Saluki Coat Color DNA Study
an update and information page for the owners of Saluki who have contributed DNA brushes to the study
This webpage was posted on August 20, 2009 and last updated on July 4, 2010 by Sheila Schmutz. Please direct all comments regarding this page directly to me.
By mid-October the original set of dogs and the official Saluki Coat Color Study was completed.
The research phase to address the gene/s causing grizzle was completed in spring 2010. The manuscript describing this will be published in Journal of Heredity in Fall, 2010 but is available as "online early". A free PDF copy of the manuscript is available for download
We are grateful to Margot Keast of Australia and Casey Gonda of the U.S. for jointly coordinating the samples, photos, consent forms, etc. for this study. I was approached by a few Saluki owners in various countries about the same time and asked that the groups work together on one project. I asked that they attempt to collect DNA brushes from Salukis registered in various countries, in all the colors recognized by at least one country, with the exception of the color they call blue. That color is not yet understood in Afghans and so I assumed it would not yet be in Saluki either.
Please note that I am apparently being said to have stated many things on chat lists and emails. Do not assume that I actually said anything that is not stated on this website. Others may choose to interpret the final data set in their own way, but that does not mean that I agree with their interpretation.
We also thank the dogs and their owners who contributed to the study. The dogs donated DNA via cheek brushes and posed for the photographs. The owners paid for the testing through contributions to this study.
Both intial batches of DNA brushes arrived in Saskatoon at our lab on August 19, 2009. Further samples arrived through the end of September. Note that the sample size is small, given the variety of coat colors in Salukis worldwide. Therefore one should not assume that a specific color of dog that resembles one shown on this webpage is always or only the genotype of the dog shown. A good example is the two different genotypes for the two cream dogs.
Call names that were provided will be used, not the registered names of the dogs. Generic information is posted on this site. Some specific information was emailed to the owner of each dog.
We have completed DNA testing of this set of Saluki for the known alleles at the loci: A, E, K, B, and S. No further dogs will be included in this full color study, at this time.
In the future, we may try to address some of the other outstanding questions discussed at the bottom of this page.
The stated coat colors of dogs sampled were:
Some dogs had white markings and others did not.
Note that there is a very good powerpoint presentation saved as a downloadable pdf file produced for AKC (see link at the bottom). Apparently AKC considers sable and grizzle as markings. On this website, white markings will be discussed as "markings".
One of the challenging questions asked was what gene causes the coloration known as grizzle in Saluki, shown by Taz at the top of the page. The term grizzle is used in a few breeds, often hound breeds, but may not refer to the same phenotype in those breeds, as it does in Saluki.
The definition of grizzle that Casey and Margot agreed upon is: "Grizzle – A pattern that is present at birth and remains throughout life, which comprises a dark overlay covering the top and sides of the body and outside of the limbs, from the top of the head to the tip of the tail. The darker color on the head gives the impression of a "widow's peak" between the eyes where it contrasts with the lighter color of the face. The underside of the dog and inside the legs comprise the lighter base color. The overlay color may be Black, Grey/Silver, red or Chocolate and can vary in intensity with the seasons and age. The base color may be various shades of red, fawn, golden, cream or silver."
So we wondered if the body coloration caused by individual reddish and black hairs intermingled or are the individuals hairs composed of alternating bands of red (phaeomelanin) and black (eumelanin) pigments, or are the hairs phaeomelanin with eumelanin tips as I'd interpret "overlay", or ??? The AKC powerpoint presentation by Walls, Hinsch and Stoner-Townsend say that all grizzles have banded hairs.
Uno, above, is a Canadian dog who was added to the grizzle portion of the study, as were several other Saluki. He would be called a deer grizzle by some and a fawn grizzle by others.
Flame, above, is one of the grizzle dogs in the original set of samples. She was described as a red grizzle.
Glitz was also part of the original set of samples. She was reported as a black grizzle.
Taz is a Canadian dog who was added to the study to help address this interesting coloration. His face is shown at the left and his full body at the top of the page.
Tyghar is another grizzle Saluki, whose face is shown at the right. His body shot is shown in the brindle section.
Mariah is another grizzle Saluki. She is shown in the section on brown below, as an adult and pup.
