This page was created by Ashley Cirimeli. The Primary focus of this page is on Borzoi but I will include some Silken Windhound examples too. These two breeds have a fantastic rainbow of colors in their respective gene pools. I have put together this page as a reference/guide for anyone interested in the magic of color genetics.
If you learn something, consider buying me a cup of coffee!
First and foremost, this page will use genetic color terms as opposed to breed color terms. There has been some confusion with the varying names (grizzle vs sable vs sabled, for example). Below is a chart that will list genotype, breed term, and genetic term. Where a genetic term for a specific color combo doesn’t exist, breed terms will be used instead. When we say "sable", we are speaking of the Ay allele (think collies and shelties), not Eg modifying Black and Tan (what borzoi folks call Sable).
| Genotype | Genetic Term | Borzoi/Silken Term |
|---|---|---|
Genetic Jargon Explained
As you read this page, you'll see a lot of words that may be unfamiliar, like homozygous, allele, locus, dominant, etc. Here is a handy guide to help!
Locus: A locus refers to the location on the chromosome where the gene is found. Example: black and tan, at/at is on the A locus
Allele: Alleles are variants of the same gene that occur on the same place on a chromosome. Example: at/at, Ay, and aw are all alleles on the A locus.
Homozygous/Heterozygous: Zygosity is the degree of similar traits in an organism. The prefixes homo and hetero respectively mean same and different. Example: at/at is homozygous whereas Ay/at is heterozygous.
Eumelanin: Black pigment
Phæomelanin: Red pigment
Dominant/Recessive: Brindle for example, is a dominant allele, meaning a dog only needs one copy of the allele in order to express the color/pattern (Kbr/-). Black and tan is a recessive allele; a dog needs two copies in order to express black and tan (at/at). We use capitals and lower case letters to differentiate between dominant and recessive alleles. On the A locus, at, aw, and a are recessive while Ay is dominant.
There is also a hierarchy of dominant and recessive alleles on each locus. On the K locus, both dominant black (KB) and brindle (Kbr) are dominant alleles-- the dog only needs one copy to express the color/pattern. However, what happens when a dog is KB/Kbr? Both are dominant alleles, but KB is dominant to Kbr, therefore the dog will be black.
Genotype: The genetic constitution of an organism.
Phenotype: The observable characteristics of an organism.
Color Guide
Dogs contain two pigments that influence their color: Eumelanin (black) and Phæomelanin (red). There are genes that can modify/dilute these pigments to look like a whole host of other colors (silver, liver, blue, etc.). White isn’t a pigment, but rather the absence of pigment.
It’s easy to comprehend canine colors when thinking about them in terms of layers. White is the top most layer, then black, then red. White hides all, black hides red. Think of white like a blanket that falls on top of everything else. Holes in the white (caused by any number of genes such as piebald or ticking) will show the color that exists underneath. Sometimes that white blanket is small and only shows up as a few hairs on the chest, or a blotch over the toes. Sometimes it covers the entire dog.
Similarly, holes/breaks in black will reveal the red underneath (black and tan, brindle, etc.).
The color of the dog still exists underneath the white! So you will still see the markings underneath if there are holes/breaks (spots) in the white pattern. In this example, you can still see the black and tan markings!
Spotted: Also called “Piebald”. Spotting is mostly random as far as we know, though some lineages seem to produce very similar markings. Heavily spotted dogs can look similar to irish marked dogs. Sometimes the spotting is restricted entirely to around the eyes and ears, often called “extreme white”.
Spotted dogs test as sp/sp.
Irish Marked: This type of white pattern describes a dog who has more white patterning up the legs, undersides, the tip of the tail, and often a partial or complete “collar”.
Most Irish Marked borzoi and silkens test as S/sp, which is often referred to as “Pseudo Irish”. An Irish marked dog who tests as S/S has true irish markings. A dog with one copy of true irish markings, and one copy of spotting (also tests as S/sp) will have much more white coverage, usually a full white collar that connects to a wide blaze on the head, high white legs, and sometimes a spot of white around the hips.
Self: The “Self” descriptor referrs to a mostly solid pigmented dog with minimal white markings, usually caused by residual white. The white is usually on the chest, toes, and tip of the nose. Residual White occurs when pigment doesn't migrate fully as the embryo develops.
