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French Bulldog Coat Color Genetics

"For Dummies"
I have titled this page "for dummies", as I would consider
myself a "dummy" or layman (non-scientist) when it comes to
the topic of coat color genetics of the French Bulldog. My
intent in presenting this information is simply to
articulate the genetics involved with breeding french
bulldogs in such a way that benefits myself and anyone else
who struggles to understand the complexities of color
genetics. All of the information I am going to summarize
based on my own understanding and way of thinking about that
which I have gathered through my own research on this
subject matter and through my personal breeding experiences.
If anyone has any challenges to anything written here please
contact me, I welcome all questions and corrections.
Most of the information found in this article below, has
come from the following major sources ... VetGen VGS, UC
Davis VGL, french-bulldog.com, Genetics of Coat Color and
Type in Dogs, Dog Coat Color Genetics, and Animal Genetics.
There are a few terms that I will explain first to help us
later. The first two are genotype and phenotype. Sounds
compicated, but just break it down like this ... geno=gene
(deals with what the dogs actual genes are), pheno (ph is
for photo, deals with what the dog physically looks like,
the physical picture of the dog). You can also consider
genotype is what is on the inside, phenotype is what is on
the outside.

D Locus - MLPH - Dilution - Blue Coat Color

The most controversial color is the Blue French Bulldog,
(also called mouse or mousegray) which appears grayish in
color. There are several genetic locations (called Loci) in
which determine a dog's color genetically. At each location,
is a pair of genes (Alleles). For Blue, the Loci, or
location is called D for dilution. The possible alleles (or
genes) at the D loci are "D" and "d". Every dog is either D-
D, D-d, or d-d. These are the genotypes. A genotype of D-D
produces a pheontype of Non-dilute, or non-blue. The
genotype D-d is what is typically called a "blue-carrier",
"blue-lined", or "blue-factored" frenchie. These dogs have a
phenotype EXACTLY the same as D-D. They are NON-dilutes, and
NON-blues. "D" is a dominant allele over "d" so only dogs
which have a d-d genotype will have a blue or dilute
phenotype. In other words, only d-d dogs are actually blue
(or gray) in color.
Blue is a dilution which acts on black pigment as well as
black colored hairs. Any pigment or hairs that would
otherwise be black are diluted from black to blue on a d-d
dog. This is what makes solid blue, blue brindle, blue fawn,
blue pied, blue with tan points, blue masked fawn, blue
sable fawn, and all other varieties of blue possible. All of
these listed colors have the same genetic make-up, d-d at
the D loci.
For breeding practices, most frenchies are D-D, non-blue.
Any frenchie that is D-D will NEVER produce a blue puppy and
any dog that HAS produced a blue puppy cannot be D-D. Only
D-d and d-d dogs can produce blue offspring. Any dog D-D,
that is bred to a blue d-d, will produce an entire litter of
D-d puppies, which are blue-carriers, but not blue. Each
puppy enherits one gene from each parent. For more
explanation on how to determine offspring percentages, here
is a great article titled, Probability of Inheritance.
Here is where the math comes into play. If a blue dog d-d is
bred to a blue-carrier D-d, then statistics tells us that
50% of puppies will be blue d-d. The other 50% of puppies
from that breeding would be blue-carriers D-d. In a perfect
world a litter of 4 from such a pair would produce 2 blue
pups and 2 non-blue blue-carriers. This is not always the
case. Firstly, we are not always dealing with litters of 4,
and secondly the percentages don't always work out when
dealing with such a small amount of data. Perhaps if we
considered data from hundreds or thousands of litters we
might find those percentages.
For our purposes it would suffice to just understand that
this breeding pair CAN create either blues, or blue-
carriers, but cannot produce any D-D pups. Learning to
 understand genetics can help a breeder predict what colors
are going to be produced in offspring, and help them make
the best decisions on which dogs to breed together.

