for it to change the Black color into Brown color.  The majority of Sporting dogs are in the Dominant Black (K/K) format and it is the Extension gene (E) responsible for changing the black to Orange and again the Brown gene (b) in double copy that changes Black to Brown.  American Brittanys can be either K/K or K/k or k/k based on whether they are Tricolor or not. The majority of American Brittanys are double copy Brown (b) hence they have the rose colored noses over their European cousins that can have single or double copy genes.   As researchers continue to map out the Dog Genome how dog breeders incorporate this information will affect dog breeding hopefully for the better.   G.C.

Inheritance of Color in the Brittany (Epagneul Breton)

(Published in American Brittany Magazine, November 2001, pages 16-19)

A 2008 revised Brittany Color Article may be read HERE

~ by   G. Coats ~

I am not a geneticist, but I am interested in the color inheritance of the Brittany.   Since I was not able to obtain accurate information from current active breeders, I sought information from books on genetics, from "old texts", and from the French breeders who have more experience in dealing with assorted colors as found in the European lines of the Brittany.    From my experiences on the various e-mail list for Brittanys, I believe most have very little knowledge concerning color in the Brittany.  I intend to provide more insights into Brittany color patterns, one of the better understood traits involved in dog genetics.

There are 5 colors admissible in the Federation Cynologique Internationale (FCI) standard, and only 3 of them recognized in the American Kennel Club (AKC) standard, with or without roan color pattern.

The Five Colors Are:

Orange and White (OW)

Liver and White (LW)

Black and White (BW) FCI only

Tricolor Liver Base (Liver,white,orange:LT)

Tricolor Black Base (Black,white,orange:BT) FCI only

Trying to explain color transmission is difficult unless you know basic information on color inheritance that apply to all breeds of dogs including mixed breeds. All dogs carry the complete sequence of color genes, including those that create colors that do not “appear” visible in the Brittany. The visible characteristics that we see in a dog (phenotype) are under the influence of genes, present within the 78 chromosomes of dogs. Each parent contributes 1/2 of the 76 somatic and 2 sex chromosomes.

Genes are always present in pairs, one from the father, and the other from the mother, so for simplicity, an example: FF= Father, MM= Mother. When the cells divide for reproduction they are halved and then recombined when sperm meets an egg, to form the new offspring which will carry the “FM” (Father + Mother) gene combination. These genes reside at the same location (locus) on a homologus chromosome pair. These two copies of the gene are called alleles. A dog with two identical alleles at a locus is called a homozygote. A dog with two alleles that are slightly different in their nucleotide sequences is called a heterozygote, or is said to be heterozygous at that locus.

Genes may have modifiers of strength which influence what is seen. An example: dark red color vs. light orange color, which is the same color gene with genetic intensity modifiers affecting how it appears to our eyes.

Genes may be either DOMINANT or RECESSIVE. By standard genetic convention dominant genes are expressed by a capital letter, while the recessive gene is reported by a small letter.

The dominant gene is ALWAYS visible, so we usually say it is expressed. The exception is when two recessives are combined together then we see them expressed. We call this double dose of the same genes either dominant or recessive as “homozygous”.

An example following our genetic rules:

Dominant + Dominant = visibility of Dominant

Dominant + recessive = visibility of Dominant

recessive + recessive = visibility of recessive

(Note: the Orange & White color of the Brittany is homozygous recessive)

Color schemes:

1. The “B” gene (the Black series)

This gene gives the black color, and also the Liver color! Both colors are due to the same pigment (eumelanin) and the chemical shape of its particles determines whether it is seen as black or liver. The gene for black is designated with “B” (= dominant), while the one for liver is designated with “b” (=recessive). All American Brittanys are of the designation “bb” and do not have the “B” (dominant) form where black is shown in the coat or the nose.

(Note: according to White, Brittanys that came to the US carrying the “B” gene were selectively removed from the gene pool by the 1960’s when Orange dogs were bred to Liver dogs and they produced Black dogs or dogs thought to be Black. Many liver colored dogs were also eliminated from the gene pool as their modifiers made them so dark that they were erroneously thought to be Black and were culled. - this culling was a loss of dogs with a valuable gene pool)

Now, all standard American Brittanys show a pinkish or deep tan nose color on Orange dogs, never any black or “a color so dark as to appear black” which meets the AKC color standard.