Most self dogs test as S/S however, some self dogs also cary for spotting and may test as S/sp. As mentioned above, S/S can also be a result on true irish marked dogs so phenotype will be your best indicator there.
Health concerns: Deafness can be caused by pigment deletion within the inner ear if it reaches the cochlear hairs. Anecdotally, this seems quite uncommon in borzoi but has popped up a few times in silkens, as silkens seem to have more occurances of “white head” and “split face” (unknown genetic markers at this time).
A genome-wide association study of deafness in three canine breeds
STRING data mining of GWAS data in canine hereditary pigment-associated deafness
White Markings
MITF and White Spotting in Dogs: A Population Study
Important Note: Only the sp allele (spotting) has been identified on the S locus, so other white patterns will all test as S/S. This includes true irish markings (often written as Si). If you color test your dog who phenotypes with irish markings but genotypes as S/S, your dog likely has at least one copy of true irish markings!
The K Locus
Dominant Black: Because it's a dominant allele, a dog only needs one copy in order to express dominatn black. In most cases, a dog carrying one copy of the Dominant Black allele will only appear black (aside from white patterns). The exceptions to this would be liver (b/b), blue (d/d), sometimes grizzle (eg), recessive red (e/e) and seal (unknown locus/allele). These alleles all modify, dilute, or hide the eumelanin in different ways.
Dominant black tests as KB
Brindle: Brindle is a dominant allele, but it is recessive to dominant black. Pheomelanin is expressed where there are breaks in the black pattern, creating a striped look. The width of these breaks can vary. Any allele(s) that dilutes or modifies black or red pigment will alter the color/appearance of the brindle; for example: above right, silver brindle. Genes that modify phæomelanin can similarly alter the look of a brindle dog.
While we write the brindle allele as “Kbr” for ease, every brindle dog will test as “KB/ky” because brindle is the result of a duplication of the KB and ky alleles. As of writing this, the only lab that can reliably test for brindle is Laboklin. All other labs will display brindle as KB/ky, whether they are KB/Kbr, Kbr/Kbr, or Kbr/ky.
The ky allele: This is the most recessive allele on the K locus, and does not alter appearance. A dog that is homozygous for ky will not express dominant black or brindle, but instead express whatever other alleles are present on the other color loci. They can still express eumelanin in their coats from the A and E loci.
The E Allele: This allele does not appear as a specific color or pattern, similar to ky. E is recessive to Em, but dominant to eg and e-- meaning a dog who is E/eg will not show the effects of grizzle and instead will show whatever color/pattern they have on the A locus, and/or K locus. When testing dogs, E is the "process of elimination" allele. When a dog tests negative for Em, Eg, and e, labs will list results as E.
Recessive Red: Also called “recessive yellow”. Recessive Red completely eliminates all black hair pigment except for somatic mutations. Sometimes it can affect the pigment of the skin on the nose, lips, and eyelids too. The intensity of this red can vary from almost white to a rich dark red/brown. It can be tricky to differentiate between recessive red and a grizzle cream or a clear sable. Usually grizzle cream will have dark silver on their ear tips that fades with age, clear sables will have at least some black whiskers, but recessive reds will have no black hairs at all, unless they express somatic mutations.
Recessive Red tests as e/e
Grizzle: Grizzle is not a specific color or pattern but rather an allele that modifies other colors and patterns. It is Recessive to Em, and E, but dominant to e. When paired with eumelanin on the K locus, it seems to fade the color to shades of silver. When paired with the A locus, it seems to restrict the expression of eumelanin markings.
Scroll down to read more about the different grizzle combinations of colors (silver, sivler brindle, classic grizzle, grizzle brindle, grizzle cream/gold).
Mask: The Mask allele is the most dominant on the E locus. Masking describes the eumelanin on the face of the dog. It can range from just a small bit around the nose to a full face. In some breeds, masking can extend beyond the head! This is commonly referred to as “extreme masking”.
Masking tests as Em
The hierarchy of dominance on this allele is as follows: Em, E, eg, e. As far as we know, additional E locus alleles such as eA and eH are not represented in either borzoi or silken gene pools.
