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Egyptian Mau Colour Genetics
by Melissa Bateson
When I first
started breeding Maus in 1996 cat colour genetics were a complete mystery to me.
I asked some other breeders for explanations and was told rules of the type
‘smoke x smoke = smoke and black’, ‘bronze x bronze = bronze and black’ and ‘silver
x silver = anything’. Although these rules are correct, it turns out that they are
too general to be very useful. I wanted to breed silvers, but although my first
two litters were from silver to silver breedings I got a complete rainbow of
kittens: a total of four silvers, two bronzes, a smoke and two blacks.....Why
was I being so unlucky when I saw other breeders having litter after litter of
silvers? Finally I began to understand when my breeder friend Melanie Morgan
sent me an article on Mau colour genetics written by Kerry Conway of Kaakhamit
cattery. I realized that with a little detective work I could have much greater
control of the colours of my future kittens. I think Melanie and I spent about
two hours on the phone excitedly trying to work out the genetics of all of our
cats. It has been well worth the effort. In 2001 I had three litters of kittens,
one of two, one of six and one eight, ALL silvers ! My intention in the article
is to introduce breeders to the genetics of the four basic mau colours, and
provide some simple rules for predicting the outcomes of different matings. I
have not included the blues or classic tabbies here because they are very rare
and hugely increase the number of possibilities that need to be
considered.
Click here for an expanded colour genetics chart.
Silver
The silver Mau is genetically a black silver spotted tabby. The
equivalent colour is also found in breeds such as the British Shorthair, the
Oriental and the Ocicat, but note that the silver Mau is genetically completely
different from the snow Bengal, whose pale colour is produced by the Siamese or
Burmese colour restriction gene, not the silver gene. The color and pattern of
the silver Mau are produced by the addition of just two dominant alleles
(variants of a gene) to the solid black Mau: the inhibitor allele, I, and the
agouti allele, A. The dominant inhibitor allele, as its name suggests, inhibits
the deposition of brown color in the hairs and is thus responsible for the
silver color. The dominant agouti allele causes pigment in the hairs to be
deposited in bands along the hair producing the effect known as ticking.
Presence of the agouti allele, A, is necessary for the tabby pattern to be
clearly seen. A Mau with AT LEAST ONE I allele AND AT LEAST ONE A allele will
be a silver. Therefore, since each cat has two copies of each gene, one
inherited from the dam and one from the sire, there are four different types of
silver Mau (known as genotypes). A cat is referred to as homozygous for a
particular gene if both alleles are the same, and heterozygous if they are
different. Hence the four genotypes of silver Mau can be described as follows:
• AAII = silver homozygous for the inhibitor and agouti alleles, i.e.
pure silver
• AAIi = silver heterozygous for the inhibitor allele, i.e. carrying bronze
• AaII = silver heterozygous for the agouti allele, i.e. carrying smoke
• AaIi = silver heterozygous for both inhibitor and agouti alleles,
i.e.carrying both smoke and bronze.
There is some debate about whether or not these four different genotypes
have different physical appearances (known as phenotypes) associated with them.
There is some evidence from other breeds (e.g. Persians) that although the I
allele is dominant, its effects are somewhat additive. Thus a cat with two
copies of the I allele might show greater inhibition of color than a cat with
just one. We might therefore expect II silvers to be less tarnished (i.e. have
less yellow or brown colouration) than their Ii littermates. This hypothesis
remains to be tested, however based on my own limited observations it does seem
to be the case that silvers with tarnish are more likely to carry bronze (i.e.
have the Ii genotype) than their cleaner coloured litter mates.
Aside from the possibility of phenotypic differences between the four types
of silvers, the genotype of a cat has important implications for breeding. By
the judicious pairing of cats of known genotype it is possible to control the
number of colours that can occur in the offspring. The colour table I have
created shows the proportions of different colors that will arise on average
from all the possible pairings of Mau genotypes. To use the table all you need
to do is (1) determine the genotype of your cat, (2) determine the genotype of
the cat it will be mated to, and (3) read off the average percentages of
different colours from the appropriate cell of the table. Neither the agouti
nor the inhibitor genes are sex-linked, thus it is not relevant which cat is
the dam and which the sire. Bear in mind that unless the percentage given is
100%, in which case you can predict exactly what you will get, these
percentages represent the AVERAGES ONLY, and you are unlikely to get exactly
the given percentage in any single litter. Although, if the mating is repeated
many times, the proportions of kittens of different colours produced should
approach the percentages in the table.