If parents with the father=BB (Black) and mother= bb(liver) are mated together, all of the offspring will be uniformly “Bb”. As explained above, the dominant gene masks the expression of the recessive, so this first generation offspring will be uniformly black, all of the pups carrying the “Bb” combination. If first generation of “Bb” pups are bred back to each other, the offspring in the second generation would statistically show the following possible combinations, tabel 1.

Table 1

BB and Bb are the “phenotypes” (what we see as black, but cannot be sure of which gene format the offspring are carrying; BB or Bb since both are black), while “bb” is what we see (Phenotype) and in this case also know the “genotype” ( liver is always and only of the “bb” form) so it is a ‘gene expressed’ appearance.

From table 1, “statistically” such a breeding of Bb to Bb will produce 75% black coated offspring, and 25% liver. But, for a small number of matings this result could be varied as the size of the litter will determine the number of possibilities seen, so it may take many repeated matings to approach the “statistical” results as given.

In the case of Liver colored dogs you may have plus or minus modifiers that also affect intensity of the liver (bb) shade, from almost black (it will NEVER be black), to a washed out, faded or “sunburned” like color appearance.

One must understand the genetic facts above, in order to understand the following concepts on how the “B” gene is further expressed.

2. The “A” gene ( the agouti series)

There are many variants or “alleles” of this gene, but the Brittany breed standard dictates that we should have only two acceptable forms (some breeders may have seen a third but this may be a variant in expression of the "at" gene).

“As” gene combination which is dominant giving solid colors (normal), and “at” combination which is recessive, give “tan point” which is a tricolor pattern (tan ticking on lower legs, around the anus, on the eyebrows, and on the cheeks). So basically “As” is non tricolor, and “at” is tricolor.

Table 2

Depending on which format the “B” gene is present in, the following phenotype is seen:

Table 3

3. The “E” gene ( the extension series)

This is the gene that gives individuals Orange color. Or explained another way the “E” dominant allows the genes in the “B” series to express themselves. The homozygous recessive form “ee” (genotype) PREVENTS the expression of black (B-) or liver (bb) in the body hair color but does not affect skin color (nose, eye rims, scrotum, lip skin). It is said the “ee” genotype is “epistatic” (an allele at one locus influencing the expression of another locus) to the “B” gene.

So basically, an orange and white Brittany is ALWAYS a carrier of the “ee” combination, regardless of what it carries on the “B” or “A” genes. So two Orange and white dogs bred together can only produce OW offspring. “Other” colors (liver and black) can produce a variety of colors as they could be “heterozygous” meaning they could have the format “Ee”.

Dogs “homozygous” in the form “EE” will only produce their self color so in this genetic form only Liver and Black are seen and PREVENTS Orange.

Remember the 'E' series does not affect skin pigment thus nose, lips, etc. would show black (B) if a dog is of the makeup BB or Bb. The Epagnuel Breton that are carrying the "B" gene in dominant format will show a very black nose if "BB" or what is termed a "Tobacco nose" (dark brown or a fading black 'winter nose', the color of purplish tobacco juice) if it is of the makeup "Bb". American standard Brittanys are of the makeup "bb" and will show a pinkish or deep tan shade of nose color.

Note: French breeders prefer a Black nose or a "dark" nose over pinkish tan "bb" noses.

While there is some benefit for a darker nose color in the prevention of "Solar dematitis & skin cancers", French breeders feel the darker nose and eye color (darker eyes are associated with the "B" gene) gives a much softer expression or they use the term "warm look", both kinds of of nose pigments are accepted.

Coat Colors of Brittanys

Classification of the dog coat colors is done by using alphabetical codes, this allows what color possibility a mating will offer. The following examples are “minimum” rule requirements to predict expression for a particular color (the use of ‘-’ is when that gene is unknown):

Table 4

Table 5

Table 6

An explaination of Orange-tricolor (OT)is given:

An Orange colored dog may be hiding the "at" gene in the form of 'As at' or 'at at', since "ee" PREVENTS the colors of Liver or Black to show, the "tan point" pattern is hidden in an Orange dog. Orange dogs mated to Liver dogs that produce a Tricolor, are now known to have at least 1 tricolor gene, as does the Liver(Black) dog it was mated with.

