Pardon our dust! This site is getting a bit of a makeover and switching hosts.
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 |
---|---|---|
KB | Dominant Black | Dominant Black |
Kbr | Brindle | Brindle |
ky | non-domonant black | - |
E | Normal Expression | - |
Em | Mask | Mask |
eg | Grizzle | Sable (when modifying at/at) |
e/e | Recessive Red | Gold / black suppressed cream |
Ay | Sable | Sabled [red intensity] |
at | Black and Tan | Black and Tan |
KB/- Eg/Eg Ay/- | - | Chinchilla/Silver |
KB/- Eg/Eg at/at | - | black and silver |
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.).
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 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!
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.
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, all white legs, and sometimes a spot of white around the hips.
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.
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
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.
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 Locus
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.
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
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).
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
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.
Silver: Also called Chinchilla in silken windhounds. Similar to silver brindle, but with dominant black on the K locus instead of brindle. Some silver dogs will look very black while others look like a much lighter silver color. The darkness of eumelanin seems to depend on how much shading is expressed from Ay. Dogs that would otherwise be shaded sable end up much darker silver. Clear sables end up being a much lighter silver color. In some cases, eg doesn’t lighten phæomelanin, and the dog will have an interesting look of a brick red smudged together with black.
Silver tests as KB/- eg/eg Ay/-
On the E locus, it can also be eg/e.
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
Grizzle, the eg allele, truly deserves its own section in a color break down because it has such a broad influence over other alleles. When combined with the K locus, it seems to lighten the eumelanin to a more silvery color. When combined with the A locus, it seems to restrict the expression of eumelanin markings. Interestingly, it also seems to lighten the intensity of Through coat color genotype testing in Borzoi, Silkens, Salukis, and Afghan Hounds, we have come to better understand the unique ways in which this allele can interact with other genes.
What we have concluded is that the phenotype of a homozygous eg hound depends entirely on what is happening on the K locus and A locus. If the dog is E/eg or Em/eg, then Grizzle will not affect the other loci, and the dog will not express any of the various grizzle looks below.
Grizzle Variants
Classic Grizzle - Alternatively can be eg/e. In this instance eg is modifying at/at, black and tan. The black markings recede a bit to create a widow's peak on the face, and a dark blanket of black covering the rest of the body. The black hairs are banded-- the tips are black while the roots are red. eg doesn't dilute eumelanin in this instance, but can dilute pheomelanin. A dog that would otherwise be a dark, rich, Irish Setter type red will typically lighten to tan, tans lighten to cream, and creams lighten to create what borzoi folk call "silver sable" (ahem, grizzle).
Classic Grizzle tests as: ky/ky eg/eg at/at or eg/e
Grizzle Brindle: Just like classic grizzle, but with the added bonus of brindle. Brindle markings are primarily visible in the tan markings, but interestingly, sometimes the tan markings extend up into the black markings as brindle stripes.
Grizzle brindle tests as KB/ky eg/eg at/at
As a reminder, brindle always tests as KB/ky. When writing out this genotype, it is best to use Kbr. On the E locus, it can also be eg/e.
Cream: This can also create other red intensities but most often we see the lighter end of the spectrum with cream and gold. When grizzle affects the Ay allele, it makes any black markings recede, similarly to how classic grizzle makes the at/at black markings recede. So much so that any remaining eumelanin is typically only visible on the ear tips, with a silvery look. Puppies with this genotype are often born with silvery markings that fade over time. This genotype can be hard to differentiate from recessive red, e/e, because there are sometimes no black hairs to be found, even on the whiskers. Note, this isn’t the only genotype for solid creams or lighter phæomelanin. Some sables without eg can be clear and have light phæomelanin, as well as the aforementioned recessive red.
Grizzle based creams test as ky/ky eg/eg Ay/-
Silver Brindle: When the A locus is sable (Ay) instead of black and tan (at/at), all eumelanin on the K locus becomes lightened. As such, the brindle stripes become a silver color. Because grizzle dilutes phæomelanin as well, the end result can sometimes look like a silver dog with slightly darker silver stripes. With darker phæomelanin, the dog might instead appear to have silver stripes on a more gold base. We call that gold brindle.
Silver Brindle tests as KB/ky eg/eg Ay/-
As a reminder, brindle always tests as KB/ky. When writing out this genotype, it is best to use Kbr. On the E locus, it can also be eg/e.
Black and Silver: Just like with Silver, eg dilutes eumelanin from the K locus but leaves eumelanin from the A locus mostly untouched. This genotype is almost identical to classic grizzle, save for the addition of dominant black (KB). Instead of modifying at/at to create a widow’s peak marking, dogs retain their at/at markings but with silver instead of tan. Sometimes varying degrees of phæomelanin remain on the legs. Interestingly enough, as dogs with this genotype grey with age, it appears to first affect the cheeks and pips, but then follows the classical grizzle pattern, eventually creating that widow’s peak.
Black and silver tests as: KB/- eg/eg at/at
On the E locus, it can also be eg/e