So now you are probably asking, ‘How do I determine the genotype of my
cat?’ Here are a few simple rules for silvers. Similar rules for bronzes and
smokes follow below.
First look at the kittens your cat has produced. This is the most
reliable method of determining a cats genotype.
• If your cat has produced a bronze kitten ever then it is carrying
bronze (Ii)
• If your cat has produced a smoke then it is carrying smoke (Aa)
• If your cat has produced both bronze and smoke offspring then it is carrying
both bronze and smoke (AaIi)
• If your cat has produced a black then it is carrying both bronze and smoke
(AaIi)
• If your cat has only had silver kittens then you know nothing certain unless
it has been bred multiple times to a cat known to be carrying smoke and bronze
in which case it is likely to be a pure silver (AAII).
Second, look at your cat’s pedigree. This is less reliable because it
depends on the pedigree being correct.
• If your cat has a bronze parent then it is carrying bronze (Ii)
• If your cat has a smoke parent then it is carrying smoke (AA)
• If your cat has one bronze and one smoke parent then it is carrying both
bronze and smoke (AaIi)
• If your cat has a black parent then it is carrying both bronze and smoke
(AaIi)
• If you know the genotypes of the cats parents and/or grandparents this can
help you determine what colours your cat has a probability of carrying
Third, look at your cat. This method is not proven to be reliable at
this stage.
• If your cat is tarnished relative to other silver litter mates it may
be carrying bronze (Ii).
Similar rules to those outlined above can also be applied to determine
the genotype of bronzes and smokes, and will be outlined in detail in the
following sections.

Bronze
Genetically the bronze Mau is a brown (i.e. black) spotted tabby. Brown
tabbies are known in may breeds such as for example the British Shorthair, the
Ocicat, and the Oriental, however, the warmer bronze colour of the Mau is only
seen in the brown Bengals and the usual Abyssinians. The bronze color is
achieved by the addition of the dominant agouti allele, A, to the solid black
Mau; thus a bronze is a Mau with AT LEAST ONE A allele. As in the silvers, the
agouti gene causes ticking on the hairs revealing the spotted tabby pattern.
Bronzes lack the dominant inhibitor allele found in the silvers and smokes.
There are two distinct bronze genotypes determined by whether the cat
has one or two copies of the dominant agouti allele:
• AAii = homozygous bronze, i.e. pure bronze
• Aaii = bronze heterozygous for the agouti allele, i.e. bronze carrying smoke
The warmness of the coat colour is determined by the presence of what
are known as ‘rufous polygenes’. As their name suggests there are many of these
genes, and the more a bronze Mau has the warmer its colour will be. The usual
Abyssinian is an example of another brown tabby (in this case of the ticked
rather than the spotted pattern) that has been selected for a large number of
rufous polygenes. The practice of breeding bronze to bronze generation after
generation (known as ‘colour breeding’) paired with the selection of the
warmest cats from each generation causes the accumulation of rufous polygenes
within a line that is necessary to produce really warm bronzes. Breeding
programs for warm bronzes and clean silvers probably cannot be successfully
combined, since the rufous polygenes necessary for warm bronzes will result in
tarnished silvers.
Determining the genotype of a bronze: First look at the kittens your cat
has produced previously
•If your cat has produced a smoke then it is carrying smoke (Aaii)
• If your cat has produced a black then it is carrying smoke (Aaii)
• If your cat has only had bronze and/or silver kittens then you know nothing
certain about its genotype unless it has been bred multiple times to a cat that
is either smoke or known to be carrying smoke, in which case it is likely that
it is a pure bronze (AAii).
Second, look at your cat’s pedigree:
• If your cat has a smoke parent then it is carrying smoke (Aaii)
• If your cat has a black parent then it is carrying smoke (Aaii)
Smoke
Genetically the smoke Mau is a non-agouti black smoke cat that has been
selected to have very prominent ghost markings. A smoke Mau has AT LEAST ONE
inhibitor, I, allele, but lacks the agouti, A, allele. Thus despite its
markings the smoke is not a tabby (i.e. agouti) cat. Egyptian Maus are unusual
in having a patterned smoke. There is no equivalent of the smoke Mau in other
breeds since although black smoke exists as a colour in many breeds, these
other smokes are preferred to show no tabby markings.