In France it is popular to breed tricolors so in the case of 2 tricolors mated together all resulting pups will be homozygous tricolor. An Orange dog that has both parents tricolor is itself a homozygous tricolor ('at at') even if we cannot see the pattern expressed on its Orange coat.

This particular dog when mated to either a Tricolor, Liver and Black dogs carrying a tricolor gene will always produce tricolor in those offspring exhibiting Black or Liver.

4. White, The “S” Gene ( the “white” Spotting series)

The white in the coat of dogs is perceived to be “color painted on white”, but it is actually a solid dark base color with “white painted on top” or more correctly allowed to filter (bleed) through the base color.  The original color of wolves was yellow(ayay), and it is only from mutations over many centuries that black appeared, and then white.

White is not like a paint color that blends with color to change the shade, it is a distinct color that is affected by a gene known as “S” (for spotting), or “s” for the other recessive alleles.

There are 4 specific alleles for “spotting”:

“S+” = Gives almost a completely colored coat, nearly without white (ie. Labrador, Gordon Setter, Vizsla)

“si” (i = the “Irish” spotting a term carried over from a color pattern in rats, not the Irish setter) Heavy color and solid heads with maybe a small white collar and white feet. Affected by plus or minus modifiers.

“sp” (p= piebald) giving the coat an irregular spotting of color with white.

“sw” (w= white) concerns coats mostly white with “invading” white spotting (West Highland White Terrier, Great Pyrenease, Samoyeds, Dalmatians) no color around eyes, may have slight color on head or edges of ears, but generally the body is fully white.

REMEMBER THAT WHITE AREAS ARE IN OPPOSITION TO COLORED AREAS.

Disregard any ticking, flecking, or roan color patterns which is under the influence of totally different gene combinations.

White color is influenced by recessive genes, which means to be expressed, they must be in a homozygous form. If white or lots of color is desired the color combinations must be extracted from the ancestors. Perhaps a very white dog would be “sp sp” a very heavily colored dog could be “si si” or maybe “si sp”. The dominance of the white genes are listed in the order as above so basically a heavily colored dog is dominant over a more white dog, solid colored heads are dominant over heads with blazes; but, there is no “mathematical” way to obtain a given result.

The French breeders believe that there are NO “S+” or “sw” in the Epagneul Breton breed (and really there should not be). “S+” being a solid colored dog is not allowed in the standard (both FCI & AKC call for “piebald”), and the French have always disqualified white dogs that have no color over the eye or any white spot in the ear, for which they blame “sw” as the culprit (although if you read Robinson, color sketches show extreme piebald can have an occasional lack of eye patch or ear color, but generally there is also color on the body, but “sw” calls for a white body with no color patches).

Note: Genetically inherited health issues are associated with extreme “piebald” (spsp) or White (swsw) dogs in general concerns lack of pigment granules of the hair in the concha of the ear, these hairs are responsible for transmitting sound.Lack of pigment in the ear contributes to a condition known as “Color deafness”. The inner ear develops from the dorsal involution of the skin to form the nervous system, if a white “spot” forms the aural cavity resulting in lack of pigment you will encounter deafness. This accounts for some forms of deafness in Brittanys and is commonly seen in English Pointers, English Setters and Dalmatians.

A solid colored Brittany or a totally white Brittany implies crossing with another breed that allows for solid or totally white patterns.

Note: In English Setters the pattern “sw sw T T” is Belton, a white body color that allows only small patches of color on the head or ears. TT allows for the heavy ticking on the body. While English Setters certainly figured into the creation of the Brittany those color traits associated with them have been diligently selected away from over the past 100 years. Remember ‘sw sw’ allows NO large color patches on the body.

5. The “T” gene ( the Ticking series)

Ticking, flecking, and such small spots of uniform color that appear in the white areas of the coat (disregarding larger true color spots) are the effects controlled by the “T” gene.

Coats that show ticking are under the influence of the “T” gene which is dominant, and coats that have no ticking are under the influence of “t” gene which is recessive. Like in the spotting series given above, there is no mathematical way to determine how much ticking is present as “T” dominance over “t” is incomplete and may have plus or minus modifiers involved.

Belton patterns as seen in the English Setter, or the ticking seen in Dalmatians are of the form “TT”. The majority of our Brittanys are of the makeup “Tt” with many variants in ticking densities, standards (FCI & AKC) says ticking is desirable in some form.