The E Locus
The A Locus
The only two alleles on the A locus that exist in Borzoi and Silken Windhoudns are Ay and at. However, UC Davis has updated their A locus reporting to separate Ay into “dominant yellow” (ASIPᵈʸ) and “shaded yellow (ASIPˢʸ), as well as giving saddle it’s own A locus allele as “black saddle” (ASIPᵇˢ). They also report 3 different variants of the at allele as “black back” 1-3 (ASIPᵇᵇ¹, ASIPᵇᵇ², ASIPᵇᵇ³). The hierarchy of dominance in borzoi and silkens is as follows: ASIPᵈʸ, ASIPˢʸ, ASIPᵇˢ, ASIPᵇᵇ¹, ASIPᵇᵇ², ASIPᵇᵇ³. Read more here.
Dominant Yellow: This phenotype can be indistinguishable from recessive red, or grizzle based creams and golds. Dominant yellow is the most dominant allele on the A locus.
Dominant Yellow tests as “Ay” with most labs and ASIPᵈʸ with UC Davis.
Shaded Yellow: This is the phenotype often called “Sabled” by borzoi and silken folks, that the rest of the dog world calls “sable”. The heavily shaded phenotype can look quite similar to a classic grizzle, though usually the widow’s peak isn’t as strong in a shaded yellow. Sometimes the shading is contained only to the ears, cheeks, and parts of the tail. Shading is recessive to dominant yellow and requires two copies of the allele in order to express.
Dominant Yellow tests as “Ay” with most labs and ASIPˢʸ with UC Davis.
Saddle, Creeping Tan: This phenotype is not present in borzoi from what I have seen, but does occur in Silken Windhounds. The UC Davis genetics lab has assigned Saddle its own allele on the A locus as ASIPᵇˢ for black saddle. They list ASIPᵇˢ as dominant to any of the “black back” alleles so a dog that is ASIPᵇˢ/ASIPᵇᵇ³ will phenotype as saddle. In other labs, they test for the presence of RALY though an at/at dog with two copies of RALY does not always phenotype as a saddle, as is the case with borzoi. As such, I would rely more on the UC Davis test for these breeds.
Saddle tests as ASIPᵇˢ with UC Davis.
Black and Tan: This phenotype is recessive to either variant of the Ay allele. At birth, puppies appear nearly solid black, especially if they are also brindle, with tan markings expanding as they grow. The tan point markings can be any shade of phæomelanin.
Black and Tan tests as at/at with most labs and ASIPᵇᵇ¹, ASIPᵇᵇ², or ASIPᵇᵇ³ with UC Davis.
Grizzle Variations
The eg allele does not have one specific phenotype but rather, this allele modifies and restricts color and markings on other loci. Below are all known grizzle variants that exist in silken windhounds, their genotypes, and how to register them. Grizzle affects the K locus by lightening any eumelanin from either dominant black or brindle, and only minimally affecting the A locus markings. Without dominant black or brindle, grizzle affects the A locus by restricting the expression of the eumelanin markings from either shaded yellow, black and tan, or saddle markings. The eg allele is recessive to both Em and E, and therefore a dog cannot be both masked and express any of the grizzle variants. The word “Grizzle” is also used to describe eg’s modification of black and tan. When the allele was originally studied in Salukis and Afghan Hounds, it was thought that this was the only other alleles that eg affected. We now know otherwise.
The list of grizzle colors are: Silver, Black and Silver, Silver Brindle, Gold Brindle, Grizzle, Grizzle Brindle, and Grizzle based creams/golds/reds.
Silver: Also called “Chinchilla”. Silver is caused by grizzle combining with dominant black and the Ay allele (either shaded yellow or dominant yellow). This genotype can range from a very light silver color to nearly black. Sometimes you can see red pigmented hairs in areas around the ears, pants, and shoulders. In a few instances, we have seen examples of silvers who have a much larger spread of red pigment and higher red intensity.
Tests as: Kb/Kb or Kb/ky, eg/eg or eg/e, Ay/Ay or Ay/at. With UC Davis the A locus can be ASIPdy or ASIPsy and we suspect that ASIPsy may result in darker silvers.
Black and Silver: This genotype is nearly identical to silver but with black and tan on the A locus instead of Ay. Unlike silver, a black and silver dog will appear to have black markings in a very similar pattern to a tan point, but with silver points instead. The edges of the black markings are sometimes not as sharp as a clear black and tan dog. Like silver, you will sometimes see swatches of red hairs in some areas.