Smoke Maus are often used in silver breeding programs. Mixing smokes and
silvers is not fraught with the same problems as mixing warm bronzes and
silvers because with both the smokes and silvers it is desirable to select for
as few of the tarnishing rufous polygenes as possible. Thus silvers and smokes
can be interbred without detriment to either colour. On the contrary, some
breeders maintain that smokes are actually useful for producing silvers with a
high level of contrast between the spots and ground color. It is certainly the
case, that some of the most black and white silvers I have seen have been
produced from smoke to silver matings, and I have been particularly intrigued
by the explanation for this phenomenon. The obvious explanation is that silvers
carrying smoke (i.e. those with AA genotypes) have better contrast than silvers
not carrying smoke (i.e. those with AA genotypes). This would imply that the
agouti gene is additive in its effects with a single A producing a cat with
higher contrast than a cat with two AAs. However, I have been able to find no
evidence from other breeds that this is the case: the agouti gene is believed
to have a simple dominant effect such that AA cats are visually
indistinguishable from AA cats. My preferred explanation for the phenomenon is
as follows. I believe that the smokes retained for breeding probably have much
better contrast relative to the pool of all smokes produced than the average
silver breeding cat does relative to the pool of all silvers, and it is this
that leads to the better contrast observed in the silver offspring of smokes.
Breeders are selecting smokes more on the basis of contrast than they are
silvers because a smoke Mau has to have exceptional contrast in order for the
spots to be visible, whereas in a silver a lesser degree of contrast may be
tolerated in a breeding or show cat if it has other desirable traits. Also, the
dark color of the smoke may mask traits that would cause a silver to be
rejected as a breeding cat, for instance, tarnish will not be visible on a
smoke, and even some pattern flaws may not be so obvious. For these reasons I
believe that contrast is a more dominant criterion in the selection of smokes for
breeding than it is in the selection of silvers. If these arguments are
correct, then the implication is that it is not necessary to breed to smokes to
get good contrast; the same effects could be achieved by breeding from silvers
with the highest contrast.
As with the bronzes, there are two distinct smoke genotypes determined
by whether the cat has one or two copies of the dominant inhibitor allele:
• aaII = homozygous smoke, i.e. pure smoke
• aaIi = smoke heterozygous for the inhibitor allele, i.e. smoke carrying
bronze
Determining the genotype of a smoke:
First look at the kittens your cat has produced previously:
• If your cat has produced a bronze then it is carrying bronze (aaIi)
• If your cat has produced a black then it is carrying bronze (aaIi)
• If your cat has only had smoke and/or silver kittens then you know nothing
certain about its genotype unless it has been bred multiple times to a cat that
is either bronze or known to be carrying bronze, in which case it is likely
that it is a pure smoke (aaII).
Second, look at your cat’s pedigree:
• If your cat has a bronze parent then it is carrying bronze (aaIi)
• If your cat has a black parent then it is carrying bronze (aaIi) 
Black
Genetically the
black Mau is a plain old non-agouti black cat with the black masking the spotted
tabby pattern. There is only one type of black Mau, aaii, and it is the most
recessive of the many Mau genotypes (excluding the blues). The fixed genotype of
the black Mau, aaii, gives it specific uses in a breeding program. Blacks can be
used to perform test crosses to reveal the genotypes of other cats. For example,
if a silver is bred to a black and produces only silver kittens time after time,
then the chances are very good that the genotype of this silver is
AAII.
The down side of
using a black in a breeding program is that it is equivalent to choosing a
breeding cat with your eyes closed! Given the importance attributed to colour,
pattern and contrast in the Mau standard, using a black Mau in a breeding
program is not something to be undertaken without some thought. The black masks
not only the pattern, making it hard to see whether the cat has any pattern
faults and how good its contrast is, but also the rufous polygenes, making it
impossible to tell how tarnished it is. It's important to remember that just
because you can't see the pattern, contrast and tarnish it does not mean that
there are not genes for these traits present; they are there, you just can't see
their consequences. Of course the black Mau’s parents can be used to get some
idea about the pattern and contrast genes it is carrying and the number of
rufous polygenes it is likely to have. A second problem of breeding from blacks
is that the probability of producing black kittens is greatly increased (see the colour table for details). In general it is not advisable to breed from a black
Mau unless it comes from irreplaceable lines or it has absolutely outstanding
health, temperament, type and eye colour.
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