Some studies also indicate that “tt” is involved in the majority of dogs in the “spsp” pattern type, over dogs of the “sisi” pattern type. But again this is some what subjective and breeders have managed to rear very clear white dogs with lots of color, and heavily ticked dogs with lots of white.

It should be remembered that ticking is independent of the extension of the colored area.

In this same category is the color pattern of ROAN. 'Rouanne', a french word, is defined to be a fine intermingling of colored and white hair. Experts have not really done much research on this color pattern except that it also appears to be a Dominant trait “R” and non-roan is recessive “r”. It is not fully understood if Roan is actually an extreme form of ticking or a separate entity.

For those who have bred litters of “true roan” it is felt this is indeed a separate gene as it seems to have an effect on base pigment at the skin and foot pad level and not just at the coat level. True roan dogs appear to have a unicolored skin (except in areas that are under a double “white spotting” effect that will appear totally clear white along with skin pigment as well). In Roans the areas that are interpreted to be white are not a clear white but a fine blend of colored and white hairs on a dark skin base. It is also affected by the “S” series in that it allows white spots to show through the roan patterned areas (roan dogs are frequently seen with a clear white spot on top of head, or on a part of the body). This gene also seems to be affected by other plus or minus modifiers that cause the roan to appear more color or more white in tone. It also appears to be independent of the “T” gene as well, as we can find areas of distinct ticking of a certain diameter over or in roan areas, or an apparent lack there of.

At birth roan pups appear clear orange and white, white and orange, liver and white, or white and liver with the darker hair becoming noticeable from approxamately 4-8 weeks, however, the time period varies. Most orange and white roan pups will have dark foot pads at birth, or a partial darkening of foot pad perimeter, thus giving indication of what color pattern they will have as young adults.

6. Other Series

There are other color series that generally do not apply to the Brittany. They include:

Merle, Chinchilla, Dilute, Creme, Brindle, Gray. While these series are present in every dog they are in a form that does not allow them to be seen in the Brittany.

However, the factor for Graying seems to be present in some Brittany lines where young dogs turn white faced or body color turns to white. Graying may also have an environmental component as has been shown in some species (mice) to be related to utilization of antioxidant agents such as vit. E and selenium.

In summary, it is prudent to remember the Brittany is a performance breed and that instincts of pointing, hunting ability, a balanced willing temperament, trainability, and physical soundness to be a performance dog is much more important than the color of its coat (with the exception of deafness found in some Brittanys that are white or mostly lacking color).

Approxamately 61% of American Brittany breeders prefer the clear orange and white color. This limits the gene pool when American Brittany breeders have so many choices of color, table 5.   A variety of colors allows broadning of the gene pool by selection of many color combinations, thus increases availability of other selectable traits.

Foot note: At the time of this writing DNA samples have been submitted to VetGen Labs of various colored Brittanys to determine if their current color tests are applicable to the Brittany.

References:

1. Willems, P. - L'Epagneul Breton ( Club L'Epagneul Breton Publication ) Spring 1991. Article- The Colours of the     Brittany ( the Genetic Transmission)

2. Willis, M - Dog Genetics, Howell Publishing,1989.

3. Whitney, L - How to Breed Dogs , Howell Publishing, 1971, 1941.

4. Robinson, R. - Genetics and the Dog, Pegamon Press, GB, 1982.

5. White M.D., F - The Brittany in America, Self published, 2nd edition 1965.

6. Munsch, R. - L'Epagneul Breton, Historique Characteristiques - Elevage-, ed. De L'Eleveur, Paris 1937                                                       Preface by R. de Kermadec.

7. American Brittany Magazine, Vol. 1, Issue 2,  October 1950.- Black noses referenced in report from Board of Directors meeting concerning ballots for standards change to Disqualify Black and writer seeing and producing several black mottled noses on Orange dogs, and giving them away to hunters.

8. Wilkie, PJ - Future Dog Breeding for Genetic Soundness. Minnesota Agricultural Experiment Station. University of Minn. St. Paul, Mn. in Cooperation with the Canine Health Foundation, 1999.

9. Genetics Defects Commitee Report Results - Section one, American Brittany Club Genetic and Health 1996 Questionaire, P.30. The American Brittany, No.11, Nov. 1998.

10. Gunther, C. - L'Epagneul Breton, ed. DeVecchi, S.A., Paris 1995

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