Puli Coat Color DNA Study
an update and information page for the owners of Puli who have contributed DNA brushes to the study
This webpage was last updated on December 3, 2007 by Sheila Schmutz
We are grateful to Linore Cleveland for coordinating the samples, photos, consent forms, etc. for this study. The first DNA brushes arrived at our lab in Saskatoon in May, 2005. Susan McConnell has agreed to present an update about this study at the 2005 specialty this fall.
DNA analysis is being conducted in the lab of Sheila Schmutz at the University or Saskatchewan, Saskatoon. Interpretation is a collaborative effort with Dr. Sue Ann Bowling, Alaska who intially suggested studying this breed.
The most common color for Puli is black. They also occur in white. Fako is a Hungarian term used to describe Puli of other pale colors. Some Puli are called apricot. Colors other than black and white are not allowed to be shown in all countries.
Useful references on Puli color were provided by Sue Hopgood of Great Britain. She had saved an article entitled "Coat Colour and Pigmentation and It's Inheritance" from a breed club magazine. A book entitled "The History of the Puli" written by Arany Csaba in 1998 was particularly helpful. This book mentions pale pups born to two black parents and black pups born to two black parents, which implies that either Puli have two forms of black or two forms of fako.
Some black Puli gray with age. Whether this is the progressive graying similar to Poodle silver or whether it is more that some white hairs gradually occur amongst the black ones, is not clear.
Libbye Miller is a vet who has worked with us on a color study in Belgian Sheep Dogs. She co-owns a white Puli too. She noted: "Since they don't shed, the cords grow continuously and it takes them about 5 years to reach the ground. The cords are like a little diary of the changes in their coat color over the years. And if you let the white Puli work in a field with those purple poke berries...you'll have to hear about that for years too!". This is certainly not typical of most other dogs breeds and may affect the way color is "seen" in Puli.
There are 3 alleles at this locus. E, e, and EM. "Melanistic Mask" is caused by the EM allele and results in eumelanin pigmented hairs on the muzzle. EM is the top dominant allele at the E locus. In other words, EM > E > e in its effect on phenotype. We have found both the E and e alleles in Puli so far.
Although both Puli pups at the right, Danka and Dalia, appear to have melanistic masks, unless these persist into adulthood, they are not considered masks. Chows and Akitas are all typically born with dark muzzles but in Chows they all fade as they age, whereas in Akitas, those dogs without an EM allele fade and those with at least one such allele, remain.
Dalia is male Puli with an E/E genotype. His coat color is known as fako, often called masko fako. Puli do not seem to have a full melanistic mask caused by the EM allele. Instead, like Chows their puppy mask fades as they grow into adulthood. Fako may encompass more than one color, but we don't know that yet. Dalia is a "fawn" dog with a genotype ay/ay. In Hungary, fako seems to be used for any pale color, including gray.
The photo at the right shows a cord from these same two dogs as adults. Note that the reddish hairs have faded to cream but the few black hairs are still black.
Sadie, the black dam shown here, had a litter of one reddish pup named Hector who is hiding behind his white sister Blizzard. They had two black littermates, Raz and Madeline. Their sire was Corin, a black Puli.
The photo at the right shows cords from these same dogs as adults. Note that the reddish hairs Hector had as a pup have also faded to cream of a shade almost, but not as white as Blizzard. Neither Blizzard nor Hector have any black hairs. This is typical of dogs with an e/e genotype.
Hector, at the left retained more reddish pigment as an adult than Blizzard, middle. However, Hector faded somewhat since he was a pup. Both have e/e genotypes. Raz, one of their black littermates, left, has an E/e genotype as does the dam, Sadie.
Dogs may be black by two different genetic mechanisms. Dogs that have two "a" alleles at the agouti locus inherit black as a recessive trait. They are unable to make red pigment since their agouti gene is not functional. The "a" allele occurs in Puli and several other herding breeds. Kasa, at the left, is "a/a" in genotype.
Black that is inherited as a dominant is caused by having at least one copy of the KB allele plus at least one copy of the E or EM allele. This black may also occur in Puli since most breeders consider black dominant to white and fako. Kasa is E/E and KB/ky
Keddi, a black dog with a white dam and black sire, tested E/e at MC1R. He is also "a/a" so would be black, even if he inherited no K. However his genotype is also KB/ky.
Proof that black can be caused by both genetic mechanisms in Puli is beautifully illustrated by Raz and Madeline, two black littermates. Raz is "a/a" but Madeline has only one copy of "a". Therefore Raz is an "a/a" black and Madeline is a KB/KB, E/E black.
Some Puli are e/e at MC1R and in the case of these dogs, one can not predict their K genotype. Such e/e dogs could be KB/KB or KB/ky and still not be black since the e/e genoytpe prevents black pigmentation of hairs in dogs (but not nose leather or pads). This is an example of "epistasis".
Macintosh, on the right, is a white Puli with an e/e genotype. Therefore one can not predict his genotype at K. He had white parents and although he has had a black pup, this pup was born to a black dam. White appears to be inherited as a recessive to black, according to breeder experience and "The History of the Puli". If e/e is required for white to occur in KB/-, E/- black dogs, then it would appear to be inherited as a recessive.
Note that Blizzard, shown earlier, is another white Puli with an e/e genotype. So far these are the only two white Puli we have tested, and both are e/e.
Linda Hall took the photos of some of her grey Puli over the years and has shared them. The photo above illustrates that Puli have "progressive greying". These dogs were all born black. She refers to them as "graying", "silver" and "black". Not all the Puli continue to go to the pale silver color. Some stay a darker grey.
Linda Hall took the photos above of Spot at 5 weeks, 12 weeks and one year of age. She says that not all Puli begin the graying process this young however.
Dog Coat Color Genetics Main Page
English translation of an article published in Unkarinpaimenkoirat (Hungarian herding dogs) magazine in April, 2002.
Sheila M. Schmutz, Ph.D.
Department of Animal and Poultry Science
University of Saskatchewan
Saskatoon, Canada S7N 5A8
phone: (306)966-4153 fax: (306)966-4151