B Locus - TYRP1 - Brown - Chocolate / Liver Color

Chocolate or liver, works the same way as blue. At the
chocolate loci, B (I say B for brown) exists the two alleles
"B" and "b". French bulldogs can either be B-B (non-
chocolate), B-b (chocolate-carriers), or b-b (chocolate).
"B" is dominant to "b" so only those dogs with a b-b
genotype will express the chocolate color. Chocolate is
similar to a dilution as it acts upon black pigment and
hairs and can create such color patterns as solid chocolate,
chocolate brindle, chocolate masked fawn, chocolate pied,
chocolate with tan points, etc.
Lilac Coat Color
The Blue Dilution mentioned above can also dilute Brown
hair. This is where the color "Lilac" (historically referred
to as "Isabella") comes from. When a dog is b-b at the "B"
loci and d-d at the "D" loci, this color will be exhibited
in the phenotype of the dog. Lilac is basically a dog which
is both blue and chocolate. They appear a very pale gray
color that many believe looks slightly purp;e in certain
lighting.
Champagne Coat Color
The Blue Dilution mentioned above can also dilute Cream
hair. This is where the color "Champagne" comes from. When a
dog that is cream is also d-d at the "D" loci, this color
will be exhibited in the phenotype of the dog. Champagne is
basically a dog which is both blue and cream.

Cream Coat Color - 3 Theories (still being

The Old Theory
The old theory puts cre
r ea
sm
eaar
sca
hesd
im
)ple recessive gene similar
to Blue and Chocolate. The supposed allele "Cch" (nicknamed
chinchilla) would be found at a supposed C loci and would be
recessive to the "C" allele. This would mean C-C is non-
cream, C-Cch is cream carrier, and Cch-Cch is cream.
The Newer Theory
The new theory puts cream as a dilution of the intensity of
red/yellow coat color. The supposed I loci (I for
instensity) would have alleles allowing for different hues
of red/yellow to cream.
 My Theory
First off let me just say this isn't techinically my theory.
I developed this way of thinking based on all of the
research articles mentioned above, but have altered my own
thinking based on what I have and haven't seen in the
frenchie population over the almost 10 years I have been
owning, breeding and fascinated with these dogs and their
vast array of colors. Nobody completely understands how the
genetics works for cream so we're all still guessing.
However, here's what I do know. I have personally never seen
a brindled or masked cream. This to me is enough proof to
rule out the newer theory. If cream was a simple red
dilution, you could have masked creams, black brindled
creams, cream brindles (reverse brindle but with cream),
black with cream points, etc.
The same would hold true if cream was caused by a distinct C
loci as in the old theory. If the evidence shows that cream
only exists is two forms, solid and of course pied/pieblad,
then my theory is that cream is controlled at the E loci,
possibly by a yet undiscovered allele. If an allele existed,
lets say Ec, which was recessive to all other alleles at E,
then this could account for the lack of masked creams, cream
brindles, etc in the frenchie population, and could account
for the fairly obvious recessive behavior of creams. By this
I mean that cream seems to work like blue and chocolate. Two
creams will produce a cream litter and you need two cream
carriers to produce cream. Hence the old theory which
accounts for breeding statistics, but not the lack of masked
creams.

K Locus - MC1R - Dominant Black / Brindle Pattern

The next Loci to talk about is K. At the K loci is where the
determination is made for brindle or fawn. Possible alleles
are "K", "Kbr", and "k". "K" is dominant (solid) black,
which is theorized not to exist in the French Bulldog
population. This would not require that no solid black
frenchies exist, but that solid black is not due to a
dominant "K" allele. If this is the case, then the only
allele options are "kbr" and "k" and we can simplify this
loci to a basic dominant-recessive relationship (similar to
blue or chocolate) with "kbr" (brindle) being dominant over
"k" (fawn or non-brindle). kbr-kbr and kbr-k would be
brindle, while only k-k would be fawn (or non-brindle). A
 Homozygous (which just means two of the same gene) brindle
kbr-kbr would always produce brindle pups. This would be
beneficial if one is trying to produce brindle-colored pups,
or detrimental if one is trying to produce non-brindles.
Though this relationship seems to be easily understood by
removing the K (dominant black) allele from the picture,
there is still much more to consider on top of the K loci in
the brindle-fawn relationship. See below notes regarding the
E loci. Going back to solid black, which obviously exists,
it is understood that these solid black frenchies are a
result of the "a" allele at the A loci which is recessive
solid black. This has obvious implications on the breed
standard because solid black is listed as a disqualification
which would make sense if it was the result of a dominant
color gene that could potentially overrun the gene pool
hiding the vast array of frenchie colors from the world.
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This disqualification makes
|sense for dominant K (if it
exists in frenchies) but not for a recessive solid black.