Tests as: Kb/Kb or Kb/ky, eg/eg or eg/e, at/at
Grizzle: This color was first studied in Salukis and Afghan hounds, and is what the g stands for in the eg allele. It was initially thought that the grizzle allele was only responsible for modifying black and tan (at/at), and as such, the word “grizzle” referrs both to the eg allele, and this specific phenotype. However, we now know that it affects both the K locus and other alleles on the A locus, as seen by this huge section!
Without the presence of dominant black, grizzle restricts the expression of black on the A locus. The black markings recede, the pips widen to create a striking widow’s peak, and the black hairs become banded showing phæomelanin at the base and eumelanin at the tip.
Puppies are born with what appear to be black and tan markings that fade into a classic grizzle appearance as they grow. There is usually a period of coat change when these dogs are a few months old, where the black hairs almost disappear, and then grow back. We see this coat change pattern on young silver and gold brindles as well.
Dogs who are eg/eg tend to have much stronger black markings that don’t recede as far. Dogs who are eg/e, carrying recessive red, typically have a more faded look to their black hairs and even more receeding of the black markings.
Tests as: ky/ky, eg/eg or eg/e, at/at
Silver Brindle, Gold Brindle: A silver brindle and gold brindle are genetically the same, save for the level of red intensity. Silver brindles have quite pale (cream or lighter) phaeomelanin, while gold brindles are gold or darker. The eumelanin stripes are lightened to a silvery color on a silver brindle. On gold brindles, the lightening sometimes makes the stripes appear more brownish than silver. A brindle dog who is E/- or Em/- should not be registered as gold brindle.
Tests as: Kb/ky(brindle) eg/eg or eg/e, Ay/Ay or Ay/at. With UC Davis, can be ASIPsy or ASIPdy.
Grizzle Brindle: This genotype looks quite similar to grizzle, but with the addition of brindle. Adding brindle to the recipe has a fascinating effect. Instead of brindle appearing only where the phæomelanin is expressed, typically you can find brindle like stripes extending up into the black markings along the topline and sides as well. There are dogs with this genotype that appear silver brindle. Typically this occurs when the dog is also saddle (ASIPbs with UC Davis) and eg/e on the e locus. That combination restricts the black mantle marking to a very small patch of silver over the topline, revealing more brindle stripes.
Tests as: KB/ky (Brindle), eg/eg or eg/e, at/at
Grizzle based red, gold, and cream: Dogs with this genotype will are easy to identify as puppies because they are born with silver hairs on their shoulders and thighs, which fades away by 2 weeks. Any eumelanin expressed from the A locus is restricted down to almost nothing. Some dogs with this genotype still have barely perceptible silvery hairs on the ends of their ears as adults. Most often, these dogs are gold or cream. A very pale cream with a high level of white markings can be registered as white.
Tests as: ky/ky, eg/eg or eg/e, Ay/- (can be ASIPdy or ASIPsy)
Liver and Blue
Liver/Chocolate/Brown: Liver affects any eumelanin and changes it to a red/brown color. Any black skin pigment, such as on the lips, eyelids, and nose leather, will be similarly brown in color. Eyes are typically a more yellow-ish color. B/B (capital letters) indicates no presence of liver, and all pigment will be black. Some have mentioned seeing liver borzoi in the past but I have yet to see evidence of it existing in borzoi. It should be noted that blue dilute can look deceptively like liver on some dogs such as Ay sable. If you have a borzoi or silken that has tested B/b or b/b, please let me know!
Liver tests as b/b
Blue/Dilute: Like liver, dilute will affect any eumelanin and dilutes it to a grey color (including nose, eyelid, and lip pigment). Eyes will usually be more of a yellow or tea color. Sometimes the pigment will be dark enough that the dog will appear to be a typical silver. However, in bright sunlight and next to a dog with dark black pigment, it is more noticeable. In some breeds, hair loss is associated with dilute (called Color Dilute Alopecia), but there hasn’t been evidence of this existing in borzoi or silken windhounds. D/D (capital letters) indicates no presence of dilute. Dilute is quite rare in borzoi, and a bit more common in silkens. Confusingly, some dilute silkens look closer to liver with a more brown-ish grey nose, and coat.
Dilute tests as d/d