The six grizzle Saluki illustrate the wide range of shades in this pattern. All of them have the widow's peak facial pattern. All except Tyghar who is also brindle, are ky/ky. None have an ay allele. None are e/e, although Uno and Tyghar both have one e allele.
The gene causing grizzle had not been previously reported. Our research suggests that a unique mutation in one of the genes known to affect coat color, MC1R, is primarily responsible for this pattern. A single copy of this mutation, which we named EG seems sufficient to cause this pattern. (Uno has only one). However, this mutation interacts with the ASIP gene for the grizzle pattern to be visible. This is shown by Bertha and Chief, both of whom have the EG mutation but not the required at/atgenotype at the A locus.
A similar pattern in Afghan hounds is called "domino" after a famous show dog by that name. Domino and grizzle are both caused by this same gene interaction.
Cream, Red, Fawn, Sable
There a two different genetic mechanisms by which a dog can be primarily pigmented with phaeomelanin. Phaeomelanin is the reddish pigment and Eumelanin is the blackish pigment.
The first mechanism is an "e/e" genotype at the MC1R gene or E locus. In dogs, this is "clear red", in that there is never a single black hair on such a dog. However the shade of red can vary from cream to deep dark mahogany red in various breeds. Pansy is an example of a cream Saluki with an "e/e" genotype. Not all Saluki with an e/e genotype will be cream or very pale, although this is the case in some breeds. Pansy and the two Saluki siblings Cygfa and Torc were the only dogs in the original studygroup that had this genotype. Among those 15 Salukis, "e" was a rather common allele.
The genotype "e/e" is epistatic or masks the colors normally produced by several other loci including A and K. For example, Pansy is not fawn. She does not have an ay allele.
As a pup, Pansy showed some reddish pigment on her ears.
Bertha is another Saluki that is cream. However Bertha does not have an e/e genotype, or even a single e allele. Her genotype at MC1R is EG/EG. Bertha has an ay allele so is not grizzle.
We do not know why Pansy is cream and not a darker shade of red. But in some ways Bertha is even more puzzling since she is not e/e. However, she does have an ay allele. There must be another gene, yet undiscovered, that causes cream in Saluki. In Afghans, dogs with e/e or with an E or EM or EG allele can all be cream. This seems to also be true in Saluki.
The second genetic mechanism for a reddish coat is having an ay allele at the A locus or ASIP gene and also being ky/ky at the K locus or beta defensin 103 gene, and at least one E or EM allele at the MC1R gene. Such dogs can have a few black hairs on their body and can have some black tipped hairs, especially as pups. Among the 15 Saluki for which genotyping has been done thus far, the ay allele was not common.
Chief is a red Saluki of the fawn/sable type. He has an ay allele and is ky/ky. He has a genotype of EG/e at MC1R. Although Chief is fawn, he has pale ear fringes.
Several people in Saluki circles seem to consider that sable dogs would always have dark tipped hairs and an "overlay" of some eumelanin color. Although this is true in some, Chief and Simba (below) demonstrate it is not true in all. Alternatively the term sable is used only for dogs with black ear fringes, even though other dogs have a fawn/sable ay allele.
Simba was reported to be a fawn Saluki. He has one ay allele and is ky/ky. He is EG/e at MC1R.
Ali is a Saluki that is sable, or black fringed red, as it is sometimes called in this breed. This is because Ali has an ay allele and is also ky/ky. Ali shows at least a hint of an "overlay" from her black tipped hairs and she has black ear fringes. Ali is E/E at MC1R.
Some dogs in some breeds have three stratified sections of pigmentation. They have eumelanin on their backs - either black, brown or grey and then phaemelanin on their legs and often on their eyebrows and muzzle although we are concentrating on body pigmentation here. Finally these dogs have white or no pigmentation on their feet and belly area.
Dogs that are "eumelanin-and-tan" would have an at/at genotype at the A locus or ASIP gene and also ky/ky at the K locus or beta-defensin 103 gene and would have at least one E or EM allele and no EG allele.