E Locus - Beta-def 103 - Extension / Masking

E, the extension loci, has the ability to hide brindle. At
the E loci, possible alleles are Em which is dominant to E,
E (also dominant to e) and e. Any frenchie with "Em-Em",
"Em-E", or "Em-e" will have a black mask, unless the mask is
hidden by white in a piebald dog. Masks can also sometimes
be undectible in very dark brindles or solid black. The
recessive "e-e" (called recessive red or recessive yellow)
is a disfunctional extension that does not permit black to
be formed. This is dominant to Kbr so a dog that is "Kbr-
Kbr" or even "Kbr-k" but which also has "e-e" at the E loci
would be fawn, but carry brindle genetically. You could even
have a homozygous (Kbr-Kbr) that is fawn due to a
disfunctional extension "e-e" but would only produce brindle
pups, even though brindle is dominant to fawn.

A Locus - ASIP - Agouti - Fawn / Tan Points

The A loci is the one responsible for the different
variations of fawn. Possible alleles are Ay, Aw, At, and a.
Ay is dominant (solid) fawn or sable. Aw is for (wild sable)
and is very similar to Ay. At is responsible for tan points
(black and tans, blue and tans, chocolate and tans, etc).
Lastly, a, is recessive black. For more information, see my
new Color Genetics Chart.
 S Locus - MITF - Spotted - Piebald Colors
The last loci to discuss is S, spotted. This is where
piebald markings come from. Alleles are S, si, sp, and sw. S
is dominant for self-colored (can have white chest or toes -
under 10 percent white). The si is for Irish spotting, which
usually is white chest, lower legs, undersides, white
collar, white blaze and can be 10-30 percent white. The sp
allele is for the typical piebald, which is random spots of
color can be 20-80 percent white and is usually non-
symmetrical. The sw allele is for extreme white. Our female
Butters is a good example of this genotype. The sw allele is
usually at least 80+ percent white with only small colored
patches on the face and near the tail. I am also still
researching how these alleles interact with one another for
breeding prediction purposes. For now, it can suffice to
think of any of the pied genes as N which would leave us
with S and N, where S is dominant. So S-S is not pied, S-N
is a pied carrier, and N-N would be a pied. S can be DNA
tested for.

M Locus - PMEL - Merle

Merle french bulldogs have recently become the newest fad or
rare color or as some call them exotic color. I have seen
several pictures of merle frenchies, and while skeptical, I
will say they look like french bulldogs as pure as any.
Perhaps, as some would assume, these dogs are not purebred
and have been crossbred to bring in the merle gene. The
question for me is when? How far back? If a dog was
crossbred 3 generations ago then I consider that not
purebred. But, what if the dogs was crossbred 10 generations
ago? That would mean that if you looked at a 10 generation
pedigree for your merle frenchie, you would see 1,024
ancestors to your dog. If 1,023 of those ancestors are fully
purebred french bulldogs and 1 great-great-great-great-
great...(you get the point)...grandparent was another breed,
and the dog appears to be a perfect frenchie according to
the breed standard, well then I would say you've got
yourself a merle french bulldog.
Personally I would love to have a merle frenchie. French
bulldogs are popular BECAUSE of the vast array of colors
they come in. If I had Merle french bulldog puppies, I would
not be able to keep up with how many buyers I would have
looking for them. That is what makes french bulldogs so
different and special. Having said that, I will say I remain
very skeptical about merle frenchies and would advise anyone
to do the same. The merle frenchies I have seen have had
very heavy 5-digit price tags which I think is just absurd.
The rare colors have become so popular over the last decade,
and it will be very interesting to see how this shapes the
breed over the next decade.