Current knowledge about the Husky

“domino” allele and its interactions

Written (hastily) by Corinne Benavides, 20 April 2017. E-Mail: corinne_gyger@yahoo.de

The currently known alleles at the E locus are:

• EM: Mask
• E: wild type
• eg: grizzle/domino in sighthounds and other breeds
• eh: Cocker “sable”
• e: recessive yellow

Other alleles have been mentioned but not yet published or linked to
coat color:
• Due to the phenotypic similarity to eg, as well as observed inheritance, it is extremely likely
that most Huskies and Malamutes have an E-locus allele that is responsible for their
distinctive color. The most likely candidate for this allele is a mutation termed R301C,
described in a 2013 paper by Olivier et al.
• Animal Genetics has identified an allele and termed it ew, but no publication is known.
• Vetgen has made a study involving a new allele and used the designations e1 and e2, but no
publication is known.

About the dominance of eg:

In the original paper, the eg allele has been described as dominant to E – this seems not to be the
case. Numerous litter records show a recessive inheritance (proof can be provided). It also makes
sense that eg is recessive to E since generally in mammals, dominant alleles at the E-locus result in a
darker phenotype while recessive alleles result in less eumelanin – which is the case in
grizzle/domino.

The original paper also implied that eg needs a genotype of at/at at the Agouti locus to express.
Observation has now shown that the eg/eg or eg/e genotype affects not only the expression of all the
Agouti alleles, but also dominant black and brindle (KB and kbr).

Similarities between the recessive E-locus alleles:

eg, eh and the proposed „Husky domino“ allele have in common that they reduce the expression of
eumelanin, especially on the legs, face and ventral side as well as the hair roots. Knowing that
recessive yellow (e/e) is able to cancel out all eumelanin even in a KB/- dog, it does not surprise that
the three mentioned alleles also can affect dominant blacks, albeit in a less thorough manner than
e/e. The eh allele in Cockers is known to result in a sable-like phenotype when paired with KB.
Phenotypic effect of the “domino” alleles:
(Shown on the example of eg – observations have shown the Husky variant to behave very similarly)

When combined with ky/ky at the K locus, an eg/eg or eg/e genotype results in:

• With Ay/-: Overall pheomelanin with a little eumelanin seen at birth, which often disappears
by adulthood. Called „newsprint cream“ and other terms by sighthound breeders.
• With aw/-: Enlarged light areas on face and legs, generally less eumelanin and light hair roots.
• With at/-: Grizzle/domino phenotype with moderately to greatly enlarged tanpoints and light
hair roots.
• With a/a: Only a handful of such individuals are known, but they show a tanpoint pattern
with grey points.

When combined with KB/- at the K-locus, an eg/eg or eg/e genotype results in:

• With Ay/-: Overall grey phenotype, often with a gradient (darker back, lighter ventral area).
May also cause the “bronze” phenotype where pheomelanin is not diluted to white. Called
“blue” in Afghans, “chinchilla” in Borzois, “black-nosed grey” in Salukis, among other names.
• With aw/-: Agouti phenotype with seemingly overlayed light to dark grey shade.
• With at/-: Tanpoint phenotype with grey instead of red/white tan points. Called “shaded
grizzle” by Saluki breeders.
• With a/a: unknown, expected to look similar to at/- eg/- KB/-.

When eg/- is combined with kbr/- at the K-locus, the result is a combined effect of the effects
described above – with Ay, the result can be a brindle looking dog with light, mostly unbrindled,
ventral and facial areas. Called “oyster brindle” by Afghan breeders.

Breeds with “domino” alleles:

To my knowledge, eg has been confirmed in Saluki, Afghan Hound, Borzoi, Azawakh, Sloughi, Chart
Polski, Silken Windhound, Lurcher, and Finnish Lapphund.

To my knowledge, eh has been confirmed in English Cocker Spaniels and Beagles.

Other breeds where a domino-like phenotype has been observed are Siberian Husky and related
breeds, Alaskan Malamute, Tibetan Mastiff, Tibetan Spaniel, South Russian Ovcharka, Chihuahua,
Lapponian Herder.

My wish:
That the Husky “domino” allele be described in a scientific paper and a commercial test made
available. This would help discover the various interactions this allele has with other colors.

My conviction is that the R301C mutation described by Olivier et al. is the most likely candidate for
this allele.
Sources:
Relevant papers:

• Dreger & Schmutz 2010: https://www.ncbi.nlm.nih.gov/pubmed/20525767

• Olivier et al 2013: http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0075110

Cocker sable: http://chestnutenglishcockers.com/ECSsable.pdf

Tested sighthounds:
https://docs.google.com/spreadsheet/ccc?key=0AhVRfk5exTRqdC1wRTdYcVZLSkJrNEtUOEhLTlZLZ0E

Good overview of domino: http://www.kwestmals.com/malcolorgenetics.html by Nichole Royer.

Pictures
If I had more time, I would make picture charts and examples, but for now this will have to do – a
simple chart with salukis. Pictures are NOT mine, so please do not distribute. Images are
approximations and do not necessarily represent the genotypes.
Examples of Black/Brindle + eg
Compiled by Arlene Dykstra from data submitted by breeders