Oasis is a Saluki that is black-and-tan. Her tan points are more evident against black than are those of Flirt who is chocolate brown, but they occur in the same places. The tan of Oasis is a deep phaeomelanin color. Although she does have one e allele and one E allele, that is not the explanation of this depth of tan. Why some dogs have a deep tan color and others a pale cream isn't known.
Smurf is a Saluki that is called black-and-silver. This is because his "tan" looks like a pale "silver" color. This is apparently very typical in Salukis. He is E/E at MC1R.
Smurf's face shows the eyebrow spots in tan or silver, typical of a dog with tan points.
See Flirt below for another eumelanin-and-tan example of a Saluki.
Brown or Chocolate
Brown is a color that is sometimes confused with red. However, it is an alteration of eumelanin, not a phaeomelain color. Tan, as used by dog people in terms like "black-and-tan" is a phaeomelanin color.
Flirt is brown-and-tan with white undersides. Her tan is not that easy to see although it is quite evident on her tail in the photo of her full body. Her face shows the classic "eyebrows" of a dog with tan points.
There are 3 different alleles that cause brown in dogs: bs, bd, and bc, in that order of incidence. Flirt is bs/ bs. Although there are 3 alleles causing brown in dogs, only the bs allele and the bd allele were detected in this group of Saluki.
In some breeds brown is even called red, unfortunately. In some breeds it is called chocolate or liver. Flirt was described as chocolate with tan points.....another way of saying brown-and-tan.
In addition, all dogs with a b/b genotype (any two b alleles) have brown nose leather and pads. No dog with a b/b genotype ever has any black pigmentation anywhere.
For tan points to show the dog must be ky/ky and Flirt has this genotype. Such dogs would also be at/at.
Mariah is another Saluki that is homozygous bs/ bs. Therefore her fringes and the tips of her hairs are brown, not black. Mariah did not have an ay allele. She is not a chocolate-and-tan like Flirt above, because she has an EG/E genotype at MC1R and is therefore grizzle.
In her pup photo, Mariah's brown on her head and tail and nose show more clearly than on her adult photo. However brown and red are not that different in color to make it obvious that her hairs have brown tips over her back.
The original locus for black versus brown described by Little was the B locus. Contrary to Flirt and Mariah above, Rasho is B/B. A single B allele would cause black instead of brown pigment.
Rasho has a KB allele which makes his alleles at the A locus, one of which is ay, not show. His other allele is ky.
At the MC1R gene or E locus Rasho has an e allele and also the top dominant allele EM for melanistic mask.
Farasha is another black Saluki, a daughter of Rasho, shown above. At the MC1R gene or E locus she has an EG allele and also the top dominant allele EM for melanistic mask. Her mask is not evident on her white muzzle however, since white is the result of the absence of melanocytes.
Note that black can also be caused by an "a/a" genotype at the ASIP gene. This allele is typically found only in herding breeds. It was not found in any of the Saluki that were tested for this allele.
Brindle is typically seen as vertical stripes on the body of the dog in a phaeomelanin color (cream, yellow, red) interspersed with stripes of a eumelanin color (black, brown, gray). It is often easier to see on long coated dogs in young pups before the coat lengthens. The amount of brindling varies widely among individuals (see the Brindle page in the main coat color series). Placement of brindle only occurs on certain areas, typically dictated by the genotype at the A locus.
Due to the complex nature of the brindle mutation, all brindle dogs test as if they have a KB allele and a ky allele. The current commercially available test does not distinguish between dogs that are black from KB and carrying a ky allele and brindle dogs. Brindle can be detected by another type of assay that is currently available only in the research labs involved in studying the K locus.
Tirgan is a Saluki that was reported as brindle with a black mask. She is EM/e at MC1R and therefore this fits with her large black facial mask.
She is what one would call a "full-body" brindle because her stripes occur on her whole body, not just her tan points. Tirgan has an ay allele which is typical of full-body brindles.
Her genotype at the K locus is compatible with being brindle.
Tyghar was reported as brindle and grizzle. His genotpe is EG/e at MC1R and at/at which explains his grizzle phenotype.
His genotype at the K locus is compatible with being brindle.
We do not have a DNA sample from a black or chocolate dog with brindle points, but such dogs could occur based on the alleles identified in this group of Saluki. Seeing brindle on tan or silver points in dogs that also have white markings can be difficult.
The e/e genotype is epistatic to both black and brindle and also to sable. Not a single black hair or part thereof can occur in dogs with an e/e genotype.
Some Salukis have white markings. These can be on the feet or up onto the legs and the underbelly. A few even have a white collar. In some breeds these markings of white collar, white legs and white undersides can be called "Irish Spotting", whereas in some other breeds it might be called "mantle". The mutation causing such markings has not yet been established.
In many breeds there is also a pattern of white markings that are random white spots on the body. This can be called piebald, but in Salukis it is called "parti" or particolor. Gala shows this pattern of white markings. In many breeds that we've studied, piebald is caused by the homozygous presence of a SINE in the promoter region of the MITF gene. However, in a few breeds this is not the case, such as Great Danes, Norbottenspets, Icelandic Sheepdogs, Whippets and Saluki. Gala has a genotype of SINE/not instead of SINE/SINE.
Gala is a tricolor and therefore as expected has no ay allele. Seeing tan points on particolor dogs can be difficult at times. Gala has them only on her face since white is because of the absence of functional melanocytes.
Sweetie is a particolor and she has a genotype of SINE/SINE. Her colored areas are a very pale red but if you look closely, you can see she is pale red with white spots. She has an EG/EG genotype at MC1R and has one ay allele.
We assume that there is another mutation in another gene involved in white spotting that interacts or interferes with MITF in some breeds that causes them to be different. In Great Danes, we suspect these are merle and Harlequin. Neither of these patterns occur in Norbottenspets, Icelandic Sheepdogs, Whippets or Saluki though so this suggests yet another gene must affect white markings in those breeds.
The only other Saluki in this group that had a SINE/SINE genotype was Lola, shown below in the "Pup to Adult Color Changes" section. As a pup Lola had an interesting colored spot on her belly, typical of a piebald or particolor dog but the rest of her markings are more similar to Irish spotting. This further confirms that Saluki do not fit the common patterns of white markings associated with this mutation.
Mojo is not a dog in this study. However I saw his photo and obtained permission from his owner to post it here because it illustrates an interesting point. At the age this photo was taken, Mojo looks like a "black-and-silver" although when he was younger he had more evident tan and also white. At this age, one way to tell that his "tan" points are now silver and the his belly is white, is that his belly has ticking. His "tan" marking across his shoulders has no ticking. Ticking only occurs on white, never tan or even pale tan.
The gene and mutation for ticking has not yet been identified, although one researcher stated it has been mapped. The presence of ticking is considered dominant to its absence, since the work of Little in 1957.
Pup to Adult Color Changes
Saluki pups do not necessarily look as pups, like they will as adults. An example of this is illustrated by Lola at 6 weeks on the left and as an adult on the right. Lola is a sable Saluki. She has one ay allele and is also ky/ky. Of course the photos were taken a long time apart in different lighting, even though both are outdoors. But notice the difference in her facial coloration. She has paled as an adult.
Some of the later questions will not be answerable as part of this study.
In order to answer this question we sought Salukis that were not part of the intial study set. This family story was very helpful. Gabriel was E/e at MC1R and Zia was EG/e. Both have one ay allele. Since they had one black-and-tan pup and one black-and-silver pup we can assume that their second allele is at. Gabriel has black ear fringes and Zia does not.
We also DNA tested two of their five pups and each received an e from each parent and also an ay allele from each parent. Torc looks a bit more reddish than Cygfa below. I'm not sure if each would be called cream by all Saluki fanciers but against the snow, it's evident that neither is white. Some people would likely call one or both gold.
Coat color testing for Salukis will be offered by HealthGene this summer. See http://www.healthgene.com
Saluki Club of America, AKC Color Registration Options Illustrated. This is a large pdf file, complete with photos and descriptions of Saluki colors recognized by the American Kennel Club prepared by Sharon Walls, Gertrude Hinsch and Montrue Stoner-Townsend. (2004).
Dog Coat Color Genetics Main Page
Sheila M. Schmutz, Ph.D.
Department of Animal and Poultry Science
University of Saskatchewan
Saskatoon, Canada S7